I/"* ^ j 5 . > v ^ > NOliniilSNi NVlNOSHlIWS^SHIHVaail LI B RAR I ES ^SMITHSON IAN 'iNSTITUTl 1 ^ v in — I LIB RAR I ES^SMITHSONIAN INSTITUTION NOlinillSNrNVINOSHilliMS^SBI BVd Bl m ~ cn — cn 2 o ^ ^ NouniiiSNi^NViNOSHims S3 lavaan librari eszsmithsonian“institutioi m C O ZZ £/) f: w LIBRARIES SMITHSONIAN INSTITUTION NOliniilSNi NVINOSHIUNS S3IHVH31 ' 2 xgyosi^ > 2 )ixnxixsm~NviN0SHXiws‘',S3 1 bjvb a n\i b rar i es^smithsonian^institution " Z \ ^ . ; — ^ -?> tn ZZ U ^*21 l£>X CO J%. ^ CO 1X1 Cl aiIV^X ■ H, x>^. a ~~ . /a x^wiik -f *>\ /S/s/ZV • z _j i BRARIES SMITHSONIAN INSTITUTION NOIXIUIXSNI NVINOSHXIWS S3IBVBan Z P" ^ r— _ r ro 70 > 70 m 'CLE^z g m ^ )ixnxixsNizNviN0SHXiws"sa I a va a n~u b rar i es^smithsonian'institution* < 2 < .-x.'' . 2 _ — - 5 jtv.- u 0 CO 1 _ BRARI ESt/>SMITHS0NIAN_INSTITUTI0NOTN0IXnXIXSNI^NVIN0SHXIWS _ 60 ~ .... tO 'V O 2 5 'W 2 IXnXIXSNI^NVINOSHXIINS S3 I B VB a IT ' L I B RAR I ES^SMITHSONIAN^INSTITUTION ^ Z a- r* m m B RAR I ES SMITHSONIAN INSTITUTION^NOimillSNI ~*NVIN0SH1IWS^$3 I dVd a 11 co ^ - in z x co O z s '" _ liniliJ.SNI__NVIN0SHllWSS3 I d Vd a nZLI B RAR I ES^SMITHSONIAN^INSTITUTION “ Z \ ln ^ CO — C/ O ~ ''•KSffii*' o B RAR I ES^SMITHS0NIAN(_INSTITUTI0N*C ' ^ ^ 5 .AfV Entomologica Americana VOLUME 45 CONTENTS C. F. dos Passos and A. B. Klots — The Systematics of Anthocharis midea Hlibner (Lepidoptera: Pieridae) R. C. Allen — A Revision of the Genus Leptoglossus Guerin (Hemiptera: Coreidae) JAMES A. SLATER, EDITOR PUBLISHED BY THE NEW YORK ENTOMOLOGICAL SOCIETY INCORPORATING THE BROOKLYN ENTOMOLOGICAL SOCIETY With the issuance of Parts 1 and 2 of Volume 45, publication of Entomologica Americana is resumed after a hiatus of several years. Further parts of this volume will be issued early in 1970. Entomologica Americana was first issued by the Brooklyn Entomo- logical Society in April 1885. Publication was suspended after six volumes but was resumed in 1926. Volume number 44, the last previous volume was issued in* 1964. The combined Societies’ purpose in resuming publication is to afford authors an opportunity to get longer papers (72 pages or more) into print, which its quarterly Journal cannot accommodate. Publication will be by Volume and frequency of publication will depend upon availability and acceptance of worthwhile papers. Your inquiries are solicited and should be directed to the attention of the Publication Committee: PUBLICATION COMMITTEE Lucy W. Clausen John B. Schmitt Asher E. Treat \ All correspondence concerning the Society or its publications, including subscriptions to Entomologica Americana should be sent to the address shown below. New York Entomological Society The American Museum of Natural History 79th Street and Central Park West New York, N.Y. 10024 Mailed December 29, 1969 Entomologica Americana is published for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. THE SYSTEMATICS OF ANTHOCHARIS MIDEA HUBNER (LEPIDOPTERA: PIERIDAE) By Cyril F. dos Passos and Alexander B. Klots Entomologica Americana Yol. 45, 1969, pp. 1-34 THE SYSTEMATICS OF ANTHOCHARIS M1DEA HUBNER (LEPIDOPTERA: PIERIDAE) By Cyril F. dos Passos* 1 and Alexander B. Klots2 Table of Contents Introduction 2 Nomenclature 2 Neotypes and type localities 5 Geographic variation 8 Distribution 9 Conclusion 11 Synonymies 12 a. Anthocharis ( Falcapica ) midea (Hiibner) 12 b. Anthocharis ( Falcapica ) midea midea (Hiibner) 15 c. Anthocharis ( Falcapica ) midea annickae n. ssp. 17 Life history 22 Foodplants 23x Parasites 23 Acknowledgments 24 Bibliography 24 Abstract: The geographic variation and systematics of Anthocharis midea (Hiibner) of eastern North America are described. The species is placed in the subgenus A. ( Falcapica ) Klots, 1930. Neotypes are designated for Maucipium vorax midea Hiibner, Pieris Iherminieri Godart and the homonomous Papilio danaus genutia Fabricius. The nominate subspecies A. m. midea occurs in the southern Coastal Plain. The northeastern subspecies is named as A. m. annickae dos Passos and Klots, type locality New Haven, Connecticut. Complete bibliog- raphies are given for the species and subspecies. Some notes on the life history and parasites are included. Accepted for publication November 19, 1968. 1 Research Associate, Department of Entomology, The American Museum of Natural History, and Research Associate, Section of Insects and Spiders, the Carnegie Museum. 2 Professor of Biology, Retired, th§ City College of New York, and Re- search Associate, Department of Entomology, The American Museum of Natural History. [1] 2 Entomologica Americana [Vol. 45 Introduction For many years the authors have recognized the need for a detailed study of the North American falcate orange-tip butterfly, Anthocharis midea Hlibner (“1806” [1809]), to clarify its geographic variation and nomenclature and have been accumulating data and specimens for this purpose. They have been materially aided by many persons in Europe and North America and are especially in- debted to Lucien Harris, Jr., Bryant Mather, and Richard Heitzman for the loan of important series of specimens from Georgia, Missis- sippi, and Missouri. Nomenclature Generic. This species should be considered (Klots, 1930, pp. 82-83; 1931, p. 254; and “1931” [1933], p. 151) a member of the genus Anthocharis Boisduval, Rambur and Graslin [1833], the type-species of which is Papilio cardamines Linnaeus (1761, p. 271). Following Hemming (1967, p. 46), we employ the spelling Anthocharis in- stead of Anthocaris in this paper because it is the original spelling of the authors Boisduval, Rambur and Graslin [1833], and no reason is apparent for its emendation. Earlier Hemming (1934, p. 132) had used the spelling Anthocaris and referred the name to the same authors. This was a lapsus calami. In a subgeneric sense we employ also Falcapica Klots, 1930, the type-species of which is Papilio genutia Fabricius ( nec Cramer “1782” [1780], 1793). Midea Herrich-Schaffer, 1867, of which F. genutia is also the type, is a preoccupied name having been proposed by Bruzelius in 1854 and by Walker in 1863, so that in effect Fal- capica is a substitute name for Midea Herrich-Schaffer although not expressly so stated in the original description. We are not unmindful of the fact that Kuznezov (1929, p. 58 footnote) proposed the subgeneric name Paramidea for a falcate orange-tip butterfly with the type-species Midea scolymus Butler, 1866. This is a Palearctic butterfly occurring in West China and Japan. While both midea and scolymus have falcate forewings, their facies are very distinct so that subjectively we refer each to separate subgenera. It should be noted in passing that Bernardi (1961, p. Ill) misspelled “Paramidae” and gave an incorrect page reference “52.” It should have been 58. The genitalia are of little use in separating species of Anthocharis ( Falcapica ) as will be seen in referring to figures of genutia and scolymus (Klots, 1930, p. 94, figs. 4 and 5). Specific. Four species-group names are available for the species: 1. Papilio danaus genutia Fabricius (1793, p. 193) 2. Mancipium vorax midea Hlibner (“1806” [1809], pi. 142) 1969] dos Passos and Klots: Systematics of Anthocharis midea 3 3 . Pieris Iherminieri Godart (1819, p. 118) 4. Anthocharis flavida Skinner (1917, p. 438) 1. Papilio danaus genutia Fabricius, although the oldest avail- able name, is a homonym of Papilio genutia Cramer (“1782” [1780]) and therefore unavailable. No type of Fabricius’ exists at the British Museum (Natural History), London; the Zoologisk Museum, Copen- hagen (Zimsen, 1964, p. 560, no. 967); the Zoological Museum of the University, Kiel; or anywhere else insofar as we have been able to ascertain. In the original description of genutia , Fabricius stated that the habitat was “in Indiis” and that the specimen was in the collection of Drury. He also referred to (the unpublished) Jones’s leones (figured picture 3, pi. 26, fig. 2). We have examined the leones at the Hope Museum, Oxford, and have a photograph of this figure kindly fur- nished by Miss Audrey Smith which we reproduce here (Fig. 1). Considering the standards of its time, it is a good and recognizable representation of the North American falcate orange tip, although it does not show the black discocellular spot on the upper side of the forewing and has the apical orange patch running down too far along the outer margin. It shows a very extensive orange patch which ex- tends basad to at least the end of the discal cell where the black discocellular spot would be. We are indebted to Prof. G. C. Varley of the Hope Museum for permission to reproduce this photograph which has not been published heretofore. The oldest published figure of genutia is in Donovan’s Insects of India (1802, [pi. 27], fig. *|*).3 This does not agree very well with the North American insect but resembles even less the Chinese and Japanese A. scolymus Butler and not at all anything from India. Since Donovan gives a reference to Fabricius’ original description, 3 The pages and plates in this work are not numbered. The title page is followed by two pages entitled “Advertisement” and 16 signatures, some of which are numbered and others lettered, some of the numbers being dupli- cated. Each signature appears to have been accompanied by one or more colored plates. There is little or no consistency to these numbers and letters. The copy of this work in the Library of the British Museum (Natural History) contains 57 plates. The one in the Library of The American Museum of Natural History contains 58 plates. That this work should contain 58 plates is evidenced by the index, which lists that many plates. Each plate bears a date between 1 Jan. 1800 and 1 Feb. 1804, but they are not numbered in the index in accordance with their dates of issue, a systematic method of number- ing having been adopted. The signatures are not dated, but most likely ap- peared with the plates. Whether they did or not, the plates bear the scientific names and the dates of their publication. An edition by Westwood (1842) con- tains 58 plates. 4 Entomologica Americana [Vol. 45 (All figures are approximately natural size) Fig. 1. Photograph of the figures of upper- and undersides of Papilio danaus genutia Fabricius in Jones’s leones. it is possible that his figure is a poor representation of Drury’s speci- men or of the leones figure. 2. Mancipium vorax midea Ffiibner (“1806” [1809]) is the next oldest available name. Hemming (1937, p. 429) states that the speci- men figured by Hlibner came from either “Georgien” German text or “Brasilia” Latin text. Since Brazil is an impossible locality for the species, we may safely assume that the specimen came from Georgia. No trace of any Hlibnerian specimen that might be his type was found in any of the European museums; it is safe to assume that none exists. 3. Pier is Iherminieri Godart (1819) is the next oldest available name. The locality given in the original description is “Charles- Town.” We feel safe in assuming that this meant Charleston, South Carolina, where the species is known to occur. Charlestown, Massa- chusetts, has been suggested as an alternative, but we regard this as highly improbable. Scudder (1889, p. 1150) mentions Boisduval’s reporting the butterfly to be found about Boston, but believes that this was erroneous. The closest records to Boston, then as now, are 1969J dos Passos and Klots: Systematics of Anthocharis midea 5 from New Haven, Connecticut in the coastal plain, and Holyoke, Massachusetts in the Connecticut River Valley, far inland from Boston and Charlestown. No possible Godart specimen of Iherminieri was found by us in the Museum d’Histoire Naturelle, Paris, where many of Godart’s types are preserved; and we have been assured by Dr. Pierre E. L. Viette of that institution that it is not there. Neither is any Godart specimen in the Edinburgh Museum of Science and Art, Edinburgh, where some of Godart’s North American types are preserved or elsewhere so far as we have been able to ascertain. It is safe to assume that the type no longer exists. 4. Anthocharis flavida Skinner (1917) was described from a single male from Savannah, Georgia (March 12, 1917, collected by W. T. Coxey). We figure this type (Figs. 2 & 3) which is in the Academy of Natural Sciences, Philadelphia. Although named as a variety, it is actually a normal specimen of the population about Savannah. Neotypes and Type Localities As we show below, there is considerable geographic variation in the populations of the species. In the study and nomenclature of this, however, the lack of types and exact type localities for the nominal species Papilio genutia Fabricius, Mancipium midea Hiibner, and Pieris Iherminieri Godart is a grave handicap preventing the exact application of specific-subspecific names. We therefore, under Article 75 of the International Code of Zoological Nomenclature, herewith designate neotypes for these nominal species. In doing so we have chosen specimens consistent with the original information and data, and coming as nearly as practicable from the original type localities as discussed above. We do this after consultation with other specialists in North American butterflies, from none of whom was any objection received. 1. Papilio danaus genutia Fabricius (1793). The locality (“in Indiis”) given in the original description is obviously wrong. Fa- bricius’ specimen may well have come from the coastal plain region of Georgia, whence John Abbot had been sending material to Europe for many years. The specimen figured in Jones’s leones ( loc . cit.) referred to by Fabricius most closely resembles those from the coastal plain of Georgia. We therefore designate as the neotype of this nominal species a male specimen from Wilmington I., near Savannah, Georgia, April 2-11, 1947, which is the property of The American Museum of Natural History (Figs. 4 & 5). 2. Mancipium vorax midea Hiibner (“1806” [1809]). As noted above, “Georgien” and not “Brasilia” is almost certainly the true 6 Entomologica Americana [Vol. 45 Figs. 2-3. Anthocharis flavida Skinner $ type, upper- and undersides, Savannah, Ga., 12 March 1917, W. T. Coxey. type locality. We therefore designate as the neotype of this nominal species the same male specimen designated as the neotype of Papilio danaus genutia Fabricius, from Wilmington I., near Savannah, Geor- gia, April 2-11, 1947, which is the property of The American Museum of Natural History (Figs. 4 & 5). 3. Pieris Iherminieri Godart (1819). For the reasons stated above we designate as the neotype of this nominal species a male specimen from McClellanville (Charleston Co.), South Carolina, 29 March 1967, leg. R. B. Domenick which is the property of The American Museum of Natural History (Figs. 6 & 7). We are in- debted to Dr. R. B. Domenick of McClellanville, South Carolina, for this specimen. 4. Anthocaris genutia flavida Skinner (1917) although named as a variety is based on a normal specimen characteristic of the Savannah, Georgia population. It must therefore be treated as a species-group name in accordance with Article 10b of the Inter- national Code of Zoological Nomenclature. We believe that it was first given this status by Clark and Clark (1951, p. 86) (Figs. 2 & 3). In accordance with the above, the restricted type localities of both Papilio danaus genutia Fabricius and Mancipium vorax midea Hiibner are Wilmington I., near Savannah, Georgia; and that of Pieris Iherminieri Godart is Charleston, South Carolina. All three of these names, and Anthocharis flavida Skinner also, then apply to the popu- lation of the Southern Coastal Plain that is characterized, as discussed 1969] dos Passos and Klots: Systematics of Anthocharis midea 7 Me CIoliSMB-srlU* South C*mtina &• 8. 33orai»lcfc ' *-*! rvte»U, !*£? Of?** y», I-. X Figs. 8-11. Anthocharis midea annickae holotype $ and allotype $, upper- and undersides, both West Rock, New Haven, Conn., S 22 April 1954, $ 16 May 1952, leg. C. L. Remington. (The specimens of Figs. 4-11 are in The American Museum of Natural History. All photographs except Fig. 1 are by the senior author.) 1969] dos Passos and Klots: Systematics of Anthocharis midea 19 Anthocharis Genutia, Boisduval (= Mancipium vorax Midea, Hiibn. = la femelle Pieris Lherminieri, God.) 1836, vol. 1, p. 565, no. 10 (Amerique Septentrionale; environs de Boston et de Charlestown). Anthocaris Genutia Boisd., Scudder, 1868, vol. 11, p. 376, no. 12 (New Haven, Connecticut, May 16). Anthocaris Genutia Boisd., Scudder, 1868, p. 4, no. 12 (New England). Anthocaris genutia, Doubleday, 1869, Scudder (ed.), vol. 1, p. 121 (Edwards County, Illinois). Anthocaris midea, Riley, 1871, p. 158 (Missouri). [Anthocharis, Boisduval] Genutia Fabricius, Edwards, “1872” [1869], p. 5, no. 1. (New York to Virginia; middle and western states; Texas). Euchloe Genutia, Butler, “1869” [1870], p. 214, no. 1 (Illinois, United States) . E[uchloe ] Hiibn. Genutia Fabr., (= Pieris Lherminieri God. = Mancipium vor. Midea Hiibn.) Scudder ( partim ), 1872, p. 43, no. 1 (southern New England to Georgia and Texas) . [Anthocharis, Bd.] Genutia, Bd., Edwards {partim), 1877, vol. 6, p. 15, no. 49 (New York to Virginia; western states, Texas). Anthocharis genutia Bois., Drury, [1878], vol. 1, p. 12, no. 14 (Cin- cinnati, Ohio). Anthocharis Genutia Edwards, “1874-1884” [1878], vol. 2, pt. 7, p. [83-84], pi. Anthocaris II, figs. S 1-2, $ 3-4 (Dallas, Texas; New Jersey; near Philadelphia; near Baltimore, Maryland; Newburgh, New York; Coalburgh, West Virginia, April; Illinois; Boston). Anthocaris Genutia, Edwards, 1881, vol. 13, p. 211 (Coalburgh, [West Virginia], 17 April-14 May). Anthocaris midea, Riley, 1881, p. 308 (Missouri). [Anthocharis Boisduval] Genutia Fab., Edwards, 1884, vol. 11, p. 261, no. 53 (New York to Virginia; western states). Anthocaris Genutia Fab., E. M. & S. F. Aaron, 1884, vol. 4, p. 172 (prairies southern Texas, 1st week April) . Anthocaris Genutia Fab., French, 1886, p. 118, no. 17 (New York to Virginia; Western States, Texas) . Anthocaris genutia Maynard, 1886, p. 48, no. 66, pi. 8, figs. 66 S, 66a $ (New England, Connecticut, western Massachusetts, 2 broods, first [sic] in July) . M[idea] Genutia Fabr., Staudinger {partim), 1888, in Staudinger & Schatz, vol. 1, p. 47 (southern states of North America). Anthocharis genutia, Scudder {partim), 1889, vol. 2, pp. 1147-1153; vol. 3, pi. 15, figs. 13, 15, pi. 26, fig. 3 (distribution), pi. 35, fig. 14, pi. 40, fig. 5, pi. 46, fig. 41, pi. 56, fig. 7 {imago), pi. 65, fig. 29 {ova), pi. 73, fig. 9, pi. 76, fig. 5, pi. 79, fig. 54 {larva), pi. 84, fig. 59 {pupa) (southern half of Alleghenian, northern half of Carolinian faunas, from Atlantic to southern Mississippi valley; Pennsylvania, central Texas at Dallas; nearly all Atlantic states from Connecticut to Georgia; Savannah, Ga.; Kanawha Co., W. Va.; Mexican border; Illinois; Ohio at Cin- cinnati; Newburgh, N. Y.; Connecticut at Greenwich, New Haven, New 20 Entomologica Americana [Vol. 45 Britain, Farmington and tops of Meriden Hills, vicinity Holyoke, Mass., May to June). Anthocharis genutia, Skinner and Aaron, 1889, vol. 21, p. 129. (West- ville, New Jersey, May 6) . Anthocharis genutia , F., Beutenmiiller, “1889-1891” [1890], vol. 5, p. 200 (Delaware Water Gap, Pennsylvania; Nyack, New York). Anthocharis genutia, Rowley, 1890, vol. 22, p. 123 (Coalburgh, West Virginia, May) . Anthocaris genutia, Maynard, 1891a, p. 48, no. 66, pi. 8, figs. 66$, 66a $ (rare in New England, taken in Connecticut and western Massa- chusetts, 2 broods, first [sic] in July). Anthocharis genutia Fab., Maynard, 1891b, p. 29, no. 52, figs. 14b, 14g, pi. 2, fig. 1 male (New York to Virginia, western states and Texas. Rare in southern New England, February, March, Texas, April further north) . [ Anthocharis ] Genutia, Edwards, 1892, vol. 24, pp. 52, 109 [Coal- burgh, W. Virginia]. Anthocharis genutia Fab., Blatchley, 1892, in Gorby, p. 372, no. 12 (53) (Vanderburgh County, Indiana). Anthocharis genutia, Skinner, 1892, vol. 3, p. 240 (Areola, Perkio- men Creek, Pennsylvania, May 9). Anthocharis genutia (Fabr.) Beutenmiiller, 1893, vol. 5, p. 248, pi. 2, fig. 5 (Nyack and Newburgh, New York; Delaware Water Gap, Pennsylvania, May). Anthocharis Genutia, Scudder ( partim ), 1893, p. 140 (eastern half of southern portion of our district, [eastern United States] even into New England; southern Illinois and Ohio, May to June). Anthocharis genutia , [Anonymous], 1895, vol. 6, p. 145 (near West- ville, New Jersey; Fox Chase, west of Quaker City, [Pennsylvania]). Anthocharis Genutia, Edwards, “1888” [1897], vol. 3, pt. 6, pp. [57-61], pi. Anthocharis 1, figs. 5 male a-h3 (Washington, D. C.). [Anthocharis Boisduval] Genutia Fab., (= midea Hub.), Skinner, 1898, p. 65, no. 397 (Connecticut, New York to Virginia, western states). Euchloe genutia Fabricius, Holland, 1898, p. 284, p. 4, fig. 6 (ova); pi. 32, figs. 37$, 38$; pi. 2, fig. 5 (larva); pi. 5, fig. 59 (pupa) (New England to Texas) . Euchloe genutia (Fabr.), Beutenmiiller, 1898, vol. 10, p. 246, pi. 14, fig. 7 (Massachusetts to Texas). Anthocharis Genutia Fabr., Smyth, 1900, vol. 11, p. 465 (Blacks- burg, Virginia, summit Alleghenies, April to May) . [Synchloe Hiibner] genutia Fabricius, (= midea Hiibner) Dyar (partim), “1902” [1903], no. 52, p. 7, no. 48 (southern Atlantic states). Anthocharis genutia, Hornig, 1903, vol. 14, p. 252 (Westville, New Jersey, May) . Anthocharis genutia Fab., Grossbeck, 1905, vol. 16, p. 131 (prepara- tory stages; Garret Mountain, Paterson, New Jersey). 1969] dos Passos and Klots: System atics of Anthocharis midea 21 A[nthocharis Bdv.] genutia Fab., Smith, 1910, p. 418 (New Jersey, April and May) . Anthocharis genutia, Skinner, 1917, vol. 28, p. 438 (Illinois). [Synchloe Hiibner] genutia Fabricius, Britton, 1920, no. 31, p. 159 (Connecticut). Anthocharis genutia, Comstock & Comstock ( partim ), 1923, p. 385 (southeastern United States not Florida; north to New Haven Con- necticut) . Synchloe genutia, ( Anthocaris genutia or Euchloe genutia), Weed {partim), 1924, pp. 94, 97, 256, pi. [fig. 2]. (east of Rocky Mountains; north to New England; Southern states south to Texas; western portion North Carolina). Anthocharis genutia, Clark, 1927, pp. 424, 428, pi. 2, figs. 10, 11 (Washington, D. C.). A[nthocharis Boisduval] genutia Fab., Forbes, “1926” [1928] in Leonard, p. 677, no. 43 (highlands of Hudson, Greenwood, Tuxedo, Newburgh, Ramapo Mts., Canarsie, [New York] Apr .-May). Synchloe genutia, Comstock & Comstock (partim), 1929, p. 82, pi. 15, figs. 1-2 (southeast United States, except Florida; north to New Haven, Connecticut) . Euchloe [(Anthocharis)] genutia (Fabricius) Holland (partim), 1931, p. 287, pi. 4, fig. 6 (ova)’, pi. 32, figs. 37 <2 38$; pi. 2, fig. 5 (larva); pi. 5, fig. 59 (pupa) (New England to Texas). Euchloe genutia Fabricius, [anonymous] [Wyss comp.] [1932], p. [24], no. 45 (Cincinnati, Ohio). Anthocharis genutia (Fabricius), Clark, 1932, pp. 2, 3, 6, 7, 9, 12, 21, 23, 26, 30, 32, 43, 44, 61, 148, 164, 167, 168, 173, 234, 238, 252, pi. 2, figs. 10, 11, pi. 29, figs. 1-4 (Maryland; District of Columbia; Vir- ginia). (Anthocharis) Subgenus Falcapica Klots genutia Fabr., Talbot, 1934, in Strand, pt. 60, p. 321 (United States [New England to Texas]). E[uchloe Hiibner] genutia Fab., Brimley, 1938, p. 259 (Raleigh, Chapel Hill, Roanoke Rapids, [North Carolina], March and April). Anthocaris medea [s/c] (Hbn.), Leussler, 1938, vol. 49, p. 77 (Crete and Omaha, Nebraska) . Anthocharis (Falcapica) midea (Hiibner), Field, 1938, vol. 39, no. 10, p. 176, no. 74 (New England south to Virginia and west to Texas and Kansas, Douglas, Greenwood and Leavenworth Counties, Kansas). Anthocharis genutia Fab., Engelhardt, fide Tulloch, 1939, vol. 34, p. 227 (Lincoln, New Jersey, April and May). [Anthocharis Boisduval] midea Hiibner, (= genutia Fabricius), W. P. Comstock, 1940, vol. 48, p. 70, no. 30 (locally common throughout state [New Jersey]). Anthocharis midea (Hiibner), Field, 1940, vol. 13, p. 28 (Douglas and Leavenworth counties, Kansas, April). Anthocharis midea (Hiibner), Macy and Shepard, [1941], p. 36, fig. 7; pp. 52, 53, pi. 4, [fig. 4] (New England westward and southward 22 Entomologica Americana [Vol. 45 to Ohio, Illinois, and Texas; common near Washington, D. C.; Oakland, New Jersey; Athens County, Ohio, March to May). Anthochris [m'c] midea, Cook, 1948, vol. 2, p. 22 (Crailhope, Ken- tucky) . Anthocaris [( Falcapica )] genutia [= midea\, Edwards, dos Passos (ed.) 1951, vol. 59, p. 163 (Washington, D. C.). Anthocaris genutia midea Hbn., Rawson, 1951, vol. 5, p. 70 (Mt. Peter, Greenwood Lake, New York). [ Anthocaris ] midea , Remington, 1951, (New Haven, Connecticut). Anthocharis genutia (Fabricius), Clark & Clark, 1951, vol. 116, no. 7, pp. 86, 87, pi. 9, figs, i-j (Maryland and Virginia). Anthocaris genutia Fabricius, Klots ( partim ), 1951, pp. 49, 181, 208, pi. 6, fig. 5 {pupa), pi. 25, fig. 1 $ (Ramapo Mts., N. Y.) (Massa- chusetts and Connecticut, s. to Georgia, w. to Illinois and Texas [Dallas]). Anthocaris midea, Muesebeck & Walkley, 1951, p. 128-130. [Euchloe Hub.] genutia Fab. = midea Hi.ib., Tietz, [1952], p. 2, no. 30 (Pennsylvania) . E[uchloe ] genutia Fabricius, Forbes {partim), 1960, pp. 108-110 (southern Massachusetts to Illinois and Arkansas, south to Georgia and Texas) . Anthocaris midea (Hiibner), Klots {partim), 1965, p. 463 (in the Piedmont, westward to Texas and northward into Connecticut and Missouri) . Life History The life history of midea is well known and all stages are beauti- fully figured and described by Edwards in the third series of the Butterflies of North America, [1888], pt. 6, pi. Anthocharis I, figs. a-h3, pp. [57]— [6 1 ]. One interesting fact that we have observed concerning the life history of this insect is that the imagines do not always emerge the year following pupation. Among those reared by the senior author in 1933 from specimens taken at Edison, New Jersey, one male did not emerge from the pupa until 26 April 1935. Others reared from specimens taken by us at Harrisville and New Gretna, both in Burl- ington County, New Jersey, 8-9 May 1953 and reared by the senior author on Arabidopsis Thaliana (Linnaeus) resulted in seven out of thirteen specimens not emerging until the second year, the record being four males and two females the first year, while three males and four females emerged the second year. All of these specimens are in the senior author’s collection except for one male, Harrisville, ex pupa, 1 March 1955 and one pupal shell which is in the junior author’s collection. The late Charles E. Rummel of Green Village, New Jersey, ob- 1969] dos Passos and Klots: Systematics of A nthocharis midea 23 served many years ago (1931, p. 268) a case where an imago did not emerge until the third year, but we have never had that experience. In the northern part of its range, the species is monogenetic. In Virginia teste Clark & Clark (1951, p. 86) there may be a partial second generation influenced by local climatic conditions, and this may well be the case in many parts of its southern range. Foodplants The larva of midea feeds on various Cruciferae (Mustard Family), among which the following have been reported : Bursa Bursa-pastdris (Linnaeus), (1753, p. 647) Britton (Shepherd’s purse, -bag, or -pouch) ; Arabidopsis Thaliana (Linnaeus), (1753, p. 665), (Mouse- ear or Thale-cress, Wall-cress); Barbarea Barbarea (Linnaeus), (1753, p. 660), MacMillan, (Yellow Rocket or Cress); Arabis glabra (Linnaeus), (1753, p. 666), Bernhardi, (Tower Mustard or Cress); and a species of Cardamine ( [Tournefort] Linnaeus), (1753, p. 654) the specific name of which does not appear to have been reported but may be the one mentioned below. We have reared the insect on A. glabra when taken in the moun- tainous section of New Jersey and on A. Thaliana when taken on the coastal plain. Dr. Charles L. Remington of the Gibbs Research Laboratories, Yale University, reports finding eggs and larvae at New Haven, Connecticut, on Arabis laevigata (Muhlenberg), (1801, p. 543), Poiret (Smooth Rock-cress); A. lyrata Linnaeus (1753, p. 665), (Lyre-leaved Rock-cress); and Cardamine parvifldra Linnaeus and C. arenicola Britton, (1892, p. 220,) (Sand Bitter-cress or Small Flowered Bitter-cress), and rearing the insect on all three. Doubt- less there are other foodplants such as Cardamine rhomboidea De Candolle (1821, p. 246), (Bulbous Cress) probably a synonym of Cardamine bulbosa (Schreber) Britton (1793, p. 174). According to Smyth (1900, p. 465), eggs of midea were laid on Dentaria laciniata Muhlenberg (1800, p. 479), (Cut-leaved Tooth- wort or Pepper-root) which seems to be the only foodplant in Blacks- burg, Virginia. Parasites From larvae collected on Arabis glabra (see above) were reared some parasitic wasps determined by Miss Luella M. Walkley of the United States National Museum, who stated that it was the first record of that parasite on midea , to be Hyposoter exiguae (Viereck) (1912, p. 638) (Ichneumonidae). One male wasp and cocoon are in that institution, and others in the collection of the senior author. This wasp has also been recorded on Heliothis armiger (Hiibner), 24 Entomologica Americana [Vol. 45 ([July 1803]— [1808]), Prodenia ornithogalli praefica Grote, 1875 (both Noctuidae), and Colias eurytheme Boisduval, 1852 (Pieridae). Also reported on Anthocharis midea are Apanteles flaviconchae Riley (1881, p. 308) (Braconidae) and A. limenitidis (Riley) (1871, p. 158) according to Muesebeck (1951, p. 128). Another reported parasite is Apanteles pergandei Grossbeck (1905, p. 133) nec Ashmead (Braconidae). We do not find that this name was ever published by Ashmead. Grossbeck (supra) stated in a footnote that, “The species is described only in MS and will appear in Dr. Ashmead’s monograph of the North American Braconidae now in preparation.” Ashmead died a few years later (1908) without however having published any such monograph or described Apanteles pergandei. The name is, therefore, a nomen nudum. Acknowledgments We again express our thanks to Messrs. Lucien Harris, Jr. of Avondale Estates, Georgia, Bryant Mather of Jackson, Mississippi, and John R. Heitzman of Independence, Missouri, for the loans of their entire collections of Anthocharis midea with permission to retain such specimens as might be necessary to enlarge the distribution records of The American Museum of Natural History; and to Pro- fessor Charles L. Remington for the gift of specimens to be used as the types of A. m. annickae as well as notes on the foodplants of that subspecies in Connecticut; also to Dr. Richard B. Domenick of McClellanville, South Carolina for the loan of 9 male specimens taken at that locality with permission to retain one male to serve as the neotype of Pieris Iherminieri Godart. We also wish to thank Miss Luella M. Walkley of the United States National Museum for identifying the parasitic wasp that was reared on midea and Dr. Pierre E. L. Viette of the Paris Museum for searching for Godart’s type of Pieris Iherminieri in that institution. Bibliography Aaron, E. M. and S. F. Aaron. 1884 [1885]. List of a collection of diurnal Lepidoptera from southern Texas. Papilio 4: 172-182. Abbot, J. ineditus. ca. 1800. Drawings of the insects of Georgia. British Museum (Natural History) 6: 20, figs. 79-81. See Smith, J. E. Anonymous. 1882 [1881]. Check list of the North American Macro- lepidoptera of America. Publication Committee, Bull. Brooklyn ent. Soc. 4: 45, [2] + 25 pp. . 1893-94. List of Pteridophyta and spermatophyta growing without cultivation in northeastern North America. Mem. Torrey Botanical Club 5: 5-9, 172-173. 1969] dos Passos and Klots: Systematics of Anthocharis midea 25 . 1895. Popular Entomology A chase for a butterfly. Ent. News 6: 145-147. . 1932. See Wyss, A. E. Arnhold, F. R. 1952 [1953]. Notes on collecting Anthocharis midea and Euchloe olympia. Lep. News 6: 99-100. Ashmead, W. H. Monograph of the North American Braconidae. [Ap- parently never published.] Barnes, W. and J. H. McDunnough. 1917. Check list of the diurnal Lepidoptera of Boreal America. Decatur, 111., Herald Press, viii + [2] + 392 + [4] pp. Barnes, W. and F. H. Benjamin. 1926. Check list of the diurnal Lepidoptera of Boreal America. Bull. So. Calif. Acad. Sci. 25: 3-27. Bernardi, G. 1961. Note synonymique sur les Pieridae. Bull. Soc. ent. Fr. 66: 111-112. Bernhardi, J. J. 1800. Systematisches Berzeichniss der Pflanzen, Welch in der Gegend um Erfurt Gefundenwerden. Octavo, Erfurt, Germany. Pt. 1, 28 + 346 pp. Beutenmuller, W. 1889-91 [1890]. Catalogue of the lepidoptera found within fifty miles of New York City, with their foodplants. Ann. N. Y. Acad. Sci. 5: 199-229. . 1893. Art. XVI Descriptive catalogue of the butterflies found within fifty miles of New York City, together with a brief account of their life histories and habits. Bull. Amer. Mus. Nat. Hist. 5: 241-310, 4 pis. . 1897. Meeting of October 16, 1897. N. Y. ent. Soc. 5: 208. . 1898. Revision of the species of Euchloe inhabiting America north of Mexico. Bull. Amer. Mus. Nat. Hist. 10: Art. XIII: 235-248, pis. 13-14. Blatchley, W. S. 1892. A catalog of the butterflies known to occur in Indiana in Gorby. Indiana Department of Geology and Natural Resources, 17th Annual Report, pp. 365-408. Boisduval, J. B. A. D. de. 1836. Histoire naturelle des insectes. Species general des lepidopteres. Paris, Deroret, pp. 4 + xii + 690 pp. + 24 pis. (colored and partly colored) . Boisduval, J. B. A. D. de, J. P. Rambur, and A. de Graslin. [1833]. Collection iconographique et historique des chenilles . . . d’Europe. Paris. 128 pis. col. with descriptive letterpress, pts. 1-10, 1832; 11-20, 1833; 21-30, 1834; 31-38, 1835. Brimley, C. S. 1938. The insects of North Carolina. Raleigh, N. C., North Carolina Department of Agriculture, 560 pp. Britton, N. L. 1892. New or noteworthy North American Phanero- gams.— VI. Bull. Torrey Botanical Club 19: 219-226. Britton, W. E. 1920. Check list of the insects of Connecticut. Hart- ford, Conn. Bull. #31, [8] + 398 pp. Bruzelius, R. M. 1854. Beskrifning “ofwer Hydrachnider, som fore- 26 Entomologica Americana [Vol. 45 komma inom Skane” akademisk afhandhing. Lund, 56 pp., 5 pis. Faf. Butler, A. G. 1866. A list of the diurnal lepodoptera recently col- lected by Mr. Whitley in Hakodadi (north Japan). J. Linn. Soc. (Zool) IX: 50-59. . 1869 [1870]. Catalogue of diurnal lepidoptera described by Fabricius in the British Museum. London, Order of the Trustees, pp. vi + 303, 3 pis. . 1899. Notes on the genus Euchloe Hiibner, A genus of the Pierinae. Entomologist, London 32: 1-3. Clark, A. H. 1927. Fragrant butterflies. Washington, Smithsonian re- port for 1926, publication 2901, pp. 421-446 + 13 pis. . 1932. The butterflies of the District of Columbia and vicinity. Washington, Smithsonian Institution, Bull. U. S. Nat. Mus. no. 157, x + 338 pp., 64 pis. Clark, A. H. and L. F. Clark. 1951. Butterflies of Virginia. Wash- ington, Smithsonian Institution Misc. collection 116: 7: publi- cation no. 4050, viii + 240 pp., 30 pis., frontispiece colored. Comstock, J. H. and A. B. Comstock. 1923. A manual of the study of insects. Ithaca, N. Y., Comstock Publishing Co., 17th ed., xii + 702 pp., 795 figs. . 1929. How to know the butterflies. Ithaca, N. Y., Com- stock Publishing Co., xii + 312 pp., 45 pis. Comstock, W. P. 1940. Butterflies of New Jersey. A list of the lepi- doptera suborder Rhopalocera occurring in the state of New Jersey; giving time of flight, foodplants, records of capture with locality and date. Jour. N. Y. ent. Soc. 48: 47-84. Cook, C. 1948. Early spring collecting in Kentucky. Lep. News 2: 22. Cramer, P. and [in part] C. Stoll. 1779 [ 1 775]— 179 1 . Papillons exotiques des trois parties du monde FAsie, l’Afrique, et l’Amer- ique. [The title and the text in Dutch and French.] Amsterdam, S. J. Baalde; Utrecht, Barthelemy Wild, “1779” [1775-1776], 1: [4] + i-xxx, 1-16, 1-155 + [4], frontispiece and pis. 1-96 colored; “1779” [1777], 2: [4] + 1-152 + [4], frontispiece and pis. 97-192 colored; “1782” [1779-1780], 3: [8] + 1-36, 73 [sic], 38-176 + [2], frontispiece and pis. 193-288 colored; “1782” [1780-1784], 4: [4] + 1-252, 1-30 + [2], pis. 289-400 colored. Supplement a Fouvrage, intitule les papillons exotiques, des trois parties du monde l’Asie, l’Afrique, et FAmerique; par M. Pierre Cramer, contenant les figures exactes des chenilles et des chrysalides de Suriname; comme celles des plusieurs rares et nouvellement decouvertes papillons et phalenes. Amsterdam, Nic. Th. Gravius “1791” [1787-1791], 5: [4] + i-viii, 1-67, 60 [sic], 69-181, “382-384” [183-184], pis. 1-42 colored. Davenport, D. and V. G. Dethier. 1937 [1938]. Bibliography of the described life histories of the Rhopalocera of America north of Mexico 1889-1937. Ent. Americana, new ser. 17: 155-194. 1969] dos Passos and Klots: Systematics of Anthocharis midea 27 Donovan, E. 1800 [1802]. Epitome of the natural history of the in- sects of India and the islands of the Indian Sea. London, T. Bensley, ii + 68, 58 pis., colored. dos Passos, C. F., editor. 1951. The entomological reminiscenses of William Henry Edwards. Jour. N. Y. ent. Soc. 59: 129-186. dos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lep. Soc., Mem. No. 1, vi + 146 pp. Doubleday, E. and [in part] J. O. Westwood [1850-52, 143-4 p]. 1846-1852. The genera of diurnal lepidoptera: comprising their generic characters, a notice of their habits and transforma- tion, and a catalogue of the species of each genus. London, Longman, Brown, Green and Longman, 1846-1850, vol. 1, containing the families Papilionidae, Pieridae, Ageronidae, Danaidae, Heliconidae, Acracidae, and part of Nymphalidae; by E. Doubleday; illus. with 86 pis., by W. C. Hewitson, xii -f 250 pp., pis. A uncolored 1, 1*, 2-4*, 5-18, 18 [sic], 19-30 colored: 1850-1852, vol. 2 containing the remainder of the family Nymphalidae, and the families Morphidae, Brassalidae, Satyridae, Libytheidae, Erycinidae, Lycaenidae, and Hesperidae: with a supplement by J. O. Westwood, pp. [2] + 251-534, pis. 31-54, 54*, 55-66, 66 [sic], 68-80, [1] colored. . 1869. See Scudder, S. H., Editor. Drury, C. 1878. Catalogue of the lepidoptera, observed in the vicinity of Cincinnati, Ohio, including diurnals, Sphingidae, Aegeridae, Zygaenidae, Bombycidae, Noctuidae, Phalaenidae, and Pyrali- dae. J. Cin. Soc. Nat. Hist. 1: 12-23. Dyar, H. G. 1894. Notes on Pieris and Anthocharis. Can. Ent. 26: 100. . 1902 [1903]. A list of North American lepidoptera and key to the literature of this order of insects. Washington, Bull. U. S. Nat. Mus. no. 52, pp. xx + 724. Edwards, W. H. [1868-1872]. Synopsis of North American butterflies. Philadelphia, The Amer. Ent. Soc. [1868]: pp. 1-4; [1869]: pp. 5-6; [1870]: pp. 7-14; [1871]: pp. 15-38. . 1874-1884 [1878]. The butterflies of North America. Sec- ond series, vol. 2, pt. 7. Boston and New York, Houghton, Mifflin & Co. Anthocaris II, 4 figs., [83-84] pp. . 1877. Catalogue of the diurnal lepidoptera of America north of Mexico. Trans. Amer. ent. Soc. 6: 1-68. . 1881. On the length of life of butterflies. Can. Ent. 13: 205-214. . 1884. Revised catalogue of the diurnal lepidoptera of Amer- ica north of Mexico. Trans. Amer. ent. Soc. 1 1 : 245-338. . 1884. List of species of the diurnal lepidoptera of America north of Mexico. Boston and New York, Houghton, Mifflin & Co. [16] pp. 28 Entomologica Americana [Vol. 45 . 1888 [1897]. The butterflies of North America. Third series, vol. 3, pt. 6. Boston and New York, Houghton, Mifflin & Co., Anthocaris I, fig. 5 $ , figs. a-h3, preparatory stages, [57-61] pp. . 1892. Miscellaneous notes on butterflies, larvae, etc. Can. Ent. 24: 49-56, 105-111. Ehrlich, P. R. and A. H. Ehrlich. [1961]. How to know the butter- flies. Dubuque, Iowa; Wm. C. Brown Co., [8] + 262 pp., 525 figs. Engelhardt, G. P. 1937. See Siepmann, C. G. . 1939. In Tulloch (see Tulloch) . Fabricius, J. C. 1793. Entomologica systematica emendata et aucta . . . adjectis synonymis, locis, observationibus, descriptionibus. Copenhagen, C. G. Proft, vol. 3, Part I, [6] + 488 + 1794 pp.; Part II, 350 pp. Fazzini, L. D. 1934. Butterflies and moths of America. Racine, Wise., Whitman Publishing Co., 96 pp., 47 illus. colored, 1 black and white fig. Field, W. D. 1938. Manual of the butterflies and skippers of Kansas. Vol. 39, no. 10. Bull. Univ. Kans. Biol. Ser., Kansas State Printing Plant, 328 pp., 1 map. . 1940. New records of butterflies for Kansas (Lepidoptera: Rhopalocera) . J. Kansas ent. Soc. 13: 28-29. Forbes, W. T. M. 1926 [1928]. Order Lepidoptera in Leonard, A list of the insects of New York wilh a list of the spiders and certain other allied groups. Ithaca, N. Y., Cornell Univ., no. 101, 532- 687 pp. . 1960. Lepidoptera of New York and neighboring states. Ithaca, N. Y., Cornell Univ. Agric. Exper. Station, State College of Agriculture, Part IV, Mem. 371, 188 pp. French, G. H. 1886. The butterflies of the eastern United States. Phila., Pa., J. B. Lippincott Co., 402 pp. + 93 figs. Glover, T. 1878. Illustrations of North American entomology in the order lepidoptera. Washington, D. C., vol. 2, 1 17 pis., 47 pp. ms. Godart, J. B. See Latrielle, P. A. Gorby, S. S. 1892. Seventeenth annual report. Indiana Department of Geology and Natural Resources, Indianapolis, William Burford, 706 pp. + expl. of 20 pis. Grossbeck, j. A. 1905. Notes on the life history of Anthocharis genutiaVab. Ent. News 16: 131—134. Grote, A. R. 1900. The descent of the Pieridae. Proc. Amer. Philos. Soc. 39: 4-67. Harris, L., Jr. 1931. A list of the butterflies of Georgia. Trans. Georgia Naturalists Club 1 : 1-28, 1 pi., 1 fig. . [1950]. The butterflies of Georgia revised. Georgia Soc. Naturalists, Bull. no. 5, [6] + vii + 29 + [4] pp. Harris, T. W. See Scudder, S. H. 1969] dos Passos and Klots: Systematics of Anthocharis midea 29 Hemming, F. A. 1934. The generic names of the Holarctic butterflies. London, The Oxford Univ. Press, viii +184 pp. . 1937. Hiibner, a bibliographical and systematic account of the entomological works of Jacob Hiibner and of the supple- ments thereto by Carl Geyer, Gottfried Franz von Frolich and Gottlieb August Wilhelm Herrich-Schaeffer. London. Royal ent. Soc. 1 : xxxiv + 606, 1 portrait. . 1967. The generic names of the butterflies and their type- species (Lepidoptera: Rhopalocera) . Bull. B. M. (N. H.) Ent., Suppl. 9, London, 509 pp. Herrich-Schaeffer, G. A. W. 1867. Prodromus systematis lepi- dopterorum. Correspondenz. Blatt. des zoologisch-minerolo- gischen Vereines in Regensburg, vol. 21, pp. 138-144. Holland, W. J. 1898. The butterfly book. New York, Doubleday, Page & Co., xx + 382 pp., 183 + [1] figs., 48 pis. colored. . 1931. The butterfly book, new and thoroughly revised edi- tion. Garden City, N. Y., Doubleday, Doran & Co., Inc., xii + 424 pp., 77 pis. colored (4 black and white), 198 + [1] fig. Hornig, H. 1903. The feeding habits of the larva of Anthocharis genutia. Ent. News 14: 252. Hubner, J. 1 806— [1 819]. Sammlung exotischer Schmetterlinge. Vol. 1 . Augsburg, II + [4] pp., 213 col. pis. . 181 6— [ 1 826]. Verzeichniss bekannter Schmettlinge [sic]. Augsburg, 432 + 72 pp. International Code of Zoological Nomenclature. 1964. [Second ed.] Adopted by the XV International Congress of Zoology, London, International Trust for Zoological Nomenclature, xx + 176 pp. Jones, William “of Chelsea.” 1783— [1791]. leones [an unpublished collection of drawings in the Hope Department of the Museum at Oxford], 7 volumes. Kirby, W. F. 1871. A synonymic catalogue of diurnal lepidoptera. London, John van Voorst, viii + 690 pp. . 1879. Catalogue of the collection of diurnal lepidoptera formed by the late William Chapman Hewitson of Oatlands, Walton-on-Thames and bequeathed by him to the British Mu- seum. London, John van Voorst, iv + 246 pp. Klots, A. B. 1930. A generic revision of the Euchloini (Lepidoptera, Pieridae). Bull. Brooklyn Ent. Soc. 25: 80-95, 6 pis., 12 figs. . 1931. The generic synonymy of the North American Pieridae (Lepidoptera). Ent. News 42: 253-256. . 1931 [1933]. A generic revision of the Pieridae (Lepi- doptera). Ent. Amer. 12: (n. ser.): 139-242, 100 figs. . 1951. A field guide to the butterflies of North America east of the Great Plains. Boston, Mass., Houghton, Mifflin Co., Riverside Press, Cambridge, xvi + 350 pp., 40 pis. (16 colored), 9 line illus., 1 map. 30 Entomologica Americana [Vol. 45 . 1965. Some glaciation-isolated populations of North Ameri- can lepidoptera. Proc. Xllth International Congress of Ento- mology, London, 841 pp. Kusnezov, N. J. 1929. Faune de l’URSS, et des pays limitrophes In- sects Lepidopteres. Vol. 2, pp. 337-599, 1-64, 205-233 figs. Lamarck, J. B. A. P. M. and J. L. M. Poiret. 1810. Encyclopedic methodique. Botanique, Suppl. vol. 1, Paris, H. Agasse, xviii + 755 pp., Add. pp. 756-761. Lambremont, E. N. 1954. The butterflies and skippers of Louisiana. Tulane Studies in Zoology, New Orleans, Tulane University 1: 10: 127-164. Latreille, P. A. 181 9— [1 924]. Encyclopedic methodique, Histoire naturelle. Entomologie, ou histoire naturelle des crustaces, des arachnides et des insectes. Paris, H. Agasse, vol. 9 and supple- ment, 828 pp. Leussler, R. A. 1938. An annotated list of the butterflies of Nebraska (Lepidoptera: Rhopalocera) . Ent. News 49: 76-80. Linnaeus, C. 1753. Species plantarum, exhibentes plantas rite cognita, ad genera relatas, cum differentiis specificis, nominibus trurali- bus, synonymis selectis, locis natalibus, secundum systeme sexuale digestas. Stockholm, Laurentii Salvii 1: [13] + 560 pp., 2: [2] + 561-1200 pp. + index generum 32 pp. . 1761. Fauna Suecica sistens animalia Sueciae regni: Mam- malia, Aves, Amphibia, Pisces, Insecta, Vermes. Distributa per classes & ordines, genera & species, cum differentiis specierum, synonymis auctorum, nominibus incolarum, locis natalium, de- scriptionibus insectorum. Editio altera. Stockholm, Laurentii Salvii, [48] + 578 pp., frontispiece + 2 pis. Lutz, F. E. 1918. Fieldbook of the insects with special reference to those of northeastern U.S. aiming to answer common questions. G. P. Putnam’s sons, New York and London. The Knicker- bocker Press, ix + 510 pp., 145 pis. (44 colored). . [1921]. Fieldbook of insects with special reference to those of northeastern U.S. aiming to answer common questions, second edition, revised and enlarged. London, New York, G. P. Put- nam’s Sons, 562 pp., 101 + [1] pis., (24 colored) + frontispiece. . 1935. Fieldbook of insects with special reference to those of northeastern U.S. aiming to answer common questions, third edition, rewritten. New York, G. P. Putnam’s Sons, [10] + 510 pp., 100 pis. (23 colored). MacMillan, C. 1892. Metaspermae of the Minnesota Valley, a list of the high seed-producing plants indigenous to the drainage-basin of the Minnesota River. Minneapolis, Minn. Geol. Nat. Hist. Survey, Bot. Series 1 : 826 pp. Macy, R. W. and H. H. Shepard. [1941]. Butterflies — -a handbook of the butterflies of the U.S. complete for the region north of the 1969] dos Passos and Klots : Systematics of Anthocharis midea 31 Potomac and Ohio Rivers and east of the Dakotas. Minneapolis, viii + 248 pp. + 4 colored pis. + 7 figs. + 38 photos. Mather, B. 1952. Anthocaris in Mississippi. Lep. News 6: 42. Maynard, C. J. 1886. The butterflies of New England. Boston, Mass. Bradlee Whidden, iv + [2] + 66 + [2] pp. + 8 colored pis. . 1891a. Butterflies of New England, second edition. New- tonville. Mass., published by author, [4] + iv + 78 + [4] pp. + 10 pis. colored. . 1891b. A manual of North American butterflies. Boston, Dewolfe, Fiske and Co., [4] + iv + 226 pp. + 10 pis. colored + [1] + 59 figs. McDunnough, J. H. 1938. Check list of the lepidoptera of Canada and the U.S. of America, Part I, Macrolepidoptera. Mem. So. Calif. Acad. Sci. 1: [4] + 4 (corrigenda) + 276 pp. Morris, J. G. 1860. Catalogue of the described lepidoptera of North America. Washington, Smithsonian Inst., viii + 68 pp. . 1862. Synopsis of the described lepidoptera of North Amer- ica, Pt. I, Diurnal and crepuscular lepidoptera. Washington, Smithsonian Inst., xxviii + 358 pp. + 28 figs. Moschler, H. B. 1878. Catalogue of the lepidoptera of America north of Mexico. Part I, Diurnals by William H. Edwards. Philadel- phia, Amer. ent. Soc., Entomologische Zeitung (Stettin) 39: 297-310. Muesebeck, C. F. W. and L. M. Walkley. 1951. Family Braconidae in Hymenoptera of America north of Mexico, Synoptic Cata- logue . . . U.S. Dept, of Agric., Agricultural Monograph No. 2, Washington, D.C., 1420 pp. Muhlenberg, H. L. 1810. In Lamarck, Encyclopedic methodique. Botanique. Suppl. 1, Paris, H. Agasse, p. 409-415. Poiret, J. L. M. 1810. See Lamarck. Rawson, G. W. 1951. Hilltops and Anthocharis. Lep. News 5: 70. Remington, C. L. 1951. Hilltops and Anthocharis note. Lep. News 5: 70. Riley, C. V. 1871. Third annual report on the . . . insects of the state of Missouri . . . Jefferson City, Missouri, 176 + 8 pp., 70 figs. . 1878-1886 [1881]. Notes on North American microgasters with descriptions of new species. Trans. St. Louis Acad. Nat. Sci. 4: 296-324. Rober, J. See Seitz, A. Rowley, R. R. 1890. Cases of long pupal periods among lepidoptera. Can. Ent. 11: 375-384. Rummel, C. E. See Siepmann, C. G. Scudder, S. H. 1868. Check list of the butterflies of New England. Boston, 8 pp. . 1868. Supplement to a list of the butterflies of New England. Proc. Boston Soc. Nat. Hist 1 1 : 375-385. 32 Entomologica Americana [Vol. 45 . 1872. A systematic revision of some of the American butter- flies, with brief notes on those known to occur in Essex Co., Mass. Report Peabody Acad. Sci. for 1871, pp. 24-82. . 1872. Abbot’s notes on Georgian butterflies. Can. Ent. 4: 73-77. . 1875. Historical sketch of the generic names proposed for butterflies. A contribution to systematic nomenclature. Proc. Amer. Acad. Arts and Sci., Second series 10: 91-294. . 1889. The butterflies of the eastern U.S. & Canada with special reference to New England. Cambridge, Third volume, [2] + vi + [2] + 1775-1956 pp. + 90 pis. (20 colored) + 3 maps colored. . 1893. Brief guide to the commoner butterflies of the north- ern U.S. and Canada. New York, Henry Holt & Co., [2] -f xii + 206 pp. + 10 figs. Scudder, S. H., Editor. 1869. Entomological correspondence of Thad- deus William Harris, M.D., edited by Samuel H. Scudder. Boston, Boston Soc. Nat. Hist., xlviii + 375 pp., frontispiece, (portrait), illus., iv pis. (3 colored), 46 figs. (Occ. papers of the Boston Soc. of Nat. Hist.) Seitz, A. 1924. [1907-1924]. The Macrolepidoptera of the world. Vol. 5, (The American Rhopalocera.) Stuttgart, Alfred Kernen, viii + [4] + 1140 pp., 203 pis. colored (American edition). Seipmann, C. G., Secretary. 1931. Proceedings of the Society. Meeting of May 14, 1931. ibid Soc. 26: 265-268. . 1937. Proceedings of the Society. Meeting of January 14, 1937. Bull. Brooklyn ent. Soc. 32: 87-89. Skinner, H. 1898. A synonymic catalogue of the North American Rhopalocera. Philadelphia, P. C. Stockhausen, xvi + 100 + xiv pp. . [1905]. A synonymic catalogue of the North American Rhopalocera. Suppl. 1, Philadelphia, Amer. ent. Soc., 34 pp. . 1917. Anthocharis genutia and a new variety (Lepidoptera). Ent. News 28: 438. Skinner, H. and E. M. Aaron. 1889. A list of the butterflies of Philadelphia, Pa. Can. Ent. 21 : 126-131. Skinner, H., Recorder. 1892. Doings of Societies, Ent. News 3: 239- 240. Smith, J. E. 1797. The natural history of the rarer lepidopterous insects of Georgia . . . collected from the observations of Mr. John Abbot. London, T. Bensley, vol. 1, [4] + xvi + 100 pp., 100 pis. colored; vol. 2, pp. 101-214, 104 pis. colored. Smith, J. B. 1910. Annual report of New Jersey State Museum in- cluding a report of the insects of New Jersey. Trenton, New York, MacCrellish & Quigley, 888 pp., 3 portraits, 1 map colored. 1969J dos Passos and Klots: Systematics of Anthocharis midea 33 Smith, J. B., H. Skinner, C. H. Fernald, G. D. Hulst, and C. V. Riley. 1891. List of the lepidoptera of boreal America. Philadelphia, P. C. Stockhausen, Amer. ent. Soc., vi + 124 pp. Smith, J. B., H. Skinner, and W. D. Kearfoot. 1903. Check list of the lepidoptera of boreal America. Philadelphia, P. C. Stock- hausen, vi + 136 pp. Smyth, E. A., Jr. 1900. Anthocharis genutia Fabr. Ent. News 11: 465-468. Staudinger, O. and E. Schatz. 1888. Exotische Schmetterlinge. Fiirth, G. Lowensohn, vol. 1, [6] + 334 pp. + 1 map colored. Strecker, H. 1878. Butterflies and moths of North America with full instructions for collecting, breeding, preparing, classifying, pack- ing for shipment, etc., a complete synonymical catalogue of Macrolepidoptera with full bibliography to which is added a glossary of terms and an alphabetical and descriptive list of localities. Reading, Pa., B. F. Owen, [4] + ii + [2] + 284 pp., + 2 pis. Talbot, G. 1932-1935. Pieridae I— III. Lepidopterorum Catalogus, vol. 23, pts. 53, 60, 66; pt. 1, 320 pp; pt. II, 321-384; pt. Ill, 385-698 pp. Tietz, H. M. [1952]. The lepidoptera of Pennsylvania, a manual. State College, Pa., xii +194 pp. + 1 fig. Torrey, J. and A. Gray. 1838-1840. Flora of North America con- taining abridged descriptions of all the known indigenous and naturalized plants growing north of Mexico, arranged accord- ing to the natural system. New York, Wiley & Putnam, 1 : 82-87. Tulloch, G. S., Secretary pro tern. 1939. Meeting of May 12, 1938. Bull. Brooklyn ent. Soc. 34: 226-227. Viereck, H. L. 1912. Contributions to our knowledge of bees and ichneumon-flies, including the description of twenty-one new genera and fifty-seven new species of ichneumon-flies. U.S. Nat. Mus. Proc. 42: 613-648. Walker, F. 1863. List of the . . . Lepidopterous insects in the . . . British Museum, pt. 27, Crambites and Tortricities, 286 pp. Weed, C. M. 1924. Butterflies worth knowing. Doubleday, Page & Co. for Nelson Doubleday, Inc., xiv + 386 pp. + 48 pis., (32 colored) . Weidemeyer, J. W. 1863-1864. Catalogue of North American butter- flies. Proc. ent. Soc. Phila. 2: 143-154. Worthington, C. E. 1880. A list of diurnal lepidoptera inhabiting the state of Illinois. Can. Ent. 12: 46-50. Wright, W. G. 1905. Complete list of the butterflies of the U.S. [In] The butterflies of the West Coast of the U.S. illustrated with 940 figures in color-photography of butterflies from the West Coast nearly all of which were captured by the author with accurate data for each specimen with colored figures and 34 Entomologica Americana [Vol. 45 descriptions of many new species and new varieties now first published. San Francisco, The Whitaker & Ray Co., Inc., pp. 47-70. Wyss, A. E. [1932]. Guide to the butterflies of Cincinnati and vicinity. Misc. publication no. 7, compiled by Junior Soc. of Nat. Sci., Cincinnati, Ohio, [38] + errata [2] pp., 4 pis. Zimsen, E. 1964. The type material of J. C. Fabricius. Copenhagen, Munksgaard. 656 pp. + frontispiece. .w t , 5£ - \3D A REVISION OF THE GENUS LEPTOGLOSSUS GUERIN (HEMIPTERA: COREIDAE) By Richard Charles Allen Entomologica Americana Vol. 45, 1969, pp. 35-140 A REVISION OF THE GENUS LEPTOGLOSSUS GUERIN (HEMIPTERA: COREIDAE)1,3 By Richard Charles Allen* 1 2 3 Table of Contents Introduction 35 Historical Review 36 Materials and Methods 37 Taxonomic Characters 38 Transverse Fascia on Corium 38 Coloration 39 Tibial Dilations 39 Male Genitalia 45 Genus Leptoglossus Guerin 45 Species Groups 51 Key to Species 54 Acknowledgments 134 Literature Cited 135 Abstract: All previously known species of the genus Leptoglossus are discussed. Five new species are described and redescriptions are given for existing species. A discussion of the taxonomic characters used, a key to all species, and 67 figures are included. Introduction A comprehensive world-wide revision of the genus Leptoglossus has not been attempted since Stal’s Enumeratio Hemipterorum in 1870. Much of the material present in museums is largely un- identified, except for the best known species, or where keys to species are available for a particular geographic area. The purpose of this work is to bring together as much information as possible about the genus, to provide a comprehensive key to all existing species, and to redescribe the species that are poorly known. It is hoped that a basis has been estabished for further research in this group. Accepted for publication October 31, 1968. 1 A dissertation submitted in partial fulfillment of the requirements for the Degree of Master of Science at the University of Connecticut, Storrs, Con- necticut. 2 Present address: Department of Entomology, University of Kansas, Law- rence, Kansas. 3 This work was supported in part by National Science Foundation Grant GB-4306X. [35] 36 Entomologica Americana [Vol. 45 Leptoglossus is chiefly a neotropical genus with 26 of the 38 species found primarily in, or restricted to South America. Of the remaining species found in North and Central America, and the Caribbean, all are clearly related to the South American members. The single Eastern Hemisphere species, australis (Fabricius), does not present a baffling zoogeographic problem in view of the relation- ship between it and gonagra (Fabricius). These two species are very closely related, with australis an obvious derivative of a gonagra like stock that has crossed the water barriers and invaded the Ethiopian, Oriental, and Australo-Papuan regions. Further remarks concerning gonagra and australis may be found in the discussion under the latter species. Historical Review Guerin first described the genus Leptoglossus with dilaticollis as the only included species in 1838; however, the generic and specific names date from 1831 as they were used in reference to a figure pub- lished by Guerin in that year (Dupuis 1952). Leptoglossus remained monotypic until 1870. In 1837, Spinola described Anisoscelis, to which all previously and subsequently named species now in Lepto- glossus were assigned for many years. Stal (1862) described lineosus and established the genus Theognis to include those anisosceline spe- cies which had the first antennal segments equal to the length of the head. Mayr (1865, 1866) described a number of new species in Theognis , many of which have since been synonymized, but he pro- vided in some instances well executed descriptions. In 1870 Stal synonymized his genus Theognis with Leptoglossus and added three new species. Walker (1871) described five species in the genus Anisoscelis and one in Malvana. All of these belong to Leptoglossus and, of these six, three are valid species. Between 1871 and 1894 four more species were added to the genus, of these Distant described one and Berg three. I have synonymized one Berg species and assigned another to subspecific status. The Lethierry and Severin Catalogue in 1894 was the last check list of species to be compiled for Leptoglossus with 35 species included. Before 1909 only three species were known from North America north of Mexico, phyllopus (Lin- naeus), oppositus (Say), and corculus (Say). Subsequently Barber and Heidemann have described between them five additional species. Kiritshenko (1935) resurrected the genus Theognis as distinct from Leptoglossus. Hussey (1953) supported Kiritshenko’s separa- tion of the two genera. For reasons discussed later, I do not agree with Kiritshenko, and have retained Theognis as a junior synonym of Leptoglossus. 1969] Allen: Revision of the Genus Leptoglossus 37 Materials and Methods The type specimens of the following workers were made available through the cooperation of the various museum curators: the Stal collection from the Stockholm Museum; Walker and Dallas types from the British Museum (Natural History) ; the Berg collection from the La Plata Museum; Blote’s types from the Leiden Museum, Holland; and the types of Heidemann and Barber at the United States National Museum. Dr. I. Lansbury at Oxford compared material and made drawings of the genitalia and tibial dilations of the Westwood types. Approximately 2500 specimens were examined for this study. Genitalia dissections were accomplished according to the methods described by Ashlock (1957 and 1967). The aedeagus required much time and effort to inflate osmotically, and most attempts re- sulted in failure. By gently pulling on the membranous portion, it is usually possible to extend the aedeagus and with further careful manipulation the various lobes and appendages can be exposed (although their exact shape can only be estimated by this process). The aedeagal drawings were made from an aedeagus extended manually and therefore are only approximations of the size and shape of the various lobes. The genital capsule drawings are of the posterior view with only the dorsal margin and a portion of the lateral margin shown; the shape of the capsule, except for the dorsal margin, is essentially the same for all species. The corresponding morphological structures were drawn to the same scale. All measurements are in millimeters. The length of the head is measured along a median line from the base of the head to the apex of the juga. Although the tylus is the furthest extension of the head, that portion of it which extends past the juga was not included since it is variable between individuals of the same species and between different species. One example of this is in clypealis Heidemann which has the apex of the tylus produced as an acute spine. Width of head is across the eyes when viewed dorsally; interocular distance is the shortest distance between the eyes on the dorsal surface; anteocular distance is measured from the anterior margin of an eye to the apex of the juga; the intersegmental node between antennal segments three and four was included in the length of the third segment; length of pronotum is measured along the median line as is that of the scutellum; width across humeri is also the greatest width of pronotum; total length is the distance from the apex of the juga to the apex of the folded wings; the length of the hind tibia is measured from an imaginary base line drawn across the tibia from the inner, almost right-angle bend at the tibiofemoral joint, to the distal end of the tibia; the lengths of the inner and outer tibial dilations are measured from the same base 38 Entomologica Americana [Vol. 45 line explained for the tibia, to the end of the dilations, regardless of where the dilations actually begin on the proximal portion of the tibia. An exact measurement is sometimes difficult because the dilation diminishes very gradually distally. The width of the dilations is measured from the center of the tibia to the margin of the dilations, with the dilations held in a horizontal plane. Complete descriptions are given for the less well known species. The North American species are described briefly, and the reader is referred to the original and readily available descriptions of Barber (1918), Heidemann (1909, 1910), and the descriptions given in Blatchley (1926). The references for each species include all synonymies, and additional papers that give useful descriptions, figures, keys, or biological information. Many older references that were cited in the Lethierry and Severin Catalogue (1894), and also those papers pertaining to locality records that were cited in the Van Duzee Catalogue (1917), are omitted here. Host plant records cited without references are taken from labels on specimens in the material examined. Taxonomic Characters Most of the characters used in this paper are those employed by previous workers on this genus. In the past, some taxonomic con- fusion has resulted due to reliance on only one or two of what have proven to be variable characters. The significant characters, their usefulness in distinguishing species, groups of species, and their variability are discussed below. (1) Transverse Fascia on Corium — This is an extremely useful character and its presence or absence, and shape can be de- termined with ease. In many species the pale fascia is as constant as any morphological character, but occasionally the fascia is absent in species that normally have it, such as ingens (Mayr), stigma (Herbst), concolor (Walker), neovexillatus n. sp. and zonatus (Dallas). In ingens the absence or presence of the fascia seems to be correlated with geographic distribution. The fascia also varies in completeness, and may be wide and brightly pigmented or narrow and faint. The terms “irregular” and “straight” pertain to the margins of the fascia: an irregular fascia more or less follows the branching of the radial and medial and the mediocubital cross-vein, this produces sharply angled saw-like margins; the straight fascia occupies the same area as an irregular one, but the area between the veins is also pig- mented, a condition producing an evenly margined fascia. L. ash- meadi Heidemann and macrophyllus Stal have an irregular fascia, but on some individuals this is difficult to determine as the margin 1969] Allen: Revision of the Genus Leptoglossus 39 may be only slightly angular and similar to a straight margined fascia. In order to avoid confusion in using the key, I have considered these two species as possessing both types of fascia. (2) Coloration — There are a number of useful and reliable color patterns that can be used in grouping species, as well as in separating closely related species. A large group has the thoracic and abdominal venter mottled with numerous small piceous spots (about the size of an ocellus ) . Another useful pattern is the presence of con- trasting yellowish maculae on the thoracic pleura; these maculae are large (about the size of an eye). In one group of species, the indi- viduals have five or six maculae on each side of the thorax; in another group the maculae number approximately ten or twelve. A third color pattern on the thoracic pleura is the shape of pale longitudinal fasciae. Members of this genus also possess various contrasting spots, fasciae, and rectangular bands on the pronotum. These color patterns are useful for distinguishing related species, but in some instances are highly variable and must be used with caution. The key contains a number of couplets which rely on pronotal color patterns, but I have made every attempt to use them only where those species involved showed consistence in expressing a particular pattern. (3) Tibial Dilations — These membranous plate-like expan- sions on the hind tibiae (Figs. 52-63) can be at times completely diagnostic in themselves as in corculus, gonagra, alatus, and lineosus, or in other instances the dilations are at best only a secondary char- acter. Some examples of the latter case are typical of the species zonatus, stigma , concolor , and chilensis. The outer dilation refers to the larger dilation, which is directed laterally when the hind tibia is in the position shown in Figs. 1-5. The inner dilation is the medially directed portion. The term “lance- olate” refers to a dilation that has the margin entire or almost entire (Figs. 57-61); and “phylliform” pertains to a dilation which is emarginate or scalloped on the margin (Figs. 52-56, 62-63). The inner dilation is in all species lanceolate with small teeth on the margin, and in all but a few species it is smaller both in width and length than the outer. The outer dilation may be either lanceolate or conspicuously phylliform. A phylliform dilation has a spine-like tooth where an edge of an emargination comes to a point. In many of the older descriptions a phylliform dilation was referred to as having “two or three teeth.” A certain amount of variation in the shape of a phylliform dilation can be expected. All species with this type of a dilation may have one to two deep emarginations, and only rarely will all the emarginations be shallow. A lanceolate outer dilation will usually have at least one shallow emargination. The width and length Entomologica Americana [Vol. 45 Fig. 1 . Leptoglossus dilaticollis , 1969] Allen: Revision of the Genus Leptoglossus 41 Fig. 2. Leptoglossus crassicornis. 42 Entomologica Americana [Vol. 45 Fig. 3. Leptoglossus grenadensis. 1969] Allen: Revision of the Genus Leptoglossus Fig. 4. Leptoglossus pallidivenosus. 44 Entomologica Americana [Vol. 45 Fig. 5. Leptoglossus alatus. 1969] Allen: Revision of the Genus Leptoglossus 45 of a dilation are also useful characters, but the length, as mentioned earlier, must be used carefully. I have used the various shapes of the dilation as a diagnostic character only when there was a considerable and constant difference between species. (4) Male Genitalia — The various parts of the male genitalia have been a rewarding character in all but a few instances. By far the most useful structure is the aedeagus. This has proven to be diagnostic between closely related species when the clasper and capsule were similar, and also has been useful in delineating groups of species. I have limited my discussion of the aedeagus to the dorsal sac (Schaefer 1968) of the conjunctiva, because this area is most useful, with its various lobes and scerotized spines. The different lobes on the dorsal sac are indicated according to their position as proximal, medial, or distal (Figs. 64-67). The dorso-lateral appendage (Fig. 67) is generally a well sclerotized flat appendage and basal to the dorsal sac. For descriptive purposes I have referred to the clasper as having three parts, a base, shank, and hook (Fig. 17). The postero-lateral dorsal margin of the capsule is an area displaying good specific char- acters. I have used the terms median notch and dorsal prongs in describing the shape of the margin (Fig. 42). Genus Leptoglossus Guerin Leptoglossus Guerin, 1831 (1838): pi. 12, fig. 9. — Guerin, 1838:174. Stal, 1870:160.— Kirkaldy, 1906:257.— Gibson and Holdridge, 1918:3. Anisoscelis Spinola, 1837:200. Theognis Stal, 1862:294. — Kiritschenko, 1935:191. — Hussey, 1953: 33. Small to large species, elongate; head porrect, longer than wide and usually shorter than length of pronotum, prolonged anterior to antenniferous tubercules, tylus slightly exceeding juga usually rounded, ocelli widely separated, distance between ocelli greater than distance from ocellus to eye; pronotum subhexagonal with anterior face decli- vent, humeral areas usually expanded, sometimes greatly so, greatest width of pronotum across humeri, posterior margin concave, in front of it an unevenly raised transverse ridge, calli present; length and width scutellum subequal; labial length variable, segments one and two subequal with third shortest, fourth variable; first antennal seg- ment curved and thickest, usually subequal to length of head, at least longer than anteocular distance, first segment shortest, with second segment longer than third, fourth segment equal to or longer than third segment; membrane slightly surpassing abdomen; all femora 46 Entomologica Americana [Vol. 45 armed beneath with two rows of distally directed teeth, individual teeth gradually increasing in size distad, tuberculate laterally and above, tubercules more or less oriented in rows, hind femora swollen and usually thicker in males than females; hind tibiae dilated, outer dilation variable in size and shape, usually wider than interocular distance. Type species — Leptoglossus dilaticollis Guerin, 1831. Monobasic. Stal established the genus Theognis in 1862, and synonymized it with Leptoglossus in 1870. The generic status remained unchanged until Kiritshenko (1935) restored Theognis as distinct. This action left only dilaticollis in Leptoglossus. The following is a discussion of the characters that Kiritshenko listed in his diagnosis of Leptoglossus. After each of his characters, I point out similar characters that can be found in species in the genus Theognis , and which in my opinion precludes the separation of the two genera. ( 1 ) Pronotum with the humeral angles produced as long an- teriorly curving processes — alatus (Walker) (Fig. 5) has similar angles, although not quite so wide. (2) All lateral margins strongly dentate. L. alatus has equally long teeth, and dentatus Berg despite its small size is strongly dentate. (3) Pronotal surface above with deeply excavated rounded foveolae. Ruhrescem (Walker) and fulvi- cornis (Say) both have rugulose and deeply pitted pronotal surfaces, although the punctations are not so deep or so prominent. (4) Body robust, very large species (length 27-39 mm). The two females of humeralis n. sp. approach the lower limit of the range for dilaticollis. It should be pointed out that the lower limit of the range for dilaticollis is of a male, and the upper limit for humeralis is of females, so that dilaticollis is actually larger. (5) The length of the head is three times the interocular distance. Not all dilaticollis specimens have the head quite this long, the lowest ratio of seven specimens was 2.5 and a single specimen of fulvicornis has the head 2.4 times the interocular distance. These differences between the two “genera” are ones of degree, not of kind. Stronger evidence for retaining Theognis species with Lepto- glossus is the similarity of the genitalia, and this is discussed under the dilaticollis group. It is true that dilaticollis is a unique species, but so is alatus. The genitalia of lineosus and subauratus Distant are as different from other species as are the external characters of dilati- collis; therefore, they too are unique. The entire genus is not a homogeneous unit, and until a complete re-evaluation of the generic limits and the relationships among genera of the Anisoscelidini is 47 1969] Allen: Revision of the Genus Leptoglossus Figs. 6-30. Right claspers, dorsal view. Fig. 6. Leptoglossus cinctus. Fig. 7. L. harpagon. Fig. 8. L. lineosus. Fig. 9. L. dilaticollis. Fig. 10. L. fulvicornis. Fig. 11. L. rubrescens. Fig. 12. L. australis. Fig. 13. L. chilensis. Fig. 14. L. quadricollis. Fig. 15. L. dentatus. Fig. 16. L. macro- phyllus. Fig. 17. L. ingens : Sh, shank; Hk, hook; M Lo, median lobe: I B Lo, inner basal lobe. Fig. 18. L. zonatus. Fig. 19. L. impictipennis. Fig. 20. L. conspersus. Fig. 21. L. stigma. Fig. 22. L. concolor. Fig. 23. L. pallidivenosus. Fig. 24. L. lonchoides. Fig. 25. L. balteatus. Fig. 26. L. corculus. Fig. 27. L. occidentalis. Fig. 28. L. clypealis. Fig. 29. L. brevirostris. Fig. 30. L. ashmeadi. 48 Entomologica Americana [Vol. 45 Figs. 31-51. Genital capsules, posterior view. Fig. 31. Leptoglossus cinctus. Fig. 32. L. harpagon. Fig. 33. L. lineosus . Fig. 34. L. dilati- collis. Fig. 35. L. fulvicornis. Fig. 36. L. rubrescens. Fig. 37. L. australis. Fig. 38. L. chilensis. Fig. 39. L. dentatus. Fig. 40. L. macrophyllus. Fig. 41. L. ingens. Fig. 42. L. zonatus, Dors Prg, dorsal prongs; M Notch, median notch. Fig. 43. L. impictipennis. Fig. 44. L. concolor. Fig. 45. L. pallidivenosus. Fig. 46. L. corculus. Fig. 47. L. occidentalis. Fig. 48. L. clypealis. Fig. 49. L. brevirostris. Fig. 50. L. ashmeadi. Fig. 51. L. balteatus. 1969] Allen: Revision of the Genus Leptoglossus 49 Figs. 52-63. Tibial dilations. Fig. 52. Leptoglossus australis. Fig. 53. L. australis. Fig. 54. L. gonagra. Fig. 55. L. ingens. Fig. 56. L. quadricollis. Fig. 57. L. corculus. Fig. 58. L. occidentalis . Fig. 59. L. clypealis. Fig. 60. L. lineosus. Fig. 61. L. lonchoides. Fig. 62. L. chilensis chilensis. Fig. 63. L. chilensis concaviusculus. 50 Entomologica Americana [Vol. 45 Figs. 64—67. Aedeagi, lateral view. Fig. 64. Leptoglossus cinctus. Fig. 65. L. harpagon. Fig. 66. L. rubrescens. Fig. 67. L. ingens, Ant D Lo, anterior distal lobe; L D Lo, lateral distal lobe; M Lo, median lobe; Prox Lo, proximal lobe; Dors Sac, dorsal sac; Gon Pr, gonoporal process; Hel Proc, helical process; Dors Lat App, dorso-lateral appendage; Ph, phallosoma. 1969] Allen: Revision of the Genus Leptoglossus 51 accomplished, there is no reason to revive or establish monotypic genera based on a few bizarre external characters. Species Groups The aedeagus and the color patterns have been used to establish species groups. Some species and groups of species may not be natural units, nor related to one another to the same degree as other groups. These groups, in turn, have been assembled into two major divisions based on the conformation of the dorsal sac on the conjunctiva. Within these two divisions, I have discussed the closely related species as species groups. Division A Comprised of 13 species which have the dorsal sac of the con- junctiva without a sclerotized anterior distal lobe (Figs. 64-66), and which, with the exception of dilaticollis and fulvicornis, lack the small piceous spots on the venter. Members of this division are more diverse than the species in Division B. Cinctus Group — This group includes cinctus (Herrich-Schaeffer), fasciatus (Westwood), and crassicornis (Dallas), and is characterized by the presence of six yellow spots on each side of the thoracic pleura, a yellow spot on the last abdominal tergum (frequently absent in females), lanceolate tibial dilations, and the aedeagus (Fig. 64) with two pairs of widely separated medial lobes. Harp agon Group — This group includes two “pairs” of species each pair only distantly related to the other. L. harpagon (Fabricius) and flavosignatus Blote are characterized by the presence of two yellow spots each, on the disk and posterior margin of the pronotum, two spots on the corium, and six yellow spots on each side of the thoracic pleura. The dorsal sac of the conjunctiva (Fig. 65) is unsclerotized, and bears three large proximal lobes. The tibial dilations are short and not deeply emarginate. L. lineosus and subauratus are char- acterized by having two pale longitudinal fascia on each side of the thoracic pleura, strongly contrasting pale veins on the clavus and corium, and lanceolate outer tibial dilations (Fig. 60). The species pairs are grouped primarily on the shape of the clasper (Figs. 7-8) and the capsules (Figs. 32-33). In both groups the clasper has the inner margin of the shank deeply scooped out and the posterior dorsal margin of the capsule is greatly produced back- wards. In addition, the dorso-lateral appendage of the conjunctiva is long and heavily sclerotized. The declivous anteocular portion of the head is conspicuously evident in lineosus and subauratus, but there is only a slight indication of this in harpagon and flavosignatus. 52 Entomologica Americana [Vol. 45 L. alatus also belongs to this group and most closely resembles lineosus and subauratus because of the declivent anteocular portion of the head. However, the shape of the pronotum (Fig. 5) and the color is so strikingly different that it is obviously a highly derived species. Dilaticollis Group — This is probably an artificial grouping com- posed of fulvicornis, rubrescens, and dilaticollis. All three species have the clasper with the base broad, the inner basal lobe obsolete and the shank without a median lobe. L. fulvicornis and dilaticollis have a long unsclerotized dorsal sac whereas in rubrescens it is short. The various lobes on the dorsal sac and the shape of the dorsal margin on the capsule are all quite different. This group shares in common the presence of a yellowish longitudinal stripe on the abdominal dorsum, a rugulose-like pronotal surface, dentate pronotal margins, and lanceo- late tibial dilations. Both fulvicornis and dilaticollis have small piceous spots on the venter as do members in Division B, but the importance of this character as an indication of relationships is diffi- cult to assess, since the aedeagi are so different. Australis Group — L. australis and gonagra are the only members of this group and are characterized by the presence of a pale arcuate transverse fascia on the pronotal disk and pale longitudinal vittae on the abdominal venter. The similar base on the clasper and the simple dorsal sac on the conjunctiva with only two short median lobes near the apex suggest that fulvicornis is somewhat related, but as men- tioned in the previous group, these species are rather specialized mem- bers of this genus. Division B This Division contains the largest number of species, and on the whole is a rather uniform complex. The twenty-four species included here may be characterized as follows : a distinctive aedeagus with the dorsal sac of the conjunctiva (Fig. 67) bearing low rounded usually sclerotized proximal lobes, a pair of median lobes which may have secondary appendages, and usually bearing sclerotized distal lobes of which one is larger and anterior to the sclerotized smaller more laterally placed lobes. With few exceptions the thoracic and ab- dominal venter is mottled with numerous small piceous spots. Chilensis Group — Members of this group lack the transverse fascia on the corium, have a short labium usually extending to between the meso- and metasterna, serrate to dentate lateral pronotal margins, and a genital capsule without dorsal prongs. Species in this group are chilensis , impictus, quadricollis, and dentatus. Stigma Group — This group of species possesses a genital capsule with dorsal prongs (Figs. 40-43), claspers with both the inner basal 1969] Allen: Revision of the Genus Leptoglossus 53 lobe and the median lobe on the shank well developed, usually a long labium, conspicuously phylliform dilations, and a transverse fascia which, when present is irregular. Most of the members are of medium size. Included species are stigma , zonatus, concolor, impictipennis , neovexillatus, ingens , macrophyllus, oppositus, grenadensis, con- spersus, lonchoides , pallidivenosus, and humeralis. Within this group ingens and macrophyllus are most closely allied and have a long outer tibial dilation, a “V” shaped median notch on the genital capsule, and low dorsal prongs. L. zonatus , neovexillatus , and impictipennis have similar genitalia and two rounded spots on the pronotum. L. grenadensis and conspersus also possess pronotal markings, but differ from the last three species in that the anterior distal lobe on the dorsal sac is larger and more heavily sclerotized, and the inner basal lobe on the clasper is longer (Fig. 20). The clasper of oppositus and stigma is very similar to that of zonatus and its relatives, although oppositus and stigma do not have pronotal markings. L. concolor and pallidivenosus lack dorsal prongs on the capsule. Nevertheless, the structure of the dorsal sac and the general habitus clearly ally these species with the other members of the stigma-group (see discussion under the individual species). L. lonchoides apparently belongs to this group, despite external differences, because of similarities in the genitalia to that of zonatus. Although males of humeralis were not available, this species will prob- ably prove to belong to this group on the basis of the long labium and phylliform tibial dilations. Phyllopus Group — L. phyllopus and balteatus are included here and this group is characterized by a straight transverse fascia on the corium, low rounded dorsal prongs on the capsule (Fig. 51), a clasper (Fig. 25) without a median lobe, and a poorly developed inner basal lobe. These two species are close to the stigma-group in general habitus. Cor cuius Group — Externally these species are very similar to one another, with the same general color pattern, broadly rounded humeral angles, a gradually declivent pronotum, an irregular transverse fascia, and lanceolate tibial dilations. The group includes three North Amer- ican species corculus, clypealis , and occidental^ . The clasper (Figs. 26-28) and capsule (Figs. 46-48) of the species differ, but the aedeagi are essentially the same. The clasper of clypealis is similar to that of zonatus, and there is an indication of dorsal prongs on the capsule. The shape of the pronotum and the tibial dilation of the corculus- group resemble lonchoides, so that there seems to be a definite link in both genitalia and general habitus to the stigma- group. The remaining two species in this Division are ashmeadi and 54 Entomologica Americana [Vol. 45 brevirostris , both appearing to be derived members of the stigma- group. These two species share in common an irregular transverse fascia; a short labium, extending between the meso- and metacoxae; small size, and, on the dorsal sac, a sclerotized area near the larger median lobe. L. ashmeadi has small dorsal prongs on the capsule, but the short labium, orange and black pigmentation, and the lack of numerous small piceous spots on the venter rather isolates this species from brevirostris and the stigma group. L. brevirostris on the other hand, resembles a diminutive stigma or concolor , but has a rounded median notch and obsolete dorsal prongs on the genital capsule. Key to Species of Leptoglossus 1. Thoracic pleura dark with at least three, usually more, strongly contrasting yellowish maculae (about the size of an eye), or yellowish maculated fasciae; thoracic and abdominal venter without numerous small piceous spots _ 34 Thoracic pleura orange to dark reddish brown without strongly contrasting yellowish markings; thoracic and abdominal venter usually with numerous small piceous spots 2 2. Width across humeri 5.5 to 6 times width across anterior pronotal margin; large species, length 26-39 mm 3 Width across humeri at most 5 times width anterior pronotal margin; smaller species less than 25 mm in length 4 3. Pronotum with humeral areas produced as large, broadly rounded anteriorly curving processes; corium with a straight transverse fascia; hind tibiae with outer dilations lanceolate (Fig. 1) dilaticollis Guerin (p. 79) Pronotum with humeral areas produced into long, gradually tapering projections; corium without a transverse fascia; hind tibiae with outer dilations distinctly phylliform humeralis n. sp. (p. 126) 4. Transverse fascia on corium always present and straight 5 Transverse fascia on corium irregular, not straight; or fascia absent 8 5. Dorsum piceous; all marginal areas of pronotum continu- ously and widely yellowish orange ashmeadi Heidemann (p. 134) Dorsum reddish brown; at most only the posterior marginal area of the pronotum yellowish 6 6. Anterior pronotal disk with a yellowish transverse rec- tangular band; inner tibial dilation considerably shorter 1969] Allen: Revision of the Genus Leptoglossus 55 than outer, occupying 65-70% the length of outer dila- tions macrophyllus Stal (p. 104) Anterior pronotal disk concolorous or with two longitudi- nally ovoid spots; inner tibial dilations slightly shorter than outer, occupying 85-90% the length of outer dila- tions 7 7. Humeral angles acuminate, acute and obliquely ascending; pronotal disk with contrasting yellow areas in the form of distinctly separated ovoid spots or larger more quadrate spot partially fusing medially, posterior marginal area of pronotum yellow; hind wings unicolorous (West Indies) balteatus (Linnaeus) (p. 97) Humeral angles more rounded, subacute; pronotum con- colorous, disk rarely with yellowish areas (if yellow areas are present, they are poorly defined, more diffused and dull), posterior marginal area of pronotum never yellow; hind wings with basal half dark, distal half clear (North and Central America). phyllopus (Linnaeus) (p. 100) 8. Pronotum with lateral and postero-lateral margins serrate or dentate 9 Pronotum with lateral margins entire; postero-lateral mar- gins serrate, dentate, or entire 18 9. Clavus and corium dark with strongly contrasting pale veins, a pale transverse fascia, or only two large pale maculae __ 10 Clavus and corium without strongly contrasting pale mark- ings, usually unicolorous 12 10. Humeral areas produced as large broadly rounded lateral projections; pronotum piceous with a wide orange band on disk; corium with two large pale maculae (Fig. 5) ____ alatus (Walker) (p. 71) Humeral areas produced as tapering lateral projections not broadly rounded; pronotal disk unicolorous; corium with a pale irregular transverse fascia 11 11. Labium short extending at most to middle of metasternum; posterior marginal area of pronotum concolorous with disk or slightly darker, never pale; small species less than 16 mm in length (North America) brevirostris Barber (p. 133) Labium longer extending to at least anterior margin of second abdominal sternum; posterior marginal area of pronotum pale, strongly contrasting with remainder of disk; larger species, 20 mm or more in length (Fig. 4) (Panama) pallidivenosus n. sp. (p. 128) 56 Entomologica Americana [Vol. 45 12. Labium long, extending to at least middle of third ab- dominal sternum; entire dorsum unicolorous dark reddish brown; abdominal dorsum piceous with a yellow median longitudinal stripe the width of scent gland scars 13 Labium shorter rarely extending to posterior margin of second abdominal sternum (usually to middle of meta- sternum); dorsum with at least some dark and lighter areas; abdominal dorsum unicolorous or with lateral areas yellow but never with a central yellow longitudinal stripe 1 4 13. Pronotal humeral area broadly rounded with humeral angles blunt, non-acute (North America) fulvicornis (Westwood) (p. 78) Pronotal humeral area expanded into large gradually taper- ing projections, humeral angles acute (Brazil) rubrescens (Walker) (p. 76) 14. Entire length of lateral pronotal margins with well developed teeth; a single large rounded piceous macula (about the size of an eye) on mesopleuron; abdominal dorsum mesally dark with lateral areas largely yellow dentatus Berg (p. 95) Anterior half of pronotal lateral margins entire, only the posterior half serrate; mesopleuron with at most small piceous maculae (about the size of an ocellus); ab- dominal dorsum without yellowish areas 1 5 15. Male genital capsule with prominent dorsal prongs (Fig. 43); anterior half of corium with conspicuously less pubescence than posterior half, this pubescence very dense on veins; hind tibiae with width of outer dilations equal to or greater than width of head impictipennis Stal (p. 116) Male genital capsule without dorsal prongs; pubescence evenly distributed over clavus and corium, if pubescence more dense on posterior half of corium then hind tibiae with width of outer dilations subequal to interocular dis- tance 1 6 16. Wider more robust species, total length less than 2.5 times width across humeri; dorsal surface of male hind femora with 15-20 prominent tubercules ____ impictus (Stal) (p. 91) Narrower more elongate species, total length greater than 2.5 times width across humeri; dorsal surface of male hind femora with at most 10 prominent tubercules 17 17. Fourth antennal segment averaging 1.2 times width of head; hind tibiae (Fig. 62) with outer dilations narrower and 1969] Allen: Revision of the Genus Leptoglossus 57 shorter occupying 62-71% the length of hind tibiae; entire venter usually with numerous small piceous spots chilensis chilensis (Spinola) (p. 85) Fourth antennal segment averaging 1.5 times width of head; hind tibiae (Fig. 63) with outer dilations wider and longer, occupying 71-80% the length of hind tibiae; nu- merous small piceous spots usually confined to abdominal midventer chilensis concaviusculus Berg (p. 88) 18. Hind tibial dilations lanceolate, without deep emarginations along outer margins (Figs. 57-59, 61) 19 Hind tibial dilations conspicuously phylliform with at least one (usually more) deep emargination (Figs. 55-56) 22 19. Tylus extending beyond juga as a porrect spine which is as long as, or longer than, length of an eye; broad irregular transverse fascia on corium ____ clypealis Heidemann (p. 132) Tylus rounded or pointed apically, but never spinosely pro- duced; transverse fascia on corium narrow, confined to veins or absent 20 20. Labium short, reaching at most to middle of third ab- dominal sternum, third labial segment never extending beyond anterior margin of metasternum; pronotal lateral margins straight, humeral angles acute (Brazil) lonchoides n. sp. (p. 124) Labium long, reaching well onto abdomen, at least to the anterior margin of fourth abdominal sternum, third labial segment extending to at least posterior margin of meta- sternum; pronotal lateral margins sinuate, humeral angles broadly rounded (North America) 21 21. Inner and outer tibial dilations of nearly equal length, dila- tions shorter, occupying less than 70% length of hind tibia (Fig. 58) occidentalis Heidemann (p. 131) Inner and outer tibial dilations not of equal length, outer dilations conspicuously longer than inner and occupy- ing 85% or more length of hind tibia (Fig. 57) cor cuius (Say) (p. 130) 22. Hind tibiae with outer dilations very long extending for 85-95% length of tibiae and inner dilations much shorter occupying 55-70% length of outer dilations (Fig. 55) __ 23 Hind tibiae with outer dilations shorter occupying 80% or less length of tibiae and inner dilations only slightly shorter than outer dilations 24 23. Dorsum light reddish brown; anterior portion of pronotal disk with a yellowish rectangular transverse band not 58 Entomologica Americana [Vol. 45 reaching lateral margins macrophyllus Stal (p. 104) Dorsum piceous to dark reddish brown; pronotal disk uni- colorous or the entire anterior half of pronotum before humeri yellow ingens (Mayr) (p. 102) 24. Pronotum dark without strongly contrasting yellowish and dark areas (specimens of quadricollis may have an en- tirely yellow pronotum) 25 Pronotum with strongly contrasting yellowish and dark areas 29 25. Labium short, not extending past metasternum; fourth labial segment shorter than first antennal segment 26 Labium longer, usually extending well onto abdomen, at least past the posterior margin of second abdominal sternum; fourth labial segment longer than first antennal segment 27 26. Transverse fascia on corium present (occasionally very faint); tibial dilations short, occupying 70% or less length of hind tibiae (North America) __ brevirostris Barber (p. 133) Transverse fascia on corium absent; tibial dilations longer, occupying 75-80% length of hind tibiae (Fig. 56) (Brazil) quadricollis (Westwood) (p. 93) 27. Postero-lateral margins of pronotum entire; first antennal segment unicolorous; transverse fascia on corium reduced to a short oblique mark on medial vein oppositus (Say) (p. 123) Postero-lateral margins of pronotum serrate; first antennal segment bicolored with a pale inner marginal area; trans- verse fascia on corium complete (rarely entirely absent) 28 28. Pronotal calli with a rough surface texture, numerous dark thick hairs interspersed with pale hairs on anterior pro- notal disk; male genital capsule with a rectangular median notch, no dorsal prongs (Fig. 44) concolor (Walker) (p. 118) Pronotal calli smooth, pale pilose hair only on anterior pronotal disk; male genital capsule with a rounded median notch, dorsal prongs present stigma (Herbst) (p. 120) 29. Dorsum piceous; all marginal areas of pronotum continu- ously and widely yellowish-orange; venter largely bright orange with few small piceous maculae ashmeadi Heidemann (p. 134) Dorsum dark reddish brown; pronotal disk with yellowish areas, or posterior marginal area yellow but never all margins of pronotum continuously yellow; venter tan with numerous small piceous spots 30 30. Second antennal segment entirely fuscous; pronotum with 1969] Allen: Revision of the Genus Leptoglossus 59 two large diffuse yellowish-orange areas occupying most of disk and anterior half of humeral area; pronotum behind humeri dark, without contrasting pale area ____ conspersus Stal (p. 106) Second antennal segments bicolored with medial portion pale and the remaining areas dark; pronotal disk with two distinctly round spots or, the posterior disk and the area behind humeri are pale yellow 31 31. Anterior portion of pronotal disk without two distinct pale round spots (Greater Antilles) __ concolor (Walker) (p. 118) Anterior portion of pronotal disk with two distinct pale round spots 32 32. Posterior marginal area of pronotum with a narrow yellow- ish band strongly contrasting with adjacent darker areas; lateral margins of pronotum nearly straight (Fig. 3) (Lesser Antilles) grenadensis n. sp. (p. 108) Pronotal area behind humeri concolorous without pale areas; lateral margins of pronotum sinuate 33 33. Left median lobe on dorsal sac of aedeagus long, without a secondary lobe; transverse fascia on corium, if present, narrow, confined to veins neovexillatus n. sp. (p. 113) Left median lobe on dorsal sac of aedeagus blunt and with a long secondary lobe; transverse fascia rarely absent, and usually wide, on both corium and veins zonatus (Dallas) (p. 110) 34. Clavus and corium dark brown with strongly contrasting pale yellow or ochraceous veins 35 Clavus and corium dark, veins usually concolorous or at most bright red, but never pale yellow or ochraceous __ 36 35. Pronotum with an ochraceous transverse fascia between humeri narrow, approximately width of fore femora; first antennal segment unicolorous, piceous; remaining seg- ments bicolored with strongly contrasting pale and dark areas lineosus (Stal) (p. 73) Pronotum with an ochraceous transverse fascia between humeri wide, covering most of anterior portion of disk; all antennal segments unicolorous, pale ochraceous ___ _ subauratus Distant (p. 75) 36. Pronotal disk with a narrow arcuate pale transverse fascia; thoracic pleura with 10 to 12 pale maculae on each side; abdominal venter with six to seven complete or macu- lated longitudinal fasciae 37 Pronotal disk without a narrow transverse fascia, usually 60 Entomologica Americana [Vol. 45 unicolorous or with round spots; thoracic pleura never with more than six yellowish maculae on each side; abdominal venter without longitudinal fasciae 38 37. Hind tibiae with outer dilations extending for 85-90% the length of tibiae (Fig. 54); abdominal dorsum with two yellowish spots on scent gland scars (Western Hemi- sphere) gonagra (Fabricius) (p. 84) Hind tibiae with outer dilations shorter, extending for at most 75% the length of tibiae (Figs. 52-53); abdominal dorsum unicolorous without yellowish spots (Eastern Hemisphere australis (Fabricius) (p. 81) 38. Pronotum with four small yellowish spots, two on anterior disk and two on posterior marginal area; lateral margins of pronotum entire, without serrations or teeth 39 Pronotum without spots, generally disk entirely yellow con- trasting with remainder of pronotum and hemelytra; lateral margins of pronotum serrate 40 39. Outer tibial dilations small with shallow emarginations, width outer dilations equal to or slightly less than inter- ocular distance; fore and middle tibiae unicolorous with- out a pale yellow annulus harpagon (Fabricius) (p. 68) Outer tibial dilations larger with deep emarginations, width outer dilations 1.5 times the interocular distance; fore and middle tibiae with a pale yellow annulus flavosignatus Blote (p. 70) 40. Outer tibial dilation narrow and entire, without prominent teeth or emarginations, its width 1.0 to 1.5 times width of inner dilations; length of fourth antennal segment 1.08 to 1.2 times length of third (Fig. 2) crassicornis (Dallas) (p. 64) Outer tibial dilations wider with one or two emarginations, its width 1.8 to 2.0 times width of inner dilations; length of fourth antennal segment 1.2 to 1.8 times length of third 41 41. Thoracic pleura with a single yellow maculae occupying almost the entire ventral third of each pleuron; outer tibial dilation occupying approximately 65% length of hind tibiae cinctus (Herrich-Schaeffer) (p. 61) Pro- and mesopleura with two small yellow maculae, meta- pleuron with one (maculae about the size of an eye or less); outer tibial dilation occupying approximately 55% length of hind tibiae fasciatus (Westwood) (p. 66) 1969] Allen: Revision of the Genus Leptoglossus 61 Leptoglossus cinctus (Herrich-Schaeffer) A nisoscelis cincta Herrich-Schaeffer, 1836:91, fig. 315. Theognis cincta Mayr, 1866:103. Leptoglossus cinctus Stal, 1870: 164. — Distant, 1881 : 125. — Lethierry and Severin, 1894:47. — Barber and Bruner, 1947:80. — Wolcott, 1948:197. Pronotum behind humeri, clavus, basal two thirds of corium, fore and middle legs light brown; head piceous with lateral areas and three narrow dorsal stripes light reddish brown; entire declivous face of pronotum including lateral margins straw yellow with remainder of pronotum, scutellum, veins of clavus and corium, and apical third of corium dull wine red; apex scutellum, straight transverse fascia on corium, single maculae on inner tibial dilations whitish yellow; basal areas of each thoracic pleuron with a large bright yellow macula; first antennal segments piceous with inner margins of first, entire second, third and basal half of fourth light reddish brown; distal half of fourth segment ochraceous; thoracic and abdominal hind tibiae including dilations dull to bright wine red; abdominal dorsum piceous with central portion metallic blue-green, last tergum with a large semicircular straw yellow spot; connexivum, large portions of thoracic and abdominal venter centrally, lateral areas of hind femora, femoral teeth and tubercles piceous; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures smooth, pronotal punctures slightly larger and farther apart; scutellum transversely rugulose with numerous punctures; thoracic pleura with numerous small patches of dense, short, ap- pressed whitish hairs; remainder of body and appendages with sparse short erect hairs. Head non-declivent with tylus and juga raised above level of antenniferous tubercles, tylus blunt forming a rounded elevated ridge slightly exceeding juga, length head 2.64, width head 2.28, inter- ocular distance 1.20, anteocular distance 1.44; pronotum gradually declivent with lateral margins serrate and straight, humeral areas not expanded or ascending, humeral angles acute, postero-lateral margins serrate, calli barely elevated, area between calli slightly higher and indistinct from calli, disk posteriorly without low median longitudinal carina, length pronotum 3.04, width across humeri 5.76, width anterior margin pronotum 1.68; length scutellum 2.52, width scutellum 2.40; labium reaching to anterior margin of third abdominal sternum, length labial segments I 2.76, II 2.52, III 1.32, IV 2.28; length antennal segments I 2.64, II 3.84, III 2.64, IV 4.32; hind tibiae with 62 Entomologica Americana [Vol. 45 outer dilations lanceolate with one extremely shallow emargination, occupying 66% length of hind tibiae, width about twice that of inner dilations; inner dilations lanceolate, slightly shorter than outer with numerous small spine-like teeth along entire margins; inner margins of undilated portions of hind tibiae with a double row of small spine- like teeth, length hind tibiae 9.84; length outer dilations 6.48, width outer dilations 1.80; length inner dilations 5.32, width inner dilations 0.84; total length 19.1. Claspers (Fig. 6) with inner basal lobe broad, long, and rounded; shank thick without a median lobe, hook strongly curved; capsule (Fig. 31) with a simple deep and rounded median notch, no dorsal prongs; aedeagus (Fig. 64) with dorsal sac of conjunctiva as follows: long proximal lobes present; two pairs of median lobes present, one pair lower and laterally situated with tips barely sclerotized, other pair medially and apically situated and heavily sclerotized and spinose; distal lobes present as two broad lateral extensions of sac, unsclerotized. The description is of a male specimen from Chapada, Brazil. All of the specimens from Cuba, Mexico, and Central America have the transverse fascia on the corium absent or reduced to a short and faint narrow mark. Also Cuban specimens have a dark brownish yellow to light brown pronotal disk, thereby giving the appearance of an almost concolorous dorsum. The above color patterns also occur in a few South American specimens. The labium varies in length extending from the anterior margin of the second to the anterior margin of the fourth abdominal sternum. L. cinctus may be distinguished from the closely related species jasciatus and crassicornis by the following characters: outer tibial dilations broader, almost twice the width of those of crassicornis and longer than the outer dilations on jasciatus \ the yellow maculae on the thoracic pleura are larger, occupying almost the entire ventral third of each pleuron; the dorsal sac of the aedeagus has the more distally placed pair of median lobes heavily sclerotized and spinose. L. cinctus has been collected on guava fruit and Byrsonia crassi- folia H.B.K. (Barber and Bruner, 1947), and on cactus species of the genera Cereus and Opuntia (Mann, 1969). Distribution : A widely distributed species occurring in southern Mexico, Cuba, through Central America and most of South America. Material Examined : MEXICO: 1 $ , Rosamorada, Nayarit, VIII- 4-1953, D. Rockefeller Mex. Exp. 1953 (C. & P. Vaurie); 19, La- Buena Ventura, Vera Cruz, 7-1909; 19, no data; 19, 10 mi. SE 1969] Allen: Revision of the Genus Leptoglossus 63 Tapanatepec Oax., VIII-8-1963 (F. D. Parker, L. A. Stange); 1$, 24 mi. west Cintalapa, Chis., VIII-12-1963 (F. D. Parker, L. A. Stange). CUBA: 2 8, 1$, 14 K. N. of Vinales, Sept. 16-22, 1913; 1 8 , same except 7 K. N. of Vinales; 1 2, same except 24 K. N. of Vinales; 1 8 , Cayamas (E. A. Schwarz); 1 2, Havana (F. Z. Cervera). HONDURAS: 1 2, Dept. Morazan Esc. Agr. Pan., Zamorano, 2700 ft. (ocotal), July 2, 1948 (T. H. Hubbell 146); 12, same except 2600 ft. (llano), July 26, 1948; 22, Minas de Oro, Comay, 1-VI (4000 ft.) (J. B. Edwards); 1 8 , Subirana Yoro, Jan. (Stadelmann) ; 1 2, same except 18-11; 1 2, La Ceiba I. de P., Sept. 1938 (Ent. No. 11014), “taken on Peralejo”; 18, Zamorano, IX-1953, “on guava fruit” (N. L. H. Krauss). COSTA RICA: 1 2 , Pacayas (C. Werckele). PANAMA: 18, Amer. centr. Chiriqui. BRITISH GUIANA: 12, E. C. Demerara, Pin. Hope, 29-VI-1932 (F. A. Squire). FRENCH GUIANA: 18, Cayenne, 4-VIII-1954 (N. L. H. Krauss); 18, no data. VENEZUELA: 18, Merida. COLOMBIA: 18, 22, Bogota (Lindig); 12, Lake Sapatoza region, Chiriguana District, VIII-XI- 1924 (C. Allen). PERU: 1 2, Vilcanota. BOLIVIA: 1 2, Rio Ivon Beni, February, Mulford Bio Expl. 1921-22 (W. M. Mann); 13, Prov. del Sara, CM Acc 5068, April 1913 (Steinbach); 1 2, Coroico; 13, Prov. Sara (Steinbach); 12, no data. BRAZIL: 18, 12, Chapada (2600 ft.) Nov. 1902 (A. Robert), 1903-96; 18, Chapada, Acc. No. 2966; 18, 12, Chapada, Acc. No. 2966, August; 22, same except June; 5 <2, same except September; 18, 3 2, same except Jan- uary; 3 2 , same except April; 5 8, 12, same except March; 1 8, 42, same except May; 38, 5 2, same except November; 28, 42, same except December; 28, 3 2, same except October; 18, 12, Chapada Campo, Oct.; 1 2, Corumba, Acc. No. 2966, June; 1 8 , 22, S. Paulo; 1$, Amazon (Stevens); 18, 12, Brasilia bor.; 18, Cuyaba, Matto- Grosso; 18, Minas geraes Ouropreto, 26-12-1898; 12, Annapolla, Goiaz, XI-24-1936 (G. Fairchild); 12, Corumba, Matto-Grosso, 1950 (H. G. Barber); 18, Para (P. R. Uhler collection); 1$, Corumba, Feb. PARAGUAY: 13, Asuncion, Sept. 1922-April 1923 (E. G. Kent), BM 1925-262. ARGENTINA: 13, Puerto Aguirre, Alto Parana, 1 1-2 1-III-1-1934 (K. J. Hayward), BM 1934- 337; 1 2, Mendoza (H. Rolle, Berlin W.). In United States National Museum (J. C. Lutz coll.), American Museum of Natural History, Stockholm Museum, Museum of Comparative Zoology (Harvard Uni- versity), Hungarian National Museum, British Museum (Natural History), Carnegie Museum, Berlin Humboldt University Museum, University of Kansas, J. A. Slater and R. C. Allen collections. 64 Entomologica Americana [Vol. 45 Leptoglossus crassicornis (Dallas) (Fig. 2) Anisoscelis crassicornis Dallas, 1852:454. Leptoglossus crassicornis Stal, 1870:164. — Lethierry and Severin, 1894:47. Pronotum behind humeri, clavus and basal two thirds of corium fuscous; head piceous; declivous face of pronotum straw yellow with anterior margins including calli, lateral margins widely and a rather broad median longitudinal band diminishing posteriorly to opposite humeral angles bright red; head piceous; following areas bright red: head laterally, dorsally with three narrow stripes, first three antennal segments entirely, scutellum, veins of clavus and corium, and apical third of corium; transverse fascia on corium straight, abbreviated not surpassing medio-cubital crossveins laterally, apex scutellum, inner tibial dilations with a single maculae whitish yellow; ventral area of pro- and mesopleura with two and metapleura with one large maculae bright yellow; fore and middle legs, undilated portion of hind tibiae, and punctation on pronotal disk light brown to ochraceous; hind femora and tibial dilations dark reddish brown to fuscous with teeth and tubercles piceous; thoracic and abdominal venter variegated bright red and fuscous; connexivum and abdominal dorsum piceous with anterior portions of connexival segments and last abdominal tergum with a single large rounded spot whitish yellow; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures smooth, pronotal punctations slightly larger and farther apart; scutellum transversely rugulose with few punctures; body moderately and appendages more densely covered with short erect, semi-decumbent pale hairs. Head non-declivent, both tylus and juga raised above level of antenniferous tubercles, tylus blunt, forming a rounded elevated ridge slightly exceeding juga, length head 2.76, width head 2.40, inter- ocular distance 1.20, anteocular distance 1.44; pronotum gradually declivent, lateral margins straight with few extremely small teeth, almost entire, humeral areas not expanded or ascending, humeral angles acute, postero-lateral margins entire, calli barely elevated, area between calli slightly higher and indistinct from calli, disk posteriorly with the low median longitudinal carina obsolete, length pronotum 3.04, width across humeri 5.60, width anterior margin of pronotum 1.68; length scutellum 2.16, width scutellum 2.28; labium reaching to middle of fifth abdominal sternum, length labial segments I 3.60, II 3.48, III 1.80, IV 3.72; length antennal segments I 2.76, II 5.28, III 2.88, IV missing; hind tibiae with outer dilations lanceolate, 1969] Allen: Revision of the Genus Leptoglossus 65 entire, occupying 71% length of hind tibiae, very narrow, about 1.2 times width of inner dilations; inner dilations lanceolate, slightly shorter than outer, with numerous small spine-like teeth along mar- gins; inner margins of undilated portions with a double row, dorsal margins for entire length of hind tibiae with a single row of numerous small spine-like teeth, length hind tibiae 10.1; length outer dilations 7.20, width outer dilations 0.72; length inner dilations 6.00, width inner dilations 0.60. Claspers and capsule as in cinctus', aedeagus with dorsal sac of conjunctiva as follows: without proximal lobes; two pairs of median lobes present, one pair lower and laterally situated with a small lobe arising from bases of each and tips barely sclerotized, the other pair medially and apically situated, unsclerotized; distal lobes present as two broad lateral extensions of sac, unsclerotized; dorso-lateral appendage appressed to wall of conjunctiva; total length 17.6. The above description is of the male holotype. Unfortunately, the fourth antennal segments are missing, but Dallas (1852) stated that the second segment was as long as the third and fourth combined, and the fourth was as long as the third; and that the fourth segment was pale brown with the apex yellow. I have tentatively identified a large series of specimens from South America as crassicornis primarily on the basis of identical hind tibial dilations, the close similarity of the genitalia, and the shortness of the fourth antennal segments. The following are differences exhibited in the series from the holotype: the pronotal lateral margins are generally prominently serrate, although a few individuals do approach the condition found in the type; the combined length of antennal seg- ments three and four is about 1.5 times the length of the second, and length of the fourth antennal segment is 1.08 to 1.20 times that of the third; the humeral angles are not so acute; there are no traces of red on the lateral margins or centrally as a longitudinal band on the pronotum, nor are any of the red areas present as bright as are those on the holotype. Due to the lack of specimens from Bolivia north- ward to Colombia (the type locality), I do not feel justified in estab- lishing a new species for the southern population at this time. This population has in the past been confused with fasciatus. L. crassicornis may be distinguished from both cinctus and fasciatus by the narrower tibial dilations and the shorter fourth antennal segments. The aedeagal differences are: from cinctus , the absence of proximal lobes and unsclerotized medial lobes; and from fasciatus , the close appression of the dorso-lateral appendage to the wall of the conjunctiva. 66 Entomologica Americana [Vol. 45 Distribution : Bolivia, Uruguay, Paraguay, and Argentina. The only Colombian record is of the male holotype. Material Examined: Holotype: 4. COLOMBIA. In the British Museum (Natural History). BOLIVIA: 14, Cochabamba, III-25- 1950 (L. Pena); IS, N.E. Cochabamba, 11-24-1950 (L. Pena). URUGUAY: 1 $ , Pinapolis, 1-25 (Doot). PARAGUAY: 14, no data (Berg). ARGENTINA: 3 4 , 2 2 , Cordoba (W. M. Davis); 14, Chilecito, La Rioja; 1 4, Campo del Cielo, 1-1934 (J. M. Bosq); 24, Poso Moza, Formosa, 2-2-1937 (Denier); 14, 1$, Ing. Juares, For- mosa, 18-V-1939 (Denier); 44, 1$, Sgo. del Estero, Rio Salado (Wagner); 4 4, 12, Andalgala, Catamarca, 3-III-1939 (Biraben- Scott); 1 4, same except 23-III-1 939 ; 1 2, Hualfin, Catamarca 5-III- 1939 (Biraben-Scott) ; 3 4, 3 2, Mendoza; 12, same except (Berg); 14, 12, Mendoza Parque, III- 1940 (J. M. Bosq). In United States National Museum, La Plata Museum, Museum of Comparative Zoology (Harvard), J. A. Slater and R. C. Allen collections. Leptoglossus fasciatus (Westwood) Anisoscelis fasciata Westwood, 1842:17. Theognis fasciatus Mayr, 1866:102. Leptoglossus fasciatus Stal, 1870: 164. — Lethierry and Severin, 1894: 47. Pronotum, scutellum, clavus and corium fuscous; head piceous with lateral areas and above with three narrow stripes light reddish brown; entire declivous face of pronotum, antennae, fore and middle legs, and undilated portion of hind tibiae straw yellow to ochraceous; veins of clavus and corium, and apical third of corium dull wine red; basal portion of third antennal segment, apex scutellum, straight transverse fascia on corium, and a single maculae on inner tibial dilations whitish yellow; pro- and mesopleura with two and meta- pleuron with one bright yellow maculae; hind femora and tibial dila- tions fuscous with teeth and tubercles piceous; thoracic and abdominal venter variegated dark reddish brown and fuscous; connexivum and abdominal dorsum piceous with a slight tinge of metallic green around scent gland scars; anterior portion of connexival segment and a semi- circular spot on last abdominal tergum whitish yellow; membrane uni- formly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures smooth, with pronotal punctures slightly larger and farther apart; scutellum transversely rugulose with nu- merous punctures; body and appendages with moderate amounts of short erect and appressed pale hairs. Head non-declivent with both tylus and juga raised above level 1969] Allen: Revision of the Genus Leptoglossus 67 of antenniferous tubercles, tylus blunt, forming a rounded elevated ridge slightly exceeding juga, length head 2.64, width head 2.16, interocular distance 1.20, anetocular distance 1.44; pronotum grad- ually declivent, lateral margins straight and prominently serrate, humeral areas not expanded and barely ascending, humeral angles subspinose, postero-lateral margins serrate, calli barely elevated, area between calli slightly higher with two low tubercles, disk posteriorly with low median longitudinal carina absent; length pronotum 2.72, width across humeri 5.44, width anterior margin pronotum 1.68; length scutellum 1.92, width scutellum 2.04; labium reaching to pos- terior margin of fourth abdominal sternum, length labial segments I 3.36, II 3.16, III 1.68, IV 3.00; length antennal segments I 2.76, II 5.04, III 3.24, IV 4.08; hind tibiae with outer dilations lanceolate with two very shallow emarginations, occupying 55% length of hind tibiae, width about twice that of inner dilations; inner dilations lanceo- late, slightly shorter than outer, with numerous small spine-like teeth along margins; inner margins of undilated portion with a double row and dorsal margin for the entire length of hind tibiae with a single row of numerous small spine-like teeth, length hind tibiae 10.8; length outer dilations 6.00, width outer dilations 1.20; length inner dilations 4.80; width inner dilations 0.60. Claspers and capsule as in cinctus ; aedeagus with dorsal sac of con- junctiva as follows: without proximal lobes; two pairs of median lobes present, one pair lower and laterally situated with a small lobe arising from the bases of each, tips barely sclerotized, the other pair medially and apically situated and unsclerotized; distal lobes present as two broad lateral extensions of sac and unsclerotized; dorso-lateral appendage of conjunctiva more lobate and not closely appressed to wall of conjunctiva, extending posteriorly; total length 16.0. I have determined the material at hand on the basis of drawings and comparative notes of the holotype which were made possible through the kindness of Dr. I. Lansbury of the University of Oxford Museum. The description is of a male specimen from Macahe, Brazil. The longer and more acute humeral angles and shorter tibial dilations will distinguish fasciatus from both crassicornis and cinctus . From cinctus , fasciatus is further distinguished by the generally longer labium, dark reddish brown to fuscous dorsum, smaller maculae on the thoracic pleura and the absence of a heavily sclerotized spinose medial lobe on the dorsal sac of the aedeagus. Additional char- acters that further separate fasciatus from crassicornis are: the longer fourth antennal segments, 1.3 to 1.4 times the length of the 68 Entomologica Americana [Vol. 45 third and the large lobate dorso-lateral appendage extending from the conjunctiva. Bosq (1940) reported fasciatus common on Baccharis and as feeding on cactus, although these host plant records from Argentina probably pertain to the southern population of crassicornis. In the material examined, there are three males from Macahe, Brazil, that were collected on cactus fruits. Distribution : The recorded distribution includes Argentina, but I believe this to be based on misidentified specimens of crassicornis. In a large series of Leptoglossus species from the La Plata Museum, there was not a single specimen of fasciatus. So far, I have seen only five specimens, all from Brazil. Material Examined : BRAZIL: 3$, Macahe, Dec., 1928, “on cactus fruit” (E. Mortensen); 1 9, 1905-100 (Fry); 1 9, Rio de Jan., Nov, Acc. No. 2966. In the Carnegie Museum, British Museum (Natural History), and United States National Museum. Leptoglossus liarpagon (Fabricius) Cimex harp agon Fabricius, 1775:101. Cimex harpator Gmelin , 1788:2188. Lygaeus harpagon Fabricius, 1794:146. Anisoscelis sexmaculata Stal, 1859:458. Theognis erythrinus Mayr, 1865:434. — Mayr, 1866:105, fig. 24. Leptoglossus harpagon Stal, 1870:165. — Lethierry and Severin, 1894: 47. — Blote, 1936:28. Head, pronotum, scutellum, clavus and corium uniformly dark reddish brown with two wide piceous stripes on head; pronotal disk anteriorly, submarginal area of posterior pronotal lobe and medio- cubital crossvein area of corium each with two dark to whitish yellow round spots; apex scutellum whitish yellow; antennal segments, legs including tibial dilations dark reddish brown, the latter with teeth and tubercles piceous; thoracic pleura dark reddish brown with the follow- ing large shining orange yellow maculae: each pleuron with one dis- tinct but poorly defined macula above coxal cavity, propleura with two maculae on posterior marginal area, mesopleura with one dorsal spot on posterior margin; abdominal venter light to dark ochraceous; connexivum and abdominal dorsum piceous with margins and an- terior third of each connexival segment and two round spots on scent gland scars yellowish orange; membrane uniformly dark; pronotum coarsely and closely punctate, surface above punctations uneven; clavus and corium closely and more finely punctate; scutellum trans- versely rugulose with few punctures; body and appendages moderately 1969] Allen: Revision of the Genus Leptoglossus 69 covered with short appressed and erect pale hairs, head below and thoracic mid-venter densely covered with thicker grayish hairs. Head non-declivent with both tylus and juga raised slightly above level of antenniferous tubercles, tylus blunt, forming a rounded ele- vated ridge slightly exceeding juga, length head 2.04, width head 2.16, interocular distance 1.08, anteocular distance 1.08; pronotum grad- ually declivent with lateral margins slightly sinuate and entire, humeral areas not expanded or ascending, humeral angles subacute, postero- lateral margins entire, posterior pronotal disk with an obvious low median longitudinal carina, pronotal calli prominently elevated, raised area between calli with two obscure small tubercles, length pronotum 2.24, width across humeri 4.16, width anterior margin pronotum 1.56; length scutellum 1.92, width scutellum 1.68; labium extending to posterior margin of third abdominal sternum, length labial segments I 2.04, II 1.92, III 1.20, IV 2.04; length antennal segments I 1.56, II 2.88, III 2.16, IV 3.24; hind tibiae with outer dilations slightly phylliform with two shallow emarginations, occupying 67% length of hind tibiae, and about one and a half times width of inner dilations; the latter lanceolate, slightly shorter than outer dilations and armed with a few small spine-like teeth; inner margins of undilated portion of hind tibae without small spine-like teeth, length hind tibiae 5.76; length outer dilations 3.84, width outer dilations 0.84; length inner dilations 3.24, width inner dilations 0.60; total length 13.9. The above description is based on the female holotype of L. sexmaculatus (Stal). The Fabricius type of harpagon was not seen. The following description of genitalia is of a male specimen from Corupa, S. Cath. Brazil. Claspers (Fig. 7) with inner basal lobe very low; hook and shank not differentiated but forming a scooped out blade which curves inward rather sharply at tip; capsule (Fig. 32) with deep median notch and strong dorsal prongs, entire posterior edge greatly produced back- wards over curvature of capsule; aedeagus (Fig. 65) with dorsal sac of conjunctiva as follows: three large unsclerotized proximal lobes; medial portion of sac without lobes; distal end of sac with a large blunt posteriorly directed area and two smaller antero-lateral lobes, all unsclerotized. I have based the description of harpagon on Stabs sexmaculatus which he himself synonymized. Stal also synonymized Mayr’s ery- thrinus, the description of which resembles in every respect Blote’s flavosignatus. At present, I feel it is best to treat Blote’s species as valid until both the type of harpagon and erythrinus can be examined. Unfortunately, I was unable to associate any male specimens with flavosignatus so that the differentiation of these two species is based 70 Entomologica Americana [Vol. 45 only on females. I have before me a number of individuals which are related closely to these species, but possess a number of differentiating characters, especially in the shape of the tibial dilation and the male genitalia. I do not feel it desirable to describe additional species until a larger series of specimens can be definitely associated with the types, in order to evaluate correctly any variation. The two species may be separated by the following characters: harpagon has a smaller tibial dilation, the width being slightly less than, or equal to, the interocular distance, in flavosignatus the width of the outer dilation is at least 1.5 times the interocular distance; the fore and middle tibiae have a pale yellow annulus and the dorsum has light and dark areas in flavosignatus , while in harpagon both the tibiae and dorsum are unicolorous. There is also considerable dif- ference in the size and shape of the corial spots and in overall size between these two species. Distribution : Brazil. Material Examined : Holotype, $ of L. sexmaculatus, Stal. In Stockholm Museum. BRAZIL: IS, Parana, 1905-163 (E. D. Jones); IS, Corupa, S. Cath., Nov. 1948 (A. Mailer). In British Museum (Natural History) and American Museum of Natural History. Leptoglossus flavosignatus Blote Leptoglossus flavosignatus Blote, 1936:28. Pronotum, scutellum, clavus and corium dark reddish-brown and fuscous with humeri and posterior submargin of pronotum entirely fuscous; head above and below piceous; pronotal disk with two large longitudinally ovoid spots, submarginal area with two smaller trans- versely ovoid spots, corium with two large somewhat rectangular spots bright yellowish orange; head striped laterally and with three short narrow stripes above; apex scutellum, median annulus on fore and middle tibiae, undilated portion of hind tibiae, and maculae on inner tibial dilation whitish yellow to ochraceous; all antennal seg- ments, fore and middle legs, basal three fourths and ventral portion of hind femora light to dark reddish brown; tibial dilations and distal portion of hind femora fuscous with teeth and tubercles piceous; thoracic pleura dark reddish brown, with following areas having large strongly contrasting bright yellowish orange spots : each pleuron with one spot above coxal cavity, propleuron with two spots on posterior marginal area, mesopleuron with one dorsal spot; abdominal venter variegated castaneous to fuscous with anterior lateral margins of each sterna whitish yellow; connexivum and abdominal dorsum piceous with anterior half of each connexival segment and a round 1969] Allen: Revision of the Genus Leptoglossus 71 spot on each scent gland scar whitish yellow; membrane uniformly dark; pronotum coarsely and closely punctate, surface above puncta- tion uneven; clavus and corium closely and more finely punctate; scutellum transversely rugulose and densely punctate, venter densely clothed with short appressed thick grayish hairs; dorsum and ap- pendages with short, and hind tibiae with longer, erect pale hairs. Head before eyes slightly declivent with tylus and juga barely below level of antenniferous tubercles, tylus blunt, forming a rounded elevated ridge, slightly exceeding juga, length head 2.40, width head 2.52, interocular distance 1.20, anteocular distance 1.20; pronotum abruptly declivent with disk tumid laterally, forming a noticeable wide trough medially, lateral margin entire and strongly sinuate, humeral areas barely expanded with humeral angles acute and not ascending, postero-lateral margins entire, the median low longi- tudinal carina on disk prominent, calli highly raised and area between calli with two low tubercles, length pronotum 3.20, width across humeri 5.60, width anterior margin pronotum 2.04; length scutellum 2.52, width scutellum 2.40; labium extending to posterior margin of third abdominal sternum, length labial segments I 2.64, II 2.40, III 1.32, IV 2.52; length antennal segments I 2.40, II 4.08, III 3.00, IV 4.80; hind tibiae with outer dilations phylliform with two deep emarginations, occupying 68% length of hind tibiae, width outer dila- tions slightly less than twice that of inner dilations; inner dilations lanceolate slightly shorter than outer dilations with distal margins possessing a few small spine-like teeth; inner marginal area of un- dilated portion of hind tibiae without small spine-like teeth; length hind tibiae 9.00; length outer dilations 6.12, width outer dilations 2.04; length inner dilations 5.76, width inner dilations 1.2; total length 17.9. For diagnostic characters see discussion under harpagon. Distribution : The holotype is without locality data, but the one additional specimen examined is from Peru. Material Examined : Holotype: $, Cat. No. 7. In Leiden Mu- seum. 1 $ , Marcapata, Peru. In Hungarian National Museum. Leptoglossus alatus (Walker) (Fig. 5) Anisoscelis alatus Walker, 1871:129. Leptoglossus alatus Lethierry and Severin, 1894:46. Medial posterior area of pronotum, most of scutellum, clavus and corium dull black with following parts metallic purple blue: head above, all antennal segments, anterior portion of pronotum including 72 Entomologica Americana [Vol. 45 calli, lateral basal areas of scutellum, punctations on dark areas of dorsum, all legs and tibial dilations; pronotum with a large con- tinuous, sharply defined orange band occupying following areas: entire pronotal disk including caudo-lateral margins and entire humeral area, this orange area interrupted by numerous small piceous spots; entire head below and three narrow stripes above, a large macula occupying most of outer tibial dilations and corium with two rounded spots on medio-cubital cross-vein ochraceous to orange tan; entire thoracic and abdominal venter orange tan with following strongly contrasting dark markings: large spot on antero-dorsal area of pro- and metapleura and a large spot above metacoxal cavities metallic purplish black, a few scattered small piceous spots on re- mainder of pleura; abdominal venter with four rows of large piceous maculae, two rows dorso-laterally situated, two rows lateral to mid- line; connexivum and abdominal dorsum piceous with anterior half of each connexival segment, anterior and lateral portions of abdominal dorsum orange to orange tan; membrane uniformly dark; pronotum, clavus and corium regularly and closely punctate, surface above punctures largely smooth; pronotal punctures posteriorly slightly larger and deeper than elsewhere; scutellum transversely rugulose with a few small punctures; entire venter and dorsum with very sparse short erect hairs; scutellum with more dense, longer, pilose hairs; appendages with dark thicker and long hairs. Head with anteocular portion slightly declivent, tylus and juga below level of antenniferous tubercles, tylus blunt, slightly exceeding juga and barely elevated, length head 3.00, width head 2.76, inter- ocular distance 1.44, anteocular distance 1.68; pronotum steeply declivent with lateral margins conspicuously dentate for entire length, humeral areas greatly expanded into large anteriorly curving and obliquely ascending processes, humeral angles acuminate, postero- lateral margins conspicuously dentate, calli prominently elevated, area between calli only slightly higher with two small tubercles, disk pos- teriorly with median longitudinal carina obsolete; length pronotum 3.84, width across humeri 9.6, width anterior margin pronotum 2.04; length scutellum 3.12, width scutellum 3.24; labium reaching pos- terior margin of fourth abdominal sternum, length labial segments I 3.36, II 3.12, III 1.92, IV 4.32; length antennal segments I 3.12, II 4.80, III 3.60, IV missing; hind tibiae with outer dilations almost lanceolate with one very shallow emargination, occupying 57% length of hind tibiae, outer dilations a little wider than inner dilations; inner dilations lanceolate, slightly shorter than outer, furnished with a few small spine-like teeth distally; undilated portions of hind tibiae without teeth, length hind tibiae 9.36; length outer dilations 5.40, width outer 1969] Allen: Revision of the Genus Leptoglossus 73 dilations 1.20; length inner dilations 4.80, width inner dilations 0.84; total length 22.0. L. alatus is a unique species and easily identified by the shape of the humeral expansions, the orange pronotal band and the orange-tan venter that contrasts strongly with the metallic purple-blue and black of the rest of the body. The placement of alatus with or near any species or species groups is difficult without a male specimen. A few characters do seem to relate alatus to lineosus and subauratus. These are: declivous condition of the head; shape of the tibial dilations; sur- face texture; amount of pubescence on the body. Distribution : Brazil. Material Examined : Holotype: 2, “Para,” Brazil. In British Museum (Natural History). Leptoglossus lineosus (Stal) Theognis lineosus Stal, 1862:295. Leptoglossus lineosus Stal, 1870:164. — Distant, 1881:126, pi. 12, fig. 17. — Lethierry and Severin, 1894:48. Pronotum, clavus and corium chocolate brown with head, first antennal segment, pronotal calli, and scutellum piceous; dorsum with strongly contrasting whitish yellow to ochraceous markings as follows : head laterally and with three narrow stripes above, all margins of pronotum, a median longitudinal stripe from collar to a transverse humeral fascia, and a pair of posteriorly tapering rays from calli well onto disk on either side of midline, tubercles on raised area between calli, a narrow median longitudinal stripe on scutellum, narrow straight transverse fascia on corium, very narrowly along claval commissure, all veins on clavus and corium except anterior half of radius; basal three fourths of antennal segments two, three and apex of fourth, fore, middle, undilated portions of hind tibiae, and bases of femora light reddish tan; all femora and hind tibial dilations fuscous with teeth and tubercles piceous; thoracic and abdominal venter fuscous to piceous with the following areas strongly contrasting ochraceous to light tan: all marginal areas of propleuron, area above coxal cavities, posterior margins of meso- and metapleura, two well defined longi- tudinal stripes across thoracic pleura, posterior and lateral marginal areas of abdominal venter, large maculae surrounding spiracular openings and the maculae on inner tibial dilations light reddish tan; connexivum and abdominal dorsum piceous with anterior third of connexival segments ochraceous; membrane uniformly dark; pro- notum, clavus and corium finely and closely punctate with surface above punctations largely smooth; scutellum transversely rugulose and 74 Entomologica Americana [Vol. 45 impunctate; body with sparse short erect and semi-decumbent hairs: femora with more dense appressed pubescense. Head before eyes declivent, tylus and juga below level of antenni- ferous tubercles, tylus blunt, slightly exceeding juga and barely ele- vated, length head 2.28, width head 2.16, interocular distance 1.08, anteocular distance 1.20; pronotum gradually declivent with lateral margins entire and straight, humeral areas not greatly expanded with humeral angles prominent and acuminate, obliquely ascending, postero-lateral margins entire, pronotal disk with a prominent low median longitudinal carina, calli slightly elevated, area between calli with two small tubercles, length pronotum 2.88, width across humeri 5.60, width anterior margin pronotum 1.80; length scutellum 2.04, width scutellum 2.16; labium extending to posterior margin of third abdominal sternum, length labial segments I 2.52, II 2.64, III 0.96, IV 2.64; length antennal segments I 2.40, II 3.84, III 2.28, IV 3.48; hind femora greatly incrassate; hind tibiae with outer dilations lance- olate, occupying 65% the length of hind tibiae, of same width as inner dilations; inner dilations lanceolate, as long as outer dilations, distally with a few strong teeth along margins; undilated portion of hind tibiae along inner margins with a double row of spine-like teeth, length hind tibiae 7.20; length outer dilations 4.68, width outer dilations 0.96; length inner dilations 4.68, width inner dilations 0.96. Claspers (Fig. 8) with base and shank little differentiated from each other, no lobes present; shank very wide in lateral view and scooped out medially, hook strongly curved dorsally; capsule (Fig. 33) with a shallow median notch, no dorsal prongs, entire posterior edge gradually produced backwards over curvature of capsule; aedeagus with dorsal sac of conjunctiva divided into main lobes with low broad sclerotized proximal lobes, dorso-lateral appendage long blade-like and heavily sclerotized. Total length 17.3. For diagnostic characters see key and discussion under subauratus. L. lineosus and subauratus are closely related species and very distinct from the other members of the genus. They have in common, together with harpagon and flavosignatus , the following characters: claspers with a deeply scooped out shank; capsule with entire pos- terior edge projecting backwards over curvature of dorsal portion; a long sclerotized dorsal lateral appendage on conjunctiva. Distribution : Mexico. Material Examined: 3 syntypes: 1 & , 2 2 , MEXICO (Salle). In Stockholm Museum. The $ specimen (described above) bearing the label “TYPUS” is selected as lectotype, and an appropriate label has been attached to the specimen. MEXICO: 1 2 , Cuernavaca Mor; 1 2, Tamazunchale, San Luis Potosi, V-20-1952 (M. Cazier, W. 1969] Allen: Revision of the Genus Leptoglossus 75 Gertsch, R. Scharammel); IS (Salle). In American Museum of Natural History. Leptoglossus subauratus Distant Leptoglossus subauratus Distant, 1881:126, pi. 12, fig. 18. — Lethierry and Severin, 1894:49. Pronotum, scutellum, clavus and corium dark chocolate brown with head piceous; dorsum with following areas strongly contrasting whitish yellow to ochraceous: head laterally and with three narrow stripes above, all margins of pronotum, with a large crescentic macula on pronotal disc, lateral margins and a central longitudinal stripe on scutellum, lateral margins of corium, claval sutures and all veins on clavus and corium and the narrow straight transverse fascia on corium; dorsum of femora and tibiae on fore and middle legs, un- dilated portion of hind tibiae, maculae on inner tibial dilations, and all antennal segments light to dark tan; the latter with distal portion of segments two and three and base of four somewhat darker; hind femora and tibial dilations dark brown to fuscous with teeth and tubercles slightly darker; thoracic and abdominal venter dark brown with following areas strongly contrasting ochraceous to tan: all mar- ginal areas of propleuron, ventral and posterior margins of meso- and metapleura, two well defined longitudinal vittae across thoracic pleura and posterior and lateral marginal area of abdominal venter; con- nexivum and abdominal dorsum dark brown to piceous with anterior third of each connexival segment ochraceous; membrane uniformly dark; pronotum, clavus and corium closely and finely punctate with surface above punctures smooth; scutellum transversely rugulose and impunctate; venter with sparse short appressed pale hairs; dorsum with short erect and semi-decumbent pale hairs; appendages with longer more dense pilose hairs. Head before eyes declivent with tylus and juga below level of antenniferous tubercles, tylus blunt, slightly exceeding juga and barely elevated, length head 2.40, width head 2.04, interocular distance 1.20, anteocular distance 1.44; pronotum gradually declivent with lateral margins entire and straight, humeral areas not greatly expanded or ascending, humeral angles acute, postero-lateral margins serrate, disk with median longitudinal carina obsolete, calli slightly elevated, area between calli higher and with two low tubercles, length pronotum 2.56, width across humeri 5.12, width anterior margin of pronotum 1.56; length scutellum 1.92, width scutellum 1.92; labium extending to posterior margin of fifth abdominal sternum, length labial seg- ments I 2.88, II 2.88, III 1.32, IV 3.24; length antennal segments 76 Entomologica Americana [Vol. 45 I 2.40, II 4.44, III 2.88, IV 3.84; hind femora strongly incrassate; hind tibiae with outer dilations lanceolate and occupying 60% the length of hind tibiae, width of outer dilations about equal to width of inner dilations; inner dilations lanceolate with a few large teeth distally and about equal in length to outer dilations; undilated portion of hind tibiae with a double row of small spine-like teeth along inner margins, length hind tibiae 7.80; length outer dilations 0.96; length inner dilations 4.32, width inner dilations 1.08. Genitalia as in lineosus but aedeagus without proximal lobes on dorsal sac, dorso-lateral appendage with only tip barely sclerotized; total length 16.0. The description is of a male specimen from Quezaltepeque, El Salvador. L. subauratus resembles lineosus very closely except for the following differences. The pronotum has the transverse band occupying most of the anterior portion of the disk in subauratus whereas in lineosus it is a very narrow band between the humeri. L. subauratus has more unicolorous antennal segments, in lineosus there are strongly contrasting light and dark areas. The postero- lateral margins of the pronotum are serrate and the humeral angles are acute in subauratus , whereas in lineosus the pronotal postero- lateral margins are entire with the humeral angles acuminate. Aedeagal differences are: for subauratus the lack of proximal lobes on the dorsal sac and a shorter less sclerotized dorso-lateral appendage; for lineosus the proximal lobes are present and the dorso-lateral append- age is heavily sclerotized and longer. It should be mentioned that despite the differences listed above, these two species are very closely related. The material examined is insufficient to assess any variation which may be present. Addi- tional collecting may prove these two species to be conspecific. Adult specimens of subauratus have been collected on prickly pear in Guatemala and El Salvador (Mann, 1969). Distribution : El Salvador. Distant (1881) lists his type series from Guatemala and Nicaragua (deposited in the British Museum). Material Examined : EL SALVADOR: 1 9, Quezaltepeque, VI- 24-1961 (M. E. Irwin); 1$, Acajutla, 28.8 (Fred K. Knab); IS, Metapan, 5-VII-1954, No. 444-34DB (M. S. V.). In United States National Museum and University of California (Davis). Leptoglossus rubrescens (Walker) Malvana rubrescens Walker, 1871:134. — Lethierry and Severin, 1894:51. Leptoglossus rubrescens Distant, 1901b :417-8. 1969] Allen: Revision of the Genus Leptoglossus 77 Head, pronotum, scutellum, clavus, corium and thoracic pleura dark reddish brown; veins of clavus and corium wine red; fore and middle legs and apex scutellum ochraceous to tan; abdominal venter rusty reddish brown; connexivum and abdominal dorsum fuscous with a percurrent longitudinal median stripe the width of scent gland scars pale yellow; membrane uniformly dark; pronotum coarsely and closely punctate with surface above punctations scabrous, puncta- tions on pronotum deeper and larger than elsewhere; clavus and corium finely and closely punctate; scutellum transversely rugulose with numerous punctations; head and thoracic pleura thickly clothed with short appressed whitish pubescence; remainder of body and appendages with sparse short erect and semi-decumbent whitish hairs. Head porrect with tylus and juga above level of antenniferous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 2.76, width head 2.40, interocular dis- tance 1.20, anteocular distance 1.44; pronotum steeply declivent almost vertical with lateral margins strongly sinuate and dentate caudally, humeral areas broadly expanded into long tapering projec- tions with humeral angles acute, postero-lateral margins strongly dentate, calli elevated and area between calli only slightly higher, disk posteriorly with the low median longitudinal carina obsolete, length pronotum 3.04, width across humeri 8.00, width anterior margin pronotum 1.68; length scutellum 2.28, width scutellum 2.28; labium reaching to posterior margin of fourth abdominal sternum, length labial segments I 3.12, II 3.12, III 1.92, IV 4.08; length antennal segments I 2.52, II 4.68, III 3.48, IV 5.64; hind legs missing. Clasper (Fig. 11) with inner basal lobe low and rounded, shank thick without a median lobe, hook completely curved around and directed laterally; capsule (Fig. 36) with median notch deep and rounded, no dorsal prongs; aedeagus (Fig. 66) with dorsal sac of conjunctiva rather short and without sclerotized areas, lobes on dorsal sac as follows: no proximal lobes present; left and right median lobes large and of equal size and shape; apex of sac somewhat trifid; total length 17.9. L. rubrescens is a rather unusual species but it does have some characteristics in common with julvicornis. The most striking simi- larities are the presence of a pale yellow longitudinal stripe on the abdominal tergum; the coarseness and shape of the pronotal punctures, the dark reddish ground color, and the lack of pale markings on the pronotum and hemelytra. The bases and shanks of the claspers are similar, but the hook in rubrescens is exceedingly different in that it bends completely around ventrally and the tip points out laterally. No other Leptoglossus species has this kind of hook. The dorsal 78 Entomologica Americana [Vol. 45 margins of the capsules bear no resemblance to each other, with fulvicornis having the unique condition. The aedeagal similarities and differences are: both species have the dorsal sac unsclerotized without proximal lobes; medial lobes are present but are smaller and more distally placed in rubrescens. The long tapering humeral areas, the steeply declivent pronotum, and the rounded and simple median notch on the genital capsule will further distinguish rubrescens from julvicornis. Distribution : Known only from the type locality. Material Examined: Holotype: $. BRAZIL. In British Museum (Natural History). Leptoglossus fulvicornis (Westwood) Anisoscelis fulvicornis Westwood, 1842:17. Leptoglossus fulvicornis Stal, 1870:161. — Distant, 1901a:330, pi. 30, fig. 4. — Van Duzee, 1917:89. — Gibson, 1917:70. — Parshley, 1923:747.— Blatchley, 1926:221.— Torre-Bueno, 1941:49. Leptoglossus magnoliae Heidemann, 1910: 191-5, pi. 7. Theognis fulvicornis Hussey, 1953:30. Dorsum and venter light to dark reddish brown without pale mark- ings on pronotum or hemelytra; thoracic and abdominal venter mottled with numerous small piceous spots; abdominal tergum with a yellowish median longitudinal fascia; pronotum coarsely punctate almost rugu- lose, with lateral and postero-lateral margins serrate to dentate; humeri broadly and obtusely rounded; outer tibial dilations lanceolate with two or three very shallow emargination. Clasper (Fig. 10) with a long broad base and inner basal lobe obsolete; shank without a median lobe; hook short and strongly curved; capsule (Fig. 35) with median notch deep, bottom of notch raised and acute; dorsal prongs broadly rounded and directed medially; aedeagus with dorsal sac of conjunctiva without sclerotized areas, and with only one pair of medial lobes. In the discussion under rubrescens , the similarities and differences between rubrescens and fulvicornis are pointed out. The relationship of these two species to dilaticollis is discussed under the latter species. The aedeagus in fulvicornis is similar in some respects to those of australis and gonagra. All three species have a long dorsal sac which is totally unsclerotized, and only the median lobes are present. In fulvicornis these median lobes are much longer than and not as close to the apex as are those of australis and gonagra. From all other North American species, fulvicornis may be easily distinguished by having lanceolate tibial dilations, a unicolorous dorsum without a pale 1969] Allen: Revision of the Genus Leptoglossus 79 transverse fascia on the corium or pale pronotal markings, dentate pronotal lateral margins, broadly rounded humeral angles and the rugulose-like surface of the pronotum. Heidemann’s description of magnoliae is based on a male and female, type No. 13228, in the United States National Museum. Both specimens bear the same type number, and I have selected as Lecto- type the male specimen; a label to that effect has been attached to the specimen. In the original description, Heidemann also gives informa- tion on the feeding habits, and includes descriptions of the eggs and nymphs. The host plant apparently is Magnolia. Distribution : Recorded from Massachusetts and New York and south to Florida and Alabama. A single specimen labeled “Mexico on bananas, Brownsville, Texas” is in the United States National Mu- seum. Material Examined : 56 specimens: Florida : Winter Park; Port Sewall. Georgia : Savannah. Massachusetts : Humarock. New Jersey : Anglesea; Lahaway. New York: Rockaway Beach; Long Beach; New York City. North Carolina: New Bern; Fort Macon; Southern Pines. Pennsylvania : Philadelphia. Virginia : Va. Beach; Shenandoah Na- tional Park; Falls Church. Washington, D.C. In United States Na- tional Museum, American Museum of Natural History, Museum Com- parative Zoology (Harvard), Stockholm Museum, J. A. Slater and R. C. Allen collections. Leptoglossus dilaticollis Guerin (Fig. 1) Leptoglossus dilaticollis Guerin, 1831 :pl. 12, fig. 9. — Guerin, 1838: 174. — Stal, 1870:161.— Distant, 1902:382.— Kirkaldy, 1906: 257. — Kiritschenko, 1935:191. — Costa Lima, 1940:85, fig. 292. — Dupuis, 1952:449. — Hussey, 1953:33. Pronotum, scutellum, clavus and corium dark reddish brown with head and teeth on margins of pronotum piceous; wide straight trans- verse fascia on corium and fourth antennal segments whitish yellow; head laterally and with two short stripes above, first three antennal segments and undilated portion of hind tibiae tan to light brown; legs light reddish brown with lateral portion of hind femora fuscous dis- tally; tibial dilations and first three antennal segments with a narrow stripe above fuscous; thoracic and abdominal venter reddish brown with numerous small piceous spots frequently fusing to form larger maculae; connexivum, abdominal dorsum, femoral and tibial teeth and tubercles piceous with a narrow median longitudinal stripe on abdominal dorsum ochraceous; membrane uniformly dark; pronotum 80 Entomologica Americana [Vol. 45 with numerous conspicuous large rounded foveolae and smaller punc- tations; clavus and corium finely and closely punctate; scutellum trans- versely rugulose with few punctures; venter with a moderate amount of pale short erect and appressed pubescence; dorsum with short ap- pressed and semi-decumbent yellowish hairs and sparse longer pilose hairs. Head porrect with tylus and juga above level of antenniferous tubercles, tylus blunt, slightly exceeding juga, forming a rounded ele- vated ridge, length head 4.56, width head 3.12, interocular distance 1.68, anteocular distance 2.76; pronotum steeply declivent, lateral and postero-lateral margins strongly dentate, some teeth almost spinose, humeral areas broadly expanded into long and wide anteriorly curv- ing slightly ascending processes, calli barely elevated, area between calli almost indistinct from calli and only slightly higher, disk pos- teriorly with median longitudinal carina obsolete, length pronotum 5.26, width across humeri 17.6, width anterior margin pronotum 2.52; length scutellum 3.36, width scutellum 3.96; labium extending to middle of seventh abdominal sternum; length labial segments I 5.40, II 5.88, III 4.44, IV 9.36; length antennal segments I 3.00, II 5.64, III 4.56, IV 6.60; hind tibiae with outer dilations lanceolate, occupying 64% length of hind tibiae, width about one and a half times width of inner dilations; inner dilations lanceolate, slightly shorter than outer, furnished with numerous small spine-like teeth along margins; inner margins of undilated portions with a double row, and dorsal margins of hind tibiae for entire length with a single row, of numerous small spine-like teeth, length hind tibiae 12.5; length outer dilations 8.32, width outer dilations 2.04; length inner dilations 7.20, width inner dilations 1.32. Claspers (Fig. 9) with inner basal lobe obsolete and width base gradually tapering to form shank, shank without median lobe, hook strongly curved; capsule (Fig. 34) with median notch shallow, dorsal prongs low, forming a sinuate posterior margin; aedeagus with dorsal sac of conjunctiva as follows: proximal lobes present; two median lobes present on left side only, one short and small below a larger and wider blunt lobe; no distal lobes present; all areas on dorsal sac un- sclerotized; total length 29.0. The above description is of a male specimen from Brasilia, Rio Grande, Brazil. There is very little variation in the small series examined. Three females and one male lack the small teeth on the dorsal edge of the hind tibiae. Labial extensions vary from the middle of the sixth to just past the posterior edge of the seventh abdominal sternum. L. fulvicornis , rubrescens, and dilaticollis resemble each other to 1969] Allen: Revision of the Genus Leptoglossus 81 some extent in having a narrow yellow median longitudinal stripe on the abdominal dorsum, claspers with a broad base and the inner basal lobe obsolete, and the shank without a median lobe. The dorsal sac of the conjunctiva is unsclerotized. Both fulvicornis and dilaticollis have lanceolate tibial dilations. The resemblance of these three species to one another is more superficial than an indication of a close relationship. L. rubrescens and dilaticollis are quite specialized members of the genus as evi- denced by the shape of the genital structures. This is especially true of dilaticollis with its unique humeral expansions and long labium. Costa Lima (1940) reported dilaticollis on Magnolia sp. Distribution : Brazil, Central America, and Mexico. Material Examined : MEXICO: 13, Rinton, Antonio, Oaxaca, Feb. 1908 (F. Hamilton). PANAMA: IS, no data. BRAZIL: IS, Corupa, S. Cath. (Hansa Humboldt), Apr. 1948 (A. Mailer); IS, same except III- 1949; 1 S , Rio Grande; 1 2, Rio Grande do Sul; 1 9, same except (Staudgr); 1$, no locality (Boucard). In Hungarian National Museum, Stockholm Museum, and American Museum of Natural History. Leptoglossus australis (Fabricius) Cimex australis Fabricius, 1775:708. Cimex membranaceus Fabricius, 1781:351. Cimex momordicae Forster, 1844:16. Cimex orientalis Dallas, 1852:454. Anisoscelis bidentatus Montrouzier, 1855:101. Anisoscelis flavopunctatus Signoret, 1863a:27, pi. 21, fig. 4. Leptoglossus membranaceus bidentatus, Schouteden, 1907:113. Leptoglossus membranaceus sumbawensis Blote 1936:29. This is the only species of Leptoglossus that occurs outside of the Western Hemisphere. It is readily recognized by its black or dark brownish color with the following reddish orange or yellowish mark- ings : a narrow arcuate transverse fascia on the pronotal disk, the medial portion of the second, third and apical half of the fourth antennal seg- ments, numerous large maculae covering the entire ventral surface, and a small spot on the mediocubital crossveins. The humeral angles vary from long and spinose to obtuse. The outer tibial dilation may be narrow and lanceolate or wide and phylliform. The claspers (Fig. 12) lack an inner basal lobe and the inner margin gradually tapers into the shank portion; the shank lacks a median lobe and the hook is strongly curved; the capsule (Fig. 37) has a deeply rounded median notch and no dorsal prongs; in the aedeagus the dorsal sac of the 82 Entomologica Americana [Vol. 45 conjunctiva is long and has only small median lobes near the apex; all areas of the dorsal sac are unsclerotized. L. australis has, until recently, been treated as one of three closely related species. The group was composed of membranaceus, biden- tatus and australis. Some authors had relegated bidentatus to sub- specific rank under membranaceus. It is true that individuals referred to as membranaceus from the Ethiopian and Oriental regions are quite distinct from specimens of “ australis ” in the South Pacific Islands. However, examination of the large number of specimens assembled in the present study confirms Gross’s (1963) opinion that only one species is involved. Individuals from Tahiti, Fiji, New Hebrides, American Samoa, and the Society Islands are small, have subacute humeral angles, and the outer tibial dilation is narrow with one or two shallow emarginations (Fig. 53). Westward to New Guinea and northern Australia specimens tend to be larger and more deeply emarginate. Also present in Australia are specimens that resemble individuals from Tahiti and Fiji. In New Guinea an oc- casional individual will have a spinose humeral angle. All 32 speci- mens examined from the Philippines have prominently spinose humeral angles, but here the tibial dilation is more variable, being as wide as African specimens and as narrow as Australian specimens. Through- out the Oriental and Ethiopian regions, specimens consistently have a spinose humeral angle, with the overall size and the width of the outer tibial dilation (Fig. 52) averaging slightly larger than indi- viduals from Australia or New Guinea. These characters seem to be clinal in nature, and a detailed study needs to be undertaken in order to establish the exact relationships. Fortunately, australis is a common insect, and large numbers of specimens are available for study. The relationship between australis and gonagra is extremely close. In fact, this relationship is not paralleled by any other pair of species in this genus, if one takes into consideration both similarity and distri- bution. L. gonagra is quite distinct from all other Leptoglossus species in the Western Hemisphere, which unmistakably makes it stand out as the ancestral stock from which australis was derived. These two spe- cies share in common the same distinctive color pattern, the same gen- eral habitus, and genitalia which are, as far as I can tell, identical. Also the vagility of australis and gonagra are of the same magnitude, with gonagra the most widely distributed species in the New World and australis having a tremendous geographic range in the Eastern Hemisphere. It should be mentioned that the present distribution of australis may have been in part helped by man, since australis is a known pest on various agricultural crops. It cannot be over-empha- 1969] Allen: Revision of the Genus Leptoglossus 83 sized that these two species are very closely allied and the characters that distinguish each from the other are not strikingly different. In fact, the validity of their being considered two species may be ques- tioned. On the other hand, selective pressures that would favor morphological distinctness may not have been in operation, since the geographic separation once established, would provide an effective reproductive barrier. Breeding and cytological information will in all likelihood provide answers to the questions concerning the exact nature of this relationship. The dispersal route into the Eastern Hemisphere may be en- visioned as occurring via Africa, for some African specimens of australis resemble more closely gonagra in color and humeral spine shape than do individuals from other parts of the range. The alterna- tive to this is a route through the oceanic islands of the Pacific, even though the australis specimens in the Pacific Islands are morpho- logically more distinct from gonagra than African specimens. Gross (1963) pointed out that the Pacific Island Heteroptera tend to be smaller, and that spines and dilations becomes somewhat smoothed as compared to individuals of the same species on adjacent continental or large land areas. With this in mind, the latter route may be thought of as a rather rapid spread through the islands to the Asian continent and then a subsequent loss of the prominent humeral spines and large tibial dilations on the smaller islands. From gonagra , australis can be distinguished by the shorter tibial dilation (65-75% the length of the hind tibiae), the absence of yellow spots on the scent gland scars, and the generally darker brown color (some specimens of australis are the same chocolate brown as is gonagra). The genitalia of these two species are essentially the same. The life history, economic importance and control of australis has been reported by Szent-Ivany and Catley (1960), Fernando (1957) and Pagden (1928). The latter two authors use the name L. mem- branaceus. A few of the host plant records are cotton, beans, citrus, banana, guava, wild passionfruit and nearly all cucurbitaceous plants. For a complete list of references and generic name changes consult Gross (1963). Schaefer (1965, 1968) describes some morphological aspects of australis in his studies on the higher classification of the Coreoidea. Distribution : Most of Africa, Madagascar, Seychelles Is., Indian subcontinent, Taiwan, South East Asia, Malay Archipelago, Indonesia, Philippines, Micronesia, New Guinea, Northern Australia, New Hebrides, New Caledonia, Solomon Islands, Fiji Islands, Samoa, Cook Island, Tahiti. 84 Entomologica Americana [Vol. 45 Material Examined : Approximately 500 specimens from the above localities. Leptoglossus gonagra (Fabricius) Cimex gonagra Fabricius, 1775:708. Cimex grallator Herbst, 1784:259. Lygaeus gonagra Fabricius, 1794:140. Anisoscelis antica Flerrich-Schaeffer, 1836:92, fig. 316. Anisoscelis gonagra Burmeister, 1835:332. Theognis gonager Mayr, 1866:103. Leptoglossus gonagra Stal, 1868:51; 1870:165. — Berg, 1879:72. — Distant, 1881:126. — Lethierry and Severin, 1894:47. — Van Du- zee, 1917:89-90.— Gibson, 1917:72.— Blatchley, 1926:224.— Barber, 1939:310-11. — Torro-Bueno, 1941:40. — Froeschner, 1942:594, 599. — Barber and Bruner, 1947:80. — Hussey, 1953: 31. Anisoscelis praecipua Walker, 1871:128. — Distant, 1901:431. L. gonagra is easily distinguishable from all other Leptoglossus species in the Western Hemisphere by the presence of the following yellow areas that strongly contrast with the dark brown color: a narrow arcuate transverse fascia on the pronotal disk, numerous large maculae on the thoracic pleura; seven yellowish longitudinal stripes on the abdominal venter which occasionally are interrupted at the intersegmental sutures, if so, the stripes are made up of a series of rectangular maculae. Also, the humeral angles are strongly spinose, although this is more variable than the color pattern. The outer tibial dilation (Fig. 54) is relatively narrow and gradually tapers to almost the end of the tibiae and occupies about 85 to 90% the length of the hind tibiae. L. gonagra can be separated from australis on the basis of the longer outer tibial dilations, the usually lighter color, and the presence of yellowish round spots on the scent gland scars. L. gonagra has been reported as a pest on cucurbitaceous plants, Passiflora, citrus, and tobacco in Argentina and Brazil (Bosq 1940, Costa Lima 1940); oranges, grapefruit, squashes, pumpkins, corn and guava in Puerto Rico (Wolcott 1948); as breeding on Luff a cylindrica M. Roem. and adults feeding on oranges, guavas, and Taonabo parviflora Krug. & Urb. in Cuba (Barber and Bruner 1947). Hussey (1952) reported gonagra on Brazilian pepper trees ( Schinus terebinthe folia Raddi) in Florida. It has also been collected on water- melons in Louisiana. Distribution : From Florida to Texas and south through Mexico 1969] Allen: Revision of the Genus Leptoglossus 85 and Central America, into the West Indies, and most of South America. Material Examined : 515, 43 2, UNITED STATES: Louisiana ; Florida. MEXICO: Veracruz; Nochixtlan; Quint. Roo. Xcan Nuevo; La Palma; Vera Cruz; Guerrero, Acapulco; Tezonapa; Chichen Itza; Temax. EL SALVADOR: Quezaltepeque. BRITISH HONDURAS: Benque Viejo; Jamastran. WEST INDIES: St. Croix; St. Vincent; Nevis; Puerto Rico; Cuba; Dominican Republic, Sanchez; Pto. Plata; Haiti; Grande Riviere; Cape Haytien. VENEZUELA: Merida; Caripito. BOLIVIA: Province Sara; Coroico. BRAZIL: Minas Geraes, S. da Caparao; Tres Lagoas, Matto Grosso; Rio Grande so Sul; Rio de Janeiro; Madeira Mamore; Corupa, S. Cath. (Hansa Humboldt); Bahia; Herculanoum; Meilen v. Cuyaba. PARAGUAY: Asuncion. ARGENTINA: Eldorado Misiones. In Hungarian Na- tional Museum, American Museum of Natural History, Stockholm Museum, Museum of Comparative Zoology (Harvard), Berlin Hum- boldt University Museum, British Museum (Natural History), Univer- sity of California (Davis), University of Kansas, Chicago Natural His- tory Museum, Iowa State University, J. A. Slater and R. C. Allen collections. Leptoglossus eliilensis chilensis (Spinola) Anisoscelis chilensis Spinola, 1852:172. — Signoret, 1863b:555. — Reed, 1900:46. Theognis chilensis Mayr, 1866:108. Leptoglossus chilensis Stal, 1870:164. — Lethierry and Severin, 1893: 46. — Pennington, 1922:132. Pronotum behind humeri, scutellum, clavus and corium a varie- gated light brown and dark reddish brown with numerous small piceous spots; head above and below, pronotal calli and outer marginal areas of first antennal segments piceous; entire declivous face of pronotum, apex scutellum and corium wine red; head laterally and with three narrow stripes above, inner margins of first and last three antennal segments entirely, fore, middle and undilated area of hind tibiae, and basal and ventral portion of all femora light reddish brown; distal portion of femora dorsad, and tibial dilations fuscous to piceous with teeth and tubercles piceous; thoracic and abdominal venter dark reddish brown with numerous small spots frequently fusing to form larger maculae; connexivum and abdominal dorsum piceous with anterior fourth of connexival segments ochraceous; mem- brane uniformly dark; pronotum, clavus and corium coarsely and closely punctate with surface above punctures uneven; scutellum 86 Entomologica Americana [Vol. 45 transversely rugulose with few punctures; body rather thickly covered with appressed whitish silvery pubescence and sparse short erect hairs; head below, thoracic and abdominal midventer, and append- ages with longer dense erect pilosity. Head non-declivent with both tylus and juga raised above level of antenniferous tubercles, tylus blunt, forming a rounded elevated ridge, slightly exceeding juga, length head 2.04, width head 1.92, interocular distance 1.08, anteocular distance 1.08; pronotum abruptly declivent, lateral margins sinuate, serrate caudally, humeral areas broadly and obliquely ascending, humeral angles subacute, postero-lateral margins serrate, disk posteriorly with median longitudinal carina obsolete, calli slightly elevated, raised area between calli with two small tuber- cles, length pronotum 2.24, width across humeri 4.32, width anterior margin of pronotum 1.56; length scutellum 1.56, width scutellum 1.68; labium extending to middle of metasternum, length labial seg- ments I 1.68, II 1.56, III 1.20, IV 1.50; length antennal segments I 2.04, II 3.12, III 2.04, IV 2.64; hind tibiae with outer dilations barely phylliform with two shallow emarginations, occupying 64% length of hind tibiae and about twice width of inner dilations; inner dilations lanceolate, shorter than outer dilations, furnished with a few small spine-like teeth on distal margins; inner margins of undilated portions of hind tibiae with a few small spine-like teeth arranged in a double row, length hind tibiae 6.00; length outer dilations 3.84, width outer dilation 1.20; length inner dilations 3.24, width inner dilations 0.60. Claspers (Fig. 13) with inner basal lobe small and rounded, shank thin with well produced median lobe, hook short and strongly curved; capsule (Fig. 38) with a rounded median notch, no dorsal prongs, lateral margins slightly flared; aedeagus with dorsal sac of con- junctiva as follows: proximal lobes present and sclerotized; median lobes present, left lobe long and wide, right lobe shorter and more narrow; sclerotized anterior distal lobe present; lateral distal lobes of same size and shape, both sclerotized; total length 13.0. The above description is of a male specimen from El Alfalfal, Province Santiago, Chile. In a large series of specimens the following variations were noted. The last antennal segment varies from entirely light reddish brown to fuscous. The entire declivous face of the pro- notum varies from bright yellow to dark tan and is with or without small piceous maculae. A few individuals have so many large and small maculae that the yellow areas are barely evident. These yellow areas may be separated medially forming two distinct spots. The labium varies in length from the middle of the metasternum to the anterior margin of the second abdominal sternum, only rarely does it extend to the anterior margin of the third abdominal sternum. Most 1969] Allen: Revision of the Genus Leptoglossus 87 specimens have very shallow emarginations on the outer tibial dila- tions, but some females have a more phylliform dilation with deep emarginations. Typical chilensis specimens seem to be absent from the north and northeastern corner of Argentina, and it is here that one finds con- caviusculus. I have also examined from this area a number of speci- mens that have characters of both chilensis and concaviusculus. I believe that the two “taxa” are conspecific, with concaviusculus a distinct subspecific unit. L. argentinus described by Bergroth (1894) from La Plata fits the description of chilensis. This was also the opinion of Pennington (1922), although he did not list chilensis from that area. I have not seen Bergroth’s type, but there seems little doubt that he had a speci- men of chilensis. L. chilensis is closely related to impictus ; for the diagnostic char- acters see the discussion under the latter species. Thus far chilensis has been collected on peaches, nectarines, plums, grapes, figs, and grapefruit. Material Examined : CHILE: 52 S , 48$, Province Santiago: El alfalfal; Los Maitenes; El Canelo; Lampa. Province Talca: Talca. Province O’Higgins: Rancagua. Province Nuble: San Carlos. Province Malleco: Angol. Province Cautin: Temuco; Araucancia. Province Bio-Bio: Antuco. Province Coquimbo: Illapel; Huanta. Province Aconcagua: Papudo; Rio Blanco. Province Valparaiso: Valparaiso. Province Concepcion: Concepcion. ARGENTINA: 6$, 3$, Mendoza; 2$, Cabana Cordoba, III- 1945 (M. Biraben); 2$, same except 8-1 1-1937; 2S ,29 , Rio Negro, 1-1943, “s/papa” (J. M. Bosq) ; IS , 1 $ , Province de San Juan, 1-1940 (J. M. Bosq) ; 1 S , San Juan (J. M. Bosq) ; 1 S , Correntoso, L. Nah Huapi, 20-X-1926 (R. C. Shannon); 1$, Tolombon, Salta, 7-III- 1939 (Biraben-Scott) ; 1$, 1200 m. Province Salta 2.05 (J. Steinbach S.V.) ; 1 $ , Catamarca, Los Angeles, X-1945 (J. M. Bosq); 1$, Catamarca, El Rodeo, 1-1942 (Schaefer); IS, Cabana, Cordoba, 10-11-1942 (M. Biraben); 1$, Isla Los Cisnes, Parana Delta, V-XI-1920 (H. E. Box); 1 S, El Bolson, Rio Negro, III-l 5-1 964 (A. Kovacs); 1$, Bariloche, Rio Negro, Nov. 1926 (R. & H. Shannon); 2S , Metan Salta, 1906-1 (Vezenyi). In Hungarian National Museum, La Plata Museum, Mu- seum of Comparative Zoology (Harvard), Deutsches Entomologisches Institut, Berlin Humboldt University Museum, Stockholm Museum, United States National Museum, California Academy of Sciences, British Museum (Natural History), American Museum Natural His- tory, J. A. Slater and R. C. Allen collections. 88 Entomologica Americana [Vol. 45 Leptoglossus chilensis concaviusculus Berg new status Leptoglossus concaviusculus Berg, 1892:70. — Lethierry and Severin, 1894:47. — Pennington, 1922:134. Pronotum behind humeri, scutellum, clavus and corium uniformly dark reddish brown with head above and mid-ventrally, outer marginal areas of first antennal segments and pronotal calli piceous; entire declivous face of pronotum, and last antennal segment yellowish brown; apex scutellum and large maculae on inner tibial dilations whitish yellow; head laterally and with three narrow stripes above, inner margins of first and entire second and third antennal segments, fore and middle legs, basal third and ventral portions of hind femora, and undilated portion of hind tibiae light reddish brown ;dorsal three fourths of hind femora distally, and tibial dilations fuscous with teeth and tubercles piceous; thoracic and abdominal venter dark reddish brown with numerous small piceous spots confined to abdominal mid- venter; connexivum and abdominal dorsum piceous with connexival incisures ochraceous; membrane uniformly dark; pronotum, clavus and corium coarsely and closely punctate with surface above punctures uneven; scutellum transversely rugulose with few punctures; body rather thickly clothed with appressed golden pubescence and sparse short erect hairs; appendages, thoracic and abdominal mid-venter also with dense longer erect pilosity. Head non-declivent with both tylus and juga raised above level of antenniferous tubercles, tylus blunt, forming a rounded elevated ridge, slightly exceeding juga, length head 1.92, width head 1.80, interocular distance 0.96, anteocular distance 1.08; pronotum abruptly declivent, lateral margins sinuate and serrate caudally, humeral areas broadly and obliquely ascending, humeral angles sub-acute, postero-lateral mar- gins serrate, disk posteriorly without a median longitudinal carina, calli slightly elevated, raised area between calli non-tuberculate, length pronotum 2.40, width across humeri 4.32, width anterior margin pronotum 1.32; length scutellum 1.68, width scutellum 1.80; labium extending to middle of metasternum, length labial segments I 1.56, II 1.44, III 0.84, IV 1.20; length antennal segments I 1.80, II 2.88, III 1.68, IV 2.52; hind tibiae with outer dilations phylliform with two deep emarginations, occupying 77% length of hind tibiae, and about two and a half times width of inner dilations; inner dilations lanceolate, shorter than outer dilations, furnished with a few small spine-like teeth on distal margins; inner margins of undilated portion of hind tibiae with numerous small spine-like teeth arranged in a double row, length hind tibiae 6.24; length outer dilations 4.80, width 1969] Allen: Revision of the Genus Leptoglossus 89 outer dilations 1.56; length inner dilations 3.72, width inner dilations 0.60; genitalia as in chilensis \ total length 12.8. This subspecies exhibits essentially the same variation in color patterns as found in chilensis. The declivous area of the pronotum may be entirely bright yellow or yellowish brown, or the entire pro- notum may be unicolorous. The pronotal disk sometimes has two large medially separated diffuse spots. The dorsum varies from light to dark reddish brown, but is essentially unicolorous not variegated. L. c. concaviusculus , originally described from Brazil, forms a population distinct from chilensis , which is more southern. Along the areas of contact between these two populations there occur a number of specimens intermediate between the two subspecies. The material examined shows the overlapping distribution to be in the northern and northeastern portion of Argentina. In concaviusculus , distinguishing characters are the lack of small piceous maculae on the dorsum and thoracic pleura, and the dorsum being lighter and uni- colorous brown. In chilensis these same areas have numerous piceous maculae and the dorsum is usually a variegated dark brown and fuscous. Concaviusculus has a somewhat longer fourth antennal seg- ment averaging 1.48 times the width of the head. In chilensis the average is 1.2. The width of the outer tibial dilation to width of head ratio is larger for concaviusculus , average 1.1 (0.8-1. 6); in chilensis the same ratio averages 0.6 (0.5-0. 9). Concaviusculus has a longer outer tibial dilation (Fig. 63) occupying 76% (71-80%) the length of the hind tibiae, whereas in chilensis the dilations (Fig. 62) occupy 65% (62-71%). Distribution : Northern and northeastern portion of Argentina; Paraguay; Uruguay; southern Brazil. Material Examined : Syntype Series: IS, 2$, Brazil. In the La Plata Museum. Berg’s type series is mixed, a 2 from Chaco which bears the label Leptoglossus concaviusculus is actually a specimen of impictus. Of the remaining two specimens, I have selected the S as Lectotype, and have placed an appropriate label on the specimen. BRAZIL: 1$, Corupa, S. Cath. (Hansa Humbolt), Apr. 1948 (A. Mailer); 1 S, same except March 1948; 1 S, 1 2, Rio Natal, S. Cath., XII- 1945 (A. Mailer); 12, Cauna, S. Cath., XII-1945 (A. Mailer); 12, Pinhal, S. Cath., Dec. 1948 (A. Mailer); IS, R.G. do Sul, Pio Busk; 12, Rio Grande do Sul; 12, Sao Paulo (A. Heine); 12, Sao Paul; 2 2, Sao Leopoldo; 2S, 12, Brazilien. PARAGUAY: 3 S, Asuncion; IS, no data (Berg). URUGUAY: 3 S, Montevideo (Colon), 12-XII-1929 (Tremoleras) ; 2 S, Montevideo, W. del Cerro, 31-XII-1932 (Tremoleras); 1 2, Cerro Largo, Canada de los Burros (Tremoleras); IS, no data (Berg); 12, no data (Boucard). AR- 90 Entomologica Americana [Vol. 45 GENTINA: 4S, 5 2, Misiones Pindapoy, X-1935 (J. M. Bosq); 4 2, same except III-1936; 1 S ,2 2, Entre Rios, Concordia, 11-1930 (J. M. Bosq); 1 2, same except IV-1931; 1 2, Corrientes, “on citrus,” 10- 1935 (J. M. Bosq); 12, Corrientes; 1 S, Corrientes, 1-1921 (De- Carlo); IS, Corrientes, San Roque, 11-1920 (J. M. Bosq); 2 S, 12, Cordoba (W. M. Davis); IS, La Granja, A. Gracia, Cordoba, 4-1- 1939 (C. Bruch); 12, Gl. Gracia; 2 2, Forosa (Ciud.), 28-IV-1936 (Denier); 12, Formosa, Dto. Pilaga, IV-1938 (Denier); 1 S, 22, Prov. Salta, 2500 m. (J. Steinbach, S.V.); 12, Santa Fe (J. Stein- bach, S.); 3 S, 12, Isla de Oro, Formosa, 30-III-1938 (Denier); IS, 1 2 , R. Arg. La Plata (C. Bruch) ; 1 S , 1 2 , San Ignacio, Misiones, 8-XII-1914 (Biraben, Bezzi); 2 2, Puerto Tirol, Chaco, 10-IV- 1936 (Denier); 22, Province de Buenos Aires (J. M. Bosq); 1$, Delta Rio Chana, 11-1922 (J. M. Bosq) ; 1 S , Delta Abra Vieja, 15-11- 1926 (J. M. Bosq); 12, Isla Los Cisnes, Parana Delta, V-XI-1920 (H. E. Box); 1 S, Resistencia, 11-1936 (Denier); 12, Cabana Cordoba, 4al., 8-II-1947 (Biraben); IS, Santiago del Estero, Rio Salado (Wagner); IS, Fontana, Chaco, XII-1935 (Denier); IS, Chaco (Berg) ; 1 S , Chaco, 22-X-1899 (S. Venturi) ; 1 2 , Iguazu; 1 2 , Province de Cordoba Coconcho, 11-1919; 12, Salta Capital, 18-IV- 1946 (R. Maldonado Bruzzone); IS , Corrientes, Corrientes, 15-30- 11- 1959 (Biraben); 1 2 , Manatiales, Corrientes, 1-10-III-1959 (Bira- ben) ; 1 S , La Aurora, Sgo. del Estero, II-VI-1945 (R. M. Bruzzone) ; 2S, 12, Jose C. Paz, Bs.As., XI-1944 (A. R. Bezzi); 12, Toma de Agua, E. Rios, 27-IV-1951 (B. A. Torres); IS, 12, Ibicuycito, Bs.As., X-1940 (S. R. Castillo); 3 2, Jose C. Paz, Bs.As., XII-1948 (A. R. Bezzi); 12, City Bell, Bs.As., III-1945 (A. Alba); IS, San Jose Misiones, 1-1953 (R. Euembuena); 1 S , Clorinda, Formosa, IV- 1940 (Denier); IS, Ituzaingo, Corrientes, 11-1947 (Biraben); 12, Pindapoy Misiones, 29-VI-1937 (Denier); 1 S, 12, Fontana, Chaco, 20-V-1936 (Denier); IS, 42, Resistencia, Chaco (Denier); IS, Chaco, Rio Araja, IV-1936 (Denier); 3 S , 12, Mburucuya, Corrientes, 16-23-XI-1957 (Biraben); 3 S, 4 2 , Manatiales, Corrientes, 11-1946 (Biraben) ; 4S ,7 9 , Manantiales, Corrientes, V-1946 (T. de Apestol) ; 22, Fontana, Chaco, 28-IV-1936 (P. Denier); IS, San Javier Missiones, 4-VII-1937 (P. Denier) ; 7 S , 8 2 , S. Jose, S. Javier, Pinda- poy Misiones, 10al., 29-1-1959 (G. Raimondo); 1 S, La Plata, no data; 1 2, Villa Maria Cordoba, III-1939 (H. Parko); 1 S , Neuquen, XII-1932 (Koehler); IS, St. Cruz (Hensel S.). In Berlin Humboldt University Museum, Hungarian National Museum, Stockholm Mu- seum, Carnegie Museum, Museum of Comparative Zoology (Har- vard), La Plata Museum, J. A. Slater and R. C. Allen collections. 1969] Allen: Revision of the Genus Leptoglossus 91 Leptoglossus impictiis (Stal) Anisoscelis impicta Stal, 1859:233. Theognis impictus Mayr, 1866: 104. Leptoglossus impictus Stal, 1870:164. — Lethierry and Severin, 1893: 48. — Berg, 1879:72. — Pennington, 1922:131-2. Pronotum, clavus and corium uniformly dark tan with head above and below and pronotal calli piceous; head with lateral areas and three narrow stripes above light reddish brown; first three antennal segments (with the fourth somewhat lighter), scutellum, veins of clavus and corium, fore and middle legs, undilated portion of hind tibiae, and entire venter dark tan to light reddish brown; apex scutellum, large maculae on inner and a few small maculae on outer tibial dilation whitish yellow to ochraceous; lateral and ventral por- tion of hind femora and bases of teeth and tubercles light reddish brown, remainder of hind femora and tibial dilations fuscous with tips of teeth and tubercles piceous; connexivum and abdominal dorsum piceous with anterior fourth of connexival segments and apical fourth of last tergum laterally ochraceous; membrane uniformly dark; hind wing concolorous; pronotum, clavus and corium closely punctate with surface above punctures uneven; scutellum transversely rugulose with few punctures; body with whitish to golden appressed, short, erect hairs; legs more densely clothed with long erect pilosity. Head non-declivent with both tylus and juga raised above level of antenniferous tubercles, tylus blunt, forming a rounded elevated ridge slightly exceeding juga, length head 2.04, width head 2.04, inter- ocular distance 1.20, anteocular distance 1.20; pronotum steeply declivent, lateral margins strongly sinuate and denticulate caudally, humeral areas broadly and obliquely ascending, humeral angles sub- spinose, postero-lateral margins serrate, disk without a median longi- tudinal carina, calli barely elevated, raised area between calli with two small tubercles laterally, length pronotum 2.72, width across humeri 6.24, width anterior margin 1.56; length scutellum 1.92, width scutellum 2.28; labium extending to posterior edge of metasternum, length labial segments I 1.80, II 1.68, III 1.08, IV 1.44; length antennal segments I 2.40, II 3.36, III 2.04, IV 2.28; hind femora strongly incrassate with a large number of well developed tubercles; outer dilations barely phylliform with two shallow emarginations, occupying 64% length of hind tibiae, and about twice width of inner dilations; inner dilations lanceolate, slightly shorter than outer dila- tions, furnished with many small spine-like teeth; undilated portion of hind tibiae on inner margins with a double row of small spine-like teeth; length hind tibiae 6.96; length outer dilations 4.44, width outer 92 Entomologica Americana [Vol. 45 dilations 1.44; length inner dilations 3.36, width inner dilations 0.60; genitalia as in chilensis', total length 14.9. In a series of 70 specimens from Piedra Pintada, Neuquen, Argen- tina, the following differences were noted: dorsum a variegated dark brown and fuscous with numerous piceous maculae; some specimens with declivous face of pronotum entirely yellow or separated medially into two yellow areas, one specimen with entire pronotum yellow; size and number of tubercles and teeth on pronotum and hind femora greater than on other impictus individuals and usually entirely piceous. The remainder of the individuals examined are lighter than the above specimens, varying from light tan to dark brown. The holotype is the only individual I have seen without piceous maculae on the abdominal mid-venter. L. chilensis may be distinguished from impictus by the narrower hind femora of the males (both sexes have approximately the same size femora) and by the smaller and fewer tubercles. This is most evident on the dorsal surface where there are only 5-10, whereas in impictus the hind femora of the males are noticeably more swollen and have many more prominent tubercles (15-20 on the dorsal surface). In chilensis the humeral areas are not greatly expanded and the hu- meral angles are sub-acute. In impictus the opposite is true with the humeral angles subspinose. L. chilensis is a narrower species, the length is usually greater than 2.5 times the width across the humeri; in impictus the length is usually less than 2.5 ( impictus average 2.4 (2. 2-2. 6) — chilensis 2.8 (2. 5-3.0)). The fourth antennal segment is slightly longer in chilensis , averaging 1.3 (1. 1-1.7) times the width of the head; in impictus the average is 1.1 (1. 0-1.3). There is some overlap in the measurements of the two previously mentioned char- acters and they are difficult to use diagnostically. Because the series from Piedra Pintada resembles so closely the color pattern of chilensis , and there is no significant genitalia dif- ference, I am led to doubt the validity of impictus as a species. Yet, chilensis and impictus seem to be sympatric in their distribution in Argentina, and impictus is a wider, more robust species. For these reasons, I think the best solution at the present is to retain impictus as a distinct species. There probably will remain some confusion regarding these two species, but until more biological and distributional data are at hand, the exact relationship cannot be understood. Distribution : Argentina and Uruguay. Material Examined : Holotype: S, Montevideo, Uruguay. In Stockholm Museum. ARGENTINA: IS, 3$, Estancia Don Ro- berto, Lavaisse, St. Luis. 12 Oct. 1942; IS, Carmen, Patagones (Berg); 1$, Santiago del Estero, Rio Salado (Wagner); 12, Santa 1969] Allen: Revision of the Genus Leptoglossus 93 Fe (J. M. Bosq); 1$, 12, Mendoza; 1$, San Pablo, 1905-X (Ve- zenyi); 1^, Mendoza (H. Rolle, Berlin W. ); 12, Rio Negro, Rio Colorado, XII-1930 (J. M. Bosq); 12, Cordoba, III-1944 (J. M. Bosq); 1 S , Cordoba (J. M. Davis); 1 2, Salta, Chicoana, “s/tabaco” (J. M. Bosq); 42 $, 28 2, Piedra Pintada, Neuquen, 9-11-1941 (R. Maldonado). In La Plata Museum, Hungarian National Museum, Museum of Comparative Zoology (Harvard), British Museum (Nat- ural History), Deutsches Entomologisches Institut, J. A. Slater and R. C. Allen collections. Leptoglossus quadricollis (Westwood) Anisoscelis quadricollis Westwood, 1842:17. Anisoscelis inconspicuus Stal, 1860:32. Theognis pulcher Mayr, 1865:434. — Mayr, 1866: 106, fig. 25. Leptoglossus quadricollis Stal, 1870:164. — Lethierry and Severin, 1894:48. Leptoglossus impressicollis Berg, 1892:69. — Pennington, 1922:134. New Synonymy. Pronotum, scutellum, clavus and corium dark reddish brown with head above and below, pronotal calli and outer areas of first antennal segment piceous; pronotum and scutellum with indistinct piceous maculae; veins of clavus and corium reddish tan; medial area of third and entire fourth antennal segments, apex scutellum, and large maculae on inner tibial dilations yellow; head laterally and with three narrow stripes above, fore and middle legs, undilated portion of hind tibiae and inner margin of first antennal segments light brown; an- tennal segments two and remainder of three somewhat darker brown; distal three fourths of hind femora and tibial dilations fuscous with femoral teeth and tubercles piceous; thoracic and abdominal venter light brown with many small piceous spots frequently fusing to form larger maculae, also large piceous areas as follows: dorsal half of meso- and metapleura, mid-venter of thorax, most of ventral area on second, third, and fourth abdominal sterna; connexivum and abdom- inal dorsum piceous with connexival incisures ochraceous; mem- brane uniformly dark; pronotum, clavus and corium coarsely and closely punctate; pronotum with surface above punctures smooth medially, uneven laterally; scutellum transversely rugulose with few punctures; body and appendages with appressed golden pubescence and erect short hairs, becoming more dense on head, anterior portion of pronotum, propleura and hind femora; anterior portion of pro- notum and scutellum with longer erect pilose hairs. Head non-declivent with both tylus and juga raised above level 94 Entomologica Americana [Vol. 45 of antenniferous tubercles, tylus blunt, forming a rounded elevated ridge, slightly exceeding juga, length head 2.04, width head 2.04, interocular distance 1.08, anteocular distance 1.20; pronotum steeply declivent, lateral margins sinuate and entire, humeral areas broadly and obliquely ascending, humeral angles subacute, postero-lateral margins barely crenulate, disk slightly tumid with a very shallow median longitudinal furrow, median longitudinal carina obsolete, calli prominently elevated, raised area between calli with two small tu- bercles, length pronotum 2.24, width across humeri 5.12, width an- terior margin pronotum 1.44; length scutellum 1.80, width scutellum 2.04; labium extending to posterior margin of second abdominal sternum, length labial segments I 2.16, II 1.92, III 1.08, IV 1.92; length antennal segments I 2.40, II 3.60, III 2.16, IV 3.12; hind tibiae with outer dilations conspicuously phylliform with one shallow and three deep emarginations, occupying 81% length of hind tibiae, about twice width of inner dilations; inner dilations lanceolate, shorter than outer dilations, armed with many small spine-like teeth; un- dilated portion of hind tibiae with few small spine-like teeth arranged in a double row on inner margins, length hind tibiae 8.16; length outer dilations 6.60, width outer dilations 2.16; length inner dilations 5.52, width inner dilations 0.96. Claspers (Fig. 14) with inner basal lobe low and rounded, shank thin without a median lobe, hook strongly curved; capsule with a rounded median notch, no dorsal prongs, lateral margins barely flared; aedeagus with dorsal sac of conjunctiva with sclerotized proximal lobes present; medial lobes present and unsclerotized, left lobe long and wide, right lobe shorter, more narrow; sclerotized anterior distal lobe present, two lateral distal lobes of same shape and size, both sclerotized; total length 14.4. The description is of the male lectotype of L. inconspicuus. In the small series of specimens examined, there is little variation in the color pattern, although one individual has the entire pronotum yellow and with small piceous spots. The labium varies in length from the posterior margin of the metasternite to the posterior margin of the second abdominal sternite. L. quadricollis is a member of the chilensis group as evidenced by the short labium, lack of the transverse fascia on the corium, the genital capsule with a rounded median notch and no dorsal prongs. The lack of a serrate lateral margin on the pronotum, the larger and more phylliform tibial dilation, and the distinctive clasper with the median lobe of the shank absent will distinguish quadricollis from the other members of the group. After examining the male holotype of impressicollis Berg, I found 1969] Allen: Revision of the Genus Leptoglossus 95 it to be conspecific with quadricollis and I hereby formally synonymize impressicollis. Distribution : Argentina and Brazil. Material Examined : Syntype series of Leptoglossus inconspicuus Stal: 1 2 , Brazil; 1 $ , Brazil (F. Sahib) ; 1 $ , Rio Janeiro (F. Sahib) ; 1 <£, Rio Jan. (Stal). In Stockholm Museum. The S specimen from Rio Jan. (Stal) is selected as Lectotype of inconspicuus, and an appropriate label has been placed on the specimen to that effect. $ holotype of Leptoglossus impressicollis Berg from Corrientes, Argen- tina. In La Plata Museum. BRAZIL: 1$, 12, Rio de Janeiro, Brasilien (Dr. Studt G.). In Berlin Humboldt University Museum. Leptoglossus dentatus Berg Leptoglossus dentatus Berg, 1892:68. — Lethierry and Severin, 1894: 47. — Pennington, 1922:133. Pronotum, clavus and corium uniformly dark reddish brown, head above piceous and veins of corium more reddish; apex scutellum, fourth antennal segments, fore and middle legs, basal third of hind femora, undilated portion of hind tibiae and maculae on tibial dilations yellowish to ochraceous; head below and with three narrow stripes above, first antennal segments (with segments two and three somewhat lighter), hind femora dorsally, and tibial dilations dark reddish brown; thoracic and abdominal venter light reddish brown with a single large piceous spot on antero-dorsal area of mesopleuron, pronotal teeth and femoral teeth and tubercles piceous; connexivum and abdominal dorsum fuscous with anterior fourth of connexival segments, large portions of antero-lateral areas of abdominal dorsum and segmental sutures bright yellow, strongly contrasting with remaining dark areas; membrane uniformly dark; pronotum, clavus and corium coarsely and closely punctate, surface above punctures uneven; scutellum trans- versely rugulose with few punctures; body with short appressed silvery white hair, very dense in patches on anterior pronotal disk; body with sparse short and long erect hairs; legs with more dense, longer, erect pilosity. Head non-declivent with tylus and juga raised above level of antenniferous tubercles, tylus forming a rounded elevated ridge, barely exceeding juga, distal end of tylus slightly produced dorsally as a low rounded knob, length head 2.28, width head 2.16, interocular distance 1.20, anteocular distance 1.20; pronotum steeply declivent, lateral margins with well spaced teeth much larger than serrations on postero- lateral margins, humeral areas broadly rounded and almost vertically ascending, humeral angles acute, calli prominently elevated, raised 96 Entomologica Americana [Vol. 45 area between calli with two small tubercles, length pronotum 2.24, width across humeri 4.48, width anterior margin pronotum 1.44; length scutellum 1.92, width scutellum 1.80; labium extending to middle of metasternum, length labial segments I 1.92, II 1.80, III 1.08, IV 1.44; first antennal segment with a few short teeth, length antennal segments I 2.52, II 3.72, III 2.28, IV 3.96; hind tibiae with outer dilations phylliform, with two deep and one shallow emargina- tion, occupying 66% length of hind tibiae and almost two and a half times width of inner dilations; inner dilations lanceolate, considerably shorter than outer dilations, distal margins furnished with a few small spine-like teeth, undilated portion of hind tibiae with a double row of small spine-like teeth on inner margins, length hind tibiae 7.44; length outer dilations 5.04, width outer dilations 2.04; length inner dilations 3.60, width inner dilations 0.60. Claspers (Fig. 15) with inner basal lobe small and rounded, shank thin with prominent median lobe, hook short and strongly curved; capsule (Fig. 39) with a rounded median notch and with prominent but low rounded dorsal prongs, lateral margins flared; aedeagus with dorsal sac of conjunctiva with sclerotized proximal lobes; unsclerotized medial lobes present, right lobe almost as long as dorsal sac, left lobe shorter and not as wide; large anterior distal lobe present, lateral lobes present — one spinose, one rounded, all distal lobes sclerotized; total length 15.0. The description is of a male specimen from Tucuman, Argentina. There is no significant variation in the material examined, except that some individuals have a few small piceous maculae on the abdominal venter. L. dentatus is an easily identifiable species by the possession of strongly dentate pronotal lateral margins, the single large black spot on the metapleura, and the yellow pattern on the abdominal dorsum. The placement of dentatus within the chilensis group is based on the short labium and lack of the transverse corial fascia. The presence of small dorsal prongs on the capsule suggests a relationship to the stigma group. L. dentatus is one species in Division B that lacks nu- merous small piceous spots on the entire venter, although a few indi- viduals have faint piceous spots confined to the abdominal mid-ventral area. Mann (1969) reports that dentatus feeds upon a variety of cactus species, primarily prickly pear ( Opuntia spp.). Distribution : Argentina and Uruguay (Mann, 1969). Material Examined: Holotype. 1 9, Cordoba, Argentina. In La Plata Museum, Argentina. ARGENTINA: 2$, Santiago del Estero, Rio Salado (Wagner); 1 S, Corodoba (W. M. Davis); IS, Cordoba; 1969] Allen: Revision of the Genus Leptoglossus 97 1 S , Tucuman, II-1-1950 (P. Wygodzinsky) ; 1 S , Guemes, IV-3-1930 (H. A. Jaynes); IS, 2 2, Santiago del Estero Fernandez, 30-1-1939 (J. M. Bosq); 2$, Santiago del Estero (Wagner); 2 S, Chaco, Las Brenas, IV-1941 (Bosq); 12, Chaco, Asila Leg. (Bosq); 12, Cor- zuela, 8-1-1936 (Denier); IS, Colonia Benitez, l-V-1939 (Denier); 12, Prov. de La Rioja (Bosq); 12, S/algodoners danos no constat, S. Pena, l-IV-1938 (Denier); 2 2, T. Formosa, Paso Angelito, S/el rio Porteno Lag, Nainec, 12-XII-1935 (Denier); 1 S , Tablillas, Salta, 11-1945 (Martinez); 12, Concaran S. Luis, 1 6-III-1 960 (Vidal S. — Trotta). In La Plata Museum, Museum of Comparative Zoology (Harvard), United States National Museum, J. A. Slater and R. C. Allen collections. Leptoglossus balteatus (Linnaeus) Cimex balteatus Linnaeus, 1771:534. Cimex auctus Fabricius, 1781:351. Lygaeus auctus Fabricius, 1794:139. Lygaeus balteatus Fabricius, 1794:142. Anisoscelis fasciatus Herrich-Schaeffer, 1851:277. Anisoscelis thoracicus Guerin, 1857:386. T heognis schaefferi Mayr , 1866:1 02-3 . Leptoglossus auctus Stal, 1868:52. Leptoglossus balteatus Stal, 1870:161-2. — Barber, 1939: 310-11. — Barber and Bruner, 1947:80. — Wolcott, 1948:197. Anisoscelis selecta Walker, 1871 : 127-8. New Synonymy. Pronotum, scutellum, clavus and corium dark reddish brown with head above, outer areas of first antennal segments and pronotal calli piceous; medial portion of antennal segments two and three and apical half of fourth, a narrow band on posterior marginal area and two large well separated, longitudinal ovoid spots on disk of pronotum ochraceous, the latter interspersed with numerous small piceous spots; apex scutellum, straight transverse fascia on corium, large maculae on inner tibial dilations, and undilated portion of hind tibiae whitish yellow; head below and with three narrow stripes above, inner margin of first antennal segment with distal portion of segments two and three and apical half of fourth segment, basal and ventral portions of all femora, fore and middle tibiae entirely, and hind tibial dilations light reddish brown; dorsal portion of femora fuscous with teeth and tu- bercles piceous; thoracic and abdominal venter light to dark reddish brown with numerous small piceous spots frequently fusing to form larger maculae; connexivum and abdominal dorsum piceous with con- nexival sutures and medial area of dorsum broadly ochraceous; mem- 98 Entomologica Americana [Vol. 45 brane uniformly dark; pronotum, clavus and corium coarsely and closely punctate with surface above punctures uneven, scutellum transversely rugulose with small punctures; venter with dense short and appressed whitish hairs; clavus and corium with short semi- decumbent golden hairs and anterior half of pronotum with thick darker hairs. Head non-declivent with tylus and juga above level of antenni- ferous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 2.64, width head 2.16, interocular distance 1.20, anteocular distance 1.32; pronotum abruptly declivent, lateral margins straight and entire, humeral areas slightly expanded with humeral angles spinose and obliquely ascending, postero-lateral margins serrate, calli prominently elevated with area between calli only slightly higher, disk posteriorly without a median longitudinal carina, length pronotum 3.04, width across humeri 5.76, width an- terior margin pronotum 1.56; length scutellum 2.16, width scutellum 2.04; labrum reaching to anterior margin of fourth abdominal ster- num, length labial segments I 2.76, II 2.52, III 1.44, IV 2.64; length antennal segments I 2.64, II 3.96, III 2.88, IV 4.56; hind tibiae with outer dilation phylliform with one shallow and two deep emargina- tions, occupying 63% length of hind tibiae, width about twice width of inner dilations; inner dilations lanceolate, shorter and with margin distally furnished with a few small spine-like teeth; undilated portion of hind tibiae armed inwardly with a double row of small spine-like teeth, length hind tibiae 8.40; length outer dilations 5.28, width outer dilations 1.92; length inner dilations 4.44, width inner dilations 0.84. Claspers (Fig. 25) with prominent and rounded inner basal lobe, median lobe on shank rounded and well developed, hook strongly curved; capsule (Fig. 51) with median notch “V” shaped, dorsal prongs low and rounded almost obsolete, lateral margins flared and extending beyond curvature of capsule; aedeagus with dorsal sac of conjunctiva bearing paired sclerotized proximal lobes, two long similar median lobes present with tips slightly sclerotized, anterior distal lobe and two lateral distal lobes present with lateral lobes spinose, all three distal lobes heavily sclerotized; total length 17.3. The above description is of a male specimen from Jamaica. The variation in color present is of a geographic nature. Specimens from the Bahamas, Cuba, and Jamaica have the two ochraceous pronotal spots rounded and well separated, the first antennal segment is uni- colorous and the pubescence on the dorsum is generally pale and long. The specimens from Puerto Rico, Dominican Republic, and Haiti have pronotal spots that are larger, quadrate, and tending to spread out laterally and medially sometimes partially fusing together, the first 1969] Allen: Revision of the Genus Leptoglossus 99 antennal segment is bicolored, and the pubescence appears to be short and sparse. Stal ( 1871 ) and Barber ( 1939) both mentioned the pronotal coloration, but neither stated the locality of specimens in- volved. I have examined Walker’s selectus from St. Domingo, and found it to be identical to balteatus. Walker probably described the species as new based on the difference in pronotal coloration since Linnaeus described balteatus from a Jamaican specimen. Leptoglossus selectus is here formally synonymized. L. balteatus is closely related to phyllopus and can be distinguished from the latter by the pronotal coloration. Although some individuals of phyllopus may develop yellow pronotal spots, the margins of these spots are not well defined, but appear diffused. The lateral angles of balteatus are acute to subspinose and ascending, while those on phyllopus are subacute and only slightly turned up. The length of the outer tibial dilation is also of diagnostic value, in phyllopus this dilation extends 76-87% the length of the hind tibiae; in balteatus it varies between 63-70%. The unicolorous hind wing of balteatus is easily distinguished from the obviously bicolored hind wing of phyl- lopus. The median notch on the genital capsule and the basal portion of the claspers are essentially the same in the two species. In balteatus the lateral posterior margins are flared and project backwards over the curvature of the capsule, and the clasper has a well developed median lobe on the shank. The lateral margins on the capsule are not flared or prominent in phyllopus and the clasper has only a small median lobe. L. balteatus has been confused with grenadensis in the past, but the two species are quite distinct. The shape and color of the pronotum is almost identical except that in grenadensis the lateral angles are not quite so acute or ascending. The clavus and corium are more uniform in color and the transverse fascia on the corium is straight, while grenadensis has the claval area and the veins lighter in color than the rest of the corium, and the transverse fascia is distinctly angular. The difference in the genital capsule is very prominent, grenadensis has distinct dorsal prongs and balteatus (Fig. 51) has a simple margin, with prongs obsolete. The claspers and aedeagus also differ con- siderably. L. balteatus has been reported as feeding on cotton, tomatoes, leguminous plants, oranges, and cowpeas (Barber and Bruner, 1947; Pagden, 1928). Barber and Bruner (1947) also state that it breeds on guava ( Psidium guafara [j/c] L. ) and Luff a species. Distribution : This species is present in the Greater Antilles, the Bahama Islands, and there is at least one record from Florida (Barber, 100 Entomologica Americana [Vol. 45 1914). The localities cited by Uhler (1893 ) were based on erroneous identifications (see discussion under grenadensis) . Material Examined : BAHAMA ISLANDS : 1<$,1$, New Provi- dence, Nassau, July 1904 (Allen, Banbom, and Ronjant) ; 1 8 , Andros Island, Fresh Creek, April 23, 1953, Van Voast, A.M.N.H. Bahama Island Exped. (E. B. Hayden, L. Giovannoli). JAMAICA: 5 8 , 2 $ , Torrington, July 18, 1960 (P. and C. Vaurie). CUBA: 12, vie. of Havana (T. Barbour) ; 1 8 , 3 2 , Habana (F. Z. Cervera) ; 1 2 , Jaronu, V-13-31, “on weeds” (L. C. Scaramuzza); 18, Guane, Sept. 24-26, 1913; 12, Soledad, 20-11-1925 (J. G. Myers); 3$, 3 2, Nov. 1874 (Gundlack) ; 1 2 , no data. DOMINICAN REPUBLIC: 1 8 , 1 2 , San Jose de las Matas 1000-2000 ft., June 1938 (Dari.) ; 1 8,2 2 , Puerto Plata (Hurst). PUERTO RICO: 12, Humacao, July 1937 (J. Carrion); 1 2 , Ensenada, June 14-19, 1915; 1 $,Nov. 1874 (Krug). SAINT THOMAS: 1 2 , Angek, 19-1-1908 (L. Eggert) ; 2 $ , 1 2 , no data. In American Museum of Natural History, Museum of Com- parative Zoology (Harvard), Berlin Humboldt University Museum, Stockholm Museum, University of Kansas (Snow collection), J. A. Slater and R. C. Allen collections. Leptoglossus phyllopus (Linnaeus) Cimex phyllopus Linnaeus, 1767:731. Lygaeus phyllopus Fabricius, 1794:139. Anisoscelis albicinctus Say, 1831:326. Anisoscelis phyllopa Westwood, 1842:16. Anisoscelis fraterna Westwood, 1842:16. — Distant, 1901a:334. Anisoscelis confusa Dallas, 1852:453. Theognis phyllopus Mayr, 1866:103. — Hussey, 1953:32. Leptoglossus albicinctus Stal, 1868:52. Leptoglossus phyllopus Stal, 1870:161. — Distant, 1881:124, 361. — Lethierry and Severin, 1894:48. — Van Duzee, 1917:88. — Gibson, 1917:72.— Parshley, 1923 : 747.— Blatchley, 1926:225-26, fig. 46. — Deay, 1928:377. — Torre-Bueno, 1941:50. — Froeschner, 1942:594, 599.— Hussey, 1953:32.— Drew and Schaefer, 1963: 114. A medium sized species; uniformly dark chocolate brown on the dorsum with occasionally yellow spots on the pronotal disk; a straight whitish yellow transverse fascia on the corium, venter light to dark tan with numerous small piceous spots; hind wings prominently bi- colored, with the basal portion clear and the apical portion with a brownish tinge; hind tibiae with outer dilation phylliform usually with two deep emarginations. Genitalia similar to balteatus. 1969] Allen: Revision of the Genus Leptoglossus 101 L. phyllopus and balteatus are closely related species as evidenced by the similarity of the genitalia and general color pattern. The ab- sence of pronotal spots, possession of a bicolored hind wing, and sub-acute to rounded humeral angles will serve to distinguish phyl- lopus from balteatus. In general, phyllopus may be separated from any member of the stigma-group by the straight transverse fascia on the corium (stigma- group members either lack the corial fascia or if present, it is irregular not straight), the rounded median notch, and the absence of dorsal prongs on the genital capsule ( concolor in the stigma-group lacks dorsal prongs; in this case the median notch is deeply rectangular in shape and not rounded). Specifically, phyl- lopus may be separated from both concolor and oppositus by the shorter labium, which usually does not extend past the second ab- dominal sternum, in stigma and oppositus the labium usually extends well onto the abdomen around the third and fourth sternum; and from oppositus by having at least the pronotal postero-lateral margins crenulate to serrate (oppositus has all lateral margins on the pro- notum entire). L. phyllopus, a very common insect in the southern United States, has been reported as a pest on a number of important agricultural crops, where occasionally large numbers of individuals inflict con- siderable damage. The following is a partial list of crops recorded as having been attacked by phyllopus : citrus fruits (Ebeling 1959, Watson and Berger 1932), peaches (Snapp 1948), sunflowers (Adams and Gaines 1950), cucumbers and squash (Kelsheimer 1949). Blatchley (1926) lists phyllopus as common on yellow- flowered thistle, Cirsium horridulum Michx. Schaefer (1965, 1968) describes some morphological aspects of phyllopus in his studies on the higher classification of the Coreoidea. Distribution : A wide ranging species occurring as far north as New York, south to Florida, west to Iowa and Kansas, and southwest through Texas to Lower California and into Central America. I have seen specimens from as far south as Costa Rica, and Distant (1881) lists Panama and Brazil as well. Material Examined : UNITED STATES : 1 9 1 3 , 1 42 $ , Alabama, Arkansas, California, Florida, Georgia, Iowa, Kansas, Louisiana, Mis- sissippi, Missouri, Nebraska, North Carolina, Oklahoma, South Caro- lina, Texas, and Virginia. MEXICO: 13, Hidalgo, Tezontepec, 21 m. S.W. Actopan, 6600 ft. 27 August 1962 (Ordway and Marston); 23, Presisio (Forrer); 1$, Salle, no data; 12, no data (Boucard). GUATEMALA: 13, 3$, Olas de Moka, Dept. Solola, 3000, Sept. 1908. COSTA RICA: 13, Zarzero (Schild-Burgdorf); 1$, San Jose (Schild-Burgdorf). In University of Kansas, Stockholm Museum, 102 Entomologica Americana [Vol. 45 Berlin Humboldt University Museum, United States National Mu- seum, Iowa State University, Museum of Comparative Zoology (Har- vard), American Museum of Natural History, Hungarian National Museum, California Academy of Sciences, J. A. Slater and R. C. Allen collections. Leptoglossus ingens (Mayr) Theognis ingens Mayr, 1865:434. — Mayr, 1866:108, fig. 26. Leptoglossus ingens Stal, 1870:166. — Lethierry and Severin, 1894: 48. Anisoscelis santaremus Walker, 1871 : 129. New Synonymy. Pronotal disk, scutellum, clavus and corium dark reddish brown with head above, outer margin of first antennal segment, humeral and lateral areas of pronotum and pronotal calli piceous; veins of clavus and corium wine red; apex scutellum, fourth antennal segment and maculae on inner tibial dilations dark yellow to ochraceous; head below and with three narrow stripes above, inner margins of first, second and third antennal segments, bases and ventral portions of all femora, fore and middle tibiae light reddish brown; dorsum of all femora and hind tibial dilations fuscous with teeth and tubercles piceous; thoracic and abdominal venter light reddish brown with nu- merous small piceous spots frequently fusing to form larger maculae; connexivum and abdominal dorsum piceous; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate, sur- face above punctures largely even; scutellum transversely rugulose with few punctures; head, anterior half of pronotum, scutellum, and legs with rather dense long and erect yellowish pilosity; venter, clavus and corium with moderate amounts of short decumbent to semi- decumbent yellowish hairs. Head non-declivent with tylus and juga above level of antenni- ferous tubercles, tylus blunt, forming a rounded elevated ridge, slightly exceeding juga, length head 2.88, width head 2.64, interocular dis- tance 1.44, anteocular distance 1.56; pronotum abruptly declivent with lateral margin entire and sinuate, humeral areas broadly and obliquely ascending, humeral angles subacute, postero-lateral margins serrate, disk posteriorly with median longitudinal carina obsolete, calli prominently elevated, area between slightly higher, length pro- notum 3.68, width across humeri 8.00, width anterior margin pro- notum 1.68; length scutellum 2.76, width scutellum 2.76; labium extending to posterior margin of third abdominal sternum, length labial segments I 3.12, II 2.88, III 1.80, IV 3.60; length antennal segments I 2.52, II 4.20, III 3.00, IV 5.40; hind tibiae with outer 1969] Allen: Revision of the Genus Leptoglossus 103 dilations phylliform with one shallow and three deep emarginations, dilations very long, occupying 92% length of hind tibiae and width about twice width of inner dilations; inner dilations lanceolate con- siderably shorter, 55% length of outer, furnished with numerous small spine-like teeth on distal half of margins; inner margins of undilated portions of hind tibiae with two rows of many small spine-like teeth, length hind tibiae 10.8; length outer dilations 9.90, width outer dila- tions 2.76; length inner dilations 5.52, width inner dilations 1.32. Claspers (Fig. 17) with lateral basal area only slightly bulbous with inner basal lobe long, thick and rounded, median lobe on shank very prominent, hook thick, blunt and not strongly curved; capsule (Fig. 41) with median notch deep and bottom of notch “V” shaped, dorsal prongs prominent, medial posterior margins mesally projecting slightly antero-dorsally; aedeagus (Fig. 67) with dorsal sac of con- junctiva bearing sclerotized proximal lobes, two median lobes of equal size and shape with tips slightly sclerotized; anterior distal lobes large, lateral distal lobes large and spinose, all distal lobes heavily sclerotized; total length 21.6. The above description is of a male specimen from Caviuna, Parana, Brazil. This species has considerable color variation in that the pronotum may be entirely dark, or the anterior half before the humeri may be yellow, and the irregular transverse fascia on the corium may or may not be present. Specimens also vary from being uniformly piceous dorsally to a dark reddish brown. I have tenta- tively identified three female specimens from Olivenca, Brazil as ingens that have longer and more spinose humeral angles, but other- wise do not differ from typical ingens specimens. I have examined the female holotype of Leptoglossus santaremus (Walker), from Santarem, Brazil, which is deposited in the British Museum (Natural History). I was unable to find any differences be- tween it and ingens other than the fact that santaremus possesses a transverse fascia on the corium. Therefore, I am formally synonymiz- ing santaremus with ingens. I should like to point out that all the speci- mens with a transverse fascia are more northern in distribution than the specimens without it. The results of additional collecting will probably establish these two forms as distinct but conspecific popu- lations. L. ingens may be distinguished from macrophyllus by its darker color, lack of the rectangular yellow fascia on the pronotum and by the presence of two equal sized median lobes on the dorsal sac of the conjunctiva. The wide, elongate outer tibial dilations will serve to distinguish both macrophyllus and ingens from other members of the genus. 104 Entomologica Americana [Vol. 45 Distribution : Brazil, Bolivia, Paraguay, and Argentina. Material Examined : (Specimens with a corial transverse fascia) : BOLIVIA: 3 5 , Yungas de la Paz, 1000 m. (O. Garlepp) ; 1 5 , same except (Linnaea V.); 15, Province Sara (Steinbach); 15, Buena Vista, Province Tchito, Dep. Santa Cruz, 400 m., 1956 (F. Stein- bach); 15, 12, Rio Cristal Mavu, 50 mi. N.E. Cochabamba, XII-8- 1949 (L. Pena); 12, Juntus, no data; 15, no data. BRAZIL: 15, 2$, Amazon, sup. Olivenca. PARAGUAY: 1$, Horqueta, 57-10, W. 23-24 N., 44 kilm. East Paraguay Riv., 1-17-1935 (A. Schulze). Specimens without a corial transverse fascia: BRAZIL: 2 5 , Caviuna, Parana, Feb. 1947 (A. Mailer); 12, same except XI- 1945; 12, Corupa (Hansa Humbolt), S. Cath., 1-1946; 15, Nova Teutonia, Santa Catarina, IV-10-1948 (F. Plaumann). PARAGUAY: 12, Horqueta, 57-10, W. 23-24 N., 44 kilm. East Paraguay Riv., XII- 1 8- 1934 (Alberto Schulze); 12, same except 1-3-1936. In Stockholm Museum, Hungarian National Museum, Berlin Humboldt University Museum, Museum of Comparative Zoology (Harvard), American Museum of Natural History, J. A. Slater and R. C. Allen collections. Leptoglossus macropliyllus Stal Leptoglossus macrophyllus Stal, 1870:162. — Lethierry and Severin, 1894:48. Pronotal disk posteriorly, scutellum, clavus and corium bright reddish brown with head, first antennal segment, marginal areas of pronotum, calli, and claval sutures piceous; pronotal disk anteriorly with one large sharply defined rectangular transverse band, irregular transverse fascia on corium, and maculae on inner tibial dilations whitish yellow; inner margins of first, medial portions of second and third and the entire fourth antennal segments ochraceous, with proxi- mal and distal portions and a narrow dorsal stripe on segments two and three fuscous; head laterally and with three narrow stripes above, fore and middle tibiae, and undilated portion of hind tibiae light reddish tan; basal half and ventral portions of all femora bright reddish brown with remainder of femora and tibial dilations dark reddish brown to fuscous; thoracic and abdominal venter dark reddish tan with numerous small piceous spots frequently fusing to form larger maculae; connexivum and abdominal dorsum fuscous to piceous; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures largely even; scutellum transversely rugulose with a few punctures; head, anterior half of pronotum, scutellum, appendages, and mid-venter with dense 1969] Allen: Revision of the Genus Leptoglossus 105 golden pilose hairs; remainder of venter and clavus and corium with short appressed and semi-decumbent golden hairs. Head non-declivent with tylus and juga above level of antenni- ferous tubercles, tylus blunt, slightly exceeding juga and forming a low rounded ridge, length head 2.76, width head 2.40, interocular dis- tance 1.32, anteocular distance 1.56; pronotum abruptly declivent with lateral margins entire and sinuate, humeral areas broadly rounded and only barely obliquely ascending, humeral angles sub-acute, postero-lateral margins crenulate, disk posteriorly with median longi- tudinal carina obsolete, calli prominently elevated, area between calli higher, length pronotum 3.36, width anterior margin pronotum 1.56; length scutellum 2.28, width scutellum 2.40; labium reaching to an- terior margin of fourth abdominal sternum, length labial segments I 2.88, II 2.76, III 1.68, IV 3.00; length antennal segments I 2.40, II 3.84, III 2.88, IV 4.80; hind tibiae with outer dilations phylliform with one shallow and two deep emarginations, dilations very long, occupying 86% length of hind tibiae, width about twice that of inner dilations; inner dilations lanceolate, considerably shorter, 68% length of outer with a few small spine-like teeth on distal half of margins; inner margins of undilated portions of hind tibiae with a double row of small spine-like teeth; length hind tibiae 8.88, length outer dila- tions 7.68, width outer dilations 2.52; length inner dilations 5.58, width inner dilations 1.08. Claspers (Fig. 16) with lateral basal area bulbous, inner basal lobe prominent, wide and rounded, median lobe on shank prominent and sharply curving back onto shank, hook thick, blunt, not strongly curved; capsule (Fig. 40) with median notch deep and bottom of notch “V” shaped, dorsal prongs prominent with inner margins straight, posterior medial margin projecting slightly antero-dorsad, aedeagus with dorsal sac of conjunctiva with sclerotized proximal lobes; unsclerotized median lobes present, left lobe short with right lobe much longer; large anterior distal lobe present, two lateral lobes large and spinose, all distal lobes heavily sclerotized; total length 19.4. The following minor color variations were noted: four specimens have a narrow reddish stripe on the yellow pronotal band, with one individual having this band almost divided in two; the abdominal dorsum may be a rusty reddish brown instead of piceous; the scutellum occasionally piceous instead of a bright reddish brown. L. macrophyllus closely resembles ingens in shape and size. The hind tibial dilations are identical, except that the outer dilation is a little longer in ingens. The difference in color between the two species is considerable. Macrophyllus is a brightly colored species and the yellow rectangular band on the pronotum will distinguish it from the 106 Entomologica Americana [Vol. 45 darker colored ingens. In macrophyllus the two unequal sized median lobes are present on the dorsal sac of the conjunctiva, in ingens the same lobes are of equal size and shape. Distribution : Venezuela and Colombia. Material Examined : Two syntypes: IS, 1$, Bogota (Lindig), Colombia. In Stockholm Museum. The S specimen bearing the label Allotypus is here selected as Lectotype, and a label has been at- tached to the insect to this effect. VENEZUELA: IS , 3 2, Merida; IS, Rancho Grande nr. Maracay, 2-IX-1946; 1$ (Breddin); 1$, Carepilo, June, 1937 (Pablo J. Anduze). COLOMBIA: 1 S, 12, Villavicencio; 12, Villa Vincencio, 1919; 1$, Caqueza, Cundi- namarca, XII-8-39 (F. J. Otoya); 1 2, Aquadita, VI-1914. In United States National Museum, California Academy of Sciences, American Museum of Natural History, University of Kansas (J. R. de la Torre - Bueno collection), Stockholm Museum, Deutsches Entomologisches Institut, J. A. Slater and R. C. Allen collections. Leptoglossus conspersus Stal Theognis vexillatus, 1866 (nee Stal) : 101. Leptoglossus conspersus Stal, 1870:163. — Lethierry and Severin, 1894:47. Pronotum posteriorly dark chocolate brown; anterior portion of corium dark reddish brown with scutellum, clavus, apex corium, and veins contrasting bright reddish brown; head, first two antennal seg- ments and pronotal calli piceous; two large medially separated yellow- ish orange areas interspersed with numerous small piceous spots on pronotum, these light areas occupying almost the entire pronotal disk, extending laterally to margins and obliquely ascending posteriorly to include anterior half of humeral areas; fourth antennal segments and a wide irregular transverse fascia on corium whitish yellow; head below and with three narrow stripes above, basal portion of third antennal segments, small scattered maculae on outer tibial dilation and larger maculae on inner dilations light reddish tan; all legs and tibial dilations fuscous with teeth and tubercles piceous; thoracic and abdominal venter light reddish tan with numerous small piceous spots frequently fusing to form larger maculae; connexivum and abdominal dorsum fuscous to dark reddish brown with anterior portion of each connexival segment light reddish brown; membrane uniformly dark; pronotum, clavus and corium closely and finely punctate, surface above punctures largely smooth; scutellum transversely rugulose with numerous small punctures; dorsum with short semi-decumbent golden hairs, with head behind eyes, anterior half of pronotum and 1969] Allen: Revision of the Genus Leptoglossus 107 appendages more densely clothed with longer pilose hairs; venter with short erect and appressed whitish hairs. Head non-declivent with tylus and juga above level of antenniferous tubercles, tylus blunt, slightly exceeding juga, forming a rounded ele- vated ridge, length head 2.16, width head 2.04, interocular distance 1.08, anteocular distance 1.08; pronotum abruptly declivent with lateral margins entire and barely sinuate, humeral areas not strongly produced and humeral angles sub-acute, postero-lateral margins finely serrate, posterior pronotal disk with an obsolete median longitudinal carina, calli prominently elevated, area between calli slightly higher and with two small tubercles, length pronotum 2.40, width across humeri 4.32, width anterior margin 1.32; length scutellum 1.80, width scutellum 1.92; labium reaching to middle of third abdominal sternum, length labial segments I 2.16, II 2.04, III 1.08, IV 1.92; length antennal segments I 2.16, II 3.12, III 2.40, IV 4.44; hind tibiae with outer dilation phylliform with one shallow and two deep emargin- ations and occupying 74% length of hind tibiae; width outer dilation about twice width of inner dilations; inner dilations shorter and lance- olate with margins distally furnished with a few small spine-like teeth; undilated portion of hind tibiae armed inwardly with several small teeth in a double row, length hind tibiae 7.56; length outer dilations 5.44, width outer dilations 1.80; length inner dilations 4.32, width inner dilations 0.84. Clasper (Fig. 20) with bulbous lateral margins on base and inner basal lobe very prominent, shank thick with median lobe hardly evi- dent, hook strongly curved with a small knob on outer curvatures; capsule with median notch deeply rounded with dorsal prongs strongly produced and acute; aedeagus with dorsal sac of conjunctiva with sclerotized proximal lobes; median lobes present with left and right lobes bifid; anterior distal lobe large and heavily sclerotized, lateral lobes spinose and heavily sclerotized; total length 14.4. In the type series two females are darker and have a more uniform color to the clavus and corium. The labium varies in length from the posterior margin of the metastemum to the third abdominal sternum. L. conspersus is closely allied to grenadensis, having darker and lighter reddish tan areas on the clavus and corium and similar geni- talia, but differing in that conspersus has large diffused pronotal mark- ings, a longer and wider outer tibial dilation, more blunt and less elevated humeral angles, and (although not completely diagnostic) the labium is generally shorter. The following characters will separate conspersus from zonatus, neovexillatus, and impictipennis : large pro- notal spots occupying most of disk, a more or less bicolored clavus and corium; clasper (Fig. 20) with a long inner basal lobe and a more 108 Entomologica Americana [Vol. 45 blunt median lobe on the shank; a very large heavily sclerotized an- terior distal lobe on the dorsal sac of the conjunctiva and the right median lobe bifid. Distribution’. Colombia and Brazil. Material Examined: Four syntypes: 1 £ , 3 2, Bogota, Colombia (Lindig). In Stockholm Museum. The $ specimen (described above) bearing the “Typus” label is here selected as Lectotype and an appropriate label attached to the specimen. One additional speci- men was available for study, a $ from Minas Geraes, Brazil, also in the Stockholm Museum. Leptoglossus grenadensis new species (Fig. 3) Leptoglossus balteatus Uhler, 1893 {nee. L.) :705. Leptoglossus zonatus Uhler, 1894 ( nec . Dallas): 178. Leptoglossus balteatus Distant, 1901b {nec. L.):417 (pt.). Pronotum, scutellum, and corum dark reddish brown with clavus a light reddish tan; head above, pronotal calli and humeral angles piceous; irregular transverse fascia on corium, posterior sub-marginal area and two large separated longitudinally ovoid spots on disk of pronotum, apex scutellum, and maculae on inner tibial dilations whitish to dark yellow; head laterally and with three narrow stripes above, medial area of antennal segments two and three and entire fourth segments light reddish tan; legs light reddish brown with femora distally, tibial dilation, entire first antennal segment and basal and distal portions of second and third segments fuscous; femoral and tibial teeth and tubercles piceous; thoracic and abdominal venter ochraceous, mottled with numerous small piceous spots occasionally fusing to form larger maculae; connexivum and abdominal dorsum piceous with anterior third of connexival segments and intersegmental sutures on dorsum ochraceous; membrane uniformly dark; pronotum, clavus and corium coarsely and closely punctate with surface above punctures on pronotum uneven; scutellum transversely rugulose with few small punctures; venter densely clothed with short appressed whitish pubescence; dorsum with short semi-decumbent golden pu- bescence with head behind eyes, anterior portion of pronotum, scu- tellum, and legs also with longer pilose hairs. Head non-declivent with tylus and juga above level of antenni- ferous tubercles, tylus blunt, slightly longer than juga and forming a low rounded elevated ridge, length head 2.40, width head 2.16, inter- ocular distance 1.20; anteocular distance 1.32; pronotum abruptly declivent with lateral margins entire and straight, humeral areas not 1969] Allen: Revision of the Genus Leptoglossus 109 expanded with humeral angles acute and obliquely ascending, postero- lateral margins serrate, calli prominently elevated with area between calli slightly higher, disk posteriorly without a median longitudinal carina, length pronotum 2.88, width across humeri 5.12, width an- terior margin pronotum 1.44; length scutellum 2.04, width scutellum 2.04; labium reaching to anterior margin of fourth abdominal ster- num, length labial segments I 2.52, II 2.40, III 1.32, IV 2.52; length antennal segments I 2.52, II 3.84, III 2.88, IV 4.68; hind tibiae with outer dilations phylliform and with two deep emarginations occupy- ing 62% length of hind tibiae, width of outer dilations about twice width of inner dilations; inner dilations slightly shorter, lanceolate with entire margin furnished with many small spine-like teeth; un- dilated portion of hind tibiae inwardly with a double row of small spine-like teeth, length hind tibiae 7.80; length outer dilations 4.80, width outer dilations 1.44; length inner dilations 4.20, width inner dilations 0.84; genitala as in conspersus^ total length 15.5. The following variations were noted in the type series : some indi- viduals with pronotum, clavus and corium a light red brown and yellowish orange areas not so strongly contrasting as in dark red brown individuals; low obscure longitudinal carina on pronotal disk evident on many specimens; scutellum varying from black to red brown and first antennal segment having a prominent inner light stripe; variation in extension of labium from the middle of third to posterior margin of fourth abdominal sternum, one specimen has labium extending to posterior margin of fifth sternum. The diagnostic characters which separate grenadensis from its most closely related species, conspersus, are as follows: the pronotal coloration in grenadensis is two well defined yellow ovoid spots and a pale posterior sub-marginal area, in conspersus these pronotal spots are large and diffuse, occupying most of the pronotum anteriorly, and the posterior sub-marginal area is concolorous with the rest of the dark ground color of the pronotum. L. grenadensis has acute, ele- vated humeral angles, whereas in conspersus these angles are more rounded and barely ascending. The length of the outer dilation is smaller in grenadensis , occupying 61-64% of the length of hind tibiae, while that of conspersus is 74-81% the length of hind tibiae. From balteatus, grenadensis differs chiefly in that it has dorsal prongs on the capsule and an irregular transverse fascia on the corium. Distribution : L. grenadensis was first reported from St. Vincent by Uhler (1893) as L. balteatus. Uhler (1894) again reported it as L. zonatus occurring on the island of Grenada. Both of these papers pertain to the collection made by H. H. Smith. Distant (1901b) after seeing the H. H. Smith collection (now in the British Museum) re- 110 Entomologica Americana [Vol. 45 ferred to all the specimens of this species from Grenada as L. balteatus. I have seen seven specimens from the British Museum that bear the labels of the H. H. Smith collection, and these are definitely gren- adensis. Holotype: 8. GRENADINES: Grenada, Balthazar, windward side (H. H. Smith). In United States National Museum, type no. 70225. Paratypes: GRENADINES: 18, 1$, Mustique I. (H. H. Smith); 18, 2$, Becquia I. (H. H. Smith); 3$, Grenada (H. E. Summers); 12, Grenada, 8-25-1891 (H. E. Summers); 1$, 12, Grenada, St. Georges, Aug. 1910 (Allen and Brues); 12, Grenada, Santeurs, Sept. 1910 (Allen and Brues); 18, Grenada, Mount Gay Est. leeward side (H. H. Smith); 1 8, Grenada, Lake Antoine Est. windward side (H. H. Smith); 18, 12, St. Vincent, Yambu River Valley, 10-3-31, “on guava” (M. Kisliuk, C. E. Cooley); 18, 12, same except 10-2-31; 18, no data. In United States National Mu- seum, Museum of Comparative Zoology (Harvard), Iowa State Uni- versity, British Museum (Natural History), and J. A. Slater collec- tions. Leptoglossus zonatus (Dallas) Anisoscelis zonata Dallas, 1852:452. Anisoscelis vexillatus Stal, 1855:185. New Synonymy. Theognis zonatus Stal, 1862:295. Theognis vexillatus Stal, 1862:295. Leptoglossus zonatus Stal, 1870:162. — Distant, 1881:125, 361, pi. 12, fig. 16. — Lethierry and Severin, 1894: 49. — Van Duzee, 1917:88.— Gibson, 1917:72.— Torre-Bueno, 1941:50-1. Leptoglossus vexillatus Stal, 1870:162. — Lethierry and Severin, 1894: 49. Pronotum, scutellum, clavus and corium dark chestnut brown with head above, outer marginal areas of first antennal segment, pro- notal calli and marginal areas of humeri piceous; apex scutellum, wide irregular transverse fascia on corium, larger maculae on inner tibial dilations and two large well separated distinctly longitudinal ovoid spots on pronotal disk whitish yellow, the latter interspersed with numerous small piceous spots; basal three fourths of antennal seg- ments two and three and entire fourth antennal segments light reddish brown with inner margins of first somewhat darker and distal portion of second and third fuscous; head below and with three narrow stripes, above reddish brown; fore, middle and undilated portion of hind tibiae, all tarsi, basal and ventral portions of femora and small 1969] Allen: Revision of the Genus Leptoglossus 11 scattered maculae on outer tibial dilation light brown to reddish brown; distal portion of femora dorsad and tibial dilations fuscous with teeth and tubercles piceous; thoracic and abdominal venter varie- gated dark and light brown with numerous small piceous spots fre- quently fusing to form irregular maculae; connexivum and abdominal dorsum piceous with anterior fourth of connexival segments and narrow areas around intersegmental sutures yellow; membrane uni- formly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures smooth; scutellum transversely rugulose with numerous small punctures; venter thickly clothed with short appressed whitish pubescence, and sparse short erect hairs; legs, anterior portion of pronotum and scutellum with more dense, longer and erect pale pilose hairs; dorsum with semi-decumbent whitish hairs. Head non-declivent with tylus and juga above level of antenni- ferous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge; length head 2.88, width head 2.64, inter- ocular distance 1.44, anteocular distance 1.56; pronotum abruptly declivent, lateral margins only slightly sinuate and entire, humeral areas moderately expanded, not obliquely ascending, humeral angles small sub-acute, postero-lateral margins serrate, calli prominently elevated, area between calli only slightly higher, disk posteriorly with median longitudinal carina obsolete, length pronotum 3.36, width across humeri 6.56, width anterior margin pronotum 1.68; length scutellum 2.52, width scutellum 2.64; labium reaching middle of fourth abdominal sternum, length labial segments I 3.24, II 3.12, III 1.80, IV 3.36; length antennal segments I 3.12, II 4.80, III 3.36, IV 4.68; hind tibiae with outer dilations phylliform with one shallow and two deep emarginations, occupying 72% length of hind tibiae, width about twice width of inner dilations; inner dilations lanceolate, slightly shorter than outer, distally furnished with numerous small spine-like teeth along margins; inner margins of undilated portions of hind tibiae with a double row of small spine-like teeth, length hind tibiae 10.1; length outer dilations 7.20, width outer dilations 2.40: length inner dilations 6.00, width inner dilations 1 .20. Claspers (Fig. 18) with inner basal lobe strongly produced and rounded, shank with median lobe prominent and rather sharply curv- ing back onto shank, hook strongly curved and outer curvature with a small knob, capsule (Fig. 42) with median notch deeply rounded, dorsal prongs prominently acute and projecting medially; aedeagus with dorsal sac of conjunctiva as follows: proximal lobes sclerotized; median lobes present, left median lobe short with tip sclerotized and a longer thin lobe arising from its base, right median lobe longer and wider, anterior distal lobe small and closely appressed to wall of sac, 112 Entomologica Americana [Vol. 45 one distal lateral lobe large and spinose the other smaller and more blunt, all three distal lobes strongly sclerotized; total length 20.8. The description is of a male specimen from Yucatan, Mexico. In the large series of specimens examined there is a considerable amount of variation present. Individuals will vary from light brown to dark brown. The transverse fascia is typically a wide irregular band, but some specimens have the narrow band confined to the area of the veins. Two individuals lack the band. There is considerable variation in the width of the outer dilation on the hind tibiae. The labium varies in length from the anterior margin of the second sternum to the posterior margin of the fourth. Stal distinguished vexillatus from zonatus on the basis of its smaller hind tibial dilations and because there were only two emarginations or “teeth” on the margin of the outer dilation. At that time, zonatus was known only from Mexico and typical zonatus specimens from Mexico have a considerably wider dilation and usually possess one shallow and two deep emarginations. After examining a large series of specimens, I feel Stal’s type series of vexillatus represents the smaller extreme of variation present in zonatus. The color pattern and other external characters as well as male genitalia are identical for these two taxa. I am at this time formally synonymizing vexillatus with zonatus. Stal’s syntype series from Colombia consists of four specimens with the following labels: 2 males, Bogota (Lindig) ; 1 male, no data; 1 female, Remidios. These are deposited in the Stockholm Museum. The male specimen bearing the “typus” label is selected as Lectotype with an appropriate label attached. L. zonatus is closely related to impictipennis and neovexillatus, and may be distinguished from the former by its lack of serrate lateral margins on the pronotum and from the latter by the less steeply declivent pronotum, straighter lateral margins and humeral areas that are almost horizontal, not obliquely ascending. The aedeagus is dis- tinct from both impictipennis and neovexillatus in that the short left median lobe has a secondary appendage. The following characters, although less reliable are worth stating because of their slight vari- ability: zonatus is generally a larger species; the labium usually reaches to the fourth abdominal sternum; the pronotal disk has two large distinctly ovoid spots; the transverse fascia on the corium is usually a wide, brightly pigmented band, only five specimens out of 143 examined lacked this fascia. L. zonatus has been reported to feed on oranges, watermelons, dates, and cotton (Cockerell 1905). It is also known to transmit the heart rot of pomegranates ( Burgess and Hawkins 1945). 1969] Allen: Revision of the Genus Leptoglossus 113 Distribution : Lower California and southwestern United States, through Central America and into the northern half of South America. Material Examined: 51^,922. California’. Calexico, Canipole; San Jose del Cabo; San Diego; Pasadena; Angeles Bay, San Pedro; Imperial Valley; Santa Cruz; El Marmol. Arizona : Huachuca Mts.; Tucson; Phoenix; Yuma Co.; Patagonia; Whetstone Mts. Texas’. Davis Mts. MEXICO: Puebla; Antiguo; Morelos; Cuernavaca; Jalapa; Salle; Tepie; Sinaloa, Los Mochis; Oaxaca, Oax.; Tehuan- tepec; Mazatlan; Telmantepec; L. Chapata; Cyn Sapopa, Sonora; Chilpancingo; Vera Cruz; Hoege; Jacaia Hidalgo; Yucatan; Tesopaco; Baja California; Las Parras. GUATEMALA: Antigua; S. Gerocimo; Los Amates. HONDURAS: no locality. EL SALVADOR: San Miguel. NICARAGUA: La Calera; Managua. COSTA RICA: Palmer, Dept. Puntarenas; Turrialba; San Jose. PANAMA: Porto Bello; Boquete; La Campana; V. de Chiriqui; Bugaba; Cerro Zunil. VENEZUELA: Merida; Pts. Cavello. COLOMBIA: Villavicencio; Cali; Villa Vieja. ECUADOR: Paramba. PERU: Satipo; Monson Valley, Tingo Maria; Cumbase; Selipo; Tarapoto. BOLIVIA: Rur- renabaque Beni; Province Sara; Rio Cristal Mavu, N.E. Cochabamba. BRAZIL: Amazon; Olivenca. In Museum of Comparative Zoology (Harvard), California Academy of Sciences, Stockholm Museum, United States National Museum, American Museum of Natural His- tory, Iowa State University, British Museum (Natural History), Berlin Humboldt University Museum, J. A. Slater and R. C. Allen collec- tions. Leptoglossus neovexillatus new species Pronotum, scutellum, clavus and corium dark reddish brown with head above, outer areas of first antennal segments, pronotal calli, and marginal areas of humeri piceous; two large poorly defined ovoid spots on pronotal disk yellow, interspersed with numerous small dark brown to piceous maculae; apex scutellum, narrow irregular transverse fascia on corium confined to veins, maculae on inner and outer tibial dila- tions, and undilated portion of hind tibiae whitish yellow; head below and with three narrow stripes above, inner margins of first antennal segment, entire second and third with distal portion slightly darker and fourth segments with apex somewhat darker, entire fore and middle legs ochraceous to light tan; distal three fourths of hind femora and tibial dilations fuscous with bases and ventral portion of femora light reddish brown; femoral and tibial teeth and tubercles piceous; thoracic and abdominal venter tan to light brown with numerous small piceous spots frequently fusing to form larger maculae; connexivum fuscous and abdominal dorsum piceous with anterior third of con- 114 Entomologica Americana [Vol. 45 nexival segments and intersegmental sutures whitish yellow; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures uneven; scutellum transversely rugulose with few punctations ; venter rather thickly clothed with short appressed whitish pubescence and sparse short erect hairs; legs also with more dense longer and erect pale pilose hairs; dorsum with short semi-decumbent golden hairs. Head non-declivent with tylus and juga above level of antenni- ferous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 2.40, width head 2.16, inter- ocular distance 1.20, anteocular distance 1.32; pronotum steeply declivent, almost vertical, lateral margins sinuate, entire, humeral area moderately expanded, rounded and obliquely ascending, humeral angles small sub-acute, postero-lateral margins barely serrate, calli prominently elevated, area between calli only slightly higher, disk posteriorly with median longitudinal carina obsolete, length pronotum 1.92, width across humeri 3.60, width anterior margin pronotum 1.44; length scutellum 1.90, width scutellum 1.90; labium reaching posterior margin of second abdominal sternum, length labial segments I 2.16, II 1.92, III 1.20, IV 1.92; length antennal segments I 2.40, II 3.48, III 2.40, IV 4.08; hind tibiae with outer dilations phylliform with one shallow and two deep emarginations, dilations occupying 68% length of hind tibiae, width about twice width of inner dilations; inner dila- tions lanceolate, slightly shorter than outer, with numerous small spine-like teeth along margins; inner margin of undilated portion of hind tibiae with a double row of small spine-like teeth, length hind tibiae 7.80; length outer dilations 5.28; width outer dilations 1.92; length inner dilations 4.44, width inner dilations 0.84. Clasper and capsule as in zonatus ; aedeagus with dorsal sac of conjunctiva as follows: sclerotized proximal lobes present; medial lobes present with left lobe small and thin, right lobe longer and wider; sclerotized anterior distal lobe present and not appressed to wall of sac; one lateral distal lobe modified into a long spine, the other obso- lete, both sclerotized; total length 15.5. There is a considerable amount of color variation in the type series. Specimens vary from light brown to a dark reddish brown. The ir- regular transverse fascia on the corium can be a sharply outlined and brightly pigmented stripe or it can be completely lacking, as well as with all intermediate conditions. The yellow pronotal markings vary from two well defined spots to a broad uninterrupted band occupying the entire anterior pronotal area. Four specimens of the 5 1 examined have this yellow pronotal pattern reduced to small maculae. The labium varies in length from the anterior margin of second to anterior 1969] Allen: Revision of the Genus Leptoglossus 115 margin of fourth abdominal sternum (only seven species possess labia that pass the posterior margin of the second sternum). Specimens of neovexillatus have been misidentified as vexillatus because many of the males have a narrower and double “toothed” tibial dilation. L. neovexillatus may be distinguished from zonatus by its generally smaller size, the almost vertical declivity of the pronotum, the more strongly sinuate pronotal lateral margins, the more rounded and obliquely ascending humeral angles, the generally shorter labium and the dorsal sac of the conjunctiva with the left median lobe simple and without any secondary appendages, and one distal lateral lobe being obsolete. It may be distinguished from impictipennis by its non- serrate lateral pronotal margin and by the unequal sized lateral lobe on the dorsal sac of the conjunctiva. Holotype: S. Brazil: Corumba, Acc. No. 2966, March. In Carnegie Museum. Paratypes: PERU: IS, Bella Vista, San Martin, Dec. 8, 1946, “alt. 1500 ft.” (J. C. Pallister); 1 S , Vilcanota, no data (J. M. Bosq). BRAZIL: IS, Corupa (Hansa Humbolt), S. Cath., 1-1945 (A. Mailer); 1$, same except 1-1946; 1$, same except XI-1945; 12, Nova Teutonia, Santa Catarina, XII-7-1948 (F. Plaumann) ; 1 S , 1 2 , same except I-II-1949; IS, 12, Rio Grande, no data; IS, Parana, Tibagye (V. Konigswald); 1$, no data; 2 2, Loreto, Missiones, II- XII-1941 (Biraben-Bezzi) ; 12, San Ignacio Missiones, 8-XII-1941 (Biraben-Bezzi) ; 12, Corumba, Acc. No. 2966, March, “lowland”; 12, same except April, “highland”; IS, 12, Matto Crosso, 96-204 (Spencer Moore); 12, Bahia, Iguassu, Sv. Amaz. Exp., 22 Aug. (Roman); 1 S, Curitiba, Para., “on berries of Schinus terebinthi- folius ” III-1954 (N. L. H. Krauss); 12, Bilbao, 1904. BOLIVIA: IS, 12, 1904-311 (J. Steinbach); IS, Prov. Sara (Steinbach). PARAGUAY: 1 & , Horqueta, XII-26-33 (Al. Schulze) ; 1 2 , V. En- carracion, 2-III (F. Schade); IS, Rio Apa, Dto Carlos entre, Punta Apa y, Bellavista, 5-XII-1935 (Denier). URUGUAY: 12, Minas Geraes, no data; 12, Colonia Benitez Chaco, l-V-1939 (Denier). ARGENTINA: 2 S , 12, Province Salta 2500 m. (J. Steinbach S.V.) ; 1 2 , Province Salta, 2.05 ( J. Steinbach S.V. ) ; 1 S , 1 2 , Salta las Canas, III-1938; IS, Fab. Chaco, 22-X-1899 (S. Venturi); 12, S. Tome, Corrientes; 12, Guemes Salta, #45 (Martinez-Bezileg) ; 12, Salta; 1 S , Jujuy; 3 2 , Prov. of Buenos Aires ( J. Bosq) ; 1 S , Prov. Tucuman, 7-IX-1900; IS, Tucuman, 11-13-1918, Est. Expt. Agric. No. 1307; 1 2, Tucuman, no data; 1 2, Bob. de Misiones (J. M. Bosq); 1 2, no locality (J. M. Bosq). In the American Museum of Natural History, United States National Museum (J. C. Lutz collection), Hungarian National Museum, La Plata Museum, Stockholm Museum, Berlin 116 Entomologica Americana [Vol. 45 Humboldt University Museum, Carnegie Museum, British Museum (Natural History), California Academy of Sciences, Museum of Com- parative Zoology (Harvard), J. A. Slater and R. C. Allen collections. Leptoglossus impictipennis Stal Leptoglossus impictipennis Stal, 1870:163. — Lethierry and Severin, 1894:47. Pronotum, clavus and corium rusty reddish brown with scutellum fuscous; head, pronotal calli and margins of humeri piceous; two in- distinct and partially separated ovoid spots on pronotal disk ochra- ceous and interspersed with numerous small piceous spots; apex scutellum and maculae on inner tibial dilations whitish yellow; head laterally and with three narrow stripes above, all antennal segments with the fourth segments somewhat lighter, and all legs light reddish brown; dorsum of hind femora distally and tibial dilations fuscous with teeth and tubercles piceous; thoracic and abdominal venter dark reddish brown with numerous small piceous spots frequently fusing to form larger maculae; connexivum fuscous with anterior fourth of each segment ochraceous; abdominal dorsum piceous; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures largely smooth; scutellum transversely rugu- lose with few punctures; body above and below with moderate amounts of short pale erect and semi-decumbent hairs; thoracic and abdominal mid-venter, legs, head dorsally and anterior portion of pronotum with more dense long pilose hairs. Head porrect with tylus and juga raised above level of antenni- ferous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 2.04, width head 1.92, interocular distance 1.08, anteocular distance 1.20; pronotum abruptly declivent with lateral margins almost straight and serrate caudally, humeral areas not expanded, humeral angles acute and obliquely ascending, postero-lateral margins serrate, calli prominently elevated with area between calli only slightly higher, disk posteriorly with median longi- tudinal carina obsolete, length pronotum 2.24, width across humeri 4.96, width anterior margin pronotum 1.44; length scutellum 1.80, width scutellum 1.92; labium reaching to anterior margin of fourth abdominal sternum, length labial segments I 1.92, II 1.92, III 1.20, IV 2.16; length antennal segments I 2.04, II 3.24, III 2.16, IV 3.48; hind tibiae with outer dilations phylliform with one shallow and two deep emarginations, dilations occupying 77% length of hind tibiae, width about twice width of inner dilations; inner dilations lanceolate slightly shorter than outer, with numerous small spine-like teeth along 1969] Allen: Revision of the Genus Leptoglossus 117 margins; inner margins of undilated portions of hind tibiae with a double row of small spine-like teeth, length hind tibiae 7.44; length outer dilations 5.76, width outer dilations 1.80; length inner dilations 5.04, width inner dilations 0.72. Claspers (Fig. 19) with median basal lobe very small, rounded, shank with median lobe prominent, rounded hook gradually curved; capsule (Fig. 43) with median notch deep, dorsal prongs prominent, rounded, and wide; aedeagus with dorsal sac of conjunctiva as follows: proximal lobes present and sclerotized; medial lobes present, right lobe large and wide, left lobe smaller and thin; sclerotized anterior distal lobe not closely appressed to wall of sac; lateral distal lobes present, of same size and shape, both sclerotized; total length 14.4. Stal described impictipennis on the following characters: shorter labium and antennae, and lack of the transverse fascia. Among Stabs four male syntypes there are three specimens that I feel belong to L. zonatus. They have similarly shaped pronotum and genitalia as do typical zonatus specimens. The remaining male syntype (labeled “Typus”) is here selected as Lectotype and an appropriate label attached. This specimen differs from the others in the type series and zonatus by the following characters : lateral margins serrate and almost straight; humeral areas not rounded but forming an acute humeral angle; left medial lobe on dorsal sac of conjunctiva a simple, long, and thin unsclerotized appendage. Additional characters which may be less reliable are as follows: the pronotal markings are not such clearly defined spots as in zonatus and are only partially separated medially; the basal median lobe on the clasper is lower and the median lobe on the shank is not so prominent. I have assigned to this species 13 other specimens that have impictipennis characters, but differ in having wider tibial dilations, a clasper similar to zonatus and a capsule with dorsal prongs project- ing vertically. There is also a considerable amount of short incumbent pubescence on the posterior portion of the corium, this is barely evident on the type specimen. Two females in this series have the entire anterior half of the pronotum yellow, and without the small black spots. Until a larger series of impictipennis- like individuals can be assembled, I do not feel justified in assigning these specimens to a new species. All of the impictipennis specimens differ from neovexillatus by the presence of serrate lateral margins on the pronotum and the lateral distal lobes on the dorsal sac of the conjunctiva are both spinose and of the same size. In neovexillatus only one lateral distal lobe is spinose, the other is obsolete. 118 Entomologica Americana [Vol. 45 Distribution : Brazil, Bolivia, British Guiana, and Colombia. Material Examined: Holotype. S. Colombia: Bogota (Lindig). In Stockholm Museum. BRITISH GUIANA: 1 $ , no locality, B.M. 1948-60 (Bartlett). BOLIVIA: 3 $, Province Sara (Steinbach). BRAZIL: 2 2, Chapada, Acc. No. 2966, March; 12, same except June; 2 2 , same except Nov., 1 S , same except April; 1 2 , Matto Crosso (Zobrys & Wolter); 12, Corumba, Matto Grosso, 1950 (H. G. Barber); 1 2, Maracaju, M. Grosso, Feb. 1937 (R. C. Shannon); 1 $ , Matto Grosso (Kolowsky). In Museum of Comparative Zoology (Harvard), British Museum (Natural History), United States Na- tional Museum, La Plata Museum, Berlin Humboldt University Mu- seum, and Carnegie Museum. Leptoglossus concolor (Walker) Anisoscelis concolor Walker, 1871:128. Leptoglossus concolor Distant, 1881:124. — Lethierry and Severin, 1894:47.— Distant, 1901b:430. Pronotum, scutellum, clavus and corium dark chestnut brown with head, outer margins of first antennal segments, pronotal calli, and margins of humeri piceous; fourth antennal segments, apex scutellum, wide irregular transverse fascia on corium, maculae on inner tibial dilations and undilated portion of hind tibiae whitish yellow; head laterally and with three narrow stripes above, inner margins of first segment and segments two and three of antennae, fore and middle legs except distal fourth of femora above, basal third and ventral area of hind femora reddish tan; distal portion of all femora above and hind tibial dilations fuscous with teeth and tubercles piceous; thoracic and abdominal venter light reddish tan mottled with numerous small piceous spots frequently fusing to form larger maculae; connexivum and abdominal dorsum piceous with anterior fourth of each connexival segment and narrow intersegmental sutures on dorsum ochraceous; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures smooth; scutellum transversely rugulose with numerous small punctures; venter with moderate amounts of short appressed whitish pubescence, and sparse short erect hairs; legs and scutellum also with more dense longer pilose hairs; dorsum with short semi-decumbent golden hairs and pronotum anteriorly in area of calli with thicker dark almost spinose hairs. Head non-declivent with tylus and juga above level of antennif- erous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 3.00, width head 2.64, interocular 1969] Allen: Revision of the Genus Leptoglossus 119 distance 1.32, anteocular distance 1.68; pronotum abruptly declivent, lateral margins slightly sinuate, entire, humeral areas moderately ex- panded, barely ascending, humeral angles sub-acute, postero-lateral margins serrate, calli prominently elevated with surface rough, area between calli slightly higher, disk posteriorly with median longitudinal carina obsolete, length pronotum 3.68, width across humeri 6.72, width anterior margin pronotum 1.80; length scutellum 2.64, width scutellum 2.52; labium reaching to middle of fourth abdominal ster- num, length labial segments I 3.72, II 3.60, III 2.04, IV 4.80; length antennal segments I 3.12, II 5.40, III 3.84, IV 6.36; hind tibiae with outer dilations phylliform with one shallow and two deep emargina- tions, occupying 77% length of hind tibiae, width about two and one half times width of inner dilations; inner dilations lanceolate, slightly shorter than outer, and distally furnished with numerous small spine- like teeth, inner margins of undilated portions of hind tibiae with few small spine-like teeth arranged in a double row, length hind tibiae 11.3; length outer dilations 8.64, width outer dilations 3.00; length inner dilations 7.44, width inner dilations 1.20. Claspers (Fig. 22) with inner basal lobe low and rounded; shank with median lobe prominent, gradually curving back onto shank, hook strongly curved; capsule (Fig. 44) with median notch deep, rectangular, no dorsal prongs; aedeagus with dorsal sac of conjunctiva as follows: proximal lobes present and sclerotized; median lobes present, left lobe short blunt and simple, right lobe longer and wide; sclerotized anterior distal lobe closely appressed to wall of sac; one distal lateral lobe sclerotized, large and spinose, the other lobe absent; total length 21.0. The above description is of a male specimen from Chuminopolis, Yucatan, Mexico. A generally uniform color pattern is present in the series before me. Lighter and darker individuals do occur, also there is variation in the appearance of the transverse fascia on the corium: of the 83 specimens examined, 22 have the fascia narrow, confined to the veins; two, including the type, have the fascia completely lacking; the remain- ing specimens have a wide and brightly pigmented fascia. The labium extension varies from the anterior portion of the fourth to the anterior portion of the sixth abdominal sternum. Fifteen specimens from the Caribbean have, on the posterior portion of the pronotal disk, a large yellow rectangular area, and the posterior marginal area of the pro- notum is also yellow. This is quite unlike any other specimens of concolor , all of which have a unicolorous pronotum. Previous to this revision, all specimens with phylliform tibial dila- tions and a pronotum without yellow spots had been referred to as 120 Entomologica Americana [Vol. 45 stigma. After dissecting the male genitalia of a number of specimens, it was apparent that two distinct species were represented. The type of stigma was described from Surinam. I have referred to stigma all specimens with dorsal prongs on the genital capsule, and to concolor the specimens with a rectangular notch on the capsule. Externally, these two species are extremely close and the only characters, other than male genitalia, which distinguish concolor from stigma are the roughness of the pronotal calli, the thicker spinulose hairs around the calli, and the less steeply declivent pronotum. The lack of definite rounded pronotal spots and the rectangular median notch on the genital capsule will separate concolor from other mem- bers of the stigma-group. Walker described his type as new based on the fact that it lacked the transverse fascia on the corium. This character is not absolutely diagnostic in this species. The type did not differ in any other respect from specimens with a transverse fascia. Mann (1969) states that adult insects questionably identified as concolor were found on prickly pear. Distribution : Mexico, Central America, and Greater Antilles. Material Examined: Holotype: 2, 58.135, Oajoca, Mexico. In British Museum (Natural History). 40 <£,42 2. MEXICO: Tampico; Catemaca; Barbarita; Aguazarca; Yucatan; Tamaulipas; Ocotlan; Chuminopolis; Oaxaca, Tehuantepec; Navarrete. GUATEMALA: Yepocapa; Morales; Antigua; Vizcaya; Panzos. BRITISH HON- DURAS: Punta Gorda; San Antonio; Tegucigalpa. COSTA RICA: Hamburg Farm; Piedras Negras. PANAMA: Chiriqui; Colon; Barro Colorado Island. CUBA: Soledad; San Juliano; San Jose; Prov. St. Clara; Santiago de las Vegas; San Bias; San Vincente Pinar del Rio; Upper Yara Valley; Camaguey. HAITI: St. Marc. DOMINICAN REPUBLIC: Pto. Plata; Sanchez. PUERTO RICO: Barranquitas; Jayuya; Mayaguez. VIRGIN ISLANDS: St. Croix. In Berlin Hum- boldt University Museum, American Museum of Natural History, Mu- seum of Comparative Zoology (Harvard), United States National Museum, Stockholm Museum, Hungarian National Museum, Cali- fornia Academy of Sciences, J. A. Slater and R. C. Allen collections. Leptoglossus stigma (Herbst) Cimex stigma Herbst, 1784:258, pi. 39B, fig. 1. Hypselonotus scriptus Hahn, 1826:5. Anisoscelis scripta Westwood, 1842:16. Anisoscelis indocta Westwood, 1842:16. — Distant, 1901a:334. Anisoscelis minor Dallas, 1852:452. 1969] Allen: Revision of the Genus Leptoglossus 121 Theognis scriptus Mayr, 1866:101. Leptoglossus stigma Stal, 1870:163. — Lethierry and Severin, 1894: 49. Pronotum, scutellum, clavus and corium dark reddish brown with head, outer area of first antennal segment, pronotal calli and margins of humeri piceous; veins of clavus and corium wine red; apex scu- tellum, wide irregular transverse fascia on corium, and maculae on inner tibial dilations whitish yellow; head laterally and with three narrow stripes above, inner margins of first segment and segment two and three of antennae, fore and middle legs and undilated por- tion of hind tibiae dark tan to light brown; fourth antennal segments with base and apex ochraceous and medial portion somewhat darker; basal fourth and ventral area of hind femora light reddish brown with femora above and hind tibial dilation fuscous; femoral and tibial teeth and tubercles piceous; thoracic and abdominal venter varie- gated rusty reddish brown and light brown with numerous small piceous spots frequently fusing to form irregular maculae; connexivum and abdominal dorsum piceous with anterior fourth of each con- nexival segment and narrow areas of intersegmental sutures on dorsum ochraceous; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate with surface above punctures largely smooth; scutellum transversely rugulose with numerous small punctures; venter with moderate amounts of short appressed and erect pale hairs and dorsum with short erect and semi-decumbent yellowish hairs; anterior pronotum, scutellum and legs with more dense longer and erect pilose hairs. Head non-declivent with tylus and juga above level of antennif- erous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 3.00, width head 2.28, interocular distance 1.20, anteocular distance 1.56; pronotum abruptly declivent, lateral margins strongly sinuate and entire, humeral areas expanded and obliquely ascending, humeral angles sub-acute, postero-lateral margins serrate, calli prominently elevated, area between calli slightly higher, without tubercles, disk posteriorly with median longitudinal carina obsolete, length pronotum 3.04, width across humeri 5.92, width anterior margin pronotum 1.68; length scutellum 2.04, width scutellum 2.16; labium reaching to posterior margins of third abdomi- nal sternum, length labial segments I 3.00, II 2.88, III 1.32, IV 3.12; length antennal segments I 2.52, II 4.44, III 3.12, IV 7.20; hind tibiae with outer dilations phylliform with one shallow and two deep emar- ginations, occupying 79% length of hind tibiae, width about twice width of inner dilations; inner dilations lanceolate, slightly shorter than outer 122 Entomologica Americana [Vol. 45 and apically furnished with few small spine-like teeth; inner margins of undilated portions of hind tibiae with few small spine-like teeth arranged in a double row, length hind tibiae 9.84; length outer dila- tions 7.80, width outer dilations 3.24, length inner dilations 6.36, width inner dilations 1.44. Claspers (Fig. 21) with basal median lobe low rounded and hardly prominent, gradually curving back onto shank, hook strongly curved; capsule with median notch deep and rounded with dorsal prongs prominent and acute; aedeagus with dorsal sac of conjunctiva as follows: sclerotized proximal lobes present; left median lobe long and wide, right median lobe short, blunt and simple; sclerotized anterior distal lobe closely appressed to wall of sac; one distal lateral lobe large, spinose and sclerotized the other lobe absent; total length 17.6. The above description is of a male specimen from Santarem, Brazil. Three specimens of the 38 examined lack the transverse fascia on the corium and all specimens have a unicolorous pronotum and are without any yellowish pigments. L. stigma may be distinguished from concolor by the smooth pronotal calli, pale pilose hairs around the area of the calli, and the more steeply declivent pronotum. The males of stigma may be readily distinguished by the distinctive dorsal prongs on the genital capsule. The females of the two species are difficult to separate, and will probably continue to be confused with one another. Fortunately, there seems to be a geographic separation, with stigma present chiefly east of the Andes in northern South America, while concolor is present in Central America, Mexico, and the southern United States. Both concolor and stigma completely lack one distal lateral lobe on the dorsal sac of the conjunctiva and have the right median lobe short. Also they do not have any yellow ovoid spots on the anterior portion of the pronotal disk. These characters will serve to distinguish concolor and stigma from the other members of the stigma- group. Barber (1939) and Wolcott (1948) list guava fruit as the normal host plant for stigma in Puerto Rico; stigma also feeds on achiote ( Bixa orellana L.) (Wolcott 1948), cashew ( Anacardium ) (Barber and Bruner 1947). Hussey lists stigma as taken on lychee in Florida. The above host plant records more than likely pertain to concolor (see discussion under concolor). Heidemann (1910) used the trinomen L. stigma var. minor Dallas for brevirostris (see discussion under brevirostris) . Distribution : Surinam, Ecuador, Brazil and Paraguay. 1969] Allen: Revision of the Genus Leptoglossus 123 Material Examined : SURINAM: 1$, St. Laurent (C. Heller S.V.). ECUADOR: 18, Rio Napo, 15-35 (Rolf Blomberg); 18, 1 2, no locality, 1942; 28 , Balzapamba (R. Haensch S.). BRAZIL: 1 2, Caviuna Parana, XI- 1945; 1 2, same except 11-1946 (A. Mailer); 12, Amazon sup., Olivenca; 12, Teffe, X-24, F 6160, Ace. 33591 (H. Bassler); 12, Amazonas, Manana, Uypiranga, Rio Negro, 14 km. from Manaus 81 m., X-1941 (Alberto Rabaut); 12, Amazon sup., Teffe; 2 2, Brasilien; 1 8 , 12, Amazon (Deyrolle); 1 2, Maran- guate Mts. (Mann); 1 8, 12, Manaos, Amazon (Roman); 18, Rio de Janeiro, 1930; 18, Santarem, Ace. 2966; 1 8, same except May 1919, Ace. 6324; 3^,22, Villarica, XI (F. Schade) ; 2 2 , Amer. mer., no data. PARAGUAY: 28 , Horqueta, 45 m. E. of Paraguay Riv., III-23-1933 (Alberto Schulze); 1 8 , same except III-24- 1933; 1 2, no locality (Vezenyi); 18, Rio Yguagu, Mar. 10 (Donald Wees). In the United States National Museum, Hungarian National Museum, Museum of Comparative Zoology (Harvard), American Museum of Natural History, Stockholm Museum, Berlin Humboldt University Museum, Carnegie Museum, J. A. Slater and R. C. Allen collections. Leptoglossus oppositus (Say) Anisoscelis oppositus Say, 1832:12. Anisoscelis tibialis Herrich-Schaffer, 1842 : 12. Leptoglossus oppositus Stal, 1870:163-4. — Lethierry and Severin, 1894:48.— Van Duzee, 1917:89.— Gibson, 1917:71.— Blatchley, 1926:223-4, fig. 44.— Deay, 1928:378.— Torre-Bueno, 1941: 50. — Froeschner, 1942:594-5. — Drew and Schaefer, 1963:114. Theognis oppositus Hussey, 1953:31. Dorsum and appendages more or less uniformly dark reddish brown; whitish yellow transverse fascia on the corium reduced and confined to medial vein above where crossvein begins; venter light brown mottled with numerous small piceous spots; pronotum with all lateral margins entire, non-serrate; humeri rounded with humeral angles obtuse; labium extending to at least third abdominal sternum; hind tibial dilations phylliform. Claspers and capsule similar to zonatus. The only difference in the aedeagus is that oppositus has the left median lobe long and bifid at the tip, without a longer and thin secondary appendage as in zonatus. In the specimens examined, there are four individuals from Texas that have two very faint and poorly defined yellowish areas and small piceous spots on the anterior portion of the pronotum. One specimen, also from Texas, has the medio-cubital crossvein yellow. None of the 124 Entomologica Americana [Vol. 45 individuals examined had a complete transverse fascia on the corium, although Blatchley (1926) states that one specimen from Florida had a complete fascia. The form of the genitalia places oppositus in the stigma-group, closest to zonatus. The lack of a serrate postero-lateral margin on the pronotum will distinguish oppositus from all members of this group. Additional characters that will separate oppositus from zonatus are: the lack of a complete transverse fascia on the corium; the lack of clearly defined pronotal spots; and the previously mentioned aedeagal difference. L. oppositus has been reported in the economic literature as attacking essentially the same crops and plants as phyllopus. The life history and habits were reported by Chittenden (1902, 1925). Mc- Cullough (1968) reported the acid and aldehyde compounds in the scent fluid. Distribution : Eastern United States from Florida to New York, west to Iowa and Wisconsin, southwest to Arizona, Texas and into Mexico. Material Examined’. 49 <2, 59$, UNITED STATES: Alabama, Arizona, Arkansas, Florida, Georgia, Illinois, Indiana, Iowa, Louisi- ana, Maryland, Mississippi, Missouri, New Jersey, New Mexico, North Carolina, Ohio, Oklahoma, Tennessee, Texas, Virginia, Washington, D.C., Wisconsin. In American Museum of Natural History, Museum of Comparative Zoology (Harvard), Stockholm Museum, Iowa State University, United States National Museum, University of California (Davis), Chicago Natural History Museum, J. A. Slater and R. C. Allen collections. Leptoglossus lonchoides new species Pronotum, scutellum, clavus and corium light reddish brown, head, calli and a few small spots on disk of pronotum piceous; two large, separated but poorly defined ovoid spots on pronotal disk, apex scutellum, and medio-cubital cross veins dark ochraceous; fourth an- tennal segments and maculae on inner tibial dilations whitish yellow; head laterally and with three narrow stripes above, inner marginal areas of first antennal segment and basal portions of segments two and three, all legs except femora above tan to light brown; outer marginal area of first antennal segment and distal portion of segments two and three, femora above and hind tibial dilations fuscous with teeth and tubercles piceous; thoracic and abdominal venter ochraceous to light brown with numerous small piceous spots frequently fusing to form larger maculae; connexivum and abdominal dorsum piceous with 1969] Allen: Revision of the Genus Leptoglossus 125 anterior margins of connexival segments and narrow area around intersegmental sutures on dorsum yellow; membrane uniformly dark; pronotum, clavus and corium coarsely and closely punctate with sur- face above punctures uneven; scutellum transversely rugulose with numerous punctures; body with moderate amounts of short appressed and sparse short erect hairs; head behind eyes anterior portion of pronotum, scutellum and legs with more dense longer pilose hairs. Head non-declivent with tylus and juga above level of antenni- ferous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 2.40, width head 2.28, inter- ocular distance 1.32, anteocular distance 1.32; pronotum moderately declivent with lateral margins straight and entire, humeral areas not expanded or ascending, the humeral angles barely prominent and acute, postero-lateral margins entire, calli prominently elevated, area between calli only slightly higher, disk caudally with median longi- tudinal carina obsolete, length pronotum 3.36, width across humeri 5.12, width anterior margin pronotum 1.56; length scutellum 2.40, width scutellum 2.28; labium reaching to posterior margin of meta- sternum, length labial segments I 2.16, II 2.16, III 1.08, IV 2.16; length antennal segments I 2.40, II 3.84, III 2.76, IV 3.84; hind tibiae with outer dilations lanceolate with only one very shallow emargination present, occupying 65% length of hind tibiae, width slightly less than twice width of inner dilations; inner dilations lance- olate, shorter than outer with a few small spine-like teeth along distal half of margins; inner margins of undilated portion of hind tibiae with a double row of small spine-like teeth, length hind tibiae 9.24; length outer dilations 6.00; width outer dilations 1.44; length inner dilations 4.68; width inner dilations 0.84. Claspers (Fig. 24) with inner basal lobe large and rounded; shank thick without a median lobe, hook thick and strongly curved; capsule with median notch deep and rounded dorsal prongs prominent and acute; aedeagus with dorsal sac of conjunctiva as follows: sclerotized proximal lobes present; median lobes present with right lobe long and wide, left lobe with tip sclerotized somewhat shorter and not as wide as right lobe and with a secondary appendage arising from approximately middle of lobe; large sclerotized anterior distal lobe present and closely appressed to wall of sac; spinose distal lateral lobes present and of same size and shape, both sclerotized; total length 17.8. Holotype: $. Brazil: Nova Teutonia, Santa Catarina, VI. 28, 1943 (F. Plaumann). In United States National Museum, type no. 70226. 126 Entomologica Americana [Vol. 45 Paratypes: BRAZIL: 12, Nova Teutonia, Santa Catarina, IX- 9-1950 (F. Plaumann); 12, same except IV-29-1948; 1 $, same except 11-8-1948; 1 $, Amazonas, Manaus, Uypiranga Rio Negro, 14 km. from Manaua, 81 m., XI- 1 5-XII-15-1941 (August Rabaut). PERU: 1 2, Rio Santiago, XI-22. 24, F. 6012. In American Museum of Natural History and the United States National Museum. The type series exhibits some important color variations. One specimen has the pronotal disk entirely dark ochraceous. Two indi- viduals have a complete irregular transverse fascia on the corium which is confined primarily to the veins. Two specimens lack the fascia completely. The tibial dilation is relatively constant in shape, although three individuals have completely lanceolate outer dilations, lacking even a shallow emargination. Labial length varies from the posterior margin of the metasternum to the middle of the third abdominal sternum. L. lonchoides is a very distinctive species externally. To some extent the tibial dilation (Fig. 61) resembles that of lineosus ; the steepness of the pronotum is of the same degree as in clypealis; the unexpanded humeral areas and the straight non-sinuate lateral margins are close to the condition found in crassicornis species from Argentina. Despite the external dissimilarity, the genitalia of lonchoides is of the same type as that found in the stigma-group, with the clasper reminis- cent of that in concolor, the capsule identical to that of zonatus, and the aedeagus close to oppositus. The lanceolate tibial dilations and the non-serrate postero-lateral margins on the pronotum will serve to distinguish lonchoides from all other members of the stigma-group. Leptoglossus humeralis new species Pronotum, scutellum, clavus and corium dark brown with head above, outer areas of first antennal segments and pronotal humeral areas piceous; veins of clavus and corium dark tan; basal two thirds of second and third segments, entire fourth antennal segment, apex scutellum, and maculae on inner tibial dilations pale yellow to ochra- ceous; head below and with three narrow stripes above, inner margins of first antennal segment, all legs except femora above tan to light brown; distal third of antennal segments two and three, and femora above fuscous with hind tibial dilations dark reddish brown; femoral teeth and tubercles piceous; thoracic and abdominal venter tan to light brown with numerous small piceous maculae frequently fusing to form larger maculae; connexivum and abdominal dorsum largely piceous with some dark reddish brown areas; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate with 1969] Allen: Revision of the Genus Leptoglossus 127 surface above punctures largely smooth; scutellum transversely rugu- lose with numerous punctures; head above, anterior half of pronotum, scutellum and legs with dense long and erect pale pilose hairs; re- mainder of dorsum, and venter with short appressed and semi- decumbent hairs. Head porrect with tylus and juga above level of antenniferous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 3.72, width head 2.76, interocular dis- tance 1.44, anteocular distance 2.16; pronotum steeply declivent almost vertical, lateral margins serrate caudally and strongly sinuate, humeral areas produced as broad long and tapering lateral projections and obliquely ascending, humeral angles sub-acuminate, postero- lateral margins serrate, calli barely elevated, area between calli only slightly higher, disk posteriorly with median longitudinal carina obso- lete, length pronotum 4.80, width across humeri 12.4, width anterior margin pronotum 1.92; length scutellum 3.36, width scutellum 3.60; labium reaching to posterior margin of fourth abdominal sternum, length labial segments I 4.56, II 4.20, III 2.64, IV 5.40; length an- tennal segments I 3.24, II 5.76, III 4.20, IV 7.08; hind tibiae with outer dilations phylliform with one shallow and two deep emargina- tions, occupying 84% length of hind tibiae, width about twice width of inner dilations; inner dilations lanceolate, considerably shorter than outer dilations, furnished with two small spine-like teeth on distal margin; undilated portion of hind tibiae without teeth, length hind tibiae 12.3; length outer dilations 10.4, width outer dilations 3.68; length inner dilations 7.04, width inner dilations 1.76; total length 26.0. Holotype : 9. BRITISH GUIANA: Kartabo, VIII- 1925 (S. H. Williams). In Carnegie Museum. Paratype: FRENCH GUIANA: 1$, Cayenne (Wm. Schaus). In United States National Museum. L. humeralis has the humeri expanded in a fashion identical with those on pallidivenosus , but considerably longer, the width across humeri is 5.5 to 6.0 times the width of the anterior pronotal margin, whereas in pallidivenosus it is 4.5 to 4.8. The hind tibial dilation in humeralis is similar to that in ingens and macrophyllus with the inner dilation much shorter than the outer. Since there were no males available, I can only tentatively place humeralis in the stigma- group based on the above similarities. The large size, prominently expanded humeri, long labium and distinctive tibial dilations as well as the serrate pronotal lateral margin and the lack of pale areas anywhere on the dorsum, will distinguish humeralis from any other Leptoglossus species. 128 Entomologica Americana [Vol. 45 Leptoglossus pallidivenosus new species (Fig. 4) Pronotum, scutellum, clavus and corium dark reddish brown with head, outer areas of first antennal segment and humeral areas piceous; basal half of fourth antennal segment, posterior margin of pronotum, apex scutellum, all veins on apical half of corium and maculae on inner tibial dilation whitish yellow; head laterally and with three narrow stripes above, inner margin of first antennal seg- ment and segments two and three and apex of fourth segment, fore and middle legs and undilated portion of hind tibiae tan to light brown; all femora above fuscous with basal and ventral portion of hind femora and tibial dilations dark reddish brown; femoral and tibial teeth and tubercles piceous; abdominal venter tan with numerous small piceous spots; connexivum fuscous with anterior margin of each segment ochraceous; abdominal dorsum piceous; membrane uniformly dark; pronotum, clavus and corium finely and closely punctate, surface above punctures largely even; scutellum transversely rugulose with numerous small punctures; head, anterior portion of pronotum, scu- tellum, legs, thoracic and abdominal mid-venter with dense long erect pale pilose hairs; remainder of venter and dorsum with dense short appressed and semi-decumbent yellowish hairs; scutellum with lateral margins and a narrow median longitudinal area densely clothed with short appressed golden pubescence. Head porrect with tylus and juga above level of antenniferous tubercles, tylus blunt, slightly exceeding juga and forming a rounded elevated ridge, length head 2.76, width head 2.52, interocular distance 1.32, anteocular distance 1.56; pronotum abruptly declivent with lateral margins serrate caudally and strongly sinuate, humeral areas broadly expanded and obliquely ascending with humeral angles sub- acuminate, postero-lateral margins dentate, calli prominently elevated, area between calli only slightly higher, disk posteriorly with median longitudinal carina obsolete, length pronotum 3.68, width across humeri 8.16, width anterior margin pronotum 1.68; length scutellum 2.52, width scutellum 2.52; labium reaching to middle of third ab- dominal sternum, length labial segments I 2.88, II 2.88, III 1.44, IV 3.24; length antennal segments I 3.00, II 5.04, III 3.84, IV 5.88; hind tibiae with outer dilations phylliform and with three deep emar- ginations, occupying 72% length of hind tibiae, width about twice width of inner dilations; inner dilations lanceolate, slightly shorter than outer dilations, with numerous small spine-like teeth along margins; inner areas of undilated portion of hind tibiae with few small spine- like teeth in a double row, length hind tibiae 11.2; length outer dila- 1969] Allen: Revision of the Genus Leptoglossus 129 tions 8.04, width outer dilations 2.52; length inner dilations 6.96, width inner dilations 1.20. Claspers (Fig. 23) with inner basal lobe very small and rounded; shank without a median lobe, hook strongly curved and short; capsule with median notch deep and rounded, lateral margin of notch sinuate and forming a wide angle, no dorsal prongs; aedeagus with dorsal sac of conjunctiva as follows: sclerotized proximal lobes present; right median lobe slightly smaller than left lobe and unequally bifid, left median lobe large and simple; sclerotized anterior distal lobe present; sclerotized distal lobes present, one lobe spinose, the other smaller and acute; total length 20.0. Holotype: S. PANAMA: Canal Zone, Barro Colorado Island, 2 May 1956, “taken at light” (Carl W. & Marian E. Rettenmeyer). In University of Kansas (Snow collection). Paratypes: PANAMA: 1 S , Canal Zone, Barro Colorado Island, 2 May 1956, “taken at light” (Carl W. & Marian E. Rettenmeyer); 1 2, same except June 14, 1956; 1 S , same except Apr. 28, 1956; 1 S , same except 4 April 1956; 2S, Canal Zone, Barro Colorado Island, 14-11-1955 (C. W. Rettenmeyer); IS, same except 1 6-III- 1955; 1 S, same except IX-X-40, “collected at light” (Jas Zetek); 12, Barro Colorado Island, 3-14-37 (S. W. Frost). In United States National Museum and University of Kansas (Snow collection). There is some color variation in the type series : only the holotype and one other male specimen have all the veins on the apical half of the corium completely pale; the remaining individuals have some portions of the veins in this area slightly darker and not as strongly contrasting with the corium; one specimen has the pale areas confined to the usual portion of the veins that are included in the transverse fascia. This species has a number of characters which relate it to the stigma-g roup. The ground color of the dorsum, the fact that pale pronotal markings are absent on the disc, and in some respects the shape of the humeral expansions are characters shared by stigma and concolor. The serrate lateral margins of the pronotum are similar to impictipennis , as well as the claspers — both pallidivenosus and im- pictipennis have the inner basal lobe low and rounded and both lack a well produced medial lobe on the shank. The shape of the dorsal margin of the genital capsule is in some ways like that of ingens and macrophyllus except that in the latter two species, dorsal prongs are present. L. pallidivenosus may be distinguished from other members of the stigma-g roup by the following combination of characters : lack of pale markings on the pronotal disc, the posterior edge of the pronotum 130 Entomologica Americana [Vol. 45 pale yellow, all margins of the pronotum serrate, the labium not extending past the third abdominal sternum, and the genital capsule having a wide median notch without dorsal prongs. Leptoglossus corculus (Say) Anisoscelis corculus Say, 1832:12. — Leconte, 1859:326. Theognis excellens Mayr, 1865:434. Leptoglossus corculus Stal, 1870:165. — Lethierry and Severin, 1894: 47.— Van Duzee, 1917:88-9.— Gibson, 1917:70, 71.— Parsh- ley, 1923:747.— Blatchley, 1926:222-3.— Deay, 1928:377-8.— Torre-Bueno, 1941:49. — Froeschner, 1942:594, 599. Theognis corculus Hussey, 1953:30. Ground color varying from light to dark brown with pronotum and venter prominently mottled with small piceous spots, pale yellow ir- regular transverse fascia on corium narrow, confined to the veins, sometimes obsolete or completely absent; pronotum with all margins entire, broadly and obtusely rounded; hind tibial dilations lanceolate with outer dilations longer than inner dilations. Clasper (Fig. 26) with inner basal lobe low and rounded, shank with a median lobe, hook strongly curved with a small knob on outer margin; capsule (Fig. 46) deep and rounded, no dorsal prongs; aedeagus with dorsal sac of conjunctiva as follows: proximal lobes present barely sclerotized; a pair of median lobes present, of similar size and shape, small sclerotized anterior distal lobe present; sclero- tized lateral distal lobes present, one spinose, one blunt. L. corculus and occidental is are strikingly similar in appearance, but the outer tibial dilations (Fig. 57) of corculus occupy a greater percentage of the length of the tibiae (85-95%) and are distinctively longer than the inner dilations; in occidentalis (Fig. 58) both the inner and outer dilations are approximately the same length and occupy 66-72% the length of the hind tibiae. The differences between the claspers are: in corculus there are prominent inner basal and medial lobes on the shank; in occidentalis both lobes are absent. The median notch of the capsule in occidentalis is more rectangular with the bottom flat, not rounded; in corculus the notch is “V” shaped, deeper and with the bottom of the notch rounded. Distribution : L. corculus has an eastern United States distribution, being found from New York south to Florida, west to Missouri, and southwest to Texas. Deay (1928) lists a single specimen without a locality from Kansas. Published records include New Mexico, Ari- zona, Colorado, and California (Van Duzee 1917; Blatchley 1926), but in all the specimens received from the various museums in the 1969] Allen: Revision of the Genus Leptoglossus 131 United States, I have not seen a single specimen from further west than Texas. It is possible that the Van Duzee records were based on Occident alis specimens. Material Examined : 45$, 53$, UNITED STATES: Arkansas; Alabama; Florida; Georgia; Maryland; Mississippi; New Jersey; North Carolina; Ohio; Pennsylvania; Texas; Virginia; Washington, D.C. In United States National Museum, Museum of Comparative Zoology (Harvard), American Museum of Natural History, Chicago Natural History Museum, Iowa State University, University of Kansas, J. A. Slater, P. D. Ashlock, and R. C. Allen collections. Leptoglossus occidentalis Heidemann Leptoglossus occidentalis Heidemann, 1910: 196-7, pi. 8, fig. 2. — Van Duzee, 1917:89.— Gibson, 1917:70, 71.— Torre-Bueno, 1941: 49. Theognis occidentalis Hussey, 1953:29-30, 31. The ground color varies from light reddish brown to dark brown with pronotum (usually) and venter mottled with numerous small piceous spots; pale yellow irregular transverse fascia on corium narrow, confined to veins, sometimes obsolete to completely absent: pronotum moderately declivent with humeri broadly rounded, all pronotal margins entire; tibial dilations lanceolate, both inner and outer dilations of approximately same size and shape. Clasper (Fig. 27) with inner basal lobe absent, shank with median lobe absent, hook strongly curved; capsule (Fig. 47) with a wide rectangular shaped median notch; aedeagus as in cor cuius. As pointed out in the discussion of corculus, these two species are closely related, and the best diagnostic characters are the tibial dila- tions. The shorter equal-sized inner and outer tibial dilations (Fig. 58) will serve to distinguish occidentalis from corculus. Koerber (1963) in his report on the biology and economic im- portance of occidentalis , lists 12 species of conifers that serve as host plants. Schaffner (1967) states that in Iowa Scotch pine, Pinus sylvestus L., appeared to be the most commonly utilized host plant. Schaefer (1965, 1968) describes some morphological aspects of occidentalis in his studies on the higher classification of the Coreoidea. Distribution : From southern British Columbia and Alberta south- ward to Arizona, New Mexico and Texas, eastward to Iowa. Material Examined : Syntypes: 1 $, Utah, Amarilla; 1 $, Cal. Placer Co. (E. C. Van Dyke). In United States National Museum, Type No. 13230. The $ specimen has been chosen as Lectotype, and an appropriate label attached. UNITED STATES: 36$, 38$, 132 Entomologica Americana [Vol. 45 California; Colorado; Idaho; Iowa; Kansas; Montana; New Mexico; Oregon; Texas; Washington. In Chicago Natural History Museum, Iowa State University, Museum of Comparative Zoology (Harvard), University of California (Davis), American Museum of Natural His- tory, United States National Museum, University of Kansas, J. A. Slater, P. D. Ashlock and R. C. Allen collections. Leptoglossus clypealis Heidemann Leptoglossus clypealis Heidemann, 1910:195-6, pi. 8, fig. 1. — Van Duzee, 1917:90.— Gibson, 1917:70, 71.— Deay, 1928:378-9. — Torre-Bueno, 1941-49. — Froeschner, 1942:593, 599, pi. 4, fig. 42. — Drew and Schaefer, 1963:114, pi. 1, fig. 17. Theognis clypealis Elussey, 1953:30. Ground color varying from yellowish tan to light brown with head and scutellum piceous; a wide irregular transverse fascia on corium and anterior half of each connexival segment whitish yellow to ochra- ceous; membrane transparent and pale; tylus produced anteriorly as a conspicuous stout spine; humeri broadly rounded and all pronotal margins entire, tibial dilations lanceolate. Clasper (Fig. 28) with a prominent rounded inner basal lobe, shank with a large median lobe, hook strongly curved; capsule (Fig. 48) with a rectangular shaped median notch, no dorsal prongs; aedeagus as in corculus except all distal lobes more heavily sclero- tized and larger. L. clypealis, a member of the corculus- group, is a distinctive spe- cies in this genus. The spine on the tylus and pale membrane will readily separate clypealis from all other Leptoglossus species. Clypealis has been recorded as injurious to plums and almonds (Heidemann 1910), ornamental pomegranate (Torre-Bueno), and aromatic sumac ( Rhus aromatica Ait.) (Froeschner 1942). Distribution : Central and southwestern United States and into Mexico. Material Examined : Holotype: £ , Colorado, Platte Canon, 5-20- 01 (Dyar and Caudell). In United States National Museum, Type No. 13229. UNITED STATES: 33 5 , 46 2, Arizona; California; Colo- rado; Iowa; Kansas; New Mexico; Texas. MEXICO: 3$, 12, Sabinas Hidalgo Nuevo Leon, 130VI-1939 (Ralph Hagg); 1 2, same except Jacala, 4-VII-1939. In Iowa State University, University of California (Davis), Museum of Comparative Zoology (Harvard), Chicago Natural History Museum, American Museum of Natural His- tory, Stockholm Museum, Hungarian National Museum, United States 1969] Allen: Revision of the Genus Leptoglossus 133 National Museum, University of Kansas, J. A. Slater, P. D. Ashlock and R. C. Allen collections. Leptoglossus brevirostris Barber Leptoglossus brevirostris Barber, 1918 : 35-6. Theognis brevirostris Hussey, 1953:30. A small species; dorsum reddish brown with piceous humeri and occasionally a few small piceous spots on pronotal disk, venter mottled with numerous small piceous spots; a pale yellow narrow transverse fascia on corium (occasionally very faint); humeri prominently ex- panded, humeral angles acute to acuminate; hind tibial dilations phylliform. Claspers (Fig. 29) with inner basal lobe low and rounded, shank with a prominent median lobe, hook strongly curved; capsule (Fig. 49) with median notch rounded; dorsal prongs obsolete; aedeagus with dorsal sac of conjunctiva as follows: sclerotized proximal lobes present; right median lobe large with its base sclerotized; left median lobe as a small sclerotized area; large sclerotized anterior distal lobe present; one spinose distal lateral lobe present. L. brevirostris closely resembles chilensis in size and color, but there are a number of differences between these two species. In brevirostris the lateral margins of the pronotum are entire (rarely crenulate) not serrate, a transverse fascia is present on the corium, and the dorsal sac of the conjunctiva has only one median lobe and one distal lobe. L. brevirostris appears to be more closely related to stigma in respect to the similarity of the habitus and the genitalia. L. ashmeadi and brevirostris are the only North American species with a short labium and short phylliform tibial dilations. Barber (1906) first referred to brevirostris as a possibly unde- scribed species from southwest Texas. Heidemann (1910) used the name Leptoglossus stigma var. minor Dallas, in a list of Leptoglossus species occurring in the United States without any further reference to the origin of the name. Barber (1918) in the original description of brevirostris states that he and Mr. Heidemann had previously been using the name stigma var. minor Dallas for brevirostris , although Barber never used the trinomen in any earlier literature. Distribution : Southwestern United States. Material Examined: Holotype: $. ARIZONA: Huachuca Mts., Aug. 6, 05. In United States National Museum, Type No. 61107. Paratypes: 6$, same data as holotype. TEXAS: 1$, St. Tomas, Brownsville (Charles Schaeffer). In United States National Museum. UNITED STATES: 11$, 15 9, Arizona; California; Texas. MEX- 134 Entomologica Americana [Vol. 45 ICO: Baja California. In American Museum of Natural History, United States National Museum; California Academy of Sciences, J. A. Slater and R. C. Allen collections. Leptoglossus ashmeadi Heidemann Leptoglossus ashmeadi Heidemann, 1909:237. — Van Duzee, 1917: 90.— Gibson, 1917:70, 72.— Blatchley, 1926:225, fig. 43.— Torre-Bueno, 1941:50. Theognis ashmeadi Hussey, 1953:31. One of the most brightly colored Leptoglossus species in North America. Dorsum and appendages piceous with following areas strongly contrasting orange-yellow: all margins of pronotum widely and confluently and irregular transverse fascia on corium; venter almost entirely orange-yellow; humeri moderately expanded with humeral angles acute; hind tibial dilations phylliform. Clasper (Fig. 30) with inner basal lobe prominent and rounded, shank thick with a low median lobe, hook strongly curved; capsule (Fig. 50) with a deep round median notch, dorsal prongs small; aedeagus as in brevirostris. Blatchley (1926) states that adults and nymphs were collected on mistletoe, Phoradendron flavescens (Pursh. ) Nutt. Three specimens that I have examined from Natchez, Mississippi were also collected on mistletoe. Distribution : Southeastern United States. Material Examined : Holotype: $. Florida: St. Nicholas. In United States National Museum, Type No. 12191. ALABAMA: 2$, Mobile, XI-18-1927 (Th. van Allen). FLORIDA: 2$, 7-Oaks, May 1, 1908 (Van Duzee); 1$, Orlando, 12-26-1908 (A. W. Morril ) ; 1$, Dune Din, 4-1-1921 (W. S. B.). MISSISSIPPI: 1$, Ocean Springs, Jan. 22, 1944, S.S. #9546. 3 9, Natchez, V-26-1909, “on Phoradendron flavescens ,” Hunter No. 1667 (E. S. Tucker). In United States National Museum, California Academy of Sciences, and Chicago Natural History Museum. Acknowledgments I wish to express my gratitude and appreciation to my major advisor Dr. James A. Slater, who first suggested this study, and made available his time, personal library and collection. I am indebted to Drs. Carl W. Schaefer and Peter D. Ashlock and Mrs. Darleen Wilcox for their generous advice, suggestions, and discussions on many matters. My thanks go to Dr. Ralph M. Wetzel for critically reading the manuscript. 1969] Allen: Revision of the Genus Leptoglossus 135 I am most grateful to the following persons and institutions for their generosity in lending material: Dr. G. Petersen (Deutsches Entomologisches Institut), Dr. Pieter H. van Doesburg, Jr. (Leiden Museum), Dr. A. Soos (Hungarian National Museum), Dr. A. Gollner-Scheiding (Berlin Humboldt University Museum), the late Dr. E. Kjellander (Stockholm Museum), Dr. W. J. Knight and Mrs. G. M. Black (British Museum (Natural History)), Dr. I. Lansbury (Hope Museum, Oxford University), Dr. George Wallace (Carnegie Museum of Natural History), Dr. Peter D. Ashlock (formerly of the Bernice P. Bishop Museum), Dr. Jean L. Laffoon (Iowa State Uni- versity), Dr. Paul H. Arnaud, Jr. (California Academy of Sciences), Dr. Richard C. Froeschner (United States National Museum), Dr. Jon L. Herring (United States Department of Agriculture), Dr. Henry Dybas (Chicago Natural History Museum), Dr. G. W. Byers (Snow Entomological Museum, University of Kansas), Dr. De Santis (La Plata Museum), Dr. J. C. Schaffner (Texas A & M University), Dr. Howard E. Evans (Museum of Comparative Zoology, Harvard Uni- versity), Dr. P. Wygodzinsky (American Museum of Natural His- tory). Special thanks are extended to Miss Karen Stoutsenberger for the preparation of the dorsal view illustrations. Literature Cited Adams, A. L. and J. C. Gaines. 1950. Sunflower insect control. J. econ. Ent. 43: 2: 181-184. Ashlock, P. D. 1957. An investigation of the taxonomic value of the phallus in the Lygaeidae (Hemiptera — Heteroptera) . Ann. ent. Soc. Amer. 50: 4: 407-426. . 1967. A generic classification of the Orsillinae of the World (Hemiptera — Heteroptera: Lygaeidae). Univ. Calif. Publ. ent. 48: 1-82. Barber, H. G. 1906. Hemiptera from southwestern Texas. Mus. News, Brooklyn 1:9: 255-289. . 1914. Insects of Florida II. Hemiptera. Bull. Amer. Mus. nat. Hist. 33: Art. 31 : 495-535. . 1918. A new species of Leptoglossus : A new Blissus and varieties. Bull. Brooklyn ent. Soc. 13: 35-39. . 1939. Scientific survey of Porto Rico and the Virgin Islands: Insects of Porto Rico and Virgin Islands: Hemiptera- Heteroptera (excepting the Miridae and Corixidae). Sci. Surv. P. Rico 14: 3: 263-441. Barber, H. G. and S. C. Bruner. 1947. The Coreidae of Cuba and the Isle of Pines with the description of a new species (He- miptera-Heteroptera) . Mem. Soc. cubana Hist. nat. 19: 76—88. 136 Entomologica Americana [Vol. 45 Berg, C. 1879. Hemiptera Argentina enumeravit speciesque novas. Bonariae: Pauli E. Coni pp. 1-316 (also 1878, An. Soc. cient. argent.). . 1892. Nova Hemiptera faunarum Argentinae et Uruguay- ensis. Bonariae: Pauli E. Coni et Filiorum pp. 1-112 (also An. Soc. cient. argent. 33: 65-72). Bergroth, E. 1894. Liste de quelques Hemipteres de la Plata. Rev. Ent., Caen 13: 164-167. Blatchley, W. S. 1926. Heteroptera or true bugs of eastern North America, with especial reference to the faunas of Indiana and Florida. Indianapolis: Nature Publishing Co. 1116 pp. Blote. H. C. 1936. Catalogue of Coreidae in the Rijksmuseum van Natuurlijke Historic. Part III. Coreinae, second part. Zool. Meded. 19:23-66. Bosq, J. M. 1940. List preliminar de los Hemipteros (Heteropteros) , especialmente relacionados con la agricultura nacional. Rev. Soc. ent. argent. 10: 399-417. Burgess, P. S. and R. S. Hawkins. 1945. Fifty-fifth Annual Report. Arizona Agric. Exp. Sta. 54 pp. Burmeister, H. 1835. Handbuch der Entomologie, Zweiter Band: II Ordnung Rhynchota. Berlin: Theod. Ehr. Friedr. Enslin i-iv, 1-400 pp. Chittenden, F. H. 1902. Some insects injurious to vegetable crops. Bull. U.S. Bur. Ent. (ns.) 33: 1-117. . 1925. Note on the behavior of Leptoglossus oppositus (Say). Bull. Brooklyn ent. Soc. 20: 148-149. Cockerell, T. D. A. 1905. (No title) Ent. News 16: 18. Costa Lima, A. da. 1940. Insectos do Brazil II. Hemipteros. Rio de Janeiro: Escola Nacional de Agronomia pp. 1-351. Dallas, W. S. 1852. List of the specimens of Hemipterous insects in the collection of the British Museum. Part II. London: Taylor and Francis Inc. 592 pp. Deay, H. O. 1928. The Coreidae of Kansas. Univ. Sci. Bull. 18: 371-415. Distant, W. L. 1880-1893. Insecta Rhynchota. Hemiptera-Heterop- tera. Vol. I. Biol, cent.-amer. London. 462 pp. . 1901a. Revision of the Rhynchota belonging to the Fam. Coreidae in the Hope collection at Oxford. Proc. zool. Soc. Lond. 1: 325-335. . 1901b. Rhynchotal notes. IX. Heteroptera: Fam. Coreidae. Ann. Mag. nat. Hist. (7) : 7: 416-432. . 1902. Fauna of British India, including Ceylon and Burma Rhynchota. Vol. I (Heteroptera). London: Taylor and Francis pp. 1-438. Drew, W. A. and K. Schaefer. 1963. The Coreidae of Oklahoma (Hemiptera). Proc. Okla. Acad. Sci. 43 : 112-122. 1969] Allen: Revision of the Genus Leptoglossus 137 Dupuis, C. 1952. Priorite de quelques noms d’ Heteropteres de Guerin Meneville (1831). Bull. Soc. Zool. France: 77: 447-454. Ebeling, W. 1959, Subtropical fruit pests. Univ. Calif. Div. Agric. Sci. 436 pp. Fabricius, J. C. 1775. Systema entomologiae. Flensburgi et Lipsiae: Korte. 832 pp. . 1781. Species insectorum II. Hamburgi et Kilonii: Ernest Bohnii. pp. 1-517. . 1794. Entomologia systematica emendata et aucta IV. Hafniae: C. G. Proft. pp. 1-472. Fernando, H. E. 1957. The biology and control of Leptoglossus mem- branaceus Fabricius. Trop. Agric. 113: 107-118. Forster, J. R. 1844. Descriptions animalium quae itinere admaris australis terras per annon 1772, 1773, et 1774 suscepto collegit observait et delineavit J.R.F. Nunc demum editae . . . curante H. Lichtenstein. Berolini. 424 pp. Froeschner, R. C. 1942. Contributions to a synopsis of the Hemiptera of Missouri, Part II. Amer. Midi. Nat. 27: 3: 591-609. Gibson, E. H. 1917. Key to the species of Leptoglossus Guer. occur- ring north of Mexico (Heteroptera, Coreidae). Psyche 24: 69-73. Gibson, E. H. and A. Holdridge. 1918. The genus Narnia Stal, and a key to the genera of Anisoscelini A. and S. (Coreidae: Heteroptera). Psyche 25: 1-4. Gmelin, J. F. 1788. Caroli a Linne systema naturae. 13th ed., Tom. I (IV) Lipsiae: Beer pp. 1517-2224. Gross, G. F. 1963. Insects of Micronesia: Coreidae (Alydini by J. C. Schaffner), Neididae, and Nabidae. Insects of Micronesia (B. P. Bishop Mus.) 7: (7) : 358-390. Guerin, F. E. 1838. In: Duperrey, Voyage autour du monde execute par ordre du roi, sur la corvette de la majeste La Coquille, pendant les annes 1822, 1823, 1824, et 1825. Zoologie. (plates X-XII, 1831). Guerin-Meneville, F. E. 1857. In: Sagra, Histoire physique, poli- tique et naturelle de l’lsle de Cuba. Animaux articules a pieds articules. Paril: Arthus Bertrand. Hahn, C. W. 1826. leones ad monographiam Cimicum. Nurnberg: Lechner. Heidemann, O. 1909. New species of Tingitidae and descriptions of a new Leptoglossus. Bull. Buffalo Soc. Nat. Sci. 9: 231-238. . 1910. New species of Leptoglossus from North America (Hemiptera-Coreidae). Proc. ent. Soc. Wash. 12: 191-197. Herbst, J. F. W. 1780-89. Gemeinnuzzige Naturgeschichte des Thier- reichs. Berlin and Stralsund. Vol. 6. Herrich-Schaeffer, G. H. W. 1836. Die Wanzenartigen Insekten. Nurnberg: C. H. Zeh’schen Buchhandlung. 1836, Vol. 3; 1842, Vol. 7, 1853; Vol. 9. 138 Entomologica Americana [Vol. 45 Hussey, R. F. 1952. Food plants and new records for some Hemiptera in Florida. Florida Ent. 35: 3 : 117-118. . 1953. Concerning some North American Coreidae. Bull. Brooklyn ent. Soc. 48: 29-34. . 1956. Additions to the United States list of Hemiptera. Florida Ent. 39: 2: 88. Kelsheimer, E. G. 1949. Control of insect pests of cucumber and squash. Bull. Fla. Univ. Agric. Exp. Sta. 465: 1-15. Kiritshenko, A. N. 1935. Ueber Die Gattungen Leptoglossus Guer. und Theognis Stal (Hemiptera, Coreidae). Konowia 14: 191. Kirkaldy, G. W. 1906. On the nomenclature of the genera of the Hemiptera. Entomologist 39: 253-257. Koerber, T. W. 1963. Leptoglossus occidentalis (Hemiptera: Corei- dae). A newly discovered pest of coniferous seed. Ann. ent. Soc. Amer. 56: 229-234. Lethierry, L. and G. Severin. 1894. General catalogue of the He- miptera-Heteropteres. Brussels: F. Hayes, Imprimeur de l’Acad- emie Royale de Belgique 2: 1-277. Linnaeus, C. 1767. Systema Naturae. 12th edition. Holmiae: Laur- entii Salvii 1 : (2): 533-1327. . 1771. Mantissa plantarum altera Generum editionis II. Holmiae, impensis direct. L. Salvi pp. 143-587. Mann, J. 1969. Cactus-feeding Insects and Mites. U. S. Nat. Mus. Bull. 256: 1-158. Mayr, G. L. 1865. Diagnosen neuer Hemipteren II. Verh. zool.-bot. Ges. Wien 15: 429-446. . 1866. Hemiptera In: Reise der Osterreicheschen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859. Zoo- logischer Theil 2, Abt. 1. Wien: Karl Gerold’s Sohn 204 pp. McCullough, T. 1968. Acid and aldehyde compounds in the scent fluid of Leptoglossus oppositus. Ann. ent. Soc. Amer. 61: 1044. Montrouzier, P. 1855. Essau sur la faune de File de Woodlark ou Moiou. Ann. Soc. Agric. Lyon 2:1: 1-114. Pagden, H. T. 1928. Leptoglossus membranaceous F. a pest of Cu- curbitaceae. Malay. Agric. Jour. 16: 387-403. Parshley, H. M. 1923. Family Coreidae: In Britton: Guide to the insects of Connecticut: Part IV. The Hemiptera or sucking insects of Connecticut. Bull. Conn. Geol. Nat. Hist. Surv. 34: 746-753. Pennington, M. S. 1922. Physis-Notas sobre coreidos Argentinos. Rev. de la Soc Ara de Ciencias Naturales, Buenos Aires pp. 125-170. Reed, E. C. 1900. Sinopsis de las Hemipteros de Chile. Valparaiso 1 07 pp. Say, T. 1831. Descriptions of new species of Heteropterous Hemiptera of North America. New Harmony Indiana. 1831: 310-368 (Leconte Edition, 1859). 1969] Allen: Revision of the Genus Leptoglossus 139 Schaefer, C. W. 1965. The morphology and higher classification of the Coreoidea (Hemiptera-Heteroptera) . Part III. The families Rhopalidae, Alydidae, and Coreidae. Misc. Pub. ent. Soc. Amer. 5:1: 1-76. . 1968. The morphology and higher classification of the Coreoidea (Hemiptera-Heteroptera). Part IV. The Acantho- cephala- group and the position of Stenoscelidea Westwood (Coreidae). Univ. Conn. Occas. Pap. (Biol. Sci. Ser.) 1: 153-199. Schaffner, J. C. 1967. The occurrence of Theognis occidentalis in the midwestern United States ( Heteroptera: Coreidae ) . J. Kansas ent. Soc. 40: 141-142. Schouteden, H. 1907. Les types d’Hemipteres de Montrouzier. Ann. Soc. ent. Belg. 51: 106-118. Signoret, V. 1863a. Hemiptera: In Maillard, Notes sur l’lle de la Reunion. (Bourbon) Insects, Paris. . 1863b. Revision des Hemipteres du Chile. Ann. Soc. ent. Fr. 4: 3: 541-588. Snapp, O. I. 1948. Sucking insects causing deformed peaches. J. econ. Ent. 41: 555-557. Spinola, M. 1837. Essai sur les genres d’insects appartenants a l’orde des Hemipteres Lin. ou Rhyngotes, Fab. et a la section Heter- opteres, Dufour. Genes: Chez Yves Gravier Imprimeur-Li- braire pp. 1-383. . 1852. Hemipteros: In Gay, Historia de Chile zoologica. 7: 113-320. Paris: Imprenta de Maulde y Renou. Stal, C. 1855. Nya Hemiptera. Ofvers. VetenskAkad Forh., Stockh. 12: 181-192. . 1859. Till Kannedomen om Coreida. Ofvers. VetenskAkad. Forh., Stockh. 16: 449-475. . 1860. Bidrag till Rio Janeiro-traktens Hemipter-fauna. K. svenska VetenskAkad. Handl. 2:7: 1-84. . 1862. Hemiptera Mexicana enumeravit speciesque novas descripsit. Stettin ent. Ztg. 23: 81-118; 273-281, 289-325, 437- 462. . 1868. Hemiptera Fabricana I. K. svenska VetenskAkad. Handl. 7: 11: 1-148. . 1870. Enumeratio Hemipterorum. Part I. K. svenska VetenskAkad. Handl. 9: (1): 1-232. Szent-Ivany, J. J. H. and A. Catley. 1960. Observations on the bi- ology of the black leaf-footed bug Leptoglossus australis (F.) of Papua and New Guinea. Papua New Guinea Agric. Jour. 13:70-75. Torre-Bueno, J. R. de la. 1941. A synopsis of the Hemiptera- Heteroptera of America north of Mexico. Part II. Families Coreidae, Alydidae, Corizidae, Neididae, Pyrrhocoridae and Thaumastotheriidae. Ent. amer. 21: (ns.): 41-122. 140 Entomologica Americana [Vol. 45 Uhler, P. R. 1893. A list of the Hemiptera-Heteroptera collected in the Island of St. Vincent by Mr. Herbert H. Smith; with descrip- tions of new genera and species. Proc. zool. Soc. Lond. 1893: 705-719. . 1894. On the Hemiptera-Heteroptera of the island of Gre- nada, West Indies. Proc. zool. Soc. Lond. 1894: 167-224. Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north of Mexico. Berkeley: Univ. Calif. Publ. Ent. 2: 1-902. Walker, F. 1871. Catalogue of Hemiptera-Heteroptera in the collec- tion of the British Museum. London: Part IV. Watson, J. R. and E. W. Berger. 1932. Citrus insects and their con- trol. Bull. Fla. Univ. Agric. Exp. Sta. 67: 3-140. Westwood, J. O. 1842. A catalogue of Hemiptera in the collection of the Rev. F. W. Hope, Part II. London: J. C. Bridgewater 26 pp. Wolcott, G. W. 1948. The insects of Puerto Rico. J. Dep. Agric. P.R. 32: 1-224. A //* i — ' Entomologica Americana A TAXONOMIC AND BIOLOGICAL STUDY OF SPECIES OF ATTAGENINI (COLEOPTERA: DERMESTIDAE) IN THE UNITED STATES AND CANADA Richard S. Beal, Jr. SPfHSG Aft { m PUBLISHED BY THE NEW YORK ENTOMOLOGICAL SOCIETY INCORPORATING THE BROOKLYN ENTOMOLOGICAL SOCIETY Vol. 45, No. 3 October 26, 1970 The New York Entomological Society (incorporating the Brooklyn Entomological Society) publishes two journals: Entomologica Ameri- cana (irregularly, according to the availability of acceptable papers) and the Journal of the New York Entomological Society (quarterly). Monographs and papers too long to be published in other journals (minimum length approximately 70 manuscript pages) are published in Entomologica Americana. Shorter papers may be submitted to the Journal. Manuscripts should be sent to the Editor, Entomologica Americana, Department of Entomology, The American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. Entomologica Americana was first published by the Brooklyn Entomolog- ical Society in April, 1885. Publication was suspended after six volumes, resumed in 1926, again suspended in 1964 with Volume 44, and resumed in 1969 with Volume 45. The subscription rate to Entomologica Ameri- cana is $9.00 per volume. Correspondence regarding subscriptions and other non-editorial matters should be addressed to Publication Business Manager, New York Entomological Society, Department of Entomology, The American Museum of Natural History. EDITORS Kumar Krishna Valerie Krishna PUBLICATION COMMITTEE John A. L. Cooke A. P. Gupta A. B. Klots Mailed October 26, 1970 Entomologica Americana is published for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Entomologica Americana Yol. 45, 1970, pp. 141-235 A TAXONOMIC AND BIOLOGICAL STUDY OF SPECIES OF ATTAGENINI (COLEOPTERA: DERMESTIDAE) IN THE UNITED STATES AND CANADA By Richard S. Beal, Jr.1 Table of Contents Introduction 142 Methods of Preparing Larvae 144 Characters and Terms 145 Selection of Generic Characters 145 Adult Characters 146 Larval Characters 147 Characters for Recognition of the Attagenini 149 Relationships 150 Keys to the Species 153 Attagenus cyphonoides Reitter 165 Attagenus megatoma (Fabricius) 167 Attagenus megatoma megatoma (Fabricius) 174 Attagenus megatoma canadensis Casey 175 Attagenus el on gat ulus Casey 176 Attagenus schaefferi (Herbst) 182 Attagenus schaefferi hypar, new subspecies 187 Attagenus schaefferi spurcus LeConte 190 Attagenus pellio (Linnaeus) 190 Attagenus fasciatus (Thunberg) 193 Attagenus bicolor Harold 195 Novelsis athlophora Beal 200 Attagenus lobatus Rosenhauer 200 Novelsis perplexa (Jayne) 203 Novelsis varicolor (Jayne) 204 Novelsis aequalis (Sharp) 207 Attagenus rufipennis LeConte 209 Novelsis uteana Casey 216 Novelsis horni (Jayne) 219 Novelsis andersoni Beal 222 Novelsis timia Beal 223 Novelsis picta Casey 225 Species Described but Unrecognized 226 Acknowledgments 226 Literature Cited 228 1 Northern Arizona University, Flagstaff, Arizona 86001. [141] 142 Entomologica Americana [Vol. 45, No. 3 Abstract: The following species and subspecies within the der- mestid beetle tribe Attagenini are recognized from the United States and Canada: Attagenus cyphonoides Reitter, A. megatoma megatoma (Fabricius), A. megatoma canadensis Casey, A. elongatulus Casey, A. schaefferi hypar new subspecies, A. schaefferi spurcus LeConte, A. pellio (Linnaeus), A. fasciatus (Thunberg), A. bicolor Harold, A. lobatus Rosenhauer, A. rufipennis LeConte, Novelsis athlophora Beal, N. perplexa (Jayne), N. varicolor (Jayne), N. aequalis (Sharp), N. uteana Casey, N. horni (Jayne), N. andersoni Beal, N. timia Beal, N. picta Casey. Keys are provided to the adults and known larvae. Also included are diagnoses of adults, redescriptions of inadequately described adults, and descriptions of known larval forms. The species are placed in six groups according to degrees of morphological similarity, and criteria for determining the groups are discussed. It is found that during the larval stages most of the species feed on a wide variety of dried proteinaceous materials. The most common habitats for the larvae are noted as sheltered bird nests and rodent nests, but larvae may also be found in bee and wasp nests and in sheltered spider webbing. Adults commonly are observed to fly to flowers, probably to mate; however, mating may occur without flight and without feeding or drinking. The extent of depredations of stored food products and fabrics is discussed. Two species, A. lobatus and A. cyphonoides, are newly recorded as apparent recent introductions into the United States from the Old World. INTRODUCTION Members of the genus Attagenus of the family Dermestidae are well known to every entomologist as common pests of households and granaries. The name Attagenus piceus is probably memorized by every student enrolled in economic entomology at our American universities. Despite this familiarity, knowledge of the systematics of the New World members of the genus has seemingly regressed rather than advanced over the past seven or so decades. The last revisionary study of the American members of Attagenus was pub- lished in 1900 by Thomas L. Casey. This famous coleopterist was usually able to discriminate between the forms he studied, even if his species definitions cannot be accepted in the light of present knowledge. Nevertheless, his key and descriptions to the species of Attagenus are completely inadequate for sorting out the species. From the arrangement of the specimens in his collection, which is preserved intact at the United States National Museum, it is evident that he himself was unable to recognize the species. In several instances the specimens that stand behind a name include two or three different species. Since Casey’s time, American entomologists, perhaps understandably frustrated, have quite generally designated 1970] Beal: Taxonomic Study of Attagenini 143 every more or less uniformly colored, mahogany, brown, or black specimen by the name Attagenus piceus. This practice has been followed without regard for some obvious morphological differences and in spite of the well-documented fact that the specific trivial name piceus is not applicable to any of the forms in question. The problem of defining the limits of the genus has not been of concern to American students, since our Attagenini appear to fall very neatly into two groups, and the number of species is not large. H. F. Jayne, who revised the Dermestidae in 1882, placed members of both groups in Attagenus. Casey, however, separated the groups and erected the genus Novelsis to accommodate one of them. This separation has been a convenience, but, unfortunately, when members of the group from other zoogeographical areas are considered, the classification proves inadequate. Several European workers have arranged many members of Attagenus under the subgenera T elopes Redtenbacher (1843) and Lanorus Mulsant and Rey (1867). Both these names have priority over Novelsis, and probably a number of our American species should be assigned to them. The difficulty lies in knowing exactly what these subgeneric names represent. The greatest number of characters available for a generic classification seem to be found in the larvae; however, the larvae of the type species of Lanorus and T elopes have not been available for study. A fairly adequate classification might be worked out for the species of Attagenini, but until the type species of these genera have been investigated, the status of any generic names that are used will necessarily remain in doubt. In view of this uncertainty, I am not attempting in this paper to rename any of the generic groups within the Attagenini or to reassign any of the species to other genera. Current generic names are used throughout the paper, even though they do not correctly represent the natural groups within the Attagenini. A possible arrangement of the species into natural groups is proposed, but a determination of the names to be applied to the groups should await a more cosmopolitan study. An alternative solution to the problem of reassigning the species of Attagenini to a number of different genera is to lump them all under Attagenus , as has been done by several European workers. Although this would be a convenience to cataloguers, it would also serve to delay growth in knowledge of the group. There are ap- proximately 160 species of the tribe scattered throughout all the zoogeographic regions of the world. It is scarcely conceivable that these species do not fall into a number of smaller, definable groups. 144 Entomologica Americana [Vol. 45, No. 3 The discovery of these groups will provide a framework around which biologists other than systematists can organize their data. Comparison of these data may in turn assist the systematist to confirm or to refine his classification. A reasonable degree of subdividing would better serve the needs of both the functional biologist and the sys- tematist. Therefore, it seems uneconomical at present to reassign the species of Novelsis to Attagenus. Very possibly and hopefully in a short time most of the species will be given other generic assignments. The primary purpose of this paper therefore is to define the Nearctic species of the tribe. A diagnosis of each species is given, together with a redescription of the adult when necessary and a description of the mature larva whenever possible. All available biological information that bears upon the definition of the species is also included. Keys for the identification of adults and known larval forms are provided to make the definitions more useful. A second purpose of the paper is to present the biological data available for each species. Extensive data have been collected in the past for a few of the species. Unfortunately, in many instances the specimens studied were not deposited in a museum and there is no way of knowing their actual identity. Where there has been access to the specimens from which the data have been gathered, the information is related in the paper to the correct name of the species. Otherwise the data are ignored. METHODS OF PREPARING LARVAE The difficulty of using microscopic characters for studying and identifying the larvae is further aggravated by the fact that there seems to be no rapid way to prepare a slide mount of a specimen and leave all the useful characters visible. Most of the specimens used in this study were cleared in KOH, dehydrated in glacial acetic acid, cleared in beachwood creosote, and mounted in a balsam substitute. While the specimen was in the creosote the mouthparts were dissected out and the body cut open the entire length along the median suture. The body was then mounted spread out with the inside surfaces of the integument against the slide. Mouthparts were mounted on the same slide under separate 6 mm. coverslips. Some of the specimens studied were cut open the same way but mounted directly in Hoyer’s medium. Although this method allows for rapid preparation, the terga tend to retain their curved shape, so that even if the specimen is generally well-flattened the edges 1970] Beal: Taxonomic Study of Attagenini 145 of the terga are usually bent under and the spiracles difficult to observe. Clearing in KOH seems to make the terga easier to flatten, leaving the spiracles and their associated sclerites flat against the slide. It was found that specimens of most species can be identified quickly by using iridectomy scissors to cut out a piece of an ab- dominal segment with the spiracle, a part of the tergum, and a part of the sternum included. This piece is then mounted on a slide in Hoyer’s medium. Often the setae covering the spiracle need to be teased away first. However, to use the key, the entire specimen needs to be mounted. CHARACTERS AND TERMS Selection of Generic Characters: The difficulty that arises when one attempts to split the tribe Attagenini into genera is that if one set of characters is used to segregate the genera, the groups are ordered differently than if another set of characters is used. There are a number of sets that might be used, and there are as many conceivably different ways of ordering the genera. This kind of problem is a familiar one in the animal kingdom. The taxonomist must apparently make an a priori decision that certain characters are of generic value and group the species accordingly. Some criterion such as “adaptive significance” or “phylogenetic stability” might be used to select the generic characters. However, if it were to be granted that one or another of these criteria had taxonomic significance, it would be of little use at this time in choosing char- acters for genera of the Attagenini. Not enough is known of the adaptive value of the characters present in the larval or adult stages or of the phylogeny of the group to make such judgments. For- tunately, there is a rational way out of the dilemma. The taxonomist can first determine the genera and afterward find generic characters to match. It is not necessary to have generic characters prior to dividing a larger taxon into its genera. A “natural” classification can be achieved if the species are first grouped on the basis of their over-all similarities and dissimilarities. This is exactly what the classical taxonomist did with his “eye.” However, there are so many characters present among the species of Attagenini when both adult and larval stages are considered, the taxonomist cannot trust his mere impressions. It is necessary to tabulate and count the differ- ences. Once this has been done, there is an objective basis for grouping the species. The species within each group are more likely to share a greater number of genetic similarities than with any species 146 Entomologica Americana [Vol. 45, No. 3 of a different group. After the discovery of the groups, the generic characters are relatively easy to find. At least with the use of this procedure in the Attagenini the generic characters have not been difficult to identify. They are those characters that are common to all members of the group. No judgment is made as to their adaptive or phyletic value; no judgment is necessary. So far as their use in classification is concerned, they become the “key” characters by which the genera are recognized. This is the procedure that has been followed in sorting the species of Attagenini into groups in the pages following. Those characters that are common to all members of a group are termed “generic,” even though for the present no nomenclatural assignments to genera are made. Adult Characters: Previous workers have usually considered the relative sizes of the segments of the male antennal club of generic significance. Casey placed in Attagenus those species with the first two segments of the club extremely short in comparison with the third segment and in Novelsis those species with the first two seg- ments elongated. However, if both adult and larval characters are tabulated and the species grouped as described above, the relative lengths of the segments of the club are seen to be of a low order of importance. On the other hand, a useful generic character is found in the shapes of the metacoxa and the metepimeron. In some species the ventral lamina of the coxa extends laterally to meet the metepimeron, which is somewhat curved behind the metepisternum, thus enclosing the metepisternum behind. In other species these do not meet behind the metepisternum, so that the metepisternum appears open behind. Each of these conditions is correlated with other characters common to the groups and accordingly is a useful generic character. The relative sparseness or density of the body setae and also the length of the setae on the antennal club are similarly correlated with other characters and are useful in recogniz- ing the species groups. Most of the other adult characters used in the key are of significance at the species level only, including the number of segments in the antenna, the convexity or concavity of the hypomeron, and patterns in the elytral pubescence. If there are significant differences in the genitalia, I have been unable to discover them. There are minor genitalic differences between some of the species, but none of such a nature that they would be of value in sorting the species into groups. Neither have I been able to find significant differences in the wing venation. 1970] Beal: Taxonomic Study of Attagenini 147 The length of the terminal segment of the male antenna is to some extent correlated with the combined length of the pronotum and elytra and a useful character for distinguishing between some of the species. A high correlation exists between the length of the terminal antennal segment and the length of the elytron alone, at least in the megatoma section (Group I in this paper). However, a regression line drawn for a number of measurements plotted on a graph passes relatively far to one side of the zero point. As a result, the correlation is difficult to use as well as to express. The regression line for correlations between the last antennal segment and the total length of the pronotum and elytron passes somewhat closer to the zero point. Accordingly, it is easier to use and has been used in the descriptions following, even if at some sacrifice of precision. Many authors have described or figured the male antennal club as an aid in distinguishing the species, assuming that the relative lengths of the first two segments are closely correlated with the length of the terminal segment. However, there is no more than a moderately low correlation between these measurements. Although the ratio is of value in separating some groups of species, it is of little worth in distinguishing between “close” species, such as A. elongatulus and A. megatoma. Relatively poor correlations are found between the interocular width and the length of the terminal antennal segment or between the interocular width and the combined length of the pronotum and elytra. Larval Characters: A number of characters useful both for dis- tinguishing between the species and for grouping them into natural assemblages are found in the larvae. Unfortunately, most of the characters are so minute they cannot be easily observed except through a compound microscope. The body setae are usually longi- tudinally ribbed. They may be broad and flat with numerous ribs or they may be linear and more or less round in cross section with many or few ribs. The number of such ribs is often significant. Most of the setae are recumbent, but near the posterior margin of each tergum is a row of erect setae. Close to the margin of the socket of each of these erect setae is one or two pits, each roughly a fourth the diameter of the socket and presumably sensory in function. Whether there is one or two pits beside each socket and, if there is a single pit, whether it is on the medial or lateral side of the socket appear to be useful characters. Since it is necessary to describe differences between setae inserted on different areas of the abdominal segments, the meaning of the following terms should be noted. By tergum is meant the entire 148 Entomologica Americana [Vol. 45, No. 3 dorsal sclerotized area of each abdominal segment. Near the anterior margin of each tergum, except the ninth and occasionally the eighth, there is a transverse suture, the antecostal suture. That part of the tergum anterior to the suture is designated the acrotergite. The term tergite is used for the sclerotized part of the tergum posterior to the suture. The shape and position of the spiracle and the nature of various structures associated with it are significant. The spiracle may be a simple, somewhat circular opening on the posterior margin of the tergum (Fig. 12) or it may be a narrow slit with thickened margins some distance in front of the posterior margin of the tergum (Fig. 11). Close to the spiracle at the lateral margin of the tergum is a small sclerite bearing several long setae. This structure is termed the spiracular sclerite in the key and descriptions that follow. The spiracle may be adjacent to the sclerite or may open a distance from it. If at a distance, there is a narrow suture-like slit between the sclerite and the spiracle. The sclerite may be almost completely enclosed by the tergum or may appear to be almost entirely free of the tergum. In the latter case there is a narrow sclerotized margin of the tergum along the anterior edge of the spiracular sclerite, but the part of the tergum in front of this margin is membranous, so that the margin appears at first glance to be part of the sclerite. For many of the species only a limited number of larvae were available for study. As a result, I am not certain whether some of the observed differences are valid species characters or whether they are merely differences that vary within the species. Possibly there are a number of “good” characters associated with the shape and setation of the femora and tibiae. However, there is considerable variation in the pattern of setation in the species for which long series were available. Consequently, it has been difficult to make reliable comparisons with species for which only one or two speci- mens were at hand. Apparently there are also some secondary sexual differences in the shape of the larval tibia in some of the species. The sex was not known for many of the larval specimens studied, so the status of these characters has had to remain undefined. I have illustrated the protibiae of three species where the differences are quite marked and have noted some apparent differences in the descriptions of other species. However, I have made no use of these characters in the key or in grouping the species. The number of setae observed on the labial palps is described, but may be of little significance. The shapes of the glabrous areas on the pronotum 1970] Beal: Taxonomic Study of Attagenini 149 appear to differ from species to species, but the differences are not pronounced and are difficult to utilize. The length of each antennal segment in relation to the lengths of the other antennal segments as well as the length of the antennal setae in relation to the lengths of the antennal segments provide useful characters for distinguishing between species in other tribes of dermestids. Little correlation was found between these measure- ments in these species. One character that appears to be of moderate value in distinguishing a few of the species is the length of the long seta at the apex of the third antennal segment in comparison with the length of the segment. Unfortunately, the seta is easily broken off in preparing a slide mount and is available for use with an occasional specimen only. CHARACTERS FOR RECOGNITION OF THE ATTAGENINI A single character that distinguishes adult members of the Attagenini from those of other tribes of Dermestidae is the short first segment of the tarsus of the hind leg. This segment is not more than half as long as the second segment. The following combination of characters will also serve to identify members of the tribe. The body is covered with subrecumbent to suberect hairs, never with scales. The head bears a distinct median ocellus. The pronotum does not have a distinct sublateral carina on each side. The hind wings are well developed. The antennal club is 3 -segmented, with the segments moderately enlarged and in the male sometimes greatly elongate. The hypomeron may be broadly and deeply concave to slightly convex but, if concave, not forming a distinct cavity for the antenna and not margined behind. The prosternal process is long and narrow and is received in a shallow groove in the mesosternum. This groove becomes more shallow posteriad, so that the mesoster- num is not completely divided and its hind margin is visible for its entire width. The abdomen has five free and externally visible stemites. The larvae of the Attagenini are readily recognized by their shape, the body being elongate, gradually tapering posteriad from the metathorax, rounded dorsally, and flattened ventrally. The terminal segment is provided with a brush of long, slender setae. Setae of the body may be scale-like or long and slender with minute imbricate scales (spicisetae), but are never spear-headed (hastisetae), clavate, basket-shaped, or branched. The antenna consists of two 150 Entomologica Americana [Vol. 45, No. 3 moderately long basal segments and a shorter terminal segment. The maxillary palp is 4-segmented, the first two segments being short, the third long and curved inward, and the fourth long but shorter than the third. Each notum and tergum, except the tergum of the ninth abdominal segment, includes at its lateroposterior angle a spiracular sclerite, which bears several long, slender setae. The ninth abdominal segment lacks urogomphi. The tenth segment is apparently un- represented by any sclerotized parts, unless the small paired sclerites inserted in the sternum of the ninth segment represent part of the tenth. Additional descriptive details are given by Rees (1943). RELATIONSHIPS With the use of the procedures described for sorting species into groups, our American Attagenini fall into the following six groups. Group I. Attagenus cyphonoides Attagenus megatoma Attagenus elongatulus Attagenus schaefferi Attagenus pellio Attagenus fasciatus The larvae of each of these species are characterized by having setae with smooth rather than serrulate or denticulate margins, spiracles that open a short distance in front of the posterior margin of the tergum rather than on the margin, and spiracular sclerites that are bounded anteriorly, medially, and usually laterally by the tergum, but not posteriorly. The tergum is entirely sclerotized in front of the spiracular sclerite. Characters common to the adults include the approximation of the metacoxa and the metepimeron, extremely short and fine setae on the male antennal club, and moderately short and dense, subrecumbent body setae. All except A. fasciatus have a male antennal club in which the first two seg- ments combined are much shorter than the last segment. A. fasciatus could justifiably be placed in a group by itself: the adults differ noticeably in having much stouter legs and an absence of secondary sexual differences in the form of the antenna. The decision to include A. fasciatus here is more or less arbitrary. It is meant to show that in total characters it is closer to this group than to any of the follow- ing. The type species of Attagenus is Dermestes pellio Linnaeus, 1970] Beal: Taxonomic Study of Attagenini 151 according to Hope (1840). Therefore, Group I would carry the name Attagenus, should the other groups be assigned generic names. Group II. Attagenus bicolor Novelsis athlophora In general fascies, adults of A. bicolor are similar to members of Group I. They differ in the structure of the hypomeron, which does not enclose the base of the procoxa but leaves the base and the trochantin fully exposed. Species of each of the other groups have an auricular-like structure enclosing the trochantin and the base of the procoxa behind (except that it is somewhat reduced in N. horni of Group VI). Adults also possess long, sparse, usually wholly black body setae. The setae may be dark brown in immature specimens but otherwise are uniformly black. The larvae differ from members of Group I in many respects. The margins of the setae are micro- serrulate, a character shared with the groups that follow. The spiracular sclerite is completely enclosed behind by a broad area of the tergum. The spiracle, though associated with the spiracular sclerite by a narrow channel, is some distance from it and some distance from the posterior margin of the tergum. The setae on the pretarsus of A. bicolor are subequal in width and length, as they are in Group I, whereas the species in the groups following have one of the two setae twice as wide as the other. However, the setae in A. bicolor are four-fifths the length of the claw, whereas the setae in members of Group I are not over half the length of the claw. N. athlophora is tentatively included in Group II in view of its similarity in adult characters to A. bicolor. A further discussion of the similarities will be found in the section on A. bicolor. Group III. Attagenus lobatus This species seems quite obviously to belong to no other group of Nearctic species, although it has more characters in common with Group I than with any other group. The larvae are unique in having ten or more setae inserted in each spiracular sclerite and in having a row of long, stout, linear setae near the anterior margin of each abdominal sternum. Another remarkable larval character is found in the extremely long and fine setae inserted on the acrotergites. N. uteana of Group IV also has such long and fine setae on the acrotergites but, unlike A. lobatus, it also has a row of ensiform setae on the acrotergites. Unique characters in the adult include the fine, thread-like carina on the dorsal margin of the 152 Entomologica Americana [Vol. 45, No. 3 anterior tibiae, the exceptionally fine dorsal pubescence, and the elongation of the basal lobe of the pronotum. Group IV. Novelsis varicolor Novelsis perplexa Novelsis aequalis Larval stages of N. perplexa are unknown, but it can be pre- sumed on the basis of adult similarities that the species belongs in the same group as N. varicolor and N. aequalis. Although the latter two species differ in a number of larval characters, they share more characters with each other than with any other species. The most noticeable characters aligning them with each other are as- sociated with the body setae. These are broadly oval and strongly ribbed, with serrulate or denticulate margins and ribs. In addition, the spiracular sclerite is enclosed by the tergum, although the tergum is not sclerotized in front of the sclerite in N. aequalis. Two distal sensory pits are present on the epipharynx, but are very minute. Adults of all three species are remarkably similar in the structure of the antennal club, the erect setae of the male antennal club, the dense, suberect setae of the dorsum, and the type of pattern formed by the tricolorous elytral setae. Attagenus varicolor Jayne is the type species of the subgenus Paranovelsis Casey by monotypy. Should the members of Group IV not be found congeneric with one of the Palearctic genera, they should be arranged under this name. However, adults of the type species of Lanorus, Dermestes vigintiguttatus Fabricius = Attagenus punctatus (Scopoli), resemble members of this group in a number of respects. Without a study of the larvae of A. punctatus it would be hazardous to affirm the distinctiveness of our Nearctic species. Group V. Attagenus rufipennis Adults of this species superficially look very much like members of Group I, particularly in the shape of the male antennal club, which has the terminal segment many times longer than the two very short preceding segments. However, the separation of the coxal plate from the metepimeron is a character distinguishing the species from the members of Group I and suggesting some other funda- mental differences. Numerous differences are found in the larvae, which are more like larvae of Group VI than those of any of the foregoing groups. They differ from members of Group VI in pos- sessing a distal series of two small sensory pits on the epipharynx, two rather than one sensory pit adjacent to the sockets of the sub- 1970] Beal: Taxonomic Study of Attagenini 153 marginal erect setae of the terga, and two small sclerites in the sternum of the ninth abdominal segment. Group VI. Novelsis uteana Novelsis horni Novelsis andersoni Novelsis timia Novelsis picta Unique characters among larvae of this group include the fol- lowing: the lack of paired sclerites in the sternum of the ninth abdominal segment, the lack of a distal series of sensory pits on the epipharynx, no more than three setae on the dorsal side of each lobe of the ligula, and only one sensory pit adjacent to each socket of the submarginal erect setae of the terga. With reference to the last character, members of all other groups have two or occasionally three sensory pits adjacent to each socket, except that A. bicolor has some sockets with one and some with two adjacent sensory pits. Members of Group VI along with A. rufipennis (Group IV) have the tergum in front of the spiracular sclerite unsclerotized, so that the sclerite appears to project laterad somewhat peninsula-like. Larvae of N. timia and N. picta are unknown, but these species are included in the group because of adult similarities. The adults share a number of characters not found in other North American Attagenini. The front coxae are elongated relative to the length of the prosternal process, so that the coxae extend posteriad beyond the apex of the process. The epipleuron opposite the middle of the metepisternum lies in nearly the same plane as the incurved side of the elytron, instead of forming an angle with it of 90 to 120 degrees. The segments of the male antennal club are greatly elongated, so that the antenna in repose extends beyond the anterior margin of the metepisternum. Attagenus horni Jayne is the type species of Novelsis (designated by Beal, 1954). KEY TO THE NEARCTIC SPECIES OF ATTAGENINI Adult 1. Dorsal setae unicolorous, black or blackish brown; setae on disc of pronotum and elytra as long as combined length of antennal segments 3, 4, and 5, or longer. Setal punctures on disc of pronotum and base of elytra separated by two and usually three to four or more times diameter of single 154 Entomologica Americana [Vol. 45, No. 3 Figs. 1-9: Fig. 1. Head of adult of Attagenus bicolor. Fig. 2. Head of adult of A. lobatus. Fig. 3. Head of adult of A. megatoma. Fig. 4. Left metasternal coxa and associated sclerites of A. elongatulus. Fig. 5. Left metasternal coxa and associated sclerites of Novelsis varicolor. Fig. 6. Left metasternal coxa and associated sclerites of A. bicolor (stn = sternum, epis = episternum, eppl = epipleuron, epim = epimeron, cox = coxa). Fig. 7. Left prosternal coxa and associated sclerites of A. bicolor. Fig. 8. Outline of dorsum of A. lobatus. Fig. 9. Left prosternal coxa and associated sclerites of A. megatoma. 1970] Beal: Taxonomic Study of Attagenini 155 puncture. Hypomeron not forming lobe behind base of procoxa; base of procoxa and trochantin exposed (Fig. 7). 2 Dorsal setae unicolorous, bicolorous or tricolorous; setae on disc of pronotum and elytra seldom longer than combined length of antennal segments 3 and 4, usually shorter, al- though setae on margins of elytra and pronotum may be longer, but if dorsal setae longer than length of antennal segments 3, 4, and 5 combined, then setae bicolorous or tricolorous. Setal punctures of disc of pronotum and base of elytra hidden by dense pubescence or, if visible, usually separated by one or two times diameter of single puncture. Hypomeron forming auricle-like lobe behind base of pro- coxa so that base of procoxa and trochantin largely hidden (Fig. 9) 3 2. Antenna 1 1-segmented; male antenna with terminal segment of club five times as long as length of two preceding segments combined. Attagenus bicolor Antenna 10-segmented; male antenna with terminal segment no more than four-fifths as long as length of two preceding segments combined. Novelsis athlophora 3. Elytron with black hairs except for prominent oval spot (about as broad as eye) of white hairs at middle of disc near suture and often also with two or three very small lateral white spots at about basal two-fifths. Attagenus pellio Elytral pubescence unicolorous or with bands of light pubes- cence or with several large spots of light pubescence, but not as described above. 4 4. Lamina of hind coxa not meeting metepimeron; coxa and epimeron appearing separated by metepisternum (Fig. 5). 5 Lamina of hind coxa extending behind metepisternum to meet metepimeron (Fig. 4). 15 5. Antenna 10-segmented. 6 Antenna 1 1-segmented. 9 6. Dorsal pubescence tricolorous: black, golden-brown, and white; ventral pubescence light golden-yellow; setae on margin of fifth abdominal sternum dark brown or black. Male anten- nal club clothed with erect hairs about three-fourths as long as width of third antennal segment. Novelsis perplexus Dorsal pubescence unicolorous or bicolorous; ventral pubes- 156 Entomologica Americana [Vol. 45, No. 3 cence whitish, including setae on margin of fifth abdominal sternum. Male antennal club clothed with closely appressed hairs not longer than one-fourth width of third antennal segment. 7 7. Elytral integument brownish black to black with contrasting light tan or reddish yellow maculae; maculae forming oblique subbasal band, sutural line from base to apex or near apex, and marginal line from base or near base to apical fourth; whitish pubescence limited almost entirely to light- maculate areas. Female of Novelsis andersoni Integument brownish black without maculae or maculae, if present, more or less diffuse and difficult to see because of dense pubescence; setae on elytron unicolorous, whitish, or brown and whitish; if bicolorous then white setae forming two irregular bands on apical half or several longitudinal lines on apical half or white setae interspersed among dark setae on apical half. 8 8. Hind tibia bearing on posterior side numerous erect setae more than three times as long as spines along shaft of tibia. Elytron with whitish setae forming irregular circle on basal half and two bands on posterior half. Male antenna in repose extending almost to hind margin of metepisternum. Novelsis picta Hind tibia without long, erect setae but with usual long, recumbent setae through which project short, stout spines. Elytron entirely covered with whitish hairs (females) or with dark hairs among which whitish hairs form basal band, subbasal oblique band, sutural line, and several somewhat distinct lateral lines on apical half (males and occasional females). Male antenna in repose extending about as far as hind margin of middle coxa. Novelsis uteana 9. Pronotum with middle basal part prolonged backward to form broad, apically truncate lobe (Fig. 8). Anterior tibia with sharp, knife-like carina along dorsal margin; short, stout spines inserted in row along posterior side of carina and on posterior face of tibia but not on anterior face of tibia. Attagenus lobatus Pronotum produced only feebly backwards at middle basal part. Anterior tibia rounded on dorsal margin without knife-like carina; short, stout spines inserted irregularly along dorsal margin as well as on posterior face, with or without spines on anterior face of tibia. 10 1970] Beal: Taxonomic Study of Attagenini 157 10. Male antennal club with terminal segment at least four times as long as length of two preceding segments combined; female antennal club with terminal segment one-eighth longer than length of two preceding segments combined. Integument of elytron colored black, mahogany, or ma- hogany with base black and with black sutural line; elytral setae entirely black or black with sparsely intermingled golden-yellow hairs or black with band of golden-yellow hairs at basal fourth. Attagenus rufipennis Male and female antennal club with terminal segment equal in length or shorter than length of two preceding segments combined. Integument of elytron colored black, reddish yellow, or light brown with or without clearly defined maculae, but if black with suffused mahogany area on disc, then bands or patches of whitish pubescence present on apical as well as on basal area of elytron. 1 1 11. Hypomeron (ventrally indexed side of pronotum) deeply con- cave, forming fossa for antenna. Male antennal club clothed with erect, white setae about three-fourths as long as width of third antennal segment. Pubescence of elytron consist- ing of black hairs and light colored hairs; light colored hairs forming thin band at base, semicircular subbasal band, irregular submedian band, and irregular subapical band. 12 Hypomeron flat or slightly convex. Male antennal club clothed with closely appressed setae about as long as one-fourth width of third antennal segment. Pubescence of elytron various. 13 12. Elytral integument with poorly defined brown or reddish brown area on basal two-thirds and occasionally with reddish sub- apical spot. Plane of disc of prosternum and plane of lateral lobe of prosternum separated by step-like bend in proster- num. Novelsis aequalis Elytron with more or less sharply-defined reddish yellow or light tan macula occupying most of disc of elytron or with two or three distinct subbasal, median, and subapical maculae. Declivity in prosternum forming transi- tion between plane of disc of prosternum and plane of lateral lobe. Novelsis varicolor 13. Elytral integument reddish yellow to brown, immaculate. Light- colored pubescence of elytron forming broad band at basal 158 Entomologica Americana [Vol. 45, No. 3 Figs. 10-15: Spiracles and spiracular sclerites from first abdominal tergum of mature larvae of Attagenini. Sockets where setae are inserted are shown as circles, although the setae are not shown. In general the size of the seta is proportionate to the size of the socket. Fig. 10. Attagenus cyphonoides. Fig. 11. A. megatoma. Fig. 12. A. rufipennis (form from Santa Barbara, California). Fig. 13. Novelsis varicolor. Fig. 14. A. bicolor. Fig. 15. A. lobatus. 1970] Beal: Taxonomic Study of Attagenini 159 fourth, submedian band, and subapical band; submedian and subapical bands connected by sutural line of light pubescence. Novelsis timia Elytral integument dark brown to black with contrasting light tan or reddish yellow maculae; light maculate areas forming oblique subbasal band and sutural line from base to apex or near apex; light line along lateral margin present or absent. Light-colored pubescence mostly limited to light maculate areas of elytron; no light pubescence present behind middle of elytron except on light maculate sutural and marginal lines. 14 14. Marginal line of light maculation present on elytron and extending from base to posterior third or fourth. Male of Novelsis andersoni Marginal line of light maculation usually appearing as short spur on oblique subbasal band, but at most extending from base no further than middle of elytron. Novelsis horni 15. Elytron covered with dark hairs except for very well-defined, subbasal band of light pubescence; band usually broad and extending continuously to median suture but occasionally reduced to two light-colored narrow bands on middle of elytron at basal two-fifths and spot of light-colored hairs on median suture at basal fourth. Attagenus jasciatus Elytron unicolorous or with diffuse subbasal or basal band of light-colored hairs; subbasal band, if present, not extending to median suture and no subbasal patch of light-colored hairs present on suture. 16 16. Antenna 10-segmented. Ultimate segment of antennal club more than 6.0 times as long as length of two previous seg- ments combined. Attagenus schaefferi Antenna 1 1 -segmented. Ultimate segment of male antennal club less than 5.8 times as long as two previous segments of club combined. 17 17. Posterior margin of lateral lobe of prosternum bent almost vertically in front of coxa. Posterior ventral carina of middle femur much weaker than anterior ventral carina and on apical half of femur two-thirds as far from dorsal margin as anterior ventral carina. Elytral integument and hairs medium to dark brown with few light golden hairs forming indistinct lateral fascia at about basal fourth. Attagenus cyphonoides Figs. 16-25: Fig. 16. Seta from disc of first abdominal tergite of Attagenus megatoma. Fig. 17. Seta from second abdominal acroter- gite of Novelsis varicolor. Fig. 18. Seta from disc of first abdominal tergite of A. rufipennis. Fig. 19. Seta from disc of first abdominal tergite of N. varicolor. Fig. 20. Lanceolate seta from eighth ab- dominal tergite of A. pellio. Fig. 22. Scale-like seta from disc of eighth abdominal tergite of A. pellio. Fig. 23. Apically bifurcated scale-like seta from first abdominal tergite of N. aequalis. Fig. 24. Spiracle and spiracular sclerite of sixth abdominal tergum of mature larva of N. aequalis. Fig. 25. Spiracle and spiracular sclerite of seventh abdominal tergum of A. pellio. 1970] Beal: Taxonomic Study of Attagenini 161 Posterior margin of lateral lobe of prosternum weakly reflected against base of procoxa with reflected portion extending ventrad at about thirty to forty degree angle. Posterior ventral carina of middle femur about as distinct as anterior ventral carina and almost in same plane. Elytral pubes- cense piceous to black; light-colored hairs, if present, form- ing fascia at base of elytron or limited to extreme lateral margin of elytron; integument of elytron mahogany to black. 1 8 18. Ratio of length of terminal segment of antenna of male to length of pronotum and elytra combined varying from 1:5.7 to 1:7.6. Pronotum with band of golden hairs along basal margin; greatest length of band (measured anteriorly to posteriorly) about equal to length of scutellum. Elytron with golden hairs absent or limited to scattered hairs or small patches of hair along basal margin and not extending posteriad beyond apex of scutellum except along extreme lateral margin. Attagenus elongatulus Ratio of length of terminal segment of antenna of male to length of pronotum and elytra combined varying from 1:7.9 to 1:11.4. Pronotum without band of golden hairs along basal margin, or band present and not longer than half length of scutellum, or latero-posterior angles and basal margin covered with golden hairs. Elytron without golden hairs or varying number of golden hairs present on basal fourth; if golden hairs present on base of elytron, these commonly extending from base well beyond scutellum. Attagenus megatoma Known Mature Larvae 1. Some broad, scale-like setae with five longitudinal ribs between margins inserted on acrotergites; most setae of tergite of eighth abdominal segment either subrectangular in outline (Fig. 22) or ovate (Figs. 19, 23). 2 All setae of acrotergites slender and usually with not more than three longitudinal ribs visible between margins; setae of tergite of eighth abdominal segment either lanceolate or linear (Figs. 16, 18). 4 2. All setae with smooth margins; scale-like setae of anterior terga broadly lanceolate (Fig. 21); scale-like setae of pos- terior terga subrectangular (Fig. 22). Attagenus pellio 162 Entomologica Americana [Vol. 45, No. 3 All setae with denticulate margins; scale-like setae of all terga ovate-lanceolate and ovate in form (Figs. 19, 23). 3 3. No antecostal suture present on eighth abdominal tergum. Setae of terga consisting of scale-like hairs only, except for elongate marginal and submarginal erect setae; all setae of discs of terga with single acute apex. Novelsis varicolor Antecostal suture present on eighth abdominal tergum. Setae of terga consisting of scale-like hairs with some intermingled linear hairs in addition to elongate marginal and submarginal erect setae; some setae on discs of terga with single acute apex and some with two acute apices (Fig. 23). Novelsis aequalis 4. No antecostal suture on eighth abdominal tergum. Spiracular sclerite closed behind by tergum; spiracle enclosed by tergum with distance between spiracle and posterior margin of tergum on first abdominal segment equal to more than one-fourth length of tergite (Fig. 14). ____ Attagenus bicolor Antecostal suture present on eighth abdominal tergum. Spirac- ular sclerite closed behind by tergum or not, but if closed then distance between spiracle and posterior margin of tergum equal at most to one-sixth length of tergite at midline. 5 5. Setae of dorsum (excluding head) grayish white (transparent in cleared specimens) and contrasting sharply with brownish black setae of abdominal sterna. Integument of each notum with large blackish lateral spot and several smaller blackish spots. Novelsis uteana Setae of dorsum and setae of abdominal sterna quite similar in color (although setae of head and thoracic sterna sometimes lighter than other setae). Integument of nota not marked with darker spots or, if darker in middle, then not differing in appearance from terga. 6 6. Some setae of eighth abdominal sternum broadly lanceolate with as many as 7 ribs between margins. 7 All setae of eighth abdominal sternum linear or narrowly lanceo- late with at most no more than 4 ribs between margins. __ 9 7. Disc of sterna with lanceolate setae only; row of stout linear setae one to two times as long as lanceolate setae near anterior margin of each abdominal sternum; numerous fine, long, simple setae present anterior to row of stout linear setae. Spiracular sclerite bearing 10 or more setae (Fig. 15). Attagenus lobatus 1970] Beal: Taxonomic Study of Attagenini 163 Anterior abdominal sterna with linear and linear-lanceolate setae; seventh and eighth abdominal sterna with inter- mingled lanceolate setae and linear setae; setae becoming increasingly smaller toward anterior area of sternum with- out anterior row of stout linear setae. Spiracular sclerite bearing no more than 6 and usually fewer than 6 setae. __ 8 8. Broadest setae of first abdominal tergite with 4 ribs between margins; broadest setae of eighth abdominal sternum usually with no more than 9 ribs between margins. Attagenus schaefferi Few broad setae of first abdominal tergite with 5 ribs between margins; broadest setae of eighth abdominal sternum in fully mature specimens with 1 1 or 12 ribs between margins. Attagenus elongatulus 9. Second antennal segment bearing 12 or more setae. Attagenus fasciatus Second antennal segment bearing no more than 3 and often 2 setae. 10 10. Tergum sclerotized continuously anterior to entire length of tergal margin of spiracular sclerite; spiracle closed behind by tergum, at least on anterior abdominal segments (Figs. 10, 11). Setae inserted on pretarsus subequal in width and length. 11 Tergum not sclerotized continuously anterior to spiracular sclerite, so that sclerite with associated tergal margin ap- pears to project peninsula-like from postero-lateral angle of tergum. Spiracle opening on margin or at apparent margin of tergum (Fig. 12). Setae inserted on pretarsus unequal, one about twice as wide as other. 12 1 1 . Setae of head identical in color with setae on disc of pronotum (although setae on lateral margins of pronotum may ap- pear darker). Integument of dorsum dark reddish brown except for teneral specimens. All setae on disc of tergites lanceolate except for submarginal erect setae and marginal linear setae. Attagenus megatoma Setae of head distinctly lighter in color than setae of nota and terga. Dorsal integument straw-colored. Disc of tergites with about equal number of lanceolate and linear setae. Attagenus cyphonoides 12. Setae of head brownish. Setae of disc of tergites linear and spindle-shaped; margins of setae minutely microserrulate 164 Entomologica Americana [Vol. 45, No. 3 Figs. 26-28: Posterior view of front tibiae and pretarsi of mature larvae. Fig. 26. Attagenus rufipennis. Fig. 27. A. fasciatus. Fig. 28. A. lobatus. (serrations scarcely visible except under high powers of compound microscope) or smooth. One sensory pit close to socket of each submarginal erect seta on abdomen. Epipharynx with two distal sensory pits lacking. Novelsis horni and Novelsis andersoni 1970] Beal: Taxonomic Study of Attagenini 165 Setae of head black. Setae of disc of tergites lanceolate; margins of setae serrulate or denticulate (visible under low powers of compound microscope). Two sensory pits, one on each side, adjacent to socket of each submarginal erect seta on abdomen. Epipharynx with two minute sensory pits distal to median series of 6 sensory pits. _ Attagenus rufipennis Attagenus cyphonoides Reitter Attagenus cyphonoides Reitter, 1880 (1881), p. 34. — Zhantiev, 1963b, p. 416. ' Attagenus alfierii Pic, 1910, p. 17. — Hinton, 1943, p. 227; 1945, pp. 322- 323. Trogoderma cyphonoides : Arrow, 1915, p. 427. Adults: Described adequately by Hinton (1945). Mature Larvae: Dorsal integument straw-colored. Setae of head golden-yellow; setae of body and legs golden-brown. Head : Terminal seta of antenna not observed; two setae present near apex of second segment. Epipharynx with distal series of two sensory pits, each subequal in diameter to sensory pits of middle series. Maxillary palp with two setae on penultimate segment. Labium with five to six setae on dorsal (inner) surface of each side of ligula; middle two to three broadly ensiform; outer two to three narrowly ensiform to terete; first segment of palp without seta. Body setae : Margins and ribs uniformly smooth. Setae inserted on acrotergites all linear, widest with no more than one rib visible between margins. Disc of tergites with linear setae and lanceolate setae; widest lanceolate setae with no more than three ribs visible between margins. Sockets of most submarginal erect setae with two adjacent sensory pits, one on either side near anterior margin. First abdominal sternum with setae at midline longer than other setae of disc; longer setae covered with minute, sharply pointed, imbricate scales; similar setae present along midline of thoracic segment*. Setae of sternum of eighth abdominal segment all linear-lanceolate; widest with no more than two ribs between margins; setae becoming smaller on anterior area of sternum; no anterior row of long, stout setae. Spiracle and associated structures : Spiracle closed behind by tergum; anterior margin more or less straight; posterior margin somewhat thickened. Spiracle somewhat separated from spiracular sclerite (Fig. 10). Spiracular sclerite subcircular to oval, bearing no more than six setae, not enclosed by tergum. Tergum sclerotized 166 Entomologica Americana [Vol. 45, No. 3 anterior to tergal margin of spiracular sclerite and continuously with it. Abdomen : Eighth abdominal tergum with well-defined ante- costal suture. Ninth abdominal tergum about half as long as length of tergite of eighth abdominal segment. Two small sclerites present in sternum of ninth abdominal segment. Legs : Femur of pro thoracic leg about one-half as wide as long; ventral margin about six-thirteenths as long as length of seg- ment. Tibia of prothoracic leg with usual two ventral rows of spines, one subdorsal apical spine, and row of one to three spines parallel to and very near ventral row. Setae inserted at base of pretarsus subequal in width, extending about as far as middle of claw. Recorded Distribution: The type locality for A. cyphonoides is southern Egypt. A. alfierii was also described from Egypt. It occurs as well in Tunisia and Morocco (Zhantiev, 1963a). New Records: Alabama : Brawley, August 27, 1954 (C. H. Rothe). Arizona : Buckeye, June 27, 1958 (W. B. Morrow); Chloride, July 16, 1957 (“L. L. G.”); Elfrida, June 23, 1954 (K. S. Rohwer); Elgin, June, 1954 (J. F. Brantlinger) ; Gila Bend, June 13, 1957 (L. L. Garrison); Mayer, June 20, 1957 (R. A. Murphy); Mesa, May 8, 1957 (A. M. Mroczkiewicz ) ; Phoenix, May 23, 1957 (A. M. Mroczkiewicz); Randolph (Pinal County), September 27, 1957 (D. A. Bercich); Safford, September 20, 1957 (“W. B. F.”); Tucson, July 28, 1954 (R. S. Beal); Tucson, August 11, 1955 (J. R. Boyer). Kansas : Dodge City, August 26, 1954 (D. L. Matthew). New Mexico : Las Cruces, August 15, 1957 (D. Lucht). Baja California : Tecate, May 19, 1955. Diagnosis: The integument of the adults varies from yellowish brown to a dark chestnut brown. Most of the dorsal hairs are dark brown. The elytra appear unicolorous, but close inspection of each shows an oblique band formed by a few light, golden hairs on the lateral half at the basal fourth. The antenna is 1 1 -segmented. The first two segments of the male antennal club are subequal, and the terminal segment is nearly five times as long as the first two com- bined. The prosternum has the lateral lobe bent almost vertically in front of the procoxa to form a narrow, transverse, razor-like edge. This character readily separates A. cyphonoides from all other species in Group I except A. fasciatus, from which it is readily distinguished by the pattern of dorsal pubescence. The middle and hind femora do not have the two ventral carinae on the same or nearly the same plane, as do the other species treated here. The 1970] Beal: Taxonomic Study of Attagenini 167 posterior ventral carina on the apical half of the middle femur lies about two-thirds as far from the dorsal margin as does the anterior ventral carina. Larvae of A. cyphonoides, A. megatoma, and A. fasciatus are generally quite similar. They may be distinguished from others in Group I by the relatively narrow setae of the abdominal tergites and of the eighth abdominal sternum. The broadest setae in these three species have no more than three ribs between the margins. Mature larvae of A. cyphonoides differ from those of A. fasciatus in that there are two setae rather than 12 or more on the second segment of the antenna. They differ from larvae of A. megatoma in having two rather than no setae on the second segment. They differ from larvae of both A. megatoma and A. fasciatus in having the setae of the head distinctly lighter in color than the setae of the nota and terga, rather than having the setae of head and dorsum nearly the same color. Attagenus megatoma (Fabricius) Adult Males: Integument of dorsal and ventral surfaces im- maculate, reddish brown to black; antennal club usually with all segments piceous; legs and flagellar shaft of antenna reddish brown to yellowish brown. Pubescence of head and dorsal surfaces sub- recumbent, entirely black or black with varying numbers of light golden-brown hairs on pronotum and base of elytra; ventral pu- bescence recumbent, light golden-brown except for black hairs on posterior half of fifth visible sternum. Eye slightly emarginate over base of antenna. Antenna 1 1 -segmented; ratio of combined length of first two segments of club to length of terminal segment varying from 1:2.8 to 1:3.4; ratio of length of terminal segment to length of pronotum and elytra combined varying from 1:9.00 to 1:11.77; club clothed with fine, erect hairs about one-third as long as width of third antennal segment. Channel below eye for reception of flagellar shaft of antenna concave with anterior margin forming carina visible from front of head (Fig. 3); carina projecting knife- like beneath head and curved behind base of maxilla to meet gular suture. Pronotum with lateral carina continued around anterolateral angle; basal lobe feebly rounded to slightly truncate and not dis- tinctly produced backward; setae on posterior margin of lobe not appreciably longer than other setae of pronotum. Punctures of pronotal disc varying in diameter from width of one to width of two diameters of facet of eye; punctures nearly contiguous laterally, 168 Entomologica Americana [Vol. 45, No. 3 especially on anterior half of pronotum, with anterior margins more sharply defined than posterior margins so that pronotum somewhat transversely rugose. Elytron with punctures of disc about twice diameter of facet of eye and separated by one to two diameters of single puncture. Hairs of disc about as long as combined length of fourth and fifth segments of antenna. Hypomeron slightly concave; forming lobe behind trochantin so that trochantin visible only through narrow slit (Fig. 9). Prosternum with posterior margin of lateral lobe reflected against procoxa; reflected part about as long as length of horizontal part of lobe and forming angle of about 45° with horizontal part; prosternal process slightly wider at apex than between coxae; ventral surface of process thin and knife-like; no thread-like carina extending from process onto disc, but disc longi- tudinally elevated at middle and broadly carina-like; no transverse thread-like carina present near anterior margin of disc, but denticle present at middle near anterior margin. Epipleuron terminating a little behind hind margin of metepimeron. Ventral plate of hind coxa forming more or less acute tooth lateral to insertion of trochanter; plate extending laterad behind posterior margin of metepisternum and meeting metepimeron (actually extending beneath inner margin of metepimeron). Protibia not carinate on dorsal margin. Mesofemur with anteroventral and posteroventral margins of crural cavity about equally produced and nearly on same plane. Length (of pronotum and elytra combined): 2.8 mm. to 4.0 mm. Adult Females: Antennal club with first two segments usu- ally reddish brown and terminal segment piceous; terminal segment subequal to one and one-half times length of ninth and tenth segments combined; tenth segment about five-sixths as long as ninth segment. Length (of pronotum and elytra combined) : 3.9 mm. to 5.0 mm. Mature Larvae : Integument of head and dorsum dark reddish brown. Setae of head and body dark golden-brown. Head : Antenna with terminal seta subequal in length to length of terminal segment; second segment without setae. Epipharynx with distal series of two sensory pits, each subequal in diameter to di- ameter of sensory pits of middle series. Maxillary palp with two and occasionally three setae on third segment. Labium usually with six setae on dorsal (inner) surface of each lobe of ligula; three inner setae ensiform; three outer setae slender; first segment of palp with- out seta. Body setae : Margins and ribs uniformly smooth. Acrotergites with small, linear-lanceolate, indistinctly ribbed setae and numerous 1970] Beal: Taxonomic Study of Attagenini 169 simple setae. Setae on tergites linear-lanceolate with two or three ribs between margins (Fig. 16). Sockets of submarginal erect setae usually with two adjacent sensory pits, one on each side. Sterna with linear-lanceolate setae with two ribs between margins and oc- casionally some with three ribs between margins; setae becoming smaller on anterior area of each sternum without row of stouter setae near anterior margin; first and second abdominal sterna with several setae along midline much longer than other setae of disc. Spiracle and associated structures : Spiracle closed behind by tergum; anterior margin more or less straight; posterior margin some- what thickened; spiracle slightly removed from spiracular sclerite. Spiracular sclerite oval, not enclosed by tergum, bearing six or fewer setae. Tergum sclerotized anterior to tergal margin of spiracular sclerite and continuous with it (Fig. 11). Abdomen : Eighth abdominal tergum with antecostal suture. Ninth abdominal tergum about two-thirds as long as tergite of eighth segment. Two small sclerites in sternum of ninth abdominal segment. Legs : Femur of prothoracic leg half as wide as long; ventral margin two-fifths as long as length of segment. Tibia of prothoracic leg with usual two rows of ventral setae; posterior face with subdorsal apical spine and row of five to seven subventral spines above posterior row of ventral spines. Setae inserted at base of pretarsus subequal in width and length, extending almost to middle of claw. Internal Anatomy: Details of the digestive tract, Malpighian tubules, bands of oenocytes, and reproductive systems have been described by Dunkel and Boush (1968a). The internal anatomy of no other species in the tribe has been investigated, so no comparative data are available for evaluating the significance of the interesting structures which these authors found. Nomenclature: Although the name A. piceus has been ap- plied to several species in the United States, it properly applies to this species alone. However, the name is unavailable, since Dermestes piceus Olivier (1790) is a primary junior homonym of Dermestes piceus Thunberg (1781) and was treated as such by early authors. The oldest available name is Dermestes unicolor Brahm (1791). This name was not used by any subsequent author until Mroczkowski (1968) recognized the identity of Brahm’s type and listed the name in his catalogue. It is unfortunate that it was not treated by Mroczkowski as a nomen oblitum. A request is currently before the International Commission on Zoological Nomenclature to suppress the name and to conserve the name used here. 170 Entomologica Americana [Vol. 45, No. 3 The next available name for the species is Dermestes megatoma Fabricius (1798). Latreille erected the genus Attagenus in 1802, and after that almost all authors used the combination A. megatoma for the species until Mulsant and Rey in 1868 resurrected the name A. piceus. Lacking a code, many nineteenth century authors simply accepted the authority of Mulsant and Rey, and the propriety of the name was not subsequently questioned by any American writer. Nevertheless, the name A. megatoma has been used by a number of recent European workers, as can be seen from the table of synonymy for A . megatoma megatoma. Distribution in North America: This is a polytypic species, probably originally Holarctic in distribution but now nearly cosmo- politan as a result of introductions by commerce. Two geograph- ically distinct forms are clearly present in the Nearctic region: one occurring mostly in Canada and in the northern tier of states but dipping as far south as Utah and Iowa, the other widespread through- out the remaining states. The latter (southern) form is indistinguish- able from the nominate European subspecies and may represent an early introduction to the United States. The northern form re- sembles the Far Eastern A. megatoma japonicus Reitter and possibly originated from it, migrating across the former land bridge between Alaska and northeastern Asia. In spite of the resemblances, how- ever, there appear to be enough differences between the forms that they should be regarded as distinct subspecies. Casey (1916) named the northern form A. canadensis , con- sidering it a distinct species. I believe it deserves no more than subspecific status, because intergrades are frequently found in areas where it comes together with the southern form and because cross- breeding experiments seem to show that gene exchange between the two forms is possible. In several experiments first generation hybrids were readily formed between a stock culture of “ megatoma ” obtained from Prescott, Arizona, and a stock culture of “ canadensis ” from Vancouver, British Columbia. When the hybrids were mated, second generation progeny were produced and developed into adults in several, but not all, of the experiments. The failures in the second generation crosses do not necessarily mean there is reduced fertility between the hybrids, since I was compelled to move my laboratory during the experiments, and the cultures were not kept in an ade- quately moist atmosphere for some length of time. At any rate, the ability of some hybrids to mate and produce viable progeny seems 1970] Beal: Taxonomic Study of Attagenini 171 established and indicates the probability of the same thing taking place in the wild state. The accompanying map (Fig. 29) shows the distribution of the two forms. Most specimens of A. megatoma megatoma have no golden-brown hairs inserted on the elytra. Most specimens of A. megatoma canadensis have golden-brown hairs inserted in some numbers on the base of the elytra back to a distance at least equal to three lengths of the scutellum. First generation hybrids between such forms have golden-brown hairs inserted on the base of the elytra no further back than one length of the scutellum. Using the hybrids as a rough standard, I have designated as an “intergrade” any specimen with golden-brown hairs on the base of the elytra where the hairs are inserted no further back than one and one-half times the length of the scutellum. Localities where such intergrades have been found are shown on the map with a triangle, even though one or both of the other forms may have been found in the same locality. The pattern of distribution exhibits a less regular line of demarcation between the forms than might be expected, but this irregularity is probably explained by the movement of the forms through commerce. If A. megatoma megatoma is an indigenous sub- species, I doubt that western Oregon and Washington are within its original range. Ecology and Life History: Hinton (1945) summarized the immense amount of work done on the life history, habits, and economic importance of the species. Regrettably, there is no way of knowing which of the studies listed refer to A. megatoma and which to A. elongatulus. Probably the life histories are very similar. Nevertheless, it would be unsafe to assert that any of the facts cited are true of A. megatoma without access to the specimens used in the experiments or without replicating the experiments with ac- curately identified and unmixed cultures. It is particularly un- fortunate that the experiments of Griswold and Greenwald (1941) on the life history of “A. piceus ” cannot at present be assigned definitely to one species or the other. Most of the reliable available information on the ecology of the species has been gathered for A . megatoma megatoma. The informa- tion that follows refers to this subspecies, unless otherwise noted. In each instance the identity of the species and subspecies has been verified. Out-of-doors the species is not uncommonly found in various bird nests. It was taken by E. G. Linsley at Madera, Cali- fornia, in the nests of cliff swallows, by Mrs. K. B. Wetherbee in a 172 Entomologica Americana [Vol. 45, No. 3 tree swallow nest in Connecticut, by W. L. McAtee in a starling nest at Bell, Maryland, and by A. T. McClay in an English sparrow nest at Medford, Oregon. J. A. Payne collected one larva in owl pellets at Clemson, South Carolina. In addition to these records, Ellis A. Hicks (1959) has assembled a long list of references to the occurrence of the species in various kinds of bird nests. Probably most of the references apply to this species, but very likely some refer to A. elongatulus or other similar-appearing species. The species is not limited to bird nests and possibly may occur as commonly in some other out-of-door situations that up to the present have not been as thoroughly canvassed. I collected the species in a Sceliphron mud dauber nest at Chino Valley, Arizona. Johansen and Eves (1966) published a report of A. megatoma canadensis (under the name A. piceus) as an occasional pest in the Pacific Northwest in nests of the alfalfa leafcutting bee, Megachile rotundata (Fabri- cius). E. S. Ross found a specimen in a Geomys (pocket gopher) burrow in Somerset, Texas. The species is not uncommonly found in human habitations, although probably not as frequently now since the advent of synthetic fibers in the manufacture of rugs, overstuffed furniture, and the like. It is probably best known as an inhabitant of granaries, seldom in any numbers in the grain bulk, unless the store is fairly old, but rather in grain dust on the stringers and on old spilled grain in corners and under floor boards. Triplehorn (1965) carried out a survey of insects in Ohio grain elevators and feed mills and found it in 110 out of 118 establishments investigated, a far higher rate of occurrence than found for any other insect. A. megatoma canadensis was found by Chao (1954) to be similarly prevalent in grain eleva- tors at Albion and Pullman, Washington. Following a survey he assigned five categories of frequency to the insects found and rated this species (under the name A. piceus) in the most frequently occurring group. The feeding habits of the larvae have been investigated in the laboratory, though not in comparison with other species. Hence, such questions as whether this species or A. elongatulus is the more important depredator of woolens and other animal fibers remain unanswered. Mallis, Miller, and Hill (1958) found that the species will feed extensively on wool and on various combinations of wool and synthetic fabrics. It will feed to a slight extent on silk crepe and on nylon but little or not at all on dynel, dacron, orlon, vicara, acetate rayon, viscose rayon, linen, and cotton percale. It will feed more extensively on nylon stained with human perspiration, but 1970] Beal: Taxonomic Study of Attagenini 173 Fig. 29. Nearctic distribution of subspecies of Attagenus megatoma : + = A. megatoma megatoma, % — A. megatoma canadensis, A = intergrades between subspecies. See text for further explanation. not on nylon stained with human urine or tomato juice (Mallis, Miller, and Hill, 1959). I have been able to rear the larvae success- fully on dried dog food and on oatmeal. The species has been found infesting cereal products including bran, barley, ground Indian corn, in-shell peanuts, cake mix, and alfalfa meal. The species was found infesting stored raisins in Fresno, California, but may have been feeding on the remains of other insect pests rather than on the raisins. Little accurate information is available on the kinds of food that will permit maturation of the organism or that might govern the rate of maturation. Boush, Dunkel, and Burkholder (1969) have found that sorbic acid incorporated in the diet does not affect larval or adult weight, survival to adult stage, or pupal or adult longevity. However, mated pairs fed the chemical produced progeny unable to survive the first larval instar. The significance of this finding is not clear at the present time. Mallis, Burton, and Miller (1962) experimented with the extent to which the larvae might be attracted to various food stains, in- cluding minerals, proteins, B vitamins, cholesterol, vitamin D, vita- min A, and cottonseed oil, and found them relatively unattracted to any. There was some positive attraction to stains made by glucose, orange juice, and tomato juice. Small but not large larvae were 174 Entomologica Americana [Vol. 45, No. 3 attracted definitely to stains made by human urine, tomato juice, and beer, but only at times to stains made by human perspiration, milk, black coffee, and beef gravy (Mallis, Miller, and Hill, 1959). No larvae were attracted to tea, cola syrup, or butter stains. Adults reared in the laboratory regularly mate and lay fertile eggs without the opportunity of flying and without being provided with water or food other than the cereal on which the larvae are reared. Out-of-doors adults have been taken on flowers of Spiraea (Mroczkowski, 1958), Achillea , and Costarica. Adults of A. mega- toma canadensis have been collected on alsike clover and on sweet clover. Whether adults feed on the pollen or nectar of the flowers has not been ascertained. It is possible that attraction to flowers is a mechanism bringing individuals together for purposes of mating. Yokoyama (1932, cited in Hinton, 1945) noted that mating usually takes place on a flower or some other place exposed to the sun. Symbionts: The eugregarine Pyxinia frenzeli Laveran and Mesnil has been found within cells of the midgut epithelium of the larvae (Dunkel and Boush, 1968b). The protozoan is not found in the adult, except that newly emerged adults contain what is presumed to be a remnant of the larval midgut in the form of an elongated tube, closed at both ends, within which spores and encysted forms are found (Dunkel and Boush, 1968a). The tube is probably eliminated in the first defecation of the adult. In all probability the protozoan should be considered a commensal rather than a parasite. Heavily infested individuals do not show differences in weight, survival, or length of life cycle compared with non-infested indi- viduals (Dunkel and Boush, 1968b). No survey has been made of the extent to which populations are infested in different parts of the range of the species. The specimens studied by Dunkel and Boush were from a laboratory strain maintained at the University of Wisconsin. The muscardine fungus Beauveria bassiana (Bals.) Vuill. has been recorded as a parasite of the species (Charles, 1941). This is a generalized parasite best known from its occurrence on the chinch bug and the European corn borer. The identity of the species of Attagenus on which it was described might be questioned. Possibly it attacks all species in the group to which A. megatoma belongs, but this theory should be verified. Attagenus megatoma megatoma (Fabricius) Dermestes piceus Olivier, 1790 ( non Thunberg, 1781), p. 10, pi. 1, fig. 4. Dermestes unicolor Brahm, 1791, p. 144. 1970] Beal: Taxonomic Study of Attagenini 175 Dermestes megatoma Fabricius, 1798, p. 313. Attagenus megatoma : Dahl, 1823, p. 30. — Laporte, 1840, p. 35. — Erich- son, 1846, p. 441. — Wollaston, 1854, p. 204. — LeConte, 1854, p. 109. — Chevrolat, 1863, p. 616. — Howe, 1952, p. 40. — Mroczkowski, 1954, p. 7; 1958, p. 4; 1962, p. 6; 1965, p. 668.— Zhantiev, 1963b, p. 421. Attagenus piceus : Mulsant and Rey, 1867 (1868), p. 69. — Jayne, 1882, p. 355. — Casey, 1900, p. 146. — Ganglbauer, 1904, p. 24. — Lutz, 1911, p. 152. — Mutchler and Weiss, 1927, pp. 10-11. — Moore and Moore, 1942, p. 288. — Rees, 1943, pp. 14-17. — Hinton, 1943, p. 227; 1945, pp. 309-319. — Korschefsky, 1944, p. 147. — Armstrong, 1945, p. 48. — Zinkernagle, 1952, p. 844. — Zinkernagle and Muller, 1952, p. 156.— Hatch, 1962, p. 283. Attagenus cylindricornis Casey, 1900 ( ?non cylindricornis Say, 1825), p. 147 (New synonymy). Attagenus deficiens Casey, 1900, p. 146 (New synonymy). Adult Males: Dorsal pubescence black except for few golden hairs at basal third along extreme lateral margin of pronotum and narrow band of short golden to golden-brown hairs along basal margin of pronotum; pronotal band not always distinguishable but when visible not more than half length of scutellum at longest point; often few dark golden hairs scattered among black hairs of pronotum, particularly at sides; rarely dorsal pubescence entirely black. Ratio of length of last segment of antennal club to combined length of pronotum and elytra varying from 1:7.9 to 1:9.9. Distribution: The distribution of the subspecies in the United States in shown in Figure 29. Diagnosis: This subspecies is easily recognized by its almost entirely black dorsal pubescence. Some golden-brown or golden hairs may be scattered among the black hairs of the pronotum, particularly on the sides, but they do not form patches that contrast with the dark discal area. The last segment of the antennal club of the male tends to be a little longer in proportion to the length of the body than does the last segment of the club in A. megatoma canadensis. Attagenus megatoma canadensis Casey, new status Attagenus canadensis Casey, 1916, p. 183. Attagenus piceus : Spencer, 1947 (1948), p. 7. Attagenus piceus ab. sordidus Hatch, 1962 ( non Heer, 1841), p. 283. Adult Males: Dorsal pubescence black except for concentra- tion of golden to golden-brown hairs on posterior margin and latero- posterior angles of pronotum and on base of elytra; light-colored 176 Entomologica Americana [Vol. 45, No. 3 hairs of pronotum occasionally covering most of sides of pronotum; light-colored hairs of elytra quite dense at base but intermingled with increasing numbers of black hairs posteriad; light-colored hairs in- serted posteriad two to four lengths of scutellum from base. Ratio of length of terminal segment of antennal club to length of pronotum and elytra combined varying from 1 :9.0 to 1:1 1.4. Type Locality: Ottawa, Canada. The specimen that has been labeled as the type in the Thomas L. Casey Collection at the United States National Museum bears a somewhat illegible locality label that might possibly be read “Ott.” Since Ottawa is one of the localities Casey indicated for the species, it is reasonable to limit it as the type locality. I herewith formally designate the specimen, which is a female, as the lectotype. Distribution: The known distribution is shown in Figure 29. Additional collecting is needed to establish the range of distribution of the subspecies northward in Canada and to clarify its points of contact with the nominate subspecies. Diagnosis: Adults of the subspecies may be distinguished from the nominate subspecies by the golden to golden-brown hairs that occupy the lateroposterior angles of the pronotum and form patches with almost no intermingled dark hairs. The light-colored patches stand out in contrast to the dark disc of the pronotum. In addition, there is a band of light-colored hairs on the base of the elytra. Posteriad it may be broken into somewhat irregular lines or may be intermixed with increasing numbers of dark hairs. Usually no light-colored hairs are inserted further back from the base than four times the length of the scutellum, often no further back than two times the length of the scutellum. Forms with light-colored hairs no further back than one and one-half times the length of the scutellum appear to occur only in the zones of intergradation between the two subspecies. No differences have been observed between larval forms of the two subspecies. Attagenus elongatulus Casey Attagenus elongatulus Casey, 1900, p. 147. Attagenus extricatus Casey, 1900, p. 146 (New synonymy). Attagenus bicolor Casey, 1900 ( non bicolor Harold, 1868), p. 147. Adult Males: Integument of body yellowish or reddish brown to black; elytra often somewhat lighter in color than pronotum; legs and antenna yellowish brown except for darker brown terminal antennal segment. Pubescence of head and dorsal surfaces sub- 1970] Beal: Taxonomic Study of Attagenini 177 Fig. 30. Nearctic distribution of Attagenus elongatulus. recumbent, black except for band of light golden-brown hairs along basal margin of pronotum; length of light-colored band at widest point (i.e., longest anterior-to-posterior distance) four-fifths to one and one-fourth times as long as scutellum; additional light golden- brown hairs either present or not along lateral margins of pronotum, along anterior margin of pronotum, along lateral margins of elytra at basal fourth, and at base of elytra; if golden brown hairs present at base of elytra these intermingled with black hairs and not inserted posteriad beyond length of scutellum; ventral pubescence recumbent, light golden-brown except for numerous black hairs on posterior half of fourth and on fifth visible sterna. Head with eye slightly emarginate over base of antenna. Antenna 11 -segmented; ratio of combined length of first two segments of club to length of terminal segment varying from 1:3.2 to 1:4.2; ratio of length of terminal segment to length of pronotum and elytra combined varying from 1:6.2 to 1:7.6; club clothed with fine, erect hairs about one-third as long as width of third antennal segment. Channel below eye for recep- tion of flagellar shaft of antenna concave with anterior margin forming carina; margin of carina barely visible from front of head; carina projecting knife-like beneath head and curved behind base of maxilla to meet gular suture. Pronotum with lateral carina continued as well-defined carina around anterolateral angle; basal lobe feebly 178 Entomologica Americana [Vol. 45, No. 3 rounded and not distinctly produced posteriad; setae on posterior margin of lobe about twice as long as light-colored setae on basal margin on either side of lobe. Punctures of pronotal disc varying in diameter from width of one to width of two facets of eye; anterior margins more sharply defined than posterior margins with punctures nearly contiguous laterally, especially on anterior half of pronotum, so that pronotum somewhat transversely rugose. Elytron with punc- tures of disc about twice diameter of facet of eye and separated by one to two diameters of single puncture. Hairs of disc of elytron no longer than combined length of third and fourth segments of antenna. Hypomeron moderately concave. Posternum with posterior margin of lateral lobe reflected against procoxa; reflected part about as long as horizontal part of lobe and forming angle of about 45° with horizontal part; prosternal process slightly wider at apex than between coxae; carina extending from apex of process onto disc and terminating at denticle near anterior margin of disc; carina broad and granulate on disc but thread-like on process; lateral carina present near anterior margin separating disc from anterior declivity. Epipleuron terminating little behind hind margin of metepimeron. Ventral plate of hind coxa somewhat expanded laterad to insertion of trochanter to form obtusely angled or somewhat rounding tooth; plate extending laterad behind posterior margin of metepisternum and meeting metepimeron (actually extending beneath inner margin of metepimeron) (Fig. 4). Protibia not carinate on dorsal margin. Mesofemur with anteroventral and posteroventral margins of crural cavity about equally produced and nearly on same plane. Length (of pronotum and elytra combined) : 2.9 mm. to 3.9 mm. Adult Females: Antennal club with first two segment yellow- ish brown to dark brown; all segments usually same color but oc- casionally terminal segment darker; terminal segment subequal to one and one-half times as long as ninth and tenth segments combined; tenth segment about five-sixths as long as ninth segment. Length (of pronotum and elytra combined): 3.6 mm. to 5.0 mm. Mature Larvae : Integument of head and dorsum light brown. Setae of head and body golden-brown. Head : Antenna with terminal segment subequal in length to length of third segment; no setae present on second segment. Epipharynx with distal series of two sensory pits, each subequal in diameter to sensory pits of middle series. Maxillary palp with two setae on penultimate segment. Labium with six to seven setae on dorsal (inner) side of each lobe of ligula; medial setae narrowly 1970] Beal: Taxonomic Study of Attagenini 179 ensiform with setae becoming progressively narrower laterad; first segment of palp without seta. Body setae : Margins and ribs uniformly smooth. Setae inserted on acrotergites all linear, most with no more than one rib between margins but some with two and rarely with three ribs between margins. Disc of tergite with linear and linear-lanceolate setae; linear-lanceolate setae with as many as five ribs between margins. Sockets of submarginal erect setae with two adjacent sensory pits, one on each side near anterior margin. First abdominal sternum with setae along midline longer than on remainder of disc. Setae of sternum of eighth abdominal segment consisting of linear-lanceo- late and lanceolate hairs; lanceolate setae with eight to twelve ribs between margins (Fig. 20); setae becoming smaller on anterior area of sternum without anterior row of stouter setae. Spiracle and associated structures : Anterior abdominal seg- ments with spiracle closed behind by tergum; anterior margin of spiracle more or less straight; posterior margin somewhat thickened; spiracle adjacent to spiracular sclerite. Spiracular sclerite oval, not enclosed by tergum; five or fewer setae inserted on sclerite. Tergum sclerotized anterior to tergal margin of spiracular sclerite and con- tinuous with it (similar to Fig. 11). Abdomen : Eighth abdominal tergum with well-defined ante- costal suture. Ninth abdominal tergum more than half as long as tergite of eighth abdominal segment. Two small sclerites present in sternum of ninth abdominal segment. Legs : Femur of prothoracic leg about 9/20 as wide as long; ventral margin about 9/20 as long as length of segment. Tibia of prothoracic leg with usual two rows of ventral setae; posterior face with subdorsal apical seta and longitudinal row of three setae along middle of segment; most proximal seta in row inserted little before middle of segment. Setae inserted at base of pretarsus subequal in width, extending about as far as middle of claw. Type Locality: Recorded as “Nebraska to Utah.” A female in the Thomas L. Casey Collection at the United States National Museum has been labeled the type. This specimen is herewith formally designated the lectotype. It bears a locality label with the word “Utah,” so the type locality should be restricted to this state. Nomenclature: Casey quite evidently did not understand the limits of this species. On the one hand, a second specimen from Utah that stands under this name in his collection is actually A. bicolor. On the other hand, two specimens standing under A. bicolor 180 Entomologica Americana [Vol. 45, No. 3 and a number of specimens standing under A. cylindricornis in his collection belong to this species. The type of A. extricatus appears to belong here, although the specimen is rubbed and identification is difficult. However, five other specimens that Casey arranged under A. extricatus , apparently after the original description, belong unquestionably to A. megatoma. A. elongatulus is quite similar to the European A. brunneus Faldermann. Nevertheless, the specimens of A . brunneus I have had available for study seem vaguely different, and I am unwilling to place A. elongatulus in synonymy without an opportunity to compare longer series and preferably to compare immature stages. Distribution: The species has an irregular distribution, sug- gesting that it is introduced and tending to confirm the suspicion that it is a synonym of A. brunneus or some other inadequately described, exotic species. Because the distribution is so widespread, it seems adequate to exhibit it on a map (Fig. 30) without detailing every locality. Diagnosis: This species is very clearly distinct from A. mega- toma, although doubtless much of the American economic literature on “A. piceus ” refers to it. Adult males can be distinguished by the relatively longer terminal segment, as indicated in the key, although this is a character requiring careful measurement. Females as well as males may be distinguished from A. megatoma megatoma by the presence of a band of light-colored hairs along the base of the prono- tum, the band being about as long at its longest point as the leneth of the scutellum. If such a band is present in A. megatoma mega- toma, at its longest point it is no more than half as long as the scutellum. A. elongatulus often has a few scattered golden-brown hairs at the base of the elytra but usually has no golden-brown hairs on the pronotum other than those mentioned. A . megatoma canaden- sis ordinarily has a large number of golden-brown hairs on the pronotum, particularly at the lateroposterior angles, and has a num- ber of golden-brown hairs on the base of the elytra. Nonetheless, some adult female specimens of A. megatoma canadensis or inter- grades between it and A. megatoma megatoma may be nearly im- possible to distinguish from A. elongatulus. A. elongatulus also closely resembles some adult forms of A. schaefferi but may be dis- tinguished from them by the 11- rather than 10-segmented antenna. Adults of A . elongatulus somewhat resemble those of A . cyphonoides, but careful inspection of the latter will always reveal the presence of at least a few lighter hairs forming a subbasal oblique band on 1970] Beal: Taxonomic Study of Attagenini 181 the elytron. No such band is found on the elytron of A. elongatulus. Furthermore, the pronotum of A. cyphonoides is covered with lighter golden-brown hairs instead of the lighter hairs being limited to a band along the basal margin. Little difficulty should be encountered in distinguishing adults of this species from other Nearctic Attagenini. The larvae of this species are easily separated from all other known Nearctic species except those of A. schaefferi. Both A. elongatulus and A. schaefferi have, inserted on the eighth abdominal sternum, a number of distinctively broad, lanceolate setae, each with eight to twelve ribs between the margins. With the exception of A. lobatus all other species have either subrectangular or ovate setae on this sternum or have linear or narrowly lanceolate setae with no more than four ribs between the margins. A. lobatus is easily distinguished from A. elongatulus by having a row of stout linear setae near the anterior margin of each sternum, in front of which are numerous fine, long, simple setae. No such row of stout setae is found on the sterna of A. elongatulus. No completely definitive characters have been found by which larvae of A. elongatulus and A. schaefferi may always be distinguished. Possibly last instar larvae of the two can always be seoarated bv the broader tergal and sternal setae of A. elongatulus. However, it is often impossible to know whether the specimens one is comparing are fully mature. Ecology: The following rather limited data are all I have been able to refer with certainty to this species. Adults have been collected from flowers of the catclaw, Acacia greggii A. Gray, in Presidio, Texas, on Spiraea in Ithaca, New York, and by sweeping celery and beets in Utah. Adults have occasionally been taken at lights. Larvae have been found feeding on woolens in Urbana, Illinois, by Carl Weinman and in Tifton, Georgia, by J. A. Payne. Adults and larvae have been taken quite frequently in homes, amount- ing on occasions to pests of minor importance, but no records have been kept of the materials on which they were feeding. L. F. Hoyt found the species infesting dried buttermilk in Buffalo, New York, and it has been found as a pest in dried milk in Omaha, Nebraska. J. A. Payne found the species infesting in-shell peanuts that had been in storage for four years at Douglas, Georgia. Life History: Nothing has been published, unless this species is indeed a synonym of another species. H. S. Barber left a note in the collection of the United States National Museum indicating that a specimen he reared to maturity passed through 24 larval instars. That this species is incapable of crossbreeding with A. megatoma 182 Entomologica Americana [Vol. 45, No. 3 was demonstrated by Moore and Moore (1942). The “yellow” larvae of the Moores’ experiments were this species and the “black” larvae were A. megatoma. W. E. Burkholder (in litt.) has verified these results in recent experiments. Attagenus schaefferi (Herbst) Adult Males: Integument of dorsal and ventral surfaces immaculate, light brown to black; legs and antennae light brown to dark brown; terminal segment of antennal club brown to black. Pubescence of head and dorsal surfaces subrecumbent, golden- yellow to black, with or without intermingled lighter-colored hairs; ventral pubescence recumbent, black with intermingled light golden- brown hairs on legs to entirely light golden-brown. Head with eye not at all emarginate to shallowly but clearly emarginate over base of antenna. Antenna 10-segmented; ratio of combined length of first two segments of club to length of terminal segment varying from 1:4.7 to 1:7.1; ratio of length of terminal segment to length of pronotum and elytra combined varying from 1:4.4 to 1:5.7; club clothed with fine, erect hairs about one-third as long as width of third antennal segment. Channel below eye for reception of flagellar shaft of antenna concave with anterior margin forming carina; margin of carina visible from front of head; carina projecting knife-like beneath head and curved behind base of maxilla to meet gular suture. Pronotum with lateral carina continued well-defined around anterolateral angle; basal lobe feebly rounded and not distinctly produced backward; setae on posterior margin of lobe as long as to not more than one-third longer than other setae of pronotum. Punctures of pronotal disc as small as or smaller than facet of eye, separated by one to three diameters of single puncture but sometimes feebly joined laterally so that pronotum somewhat transversely rugose. Elytron with punctures of disc one to one and one-half times diameter of facet of eye and separated by one to one and one-half times diameter of single puncture. Hairs of disc of elytron about as long as or slightly longer than third segment of antenna. Hypomeron slightly concave. Prostemum with posterior margin of lateral lobe reflected against procoxa; reflected part not as long as horizontal part and curving to form angle of about 30° with horizontal part; prosternal process slightly wider at apex than between coxae; carina of process not extended onto disc of pro- sternum; anterior declivity of prosternum sometimes separated from disc by thin transverse thread-like carina; denticle at anterior margin 1970] Beal: Taxonomic Study of Attagenini 183 of disc present or not. Epipleuron terminating gradually about middle of first abdominal sternum. Ventral plate of hind coxa forming obtuse or acute tooth lateral to insertion of trochanter and extending laterad beneath inner margin of metepimeron (i.e., ap- pearing to meet metepimeron behind metepisternum ) . Protibia not carinate on dorsal margin. Mesofemur with anteroventral and posteroventral margins of crural cavity about equally produced and nearly on same plane. Length (of pronotum and elytra combined) : 3.2 mm. to 4.2 mm. Adult Females: Terminal segment of antennal club 1.3 to 1.7 times length of eighth and ninth segments combined; ninth seg- ment subequal in length to eighth segment. Length (of pronotum and elytra combined) : 3.8 mm. to 5.3 mm. Mature Larvae : Integument of head and dorsum light brown. Setae of head and body golden-brown. Head : Antenna with terminal seta subequal in length to length of third segment; no setae present on second segment. Epipharynx with distal series of two sensory pits, each subequal in diameter to sensory pits of middle series. Maxillary palp with two or three setae on penultimate segment. Labium with five to six setae on dorsal (inner) side of each lobe of ligula; middle setae narrowly ensiform with setae becoming progressively narrower laterad; first segment of palp without seta. Body setae : Margins and ribs uniformly smooth. Setae inserted on acrotergites all linear, most with no more than one rib between margins but few sometimes present with two ribs between margins. Disc of tergite with linear and linear-lanceolate setae; linear-lanceo- late setae with as many as four ribs between margins. Sockets of submarginal erect setae with two adjacent sensory pits, one on each side near anterior margin. First abdominal sternum with longer setae along midline than on remainder of disc. Setae of sternum of eighth abdominal segment consisting of linear-lanceolate and lanceolate hairs; lanceolate setae with seven to nine ribs between margins; setae becoming smaller on anterior area of sternum without row of stouter setae near anterior margin. Spiracle and associated structures : Anterior abdominal seg- ments with spiracle closed behind by tergum; anterior margin of spiracle more or less straight; posterior margin somewhat thickened; spiracle adjacent to spiracular sclerite. Spiracular sclerite oval, not enclosed by tergum; five or fewer setae inserted on sclerite. Tergum sclerotized anterior to tergal margin of spiracular sclerite. 184 Entomologica Americana [Vol. 45, No. 3 Abdomen : Eighth abdominal tergum with well-defined ante- costal suture. Ninth abdominal tergum more than two-thirds as long as tergite of eighth abdominal segment. Two small sclerites present in sternum of ninth abdominal segment. Legs : Femur of prothoracic leg half or slightly more than half as wide as long; ventral margin about two-fifths as long as length of segment. Tibia of prothoracic leg with usual two rows of ventral setae; posterior face with subdorsal apical seta and longitudinal row of three setae along middle of segment; most proximal seta in row inserted slightly before middle of segment. Setae inserted at base of pretarsus subequal in width, extending about as far as middle of claw. Systematics: Varieties of A. schaefferi (Herbst, 1792) have long been known to exist in the United States but have been con- sidered by all previous workers as no more than accidental intro- ductions of the species from Europe. A careful study of the populations in the United States seems to indicate that this assump- tion is incorrect. The Nearctic forms are seldom found as pests in stored products or houses, yet occur commonly in the most remote areas of the West, often far from human habitation. If the forms had been introduced, they would be expected in centers of commerce or human activity. Furthermore, a number of geographic segregates are discernible in different areas of the West, a phenomenon not expected in a recently introduced species. Finally, the forms that are present in most areas of the United States appear to be dis- tinguishably different from forms that have been described for the Palearctic region. A measurable difference between European and Nearctic forms is found in the ratio of the length of the last antennal segment of the male to the total length of the pronotum and elytra. A large number of Nearctic specimens has been measured and the ratio found always to vary within the range of 1:4.67 to 1:5.67. Only four European males were available for study, but these all fell outside this range, varying from 1:4.40 to 1:4.65. These differences are small and, if a large number of European forms should be measured, no doubt it would be found that some overlap occurs in the ranges of variation. Another difference, but one difficult to measure, appears to be the degree of concavity of the epipleuron. In the Palearctic specimens I have examined the epipleuron is quite concave, but in most Nearctic specimens it is nearly flat or only slightly concave. These differences, taken with the evidence that the Nearctic forms are indigenous, 1970] Beal: Taxonomic Study of Attagenini 185 make it desirable to recognize the Nearctic forms as taxonomically distinct. Accordingly, I am designating the American forms as subspecies of the Palearctic form. The subspecies category preserves the apparent relationship with the Palearctic form while providing suitable names for reference to the American forms. The various Nearctic populations of the species, so far as visible differences are concerned, are distinguished principally by the color of the setae of the adults. Specimens from British Columbia south along the Sierra Nevada are uniformly clothed with dark hairs on the dorsal surfaces and on the abdomen. Within this range specimens from the north tend to be more chocolate brown, and from the south darker, almost black. Specimens collected in the Upper Sonoran zone across Northern Arizona have elytra covered with dark, golden- brown hairs with somewhat lighter golden-brown hairs scattered about the basal area of the elytra and on the pronotum. The type of A. spurcus LeConte from Santa Fe, New Mexico, is covered with light golden-yellow hairs. A series from Bakersfield, California, resembles the Arizona forms more than those from the higher eleva- tions of the adjacent Sierra Nevada. A short series from Tooele County, Utah, is covered with golden-brown hairs but has a band of lighter golden hairs on the basal margin of the pronotum that makes it very similar in appearance to A . elongatulus. Possibly the color differences between the forms could be de- scribed without resorting to subspecific names, except that there appears to be some kind of reproductive barrier between the Arizona forms and those from the Sierra Nevada. Five experiments were conducted to obtain crosses between specimens collected in Sequoia National Park and specimens collected near the eastern edge of the Grand Canyon in Arizona. All resulted in failure. Under identical conditions, as far as these could be regulated, specimens from the original cultures reproduced freely. Unfortunately, specimens from one culture for the most part did not mature at the same time as specimens from the other culture. As a consequence few members of the opposite sex were available from the different cultures for mating and the experiment could not be replicated as many times and under as many different conditions as would have been desirable. Assuming that the experiments are conclusive in establishing a reproductive barrier between the two populations, two explanations are possible. It might be that an absolute barrier exists and that each population should be considered a distinct species. On the other hand there might be continuous crossbreeding between adjacent 186 Entomologica Americana [Vol. 45, No. 3 populations and the extremes only might be intersterile. Of course, there is always the possibility that specimens available for crossbreeding happened to be defective in one way or another and were not representative of the populations. Obviously a great deal more study is needed to elucidate the relationships. However, until further evidence is available, it seems best to take a middle ground and designate two subspecies, one including the forms from the Sierra Nevada north to British Columbia and the other including the Arizona and New Mexico forms. Tentatively the forms from Bakersfield are placed with the Arizona subspecies. Some specimens from Little Granite Mountain, Tooele County, Utah, taken February 3, 1955, as larvae and pupae (D. M. Allred), are not assigned to a subspecies. They seem quite distinct from either of the other two subspecies and probably deserve a separate name. Nevertheless, it would be inadvisable to name them without more extensive collections from the state of Utah. Whether the nominate subspecies occurs in North America is a question that at present cannot be answered with certainty. Two specimens might be assigned to it, one collected in Atlanta, Georgia, in 1944 by P. W. Fattig and the other labeled simply “111.” from the Stromberg collection of the Illinois Natural History Survey. Diagnosis: A. schaefferi is obviously more closely related to A. eiongatulus than to any other Nearctic species. Adults can al- ways be distinguished from A. eiongatulus by the 10-segmented antenna, but otherwise some forms of the two species are almost impossible to separate. Usually, however, A. schaefferi lacks a band of light-colored setae along the base of the pronotum, a char- acter always present in A. eiongatulus. Larvae of comparable size of the two species can usually be distinguished by the wider tergal and sternal setae of A. eiongatulus , but a character that will abso- lutely separate all larval specimens has not been found. The same differences that distinguish A. eiongatulus from the somewhat similar A. cyphonoides , A. megatoma, and A. fasciatus serve to distinguish A. schaefferi from them. These differences are discussed under A. eiongatulus. Ecology: The species is apparently a general scavenger on a variety of protein materials. Specimens from Little Granite Moun- tain, Tooele County, Utah, were taken in a Neotoma nest. I col- lected larvae in an unidentified rodent’s nest in a feed storage shed at Sequoia National Park Headquarters, California. Linsley (1944) recorded it in the nests of the English sparrow and the house finch. 1970] Beal: Taxonomic Study of Attagenini 187 He also collected the species in a black phoebe’s nest at Fresno, California, where I collected it a few years later. I collected larvae in insect-infested barley and on an old deer hide in a ranch granary 20 miles west of Cameron, Arizona. To my knowledge the species has not previously been recorded as a pest of stored products. It was taken at Chloride, Arizona, by “L. L. G.” in bran and barley, but may have been feeding in grain badly infested with other insects. It seems surprising that it has not been more commonly found as a pest of stored cereals, since I have been able to rear it quite successfully in the laboratory on a diet of oatmeal. Attagenus schaefferi hypar, new subspecies Attagenus schaefferi : Casey, 1900, p. 146. — Hatch, 1962, p. 284. Adult Males: Integument dark brown. Dorsal pubescence brownish black; pubescence of abdomen chocolate brown. Ratio of length of terminal antennal segment to combined length of prono- tum and elytra 1:4.8. Epipleuron slightly inflated at base becoming slightly concave for most of length. Range of Observed Variations: Integument black to dark brown. Dorsal pubescence uniformly black to dark brown. Pubes- cence of abdomen black to chocolate brown, occasionally with few intermingled dark golden-brown hairs. Ratio of length of terminal antennal segment to combined length of pronotum and elytra vary- ing from 1:4.7 to 1:5.7. Females like males except for much shorter length of antennal club. Primary Types: Holotype male: Fallen Leaf, El Dorado County, California, July 19, 1935 (F. E. Blaisdell); allotype female: Strawberry Valley, El Dorado County, California, August 6, 1912 (Van Dyke Collection). Specimens deposited in the collection of the California Academy of Sciences. Paratypes: British Columbia : Genoa Bay, Duncan, July 10, 1928 (W. Mathers); Mara Lake, July 20, 1932 (A. C. Thrupp); Midday Valley, Merritt, July 24, 1924 (K. F. Auden); Robson, August, 1949 (H. R. Foxlee); Salmon Arm, May 9, 1936 (D. H. Leech); Salmon Arm, July, 1932 (O. R. Leech); Salmon Arm, July, 1935 (D. H. Leech); Taft, August 2, 1932 (A. C. Thrupp); Trinity Valley, July 1, 10, 11, 12, 1928, July 10, 18, 21, 23, 1929, July 27, 1927, August 18, 1929, August 24, 1930 (J. R. L. Howell); Trinity Valley, July 27, 1927 (E. A. Rendell); Vernon, June 28, 1920 (Ralph Hopping). California : Calaveras County: Big Trees, 188 Entomologica Americana [Vol. 45, No. 3 July 19, 1907 (F. E. Blaisdell). El Dorado County: Fallen Leaf, July 1, 3, 11, 28, 1935 (F. E. Blaisdell) ; Fallen Leaf, July 10, 1915 (R. Hopping); Kirkwood Lake, July 18, 1946 (A. T. McClay); McKinneys, July, 1906 (Van Dyke Collection); Strawberry Valley, August 5, 1912 (Van Dyke Collection); Tallac, July 7, 1925 (F. E. Blaisdell). Fresno County: Badger Flat, Huntington Lake, August 8, 1939 (A. T. McClay); Bubb’s Creek, 8,000 ft. elev., July 29 (Wickham Collection); Bubb’s Creek, Kings River, 9,700 ft. elev., July 7, 1910 (Van Dyke Collection); Camp Greeley, 2,800 ft. elev., May 22, 1910 (R. Hopping); Fresno, July 2, 1942; Home Creek, Huntington Lake, July 21, 1936 (A. T. McClay); Hunting- ton Lake, 7,000 ft. elev., July 20, 1919 (F. E. Blaisdell); Hunting- ton Lake, June 21, July 21, 1936, July 22, 1937 (A. T. McClay); Kaiser Pass, Huntington Lake, July 21, 1937 (A. T. McClay); Paradise Valley, Kings River, 7,000 ft. elev., July 17, 1910 (Van Dyke Collection). Inyo County: Independence, May 26, 1919, June 19, 1918 (L. L. Muchmore); Lone Pine, June 18, 1937 (C. A. Hamsher). Lassen County: Blue Lake, July 19-20, 1947 (T. F. Leigh); Butte Creek, July 26, 1948 (A. T. McClay); Facht, July 17, 1921, August 2, 1921 (J. O. Martin); Martin Springs, Sec. 14, T31N, R9E, July 23, 1922 (J. O. Martin); Susan River Camp, July 4, 9, 10, 1949 (A. T. McClay); Susan River Camp, July 10, 1949 (J. E. Gillaspy). Madera County: Biledo Meadow, July 27, 1946 (H. Chandler); Buck Camp, August 1, 3, 1958 (M. E. Irwin); Upper East Fork, Chiquito Creek, August 24, 1958 (M. E. Irwin). Mono County: Cottonwood Creek, July 11, 1961 (J. S. Buckett); Mono Lake, July, 1922 (Coriane Hilton); 7 miles east of Tioga Pass, July 15, 1961 (J. S. Buckett). Modoc County: Buck Creek, July 21, 1922 (C. L. Fox); Cedarville, July 9, 1946 (P. D. Hurd and Ray F. Smith); Warner Mountains, July 10, 11, 1919 (R. Hopping). Nevada County: 3 miles north of Boca, July 23, 1961 (F. D. Parker). Placer County: Brockway, July 2 (George Mansfield); Lake Tahoe, August 21. Plumas County: Meadow Valley, 4,500 to 5,000 ft. elev., June 13, 1924. San Bernardino County: Barton Flats, July 6, 1938 (E. Herald); Camp Baldy, June 14 to 21, 1926 (L. J. Muchmore). Shasta County: Dry Lake, June 24, 1947 (T. F. Leigh); Hat Creek, July 17, 1952 (G. Pronin); Mount Lassen National Park, July 27, 1945, July 28, 1941 (A. T. McClay). Sierra County: Gold Lake, July 8, 1934 (L. S. Ross); Yuba Pass, July 30, 1958 (A. A. Grigarick). Trinity County: Scott Mountain, 5,358 ft. elev., July 14, 1949 (A. T. 1970] Beal: Taxonomic Study of Attagenini 189 McClay). Tulare County: Dorset Creek, August 2, 1917 (R. Hopping); Kern River, 7,000 ft. elev., July 27, 1899; Kern River, 6,000 ft. elev., July 29, 1899; Rattlesnake Creek, July 26, 1917 (R. Hopping); Round Meadow, Giant Forest (Sequoia National Park), August 7, 1903 (R. Hopping); Sequoia National Park, 7,000 to 9,000 ft. elev., July 2, 1929 (A. T. McClay); Park Headquarters, Sequoia National Park, as larvae August 6, 1960 (R. S. Beal). Tuolumne County: Eleanor Lake, July 2, 1951 (S. M. Kappos); Pine Crest, July 3, 1951 (A. T. McClay); Strawberry, June 20, 1951 (S. M. Kappos); Strawberry, June 30, 1951 (R. W. Morgan). Yosemite National Park: Beehive, July 25, 1937 (E. Herald); Boot Trail, July 26, 1936 (A. T. McClay); Givens Creek, August 20, 1958 (M. E. Irwin); Miguel Meadows, 5,300 ft. elev., July 21, 1937 (E. Herald); Yosemite, 3,880 to 4,000 ft. elev., June 7, 1931 (E. O. Essig); Yosemite, 8,000 ft. elev., July 26, 1936 (A. T. McClay); Yosemite Valley, July 12, 1921 (E. C. Van Dyke). Idaho : Beaver Canyon (Hubbard and Schwartz). Coola, Priest Lake, July 24, 1927 (E. C. Van Dyke). Bonner County: Sagle, July 2, 1950 (N. M. Downie). Wardner, July 18, 1929 (R. A. Flock). Nevada : Gardnerville, 15 miles south, July 21, 1962 (E. J. Montgomery); Lake Tahoe, June 25, 1953 (G. F. Knowlton); Yerington, June, 1909. Oregon : Ashland Mountain Road, 4,500 to 5,000 ft. elev., August 10, 1953 (M. C. Lane); Crater Lake National Park, Sun Creek Meadows, 6,500 to 7,000 ft. elev., August 8, 1930 (H. A. Scullen); Crater Lake National Park near Head- quarters, 6,400 to 6,600 ft. elev., August 4, 1930 (H. A. Scullen); The Dalles (Wickham Collection); Deming Creek, 11 miles north of Bly (Klamath County), August 12, 1956 (Joe Schuh); Elgin, June 20, 1922 (A. L. Lovett); Klamath Falls, February 18, 1956 (Joe Schuh); Lake of the Woods (Klamath County), July 10, 1934 (Van Dyke Collection); McMinnville, July 26, 1938 (K. M. and D. M. Fender); Medford, August 8, 1946 (A. T. McClay); Mount Ashland, July 18, 1951 (M. F. McClay). Washington : Easton (A. Koebele); North Fork Cipsus River (Skamania County), June 25, 1934 (J. L. Wilson); King County, July 14, 1928 (M. H. Hatch); Kooskooskie, Walla Walla, August 1, 1932 (M. C. Lane); Lewis and Clark State Park, June 12, 1938 (M. H. Hatch); Mount Adams, West Klickitat, July 8, 1925 (L. A. Morky); Quilcene, July 27, 1915; Seattle, April 5, 1929; same locality, May 3, 1931 (M. H. Hatch); Sullivan Lake, July 23, 1934 (G. H. and R. Hopping); 190 Entomologica Americana [Vol. 45, No. 3 Tacoma, May 23, 1948 (N. Fuhr); Vaughn, May 17, 1940 (F. Bjorkmann) ; Whidby Island, July 30, 1898. Attagenus schaefferi spurcus LeConte, new status Attagenus spurcus LeConte, 1854, p. 109. Attagenus schaefferi: Crotch, 1873, p. 41. Attagenus piceus : Jayne, 1882, p. 355. Attagenus cylindricornis : Casey, 1916, p. 183 (see section at end on species described but unrecognized). Adult Males: Integument light brown to dark brown; when elytra light brown, pronotum usually darker. Dorsal pubescence piceous with light golden-brown hairs along lateral and basal margins of pronotum and scattered through basal area of elytra to uniformly light golden-yellow. Ratio of length of terminal antennal segment to combined length of pronotum and elytra varying from 1:4.8 to 1:5.3. Epipleuron flat at base becoming slightly concave along most of length. Type Locality: Santa Fe, New Mexico. Type No. 6868, Museum of Comparative Zoology, Cambridge, Massachusetts. New Records: Arizona : Coconino County: Basin Camp, 77 Bar Ranch, 20 miles west of Cameron, as larvae September 27, 1964 (R. S. Beal). Mohave County: Chloride, July 16, 1957 (L. L. G.). Navajo County: Linden, as larvae August 20, 1962 (R. S. Beal). California : Bakersfield, April 12, 1944 (E. G. Linsley); Bakersfield, as larvae April 13, 1963 (R. S. Beal). Attagenus pellio (Linnaeus) Dermestes pellio Linnaeus, 1758, p. 355. — Panzer, 1795, p. 96. Dermestes bipunctatus DeGeer, 1774, p. 197. Attagenus pellio: Latreille, 1807, p. 32. — Dahl, 1823, p. 30. — Stephens, 1830, p. 126. — Kirby, 1837, p. 114. — Laporte, 1840, p. 35. — Le- Conte, 1854, p. 109. — Wollaston, 1864, pp. 155-156. — Mulsant and Rey, 1867 (1868), pp. 68, 77-80.— Jayne, 1882, p. 355.— Reitter, 1889, p. 557. — Casey, 1900, p. 146. — Ganglbauer, 1904, p. 24. — Lutz, 1911, p. 152. — Wradatsch, 1914, p. 152. — Kempers, 1923, pp. 87-88. — Mutchler and Weiss, 1927, p. 10.— Lepesme and Paulian, 1939, p. 165. — Hinton, 1943, p. 227. — Korschefsky, 1944, p. 147. — Armstrong, 1945, p. 48. — Hinton, 1945, pp. 306-309; 1946, p. 485. — Kalik, 1948, Mroczkowski, 1954, p. 22. — Hatch, 1962, p. 284. — Zhantiev, 1963, p. 421. Megatoma ater Herbst, 1792, p. 95. Megatoma schrankii Kugelann, 1792, p. 480. 1970] Beal: Taxonomic Study of Attagenini 191 Megatoma pellio : Illiger, 1798, p. 316. — Brulle, 1832, p. 137. — Reitter, 1887, p. 47. Adults: Adequately described by Hinton (1945). Mature Larvae: Integument of head and dorsum brown. Setae of head dark golden-brown; setae of body consisting of inter- mingled dark golden-brown narrow hairs and light golden-brown broad hairs. Head : Antenna with terminal seta subequal to or slightly longer than third segment; no setae present on second segment. Epipharynx with distal series of two sensory pits, each subequal in diameter to sensory pits of middle series. Maxillary palp with two apical setae on penultimate segment. Labium with four to five setae on dorsal (inner) side of each lobe of ligula; medial two setae narrowly ensiform; lateral setae nearly terete; first segment of palp without setae. Body setae : Margins and ribs uniformly smooth. Setae in- serted on acrotergite of first abdominal segment consisting of linear and broadly lanceolate hairs; widest lanceolate hairs with three to six ribs between margins (Fig. 21); broad setae becoming increas- ingly wide on acrotergites of succeeding segments; broad setae on acrotergites of posterior segments with as many as twelve ribs be- tween margins. Disc of tergite of first abdominal segment with intermingled linear-lanceolate setae with two or three ribs between margins and ovate to subquadrate setae with as many as six ribs between margins; setae of succeeding tergites similar except scale- like setae becoming more rectangular although retaining acutely produced apices; scale-like setae on eighth segment with six to 14 ribs between margins (Fig. 22). Sockets of submarginal erect setae with two sensory pits, one on either side near anterior margin. First abdominal sternum with setae along midline longer than other setae of disc. Setae of sternum of eighth abdominal segment identical with setae of tergite. Spiracle and associated structures: Anterior abdominal seg- ments with spiracle closed behind by tergum; anterior margin of spiracle projecting ventrad over posterior margin and bearing acute denticle (Fig. 25); spiracle adjacent to spiracular sclerite. Spiracular sclerite irregularly oval, not enclosed by tergum. Tergum sclerotized continuously with tergal margin of spiracular sclerite. Abdomen: Eighth abdominal tergum with well-defined ante- costal suture. Ninth abdominal tergum about one-half as long as 192 Entomologica Americana [Vol. 45, No. 3 tergite of eighth abdominal segment. Two small sclerites present in sternum of ninth abdominal segment. Legs : Femur of prothoracic leg about 8/17 as wide as long; ventral margin about 4/11 as long as length of segment. Tibia of prothoracic leg with usual two rows of ventral setae; posterior face with subdorsal apical seta and longitudinal row of four seta along middle of segment; most proximal seta of row inserted at about basal third of segment. Setae inserted at base of pretarsus subequal in width and extending about as far as middle of claw. Recorded Distribution: Widespread throughout Palearctic Region; in North America in Nova Scotia, New Hampshire, Massa- chusetts, Michigan, New Jersey, New York, Rhode Island, British Columbia. New Record: Rockhaven, Kentucky (H. Soltau). Diagnosis: Adults are easily recognized by the two small patches of white or light golden-yellow hairs on the middle of the elytra near the suture. These contrast sharply with the dark brown to black hairs covering most of the dorsal surface. The male antennal club has the first two segments short and subequal. The terminal segment is four to five times as long as the combined length of the first two. The larvae are dark golden-brown. They may be distinguished from all other species described here by the smoothly margined, more or less rectangular setae inserted on the posterior abdominal terga and sterna. On the eighth abdominal segment these scale-like setae have six to 14 ribs between the margins. Setae of the anterior segments are ovate to subquadrate. All other species with scale-like setae have ovate or ovate-lanceolate setae, and the setae have dentic- ulate instead of smooth margins. Ecology: Larvae of this species, like larvae of most other species of the genus, appear to be general scavengers on dried protein materials. Out-of-doors they are commonly found in swallow nests, where they feed on feathers and the remains of other insects (Hinton, 1945). In England Woodroffe (1953) found them to be regular inhabitants of dry bird nests, particularly of pigeon nests and of jackdaw nests in hollow trees. Hicks (1959) has collected numerous additional references to the occurrence of the species in bird nests, which include, in addition to those above, nests of the house martin, swift, owl, English sparrow, titmouse, flycatcher, and white wagtail. Hinton (1945) summarizes the reported occurrences of this 1970] Beal: Taxonomic Study of Attagenini 193 species as an economic pest, listing the following products on which it has been found, presumably feeding: furs, skins, woolens, carpets, grain, flour, maize, meal, cattle food, rye bran, and sugar. Whether it can mature feeding on cereal products alone has not been de- termined. Probably it maintains itself in cereal products, at least to some extent, on the remains of other insects. It also attacks dried insects in museums. R. W. Howe (1962) states that the species shows diapause, but the factors that initiate it are unknown. Parasites: The larvae have been found parasitized by the gregarine, Pyxinia mobuszi Leger and Duboscq (Foerster, 1938, Eichler, 1939), a parasite also found in Anthrenus verbasci (Lin- naeus). Attagenus fasciatus (Thunberg) Anthrenus fasciatus Thunberg, 1795, p. 105. Anthrenus gloriosae Fabricius, 1798, p. 76. — Fabricius, 1801, p. 107. Dermestes fasciatus: Schonherr, 1808, p. 88. Attagenus fasciatus : Dejean, 1837, p. 139. — Mroczkowski, 1964, pp. 179-180. Attagenus annulifer Laporte, 1840, p. 36. Attagenus gloriosae: Lacordaire, 1854, p. 464. Aethriostoma gloriosae: Motschulsky, 1858, p. 146. Trogoderma subfasciatum Chevrolat, 1863, p. 617. Attagenus plebeius Sharp, 1885, p .47. Attagenus gossypiatus Fauvel, 1903, p. 335. Attagenus gloriosae: Arrow, 1915, p. 427. — Hinton, 1943, p. 224; 1945, pp. 320-322. — Armstrong, 1945, p. 48. — Hinton, 1946, p. 19. — Kalik, 1955, pp. 307-308. Adults: Adequately described by Hinton (1945) under the name A. gloriosae. Mature Larvae: Dorsal integument reddish-brown. Setae on dorsal and ventral surfaces dark golden-brown except for light golden-brown setae on legs. Head : Antenna with terminal seta one and one-eighth times as long as terminal segment; second segment bearing 12 or more setae. (Epipharynx not observed.) Maxillary palp with two setae on third segment. Labium with series of two medial lanceolate setae and three or four slender ensiform lateral setae on dorsal (inner) side of each lobe of ligula; first segment of palp without seta. Body setae : Margins and ribs uniformly smooth. Acrotergites with some linear setae with one rib between margins and some simple 194 Entomologica Americana [Vol. 45, No. 3 setae. Tergites with linear setae with two to five ribs present be- tween margins. Sockets of submarginal erect setae usually with two adjacent sensory pits, one on each side. Sterna with linear setae; setae of disc with two ribs between margins; setae smaller and with- out ribs on anterior area of each sternum; anterior margin without row of stouter, longer setae; setae at midline of first two abdominal sterna not appreciably longer than other sternal setae. Spiracle and associated structures : Spiracle closed behind by tergum; anterior margin of spiracle slightly curved; width of spiracle less than one-third width of spiracular sclerite; spiracle removed from spiracular sclerite by distance equal to about two-thirds its width. Spiracular sclerite oval, bearing six or fewer setae, not en- closed by tergum; area of tergum in front of spiracular sclerite well sclerotized and continuous with tergal margin overlying anterior margin of spiracular sclerite. Abdomen : Eighth abdominal tergum with antecostal suture. Ninth abdominal tergum about five-ninths as long as tergite of eighth segment. Two small sclerites present in sternum of ninth abdominal segment. Legs : Femur of prothoracic leg half as wide as long; ventral margin little less than half as long as length of segment. Tibia of prothoracic leg as illustrated (Fig. 27). Pretarsus with setae nar- rowly ensiform, subequal in width and length. Distribution: The species occurs nearly throughout the tropi- cal and subtropical areas of the world. In the continental United States it is apparently established only in Florida, although it has been taken infesting stored products in Buffalo, New York, and Indianapolis, Indiana. Records for the species in Florida are the following: Deland, April 21, 1960 (G. W. Desin); Miami, March 3 and April 14, 1960 (E. M. Collins); Orlando, May 27, 1935, July 10, 1935, December 1, 1936; Tampa, May 20, 1960 (E. M. Collins, Jr.). Diagnosis: Adults are immediately distinguished from all other Nearctic species by the distinct subbasal elytral band of light golden hairs. The remainder of the elytra is covered with brownish black hairs, which contrast sharply with the color of the elytral band. Usually the band is fairly broad and extends from the lateral margin of each elytron to the median suture. At the median suture it is narrower and projects a little basad. Rarely the band is discontinuous, but when it is there is always a distinct spot of light-colored hairs on the median suture posterior to the scutellum. Any other species 1970] Beal: Taxonomic Study of Attagenini 195 of Attagenus or Novelsis in the United States that has a distinct subbasal elytral band also has a light-colored line along the entire basal three-fourths of the median suture, or has more than one elytral band, or the subbasal band does not extend to the median suture and there is no distinct spot of light-colored hairs on the suture just posterior to the scutellum. A number of other differences seem to make this species unique among the Nearctic Attagenini, one of which is the short terminal segment of the male antennal club. Males of the species cannot be distinguished from the females unless the genitalia are exposed. It is also relatively broader than other Nearctic species, has stouter front coxae, and stouter femora. Be- cause of these traits it was placed in the genus Aethriostoma by Motschulsky. Nevertheless, larval characters show it to be much more closely allied to Attagenus and to the cluster of species des- ignated Group I. The larvae are quite similar to larvae of A. megatoma. The one distinctive character that separates them from larvae of A. megatoma , as well as from larvae of all other Nearctic species, is the number of setae on the second segment of the antenna. Twelve or more are present on this segment in mature specimens. Mature larvae of A. megatoma lack setae on the second segment, and no other species I have studied has more than three setae on the segment. Ecology and Life History: Hinton (1945) has collected and published information on these aspects of the species. If there is additional published information, I am unaware of it. In addition to those products which it has been recorded as infesting, it was found in dried buttermilk in Buffalo, New York (L. F. Hoyt) and in crude drugs in Indianapolis, Indiana (J. J. Favinger). The species also has been intercepted at Tampa, Florida, in Caesalpinia coriaria imported from Columbia. Attagenus bicolor Harold Attagenus dichrous LeConte, 1854 ( non dichrous Roth, 1851), p. 110. Attagenus bicolor Harold, 1868, p. 104. — Reitter, 1880 (1881), p. 34. Attagenus piceus: Jayne, 1882, p. 355. Attagenus schaefferi: Dalla Torre, 1911, p. 57. Attagenus sparsus Casey, 1916, p. 184 (New synonymy). Adult Males: Integument of dorsal and ventral surfaces im- maculate; head dark brown to black; pronotum reddish brown to dark brown; elytra reddish brown; venter reddish brown to dark brown; legs and antennae yellowish to reddish brown; terminal seg- 196 Entomologica Americana [Vol. 45, No. 3 ment of club not darker than preceding segments of club. Pubescence of head and dorsal surfaces subrecumbent to suberect; pubescence of ventral surfaces subrecumbent; all hairs black or hairs black on dorsum and reddish brown on venter and legs. Head with eye entire. Antenna 1 1-segmented; ratio of combined length of first two segments of club to length of terminal segment varying from 1:3.6 to 1:5.2; ratio of length of terminal segment to length of pronotum and elytra combined varying from 1:6.7 to 1:8.8; club clothed with very fine subrecumbent hairs less than one-fourth as long as width of third antennal segment; hairs slightly longer at apex of terminal segment. Channel below eye for reception of flagellar shaft of antenna concave with anterior margin forming carina visible from front of head (Fig. 1); carina projecting knife- like beneath head, curved very little behind base of maxilla and not meeting gular suture. Pronotum with lateral carina continued around anterolateral angle; basal lobe feebly rounded to slightly truncate and not distinctly produced backward; setae on posterior margin of lobe subequal in length to other setae of pronotum. Punctures of pronotal disc varying in width from two-thirds as wide to equal to width of facet of eye and separated by two to five diameters of single puncture. Elytron with hairs of disc about as long as com- bined length of antennal segments three through seven. Hypomeron nearly flat at middle, sometimes slightly concave toward base; without auricle-like lobe covering trochantin so that trochantin fully exposed (Fig. 7). Prosternum with posterior margin of lateral lobe flat and not reflected against middle of coxa but reflected against lateral part of coxa and against trochantin; reflected part short, about one-third as long as length of horizontal part of lateral edge of lobe; anterior margin of prosternum with distinct thread-like lateral carina but without denticle at middle of anterior margin; posterior process scarcely wider at apex than between coxa and without median carina extending onto disc. Epipleuron terminating at or little before hind margin of metepimeron. Ventral plate of hind coxa not forming distinct tooth lateral to insertion of trochanter but becoming gradually narrower laterally; plate terminating laterally behind metepisternum at inner margin of metepimeron (appearing to meet metepimeron behind metepisternum) (Fig. 6). Protibia not carinate on dorsal margin. Mesofemur with anteroventral and posteroventral margins of crural cavity about equally produced and on same plane. Length (of pronotum and elytra combined): 3.8 mm. to 4.4 mm. 1970] Beal: Taxonomic Study of Attagenini 197 Adult Females: Antennal club with terminal segment one and one-fifth to one and one-fourth times as long as length of ninth and tenth segments combined; tenth segment equal in length to ninth segment. Length (of pronotum and elytra combined): 4.9 mm. to 5.3 mm. (based on reared specimens alone). Mature Larvae: Dorsal integument brown. Setae of head golden-brown; setae of body black except for few dark golden-brown hairs on nota intermingled with black hairs; setae of ventral thoracic segments and legs golden-brown; setae of ventral abdominal segments black; terminal abdominal setae dark brown and black. Head: Antenna with terminal seta subequal in length to third segment; no setae present on second segment. Epipharynx with distal series of two sensory pits, each subequal in diameter to sensory pits of middle series. Maxillary palp with three setae on penultimate segment. Labium with four to six setae on dorsal (inner) side of each lobe of ligula; medial two setae narrowly ensiform; lateral setae nearly terete; first segment of palp with two setae. Body setae : Very minutely serrulate along ribs (serrulations visible only under high power of compound microscope). Setae in- serted on acrotergites all more or less terete and linear, with as manv as six ribs distinguishable between margins' (ribs visible only in speci- mens treated to remove pigments from setae and then difficult to demonstrate except under oil immersion lens). Disc of tergite with setae similar to those on acrotergite but with as many as eight ribs distinguishable between margins of larger setae. Three submarginal erect setae on each side of each abdominal segment; sockets usually with one adjacent sensory pit on either medial or lateral side but rarely with two adjacent sensory pits, one on each side. First abdominal sternum with few setae somewhat longer than other setae of disc. Eighth abdominal sternum with setae nearly identical to setae of tergum; setae becoming smaller anteriad without row of longer, stouter setae near anterior margin of segment. Spiracle and associated structures : Abdominal segments with spiracle enclosed by tergum; anterior abdominal segments with distance from posterior margin of tergite to spiracle about one-fifth length of tergite; anterior margin of spiracle more or less straight; posterior margin somewhat thickened; spiracle separated from spirac- ular sclerite by distance about equal to width of trachea entering spiracle (Fig. 14). Spiracular sclerite oval, completely enclosed by tergum; three setae inserted on sclerite of each segment. Abdomen : Eighth abdominal segment without antecostal suture 198 Entomologica Americana [Vol. 45, No. 3 on tergum. Ninth abdominal tergum about five-sevenths as long as eighth abdominal tergum. Two small sclerites present within sternum of ninth abdominal segment. Legs : Femur of pro thoracic leg about half as wide as long; ventral margin about half as long as length of segment. Setae in- serted at base of pretarsus subequal in width and extending to about apical third of claw. Type Localities: The type locality for A. dichrous is New Mexico (type specimen No. 6870 in the Museum of Comparative Zoology, Cambridge, Massachusetts). The type locality for A. sparsus is Jemez Springs, New Mexico (holotype in the Casey Collection, United States National Museum). New Records: Arizona : North Rim of the Grand Canyon, 8,150 ft. elev., as larvae August 30, 1964 (R. S. Beal). Colorado : Colorado Springs, 6-7,000 ft. elev., June 15-30, 1896 (H. F. Wickham); Rico, May 22, 1950 (Hopkins U. S. No. 34216-W). New Mexico : Fort Wingate (J. D. Sherman); Jemez Mountains, June 11, 22, 27, July 3 (John Woodgate); Santa Fe (Boyle). Nevada : Baker, Sec. 15, T13N, R69E. Mt. Diablo Meridian, May 23, 1939 (T. O. Thatcher). Utah : Callao, June 2, 1922 (specimen in the Casey Collection at the United States National Museum stand- ing under the name elongatulus) ; Circleville, June 27, 1933 (G. F. Knowlton); South Creek, Beaver County, June 22. Wyoming : Cokeville, April 20, 1952 (William May). Dead larvae and adults have also been found in matted vegetation and fill in the Mesa Verde Ruins in southwestern Colorado. Al- though the adults were badly fragmented, enough remained for positive identification. The larvae that were found were surprisingly well-preserved, with most of the setae intact. They had apparently died in ecdysis, a phenomenon often associated with starvation. All were recovered in Site No. 1285 (S. A. Graham). Diagnosis: A. bicolor and the allied N . athlophora are readily distinguished from all other Nearctic Attagenini by a number of characters. The most noticeable distinguishing feature is the long, sparse, completely black pubescence of each. Almost all other members of the tribe have light golden-brown or whitish ventral pubescence, and many have light-colored dorsal hairs. A. schaefferi hypar has predominately chocolate-colored or blackish ventral setae, but it does have at least a few golden-brown hairs on the front coxae, the setae are moderately dense and short, the dorsal setae are sub- recumbent, and the ventral setae are recumbent. A. bicolor and 1970] Beal: Taxonomic Study of Attagenini 199 N. athlophora also share the following combination of characters not found in other species treated here: the hind coxa lacks a definite tooth lateral to the insertion of the trochanter and is wider at the base of the trochanter than at any point lateral to it; the epipleuron terminates at about the hind margin of the metepimeron; the hypomeron is more or less flat or very slightly concave; the lateral lobe of the prosternum is somewhat longitudinally concave and reflected ventrad in front of the trochantin but lies mostly in a horizontal plane and is not reflected in front of the middle of the procoxa. Both males and females of A. bicolor are separable from N. athlophora by the number of segments in the antenna: A. bicolor has 11 and N. athlophora 10 segments. The male antennal club of A. bicolor is that of a typical Attagenus. The terminal segment is about five times as long as the two preceding segments combined. The male antennal club of N. athlophora has all segments elongated so that the terminal segment is no more than four-fifths as long as the two preceding combined. The dorsal integument of A. bicolor is a medium reddish brown with the pronotum and head often a dark brown. The known specimens of N. athlophora are a very dark brown, except that one has a light brown sutural line. The larvae of A. bicolor are relatively sparsely clothed with linear black hairs. They are easily distinguished from other larval Attagenini by the nearly completely enclosed spiracular sclerite and the remarkable position of the spiracle, which is some distance from the posterior margin of the tergum. Only N. varicolor has a similarly enclosed sclerite with the spiracle so remotely separated from the posterior margin of the tergum. However, N. varicolor is densely covered with ovate setae, which obscure the structures so that they cannot be seen without first teasing the setae away. Possibly N. athlophora will prove to have similar larval structures, but at present the immature stages of this species are unknown. Ecology: I collected numerous larvae of this species in an insect-infested mixture of grain and straw in an unidentified rodent’s nest (probably chipmunk) in a small granary at the North Rim of the Grand Canyon of Arizona. Adults breed in the laboratory without the necessity of flying or of feeding on nectar or pollen. Larvae mature to adults on dried dog food or on oatmeal, as well as on grain trash. Adults emerged in the laboratory in summer, fall, and winter months and have been collected in the field in April, May, and June, suggesting that emergence is not controlled by 200 Entomologica Americana [Vol. 45, No. 3 photoperiodicity. Further study might reveal some interesting dif- ferences in the physiological responses of this and other members of the tribe. Novelsis athlophora Beal Novelsis athlophora Beal, 1954, pp. 81, 89. Adults : Male described by Beal ( 1954 ) . Mature Larvae: Unknown. Type: Type locality: Beaver Canyon, Idaho. Holotype in the collection of the United States National Museum. No other localities recorded in description. New Records: British Columbia : Aspen Grove, June 17, 1931 (H. Richmond); Midday Valley, Merritt, June 4, 1924 (K. F. Auden). Idaho : Rocky Point, Benewah County. Washington : Dayton, July (L. Turner). Diagnosis: This is a dark brown species sparsely clothed with long, suberect, black hairs. There are no golden-brown or other light-colored hairs on the dorsal or ventral surfaces. Both male and female antennae are 10-segmented, a character that distinguishes the species from all other Nearctic species with uniformly dark dorsal pubescence, except A. schaefferi. The shape of the antennal club of the male, in which the length of the first two segments combined exceeds the length of the terminal segment by at least a fourth, separates the species from A. schaefferi , as also does the long, sparse setae of the body. Attagenus lobatus Rosenhauer Attagenus lobatus Rosenhauer, 1856, p. 108. — Jacquelin du Val, 1859, p. 254. — Mulsant and Rey, 1868, pp. 100-101. — Hinton, 1945, pp. 323-324.— Zhantiev, 1963b, p. 415. Attagenus byturoides Solsky, 1876, pp. 272-273. — Reitter, 1889, p. 557. — Solodovnikova, 1938, pp. 1-20. Attagenus sericeus Reitter, 1880 (1881) ( non sericeus Guerin, 1829), p. 79.— Reitter, 1887, p. 51. Adults: Adequately described by Hinton (1945). Adult head shown in Figure 2; dorsal outline in Figure 8. Mature Larvae: Dorsal integument light brownish yellow. Setae uniformly light golden-brown. Head : Antenna with terminal seta about twice as long as length of terminal segment; no setae present on second segment. Epipharynx with distal series of two sensory pits, each slightly larger in diameter than diameter of sensory pits of middle series. 1970] Beal: Taxonomic Study of Attagenini 201 Maxillary palp with two setae on penultimate segment. Labium with about four ensiform and four slender setae on dorsal (inner) side of each lobe of ligula; palp without seta on first segment. Body setae : Margins and ribs uniformly smooth. Setae inserted on acrotergites about half as long as length of tergum or longer and gradually becoming extremely fine (almost beyond point of resolu- tion except with oil-immersion lens); proximal part of shaft with very fine ribs (also difficult to observe except with high magnifica- tion). Setae on tergites consisting of linear marginal and submarginal hairs and of lanceolate hairs with five to seven ribs between margins; only lanceolate setae present on disc of tergite. Submarginal setae suberect, inserted nearly at same level as marginal setae. Sockets of submarginal, suberect setae usually with two adjacent sensory pits. Sterna with lanceolate setae only on disc; lanceolate setae with seven to eight ribs between margins; anterior margin of each sternum with row of stout linear setae one to two times as long as lanceolate setae; numerous fine, long, linear, simple setae inserted anterior to row of stout setae. Spiracle and associated structures : Spiracle closed behind by tergum; anterior margin of spiracle more or less straight; spiracle adjacent to spiracular sclerite. Spiracular sclerite elongate-oval, bearing eight to 10 or more setae, closed behind by narrow margin of tergum, but often not appearing so, since spiracular sclerite may overlap margin; area of tergum in front of spiracular sclerite well- sclerotized and continuous with tergal margin overlying anterior edge of sclerite (Fig. 15). Abdomen : Eighth abdominal tergum with antecostal suture. Ninth abdominal tergum about half as long as tergite of eighth abdominal segment. Two small sclerites present in sternum of ninth abdominal segment. Legs’. Femur of prothoracic leg about three-sevenths as wide as long; ventral margin about two-thirds as long as length of seg- ment. Tibia and pretarsus as illustrated (Fig. 28); setae at base of pretarsus slender, subequal in width and length. Distribution: The Palearctic distribution extends from Spain to Mongolia including Bulgaria and Uzbek, U.S.S.R., on the north and Tunisia, Egypt, Arabia, and Afghanistan on the south (Mroczkowski, 1968). It is apparently established in the cities of Detroit, Michigan (February 2, 1960; J. G. Hunter) and New York, New York (May 15, 1964; Jack Lipes). Diagnosis: Adults of A. lobatus are uniformly clothed with 202 Entomologica Americana [Vol. 45, No. 3 golden-brown pubescence. The pubescence is quite short except for noticeably longer tufts of hairs on the margin of the basal lobe and each lateroposterior angle of the pronotum. The integument of the elytra is reddish brown. The integument of the pronotum is similarly colored, or at most only a little darker. The most distinctive char- acter of the adults and one that immediately distinguishes it from all other Nearctic species is the elongated and truncated basal lobe of the pronotum (Fig. 8). The species is also unique among Nearctic forms in possessing a carina along the dorsal margin of the front tibia. The antenna is 1 1 -segmented. The form of the antennal club is somewhat similar to that of N. uteana , with the terminal segment subequal in length to the combined length of the first two segments. Superficially the adults resemble some females of N. uteana more than any other Nearctic species. However, the dorsal pubes- cence of N. uteana is definitely lighter in color, the pronotal integu- ment is brownish black and contrasts sharply with the reddish yellow elytral integument, the dorsal hairs are much longer, particularly along the margins of the pronotum and elytra where many hairs are as long as the width of the eyes, the front tibia lacks a dorsal carina, and the antenna is 10-segmented. Mature and semi-mature larvae are easily distinguished from all other known larval forms by the fact that there are no linear or linear-lanceolate setae intermingled with the broad lanceolate setae of the sterna. A. elongatulus, A. schaefferi, and A. pellio have similarly broad sternal setae, but each has linear setae inserted among the lanceolate setae, at least on the sides of each sternum. N. varicolor and N. aequalis both have lanceolate setae with as many ribs as are found in the setae of A. lobatus , but the margins of the setae and the ribs are strongly serrulate or denticulate and do not at all resemble the smoothly margined setae of A. lobatus. A further distinguishing character is the presence of a row of moderately stout linear setae inserted on each sternum in front of the lanceolate setae. The linear setae are longer than the lanceolate setae. Anterior to these a number of fine simple setae is inserted on each sternum. No other Nearctic species has a row of stout linear setae near the anterior margin of each sternum. In other species the sternal setae become gradually smaller toward the an- terior end of the segment. The larvae of A. lobatus are also unique in that the spiracular sclerite bears eight to ten or more setae instead of six or fewer. Ecology: Zhantiev (1963a) found this species in nests of 1970] Beal: Taxonomic Study of Attagenini 203 desert owls ( Athene noctua bactriana Hutt.), Old World jumping rats or gerbils (Gerbillinae), long-clawed ground squirrels ( Sper - mophilopsis leptodactylus Licht.), and various predatory mammals such as foxes and badgers in sandy and clayey desert regions of Turkmenia and Tadzhikistan in the U.S.S.R. He observed there is a tendency for this species to select nests of desert owls found at the base of cliffs, while other species of Dermestidae more commonly occupy nests constructed in crevices higher on the cliffs and still different species occupy nests in sandy holes. Hinton (1945) summarizes reported occurrences of this species as an economic pest, listing the following products on which it feeds or in which it has been found: skins, furs, feathers, woolen goods, museum specimens, grain and refuse in granaries, and red pepper. In the United States the species has been found in buildings. Larvae were reported by J. G. Hunter to bite the skin of humans and the bites were followed by a slight itching. Novelsis perplexa (Jayne) Attagenus perplexus Jayne, 1882, p. 35. Novelsis perplexa : Casey, 1900, p. 194. — Beal, 1954, p. 77.— Hatch, 1962, p. 284. Adults: Redescribed by Beal (1954). Other stages remain unknown. Recorded Distribution: The type locality is Nevada. Other records include scattered localities from Southern California north to British Columbia. Additional Records: British Columbia : Vernon, August 2, 1929 (R. Hopping). California : Fresno County: Tollhouse, June 30, 1961 (C. D. Johnson). Madera County: North Fork, July 19, 1963 (C. D. Johnson). Nevada County: 3 miles north of Boca, September 4, 1960 (F. D. Parker). Plumas County: Meadow Valley, July 22, 1961 (T. Gantenbein). Riverside County: San Jacinto Mountain Trail, July 1, 1952 (J. F. Powers). San Diego County: Mount Palomar, June 21, 1959 (M. E. Irwin). Tulare County: Kern River, 6,000 ft. elev., July 29, 1899 (probably L. L. Muchmore). Yolo County: Winters, August, 1961 (J. L. Camp- bell). Idaho : Krassel Ranger Station, Martin Creek, July 26, 1956 (S. Stevens). Oregon : Klamath County: Bly, July 20, 1956 (M. Wasbauer); Swan Lake, August 8, 1953 (Joe Schuh). Taxonomic Position: Judging by similarities of adult char- acters, this species is closely related to N. varicolor and N. aequalis. The allopatric pattern of distribution of the three species further 204 Entomologica Americana [Vol. 45, No. 3 suggests their relatively recent divergence from a common origin. Whether a mechanism for reproductive isolation exists between the forms is a question that remains unanswered. Diagnosis: The following characters are common to adults of N. perplexa, N. varicolor, and N. aequalis. The dorsal pubescence consists of black, golden-brown, and white hairs. The light-colored hairs form a thin basal band or basal patches, a semicircular subbasal band, an irregular and often interrupted submedian band, and an irregular subapical band or patch. The hairs are subrecumbent on the disc but suberect on the lateral margins. The hairs are also much longer than for most other Attagenini, the longest being longer than the scutellum, giving the specimens a somewhat shaggy ap- pearance. The segments of the male antennal club are densely clothed with very fine, erect pubescence with the individual hairs about two-thirds as long as the width of the third segment of the antenna. N. perplexa may be distinguished from N. varicolor and N. aequalis by the following characters. The antenna is 10-segmented. The first two segments of the male antennal club combined are about one-sixth longer than the terminal segment. The other two species have an 11 -segmented club and the first two segments of the male club combined are about one-third longer than the terminal segment. The elytral integument of N. perplexa is entirely black or black with reddish maculae corresponding in pattern to the areas of light pubescence. N. aequalis may rarely have the elytral integu- ment entirely black, but ordinarily it has a suffused reddish macula near the middle of the elytra. N. varicolor has yellowish brown maculae that may correspond to areas of light-colored pubescence or that may coalesce to varying degrees. Occasionally each elytron may have one large yellowish macula that extends over the entire elytron except for the black margins. Ecology: Virtually nothing is known of the ecology of the species. Adults have been collected at light and on flowers of yarrow ( Achillea ) and milkweed ( Asclepias ). An adult has also been collected on Douglas-fir ( Pseudotsuga Menziesii (Mirb.) Franco). Novelsis varicolor (Jayne) Attagenus varicolor Jayne, 1882, p. 357. Novelsis varicolor: Casey, 1900, p. 149. — Beal, 1954, pp. 78—80. Adults : Redescribed by Beal ( 1 954 ) . 1970] Beal: Taxonomic Study of Attagenini 205 Mature Larvae: Dorsal integument brown. Setae of head and dorsal surfaces brownish black; setae of ventral surfaces choco- late brown. Head : Antenna with terminal seta five-ninths as long as length of terminal segment; no setae present on second segment. Epipharynx with distal series of two sensory pits, each with diameter about half diameter of sensory pits of middle series. Maxillary palp with nine to 14 setae inserted on third segment. Labium with six setae on dorsal (inner) side of each lobe of ligula; medial two setae narrowly ensiform with lateral setae becoming nearly terete; first segment of palp without seta. Body setae : Margins and ribs serrulate to denticulate. Acro- tergites of anterior abdominal segments with lanceolate and ovate- lanceolate setae with three to six ribs between margins (Fig. 17). Disc of tergites with setae regularly or irregularly ovate with acute to obtuse apices and with as many as 11 to 13 ribs between margins (Fig. 19). Sockets of submarginal erect setae with two adjacent sensory pits, one on each side. Setae of sterna lanceolate and ovate- lanceolate with elongated apices. First abdominal sternum with setae at midline not appreciably longer than other setae of disc. Eighth abdominal sternum with broadest setae having six to eight ribs between margins; setae becoming slightly smaller toward anterior margin of sternum without row of longer linear setae. Spiracle and associated structures : Spiracle enclosed by tergum; anterior margin more or less straight and usually projecting over posterior margin; distance between spiracle and spiracular sclerite roughly equal to width of trachea (Fig. 13). Spiracular sclerite nearly round, enclosed by tergum; tergum on lateroposterior side of sclerite reduced to narrow margin. Abdomen : Eighth abdominal tergum without antecostal suture. Ninth abdominal tergum about two-sevenths as long as eighth ab- dominal tergum. Two small sclerites present in sternum of ninth abdominal segment. Legs'. (Ratio of length to width of femur not observed); tibia of prothoracic leg with usual two rows of stout ventral setae; posterior face with stout subdorsal apical seta and submedian group of ir- regularly arranged four or five stout setae and several slender setae. Pretarsus with posterior seta inserted at base about twice as wide as anterior seta. Distribution: Previously described (Beal, 1954) as extending from El Centro, California, to Presidio, Texas; on the south from 206 Entomologica Americana [Vol. 45, No. 3 Carr Canyon in the Huachucha Mountains, Arizona, north to Hoover Dam and to Globe, Arizona. New records outside this range are the following: Arizona: Sedona, Coconino County, June 20, 1964 (R. S. Beal). California : Indian Wells, Riverside County, April 19, 1961 (G. H. Nelson); Mitchell’s Cavern, San Bernardino County, April 16, 1962 (G. H. Nelson). Sonora, Mexico : Masiaca Huata- bampo, May 28, 1955 (F. Pacheco M. ). Diagnosis: The following characters distinguish adults of N. varicolor from those of N. perplexa and N. aequalis. The integu- ment of the elytra is black or piceous and marked with yellowish tan or yellowish brown maculae. The maculae rarely correspond in extent to the subbasal, submedian, and subapical areas of light pubescence, but commonly the subbasal and submedian maculae coalesce to form one spot. At times there is a single large macula that occupies the entire elytron except for a narrow margin of black. The pubescence is always tricolorous, although the light-colored hairs are predominately white and the golden-yellow hairs may be limited to a few near the margins of the bands of light-colored pubescence. The species may be distinguished definitely from N. perplexa by its 1 1-segmented rather than 10-segmented antenna. Characters that adults of this species have in common with N. perplexa and N. aequalis and that distinguish them as a group from other groups within the tribe are discussed under the diagnosis of N. perplexa. Larval characters shared by this species and N. aequalis are listed under Relationships, Group IV. Characters distinguishing larvae of N. varicolor and N. aequalis are discussed under the diagnosis of the latter. Ecology: Larvae have been found in Sceliphron mud dauber nests (Beal, 1951), in pigeon droppings, in hens’ nests, in nests of English sparrows, in a cactus wren’s nest, and in the nest of an unidentified spider in a chicken house. I collected the species in Tucson, Arizona, in moderate numbers from sacks of feed in a granary. The feed was infested with a number of other insects, and it was probably feeding on their dead remains rather than on the grain. It was also collected by Gus A. Amado in a seed and feed mill in Nogales, Arizona. I have reared the species through several generations on dried dog food. Adults have been collected on mesquite [Prosopis juliflora (Swartz) DC.] and on catclaw ( Acacia greggii A. Gray). At room temperatures the species matures in a year. Out-of- doors adults appear from May through July. In the laboratory adults 1970] Beal: Taxonomic Study of Attagenini 207 may emerge as early as February and March but not after June. This pattern suggests that emergence is governed by photoperiodicity but may be accelerated by warm temperatures. Novelsis aequalis (Sharp) Genattus aequalis Sharp, 1902, p. 646. Attagenus aequalis : Hinton, 1945, p. 306. Novelsis aequalis: Beal, 1954, p. 80. Adults : Redescribed by Beal ( 1954 ) . Mature Larvae: Dorsal integument brown. Setae of all sur- faces black except for some golden hairs on legs. Head : Antenna with terminal seta subequal in length to length of terminal segment; one or no seta present on second segment. Epipharynx with distal series of two sensory pits, each with diameter about half diameter of sensory pits of middle series. Maxillary palp with two setae inserted on third segment. Labium with six setae on dorsal (inner) side of each lobe of ligula; medial four or five setae narrowly ensiform; lateral one or two setae terete; first segment of labial palp without seta. Body setae : Margins and ribs serrulate to denticulate. Acro- tergites of anterior abdominal segments with linear and ovate-lanceo- late setae; largest setae with 10 to 11 ribs between margins. Tergites with linear and irregularly rounding to ovate setae on disc; rounding and ovate setae terminating in single acute apex or two acute apices with as many as 15 to 16 ribs between margins of widest setae (Fig. 23); setae on posterior margins broadly linear with acute apices and eight to 10 ribs present between margins on dorsal side and 16 to 22 ribs present between margins on ventral side. Sockets of submarginal erect setae with two adjacent sensory pits, one on either side. Sterna with linear and linear-lanceolate setae; linear- lanceolate setae with as many as nine ribs present between margins; setae becoming smaller toward anterior margin of each sternum; setae on midline of first abdominal sternum longer than other setae of disc. Spiracle and associated structures : Spiracle closed behind by tergum, removed from spiracular sclerite by distance equal to about half width of spiracle; opening simple, somewhat oval. Spiracular sclerite rhomboidal to ovate, encircled by rim-like extension of sclerotized part of tergum; area of tergum immediately anterior to front margin of rim unsclerotized; entire structure appearing to project peninsula-like from posterolateral margin of tergum (Fig. 24). 208 Entomologica Americana [Vol. 45, No. 3 Abdomen'. Eighth abdominal tergum with antecostal suture. Ninth abdominal tergum about one-third as long as tergite of eighth abdominal segment. Legs : Femur of prothoracic leg about 10/17 as wide as long; ventral margin about 8/17 as long as length of segment. Posterior face of front tibia with stout subdorsal apical seta and subventral row of four stout setae a little above ventral two rows of stout setae; most proximal stout seta at about basal two-fifths. Pretarsus with posterior seta about twice as wide as anterior seta. Recorded Distribution: Lee and Kerr counties in Texas south to Veracruz, Mexico. The type locality is Mexico without being further specified. New Records: Maryland : Baltimore, July 10, 1940 (H. Howden); College Park, as larvae October 11, 1963 (J. H. Fales). Pennsylvania : One specimen with locality not further specified (Horn Collection). Texas : Burnett County, June 4, 1953 (D. J. and J. N. Knull); Randall County, July 7, 1950 (D. J. and J. N. Knull). Diagnosis: Both adult and larval characters associate the species with N. varicolor and N. perplexa. Characters grouping these three species are discussed under Relationships. Adults may be separated from N. perplexa by the 11 -segmented rather than 10-segmented antenna. Adults are sometimes difficult to separate from N. varicolor , although usually N. aequalis lacks a clearly defined light maculate spot on the elytron, having only a suffused reddish spot at about the basal third, in contrast to the more or less sharply defined, yellowish tan maculae of N. varicolor . Occasional specimens of N. aequalis have a completely black or brown elytral integument. Some specimens of N. varicolor have a yellowish brown macula that occupies most of the elytron, making them appear quite similar to N. aequalis. However, the elytra of these specimens have definitely dark margins and a sutural line. Adults may also be distinguished by the differences in the form of the prosternum described in the key, but the prosternum is often difficult to observe without relaxing the specimen and moving the antennae and legs out of the way. In contrast to the adults, the larvae of the two species are marked by a number of obvious differences. The maxillary palp of N. aequalis has but two setae on the third segment in contrast to the nine or more of N. varicolor. The spiracular sclerite of N. aequalis appears detached from the tergum, since the tergum is unsclerotized in front of the rim surrounding the spiracular sclerite. Conversely, 1970] Beal: Taxonomic Study of Attagenini 209 the spiracular sclerite of N. varicolor appears to be completely im- bedded within the lateroposterior corner of the tergum. The setae of the acrotergites of N. aequalis consist of both broad, ovate- lanceolate hairs and very long and narrow, almost simple hairs. The setae of the acrotergites of N. varicolor are all linear-lanceolate to ovate-lanceolate, with three or more ribs present between the margins. An interesting although perhaps not invariable difference between the species is found in the shape of some of the dorsal setae of N. aequalis. The ovate hairs of this species, instead of tapering to a single acute point, frequently terminate in two acute apices (Fig. 23). Ecology: Almost nothing is known of the ecology of this species. Specimens have been intercepted at the Brownsville, Texas, port of entry on tomatoes and on orchid flowers, but these are no doubt accidental associations. A single specimen was found at Brownsville in milled wheat flour from Mexico. The species has apparently become established in Maryland, where it is probably able to survive in heated buildings. Attagenus rufipennis LeConte Attagenus rufipennis LeConte, 1859, p. 71. — Casey, 1900, p. 147. Attagenus nigripes Casey, 1916 ( non nigripes Fabricius, 1792), p. 184 (New synonymy). Attagenus atrolucens Casey, 1916, p. 183 (New synonymy). Adult Males: Integument of head and pronotum brownish black to black; elytra reddish, reddish brown, or black and if reddish or reddish brown usually but not invariably with blackish base and diffused blackish sutural line; ventral surfaces dark reddish brown to black; legs and flagellar shaft of antennae usually some- what lighter than ventral surfaces; antennal club brownish black to black. Pubescence of head and dorsal surfaces subrecumbent, entirely black, or black with oblique subbasal band of silver or golden-yellow hairs, or black with silver or golden-yellow hairs scattered among black hairs on lateral margins of pronotum and on posterior four-fifths of elytra. Eye emarginate over base of antenna. Antenna 1 1 -segmented; ratio of combined length of first two seg- ments of club to length of terminal segment varying from 1:3.7 to 1:4.8; ratio of length of terminal segment to length of pronotum and elytra combined varying from 1:6.5 to 1:7.9. Channel below eye for reception of flagellar shaft of antenna deeply concave with anterior margin forming carina; margin of carina visible from front 210 Entomologica Americana [Vol. 45, No. 3 of head; carina projecting knife-like beneath head and curved behind base of maxilla to meet gular suture. Pronotum with lateral margin continued around anterolateral angle; basal lobe gradually rounding or truncate but not produced abruptly posteriad. Punctures of disc about three times diameter of facet of eye and separated by one to one-half times diameter of single puncture; punctures toward sides becoming larger and contiguous. Setae on basal lobe and latero- posterior angles of pronotum subequal in length to setae of disc. Elytron with hairs of disc about as long as combined length of third and fourth segments of antenna. Hypomeron a little inflated or flat on anterior half, occasionally slightly concave on posterior half. Posternum with posterior margin of lateral lobe slightly re- flected against lateral half of procoxa; reflected part less than one- third as long as length of horizontal part of lobe and forming angle of 45° or less with horizontal part; prosternal process slightly wider at apex than between coxae; thread-like carina present on process but not extending onto disc of prosternum; thread-like transverse carina separating anterior declivity of prosternum from disc; no denticle present at middle of anterior margin of disc. Epipleuron terminating gradually behind middle of first sternum. Ventral plate of hind coxa forming distinct tooth lateral to insertion of trochanter; plate terminating at posteromedial angle of metepisternum and not meeting metepimeron behind metepisternum. Protibia not carinate on dorsal margin. Mesofemur with anteroventral and posteroventral margins of crural cavity about equally produced and on same plane. Length (of pronotum and elytra combined): 3.0 mm. to 4.1 mm. Adult Females: Antennal club with terminal segment sub- equal to one and two-fifths as long as combined length of two pre- ceding segments; tenth segment equal in length to ninth segment. Length (of pronotum and elytra combined): 3.5 mm. to 4.9 mm. Mature Larvae: Dorsal integument brown. Setae of head golden-brown; setae of nota and terga black; setae of venter of thorax and legs golden-brown; setae of abdominal sterna black to brownish black. Head : Antenna with terminal seta subequal in length to length of terminal segment; no setae present on second segment. Epipharynx with distal series of two sensory pits, each with diameter about half diameter of sensory pit of middle series. Maxillary palp with two setae on penultimate segment. Labium with four to five narrowly ensiform setae on dorsal (inner) side of each lobe of ligula; setae becoming narrower laterad; first segment of labial palp without seta. 1970] Beal: Taxonomic Study of Attagenini 211 Body setae : Margins and ribs denticulate (Fig. 18). Acro- tergite of first abdominal segment with linear and linear-lanceolate setae; widest setae with three ribs between margins. Disc of tergite with intermingled linear and lanceolate setae; widest lanceolate setae usually with four but sometimes five ribs between margins. Sockets of three medial submarginal erect setae with two adjacent sensory pits, one one either side; socket of most lateral erect seta often with one, occasionally two, adjacent sensory pits. First abdominal ster- num without longer setae along midline. Sternum of eighth abdominal segment with setae all linear; widest with three, occasionally four, ribs between margins; setae becoming smaller on anterior area of sternum without row of stouter, longer setae near anterior margin. Spiracle and associated structures : Anterior abdominal seg- ments with spiracle not closed behind by tergum; anterior margin of spiracle rounding; spiracle adjacent to spiracular sclerite (Fig. 12). Spiracular sclerite oval, not enclosed by tergum; four setae ordinarily inserted on sclerite. Tergum forming sclerotized ridge overlying anterior margin of sclerite but tergum not sclerotized in front of ridge, so that sclerite and ridge appear to project peninsula- like beyond lateral margin of tergum. Abdomen : Eighth abdominal tergum with antecostal suture. Ninth abdominal tergum about two-sevenths as long as tergite of eighth abdominal segment. Two small sclerites present in sternum of ninth abdominal segment. Legs : Femur of prothoracic leg about half as wide as long; ventral margin about three-eights as long as length of segment. Tibia with spines arranged as illustrated (Fig. 26). Base of pre- tarsus with lateral seta twice as wide as medial seta; medial seta extending to about middle of claw. Type Locality: Fort Tejon, California (Lebec, Kern County). The type locality for A. nigripes is Milpitas, Santa Clara County, California. The type locality for A. atrolucens is Indiana. The type specimen of A. atrolucens was from the Levette Collection, which contained many erroneously labeled specimens. I can find no differences between the type specimen of A. atrolucens and many individual specimens from California. Since the species has not subsequently been collected in the Midwest, it must be concluded that the locality is in error. New Records: Arizona : Cochise County: Cave Creek, Chiricahua Mts., as larva January 5, 1963 (Vincent Roth). Coco- nino County: Basin Camp, Bar 77 Ranch, 20 miles west of Cameron, 212 Entomologica Americana [Vol. 45, No. 3 as larvae September 27, 1964 (R. S. Beal); Williams (Barber and Schwarz). Gila County: Globe, April 2, 8, 10, 23, 1933, April 7, 1948 (F. H. Parker); base of Pinal Mt. (D. K. Duncan). Maricopa County: Canyon Lake, upriver, March 31, 1959 (Ivan Jennings); Wickenburg, March 5, 1958 (G. G. Gose). Navajo County: Linden, as larvae August 27, 1957 and August 20, 1962 (R. S. Beal). Yavapai County: Cottonwood, as larvae March 30, 1963 (R. S. Beal). California : Alameda County: Hills back of Oakland, April 12, 1908 (Van Dyke Collection). Butte County: Oroville, March 7, 1928 (H. H. Keifer). Calaveras County: Murphys, 2,500 ft. elev., May 14, 1937 (F. E. Blaisdell). Colusa County: Arbuckle, March 30, 1962, April 2, 6, 9, 1962 (P. M. Marsh). Contra Costa County: Berkeley, January 27, 1947 (A. J. Walz); Berkeley, February 27, 1932 (R. P. Allen); Danville, March 23, 1952 (F. X. Williams); Orinda, March 15, 1955 (R. H. Goodwin); Walnut Creek, April 13, 1932 (F. E. Blaisdell); Vine Hill, November 25, 1910 (F. E. Blaisdell). El Dorado County: Fallen Leaf, 6,500 ft. elev., July 3, 1935 (F. E. Blaisdell); Placerville, February 23, 1916 (E. T. Armstrong); Pyramid Ranger Station, July 12, 1948 (P. D. Hurd). Fresno County: Fresno, March 9, 1936 (H. C. Donohoe); Huckleberry Meadow, 6,500 ft. elev., May 10, 1910 (Hopping). Humboldt County: Greenpoint, June 14, 1916 (F. E. Blaisdell). Inyo County: Argus Mountains, April, 1891 (A. Koebele); Inde- pendence, April 30 and May 8, 11, 1918 (L. L. Muchmore); Gray’s Camp, Independence, May 3, 1919 (L. L. Muchmore); Lone Pine, May 23, 1930 (R. Hopping); Westgard Pass, June 18, 1955 (M. E. Irwin). Lake County: Anderson Springs, May 27, 1951 (W. R. Bauer); Kelseyville, May 15, 1922 (E. P. Van Duzee). Los Angeles County: Crystal Lake, June 29, 1950 (A. T. McClay); Lancaster, April 7, 1927 (L. L. Muchmore); Los Angeles (Coquillett) ; Mount Wilson, April 30, 1916 (J. O. Martin collection); Pasadena (A. Fenyes); Tujunga, May 20, 1944 (L. R. Gillogly); Whittier, March 26, 1916, and March 26, 1919 (L. L. Muchmore). Marin County: Mill Valley, March 25, 1957, April 5, 1959, April 6, 1954, May 4, 1959, May 13, 1956, May 28, 1951, June 5, 1949 (H. B. Leech); Novato, September 22, 1959 (T. R. Haig). Mariposa County: Yosemite National Park (probably Yosemite Valley), May 16, 20, 1934 (O. Bryant); same locality, May 22, 1908 (E. T. Cresson, Jr.); same locality, May 31, 1952 (R. Schuster); same locality, June 7, 9, 1930 (F. E. Blaisdell). Mendocino County: locality not further specified, October 20, 1918 (E. R. Leach). Modoc County: 1970] Beal: Taxonomic Study of Attagenini 213 Warner Mountains, July 10, 1919 (R. Hopping collection). Monte- rey County: Carmel, November 8, 1911 (L. S. Slevin collection); Paraiso Springs, April 24, 1914 (L. S. Slevin collection). Nevada County: Truckee, June 17, 1927 (E. P. Van Duzee). Plumas County: Chester, June 25, 1951 (D. J. and J. N. Knull); Meadow Valley, 4-5,000 ft. elev., June 4, 1924 (E. C. Van Dyke); Onion Valley, July 6, 1952 (E. I. Schlinger); Sunnyside near Seneca, June I, 1923 (V. S. Barber). Riverside County: Banning, April 6, 1933 (R. Hopping); 4 miles west of Forest Home, as larva September 16, 1951 (Ray Ryckman); Hemet, March 30, 1937 (L. D. Christen- son); 7 miles north of Idyllwild, San Jacinto Mountains, 5,500 ft. elev., May 8, 1954 (G. H. Nelson); Keen Camp, June 6-12, 1917 (E. P. Van Duzee). Sacramento County: Folsom, May 7, 1941 (A. T. McClay). San Benito County: Pinnacles National Monu- ment, April 24, 1948 (W. W. Middlekauff ) ; Pinnacles National Monument, April 28, 1914 (L. S. Slevin). San Bernardino County: Camp Baldy (Mt. Baldy), June 26, 1950 (M. J. Stebbins); 20 miles northeast of Redlands, 9,000 ft. elev., June 26, 1954 (G. H. Nelson); Victorville, May 2, 1953 (G. A. Marsh and R. O. Schuster). San Diego County: San Diego, February (F. E. Blaisdell); Poway (F. E. Blaisdell). San Francisco County: San Francisco, April 24, 1959 (MacNeill). San Luis Obispo County: Paso Robles, as larvae August 2, 1960 (R. S. Beal). Santa Barbara Countv: San Marcos Pass northwest of Santa Barbara, as larvae August 1, 1960 (R. S. Beal). Santa Clara County: Pacheco Pass, April 14, 1949, and as larvae August 2, 1960 (R. S. Beal); San Jose, April 29, 1928 (L. S. Slevin). Santa Cruz County: Santa Cruz Mountains (Koebele collection). Sonoma County: Petaluma, April 2, 1961 (G. M. Trenam); Santa Rosa (Lois B. Stiles). Stanislaus County; New- man, March 9, 1955 (C. G. Moore). Tehama County: Red Bluff, May 1, 1922 (V. S. Brown). Tulare County: Kaweah (R. Hop- ping); Lodgepole Camp, Sequoia National Park, June 24, 1945 (A. T. McClay); Marble Fork Bridge, Sequoia National Park, June II, 1929 (Van Dyke); Wolverton, Sequoia National Park, June 25, 1929 (Van Dyke). Ventura County: Ozena Station (Cuyama River at State Route 33), as larvae April 13, 1963 (R. S. Beal). Yolo County: Davis, March 15, 1950 (A. T. McClay); same locality, April 22, 1947 (Brad Stevens); Davis, May 2, 1949 (E. I. Schlinger); Davis, June 7, 1949 (R. C. Bechtel); Putah Canyon, March 26, 1960 (M. E. Irwin); Rumsey, March 31, 1962 (C. G. Moore); Winters, March 30, 1962 (P. M. Marsh). Yuba County: 214 Entomologica Americana [Vol. 45, No. 3 10 miles south of Marysville, April 19, 1956 (W. W. Middlekauff). New Mexico : Hidalgo County: Granite Pass, April 6, 1965 (F. D. Parker). Oregon : Jackson County: Green Spring Pass, June 15, 1938 (M. H. Hatch); Griffin Creek, June 6, 1957 (C. Fitch); Medford, as larvae January 12, 1947 (A. T. McClay); Medford, February 9, 1946 (A. T. McClay); Medford, March 1, 1936 (Lawrence); Medford, May 2, 1941, May 2, 1945, May 6, 1945 (C. Fitch); Medford, as larvae October 1, 1946 (A. T. McClay); Talent, March 15, 1932, May 13, 1932 (L. G. Gentner). Josephine County: Grants Pass, April 12, 1934 (Frank M. Beer). Wasco County: Bear Springs (nine miles west of Pine Grove), July 20, 1940 (K. M. and D. M. Fender); Bear Springs Ranger Station, September 6, 1952 (P. W. Orr); Maupin, June 6, 1937 (K. M. Fender). Yamhill County: Dayton, April 24, 1945 (K. M. Fender). Utah : Washington County: May 21, 1951 (G. F. Rnowlton). Washington : Kittitas County: Vantage, May 13, 1956 (H. M. Hatch). Baja California : Tecate, March 12, 1957 (Burciaga and Valdez). A specimen collected at San Antonio, Texas, by H. Soltau is deposited at the United States National Museum. This locality ap- pears to be outside the natural range of the species. The species was probably introduced into the area by commerce, and there is at present no evidence that it has become established there. Diagnosis: In size and shape and often in dorsal coloration adults of this species resemble A. megatoma. Males further resemble A. megatoma in the general form of the antennal club, in which the terminal segment is much longer than the first two segments com- bined. However, the fact that the hind coxa and the metepimeron do not join behind the metepisternum distinguishes this species from A. megatoma and all other members of Group I. Adults are separated from species of other groups by the form of the male antennal club and by one or several of the following characters: the antenna is 1 1 -segmented; a small auricle-like lobe is present on the hypomeron behind the base of the procoxa; there is no knife-like carina along the dorsal margin of the anterior tibia; although there may or may not be a subbasal elytral band of light pubescence, there are no distinct submedian or subapical spots, lines, or bands of light pubes- cence on the elytra. Differences between the larvae of this species and those of some- what similar species are listed under Relationships, Group V. Infraspecific Forms: This is a highly polytypic species that 1970] Beal: Taxonomic Study of Attagenini 215 cannot conveniently be divided into a few manageable subspecies. A consistent treatment would require that either the subspecies be ignored or that a considerable number of subspecies be designated. If only the color of the elytra were to be considered, the species could easily be separated into the black northwestern form that Casey (1900) named A. nigripes , the nominate southern California form with red elytra, and an Arizona and New Mexico form with mahogany elytra. However, there is no reason why the presence or absence of a subbasal elytral fascia of light-colored hairs and the presence or absence of light-colored hairs on the pronotum should not be given equal weight in determining subspecific limits, since these characters are also geographically ordered. The difficulty is that they are not geographically concordant with the distribution of color variations in the elytra. Black specimens from Merced County, California, and northward generally lack light-colored hairs on the pronotum and elytra. Southward through central and coastal California the black forms commonly have a short subbasal elytral fascia of silvery hairs. However, around San Diego, California, black forms are again found without light-colored hairs. Forms with red elytra occur in the California desert areas, but are also found on the east side of the coastal ranges as far north as San Benito County and along the coast from Los Angeles south to San Diego. One red specimen has been taken at Berkeley, California, and two red specimens in Wasco County, Oregon. Ordinarily the red California specimens have at least a few silvery hairs forming a subbasal elytral fascia. Inter- mediates between the red and the black forms are not uncommon. A long series from Ozena Station on the north side of the San Rafael Mountains in Ventura County includes numerous intermediates between the red and the black forms as well as specimens with and without elytral fasciae. So also does a shorter series from Lancaster in Los Angeles County. No collections of the species have been made between Victorville, California, and Wickenburg, Arizona. Nevertheless, the species is probably continuous; the single specimen from Wickenburg is identical to many Mojave Desert specimens. A moderate number of specimens has been taken in the neighborhood of Globe, Arizona, and at Linden in Navajo County, Arizona. These all have mahogany elytra with a broad subbasal elytral fascia of golden-yellow hairs. Another long series collected a relatively short distance north near the eastern end of the Grand Canyon but sep- arated from the southern Navajo County population by an interven- ing stretch of desert has mahogany elytra but no elytral fasciae. 216 Entomologica Americana [Vol. 45, No. 3 In view of this somewhat confusing picture, it seems inadvisable to name any of the infraspecific forms until much more extensive collections have been made and until the population dynamics are better understood. Ecology: Larvae have been taken in nests of the wood rat (Neotoma) , in webs of the spider Physocyclus tanneri Chamberlin, and in insect-infested grain trash in a granary. Linsley (1944, 1946) has recorded it in nests of the English sparrow, house finch, black phoebe, barn swallow, and cliff swallow. It is evidently a general scavenger on dried protein materials, as are most other members of the genus. H. B. Leech found adults emerging from stems of dead Ceanothus thrysiflorus Eschscholtz, but in all probability they were feeding on the dead remains of other insects in burrows in the stems. The species has been found on one occasion infesting an insect collection. Specimens from Talent, Oregon, are labeled by L. G. Gentner as “breeding in casein.” Probably adults ordinarily fly to flowers, where they feed on nectar and pollen. They have been collected on apple blossoms, lupine, Ceanothus cordulatus Kellogg and Ceanothus divaricatus Nuttall. Flying to flowers does not appear to be necessary for reproduction for Arizona forms of the species, since I have been able to carry them through several generations without it. On the other hand, although I have been able to rear larvae of the black forms to maturity in the laboratory, the adults have never produced a second generation in the laboratory. It may be that in contrast to the Arizona forms they need to feed on nectar or pollen before being able to mate. Novelsis uteana Casey Novelsis uteana Casey, 1900, p. 148. — Beal, 1954, pp. 82-84. Novelsis byturoides Casey, 1900, p. 148. Adults : Redescribed by Beal ( 1 954 ) . Mature Larvae: Integument of head brownish black; integu- ment of nota and terga light reddish brown with prominent blackish brown macula on side of each notum and smaller, somewhat diffused, blackish brown macula on side of each tergum. Setae of head dark brown; setae of dorsal surfaces white except for dark brown setae on spiracular sclerites, dark brown setae on margins of eighth and ninth terga, and dark brown terminal setae; setae of ventral surfaces black; setae of legs light golden-brown; stout setae of legs brownish black. 1970] Beal: Taxonomic Study of Attagenini 217 Head : Antenna with terminal seta about one-fourth longer than length of terminal segment; second segment without setae. Epi- pharynx with usual proximal series of about 16 sensory pits and middle series of six sensory pits but without usual distal sensory pits. Maxillary palp with two setae inserted on third segment. Labium with two ensiform setae only on dorsal (inner) surface of each lobe of ligula; palp with or without two setae on first segment. Body setae : Long, fine setae of acrotergites smooth; setae of dorsal surfaces denticulate on margins and ribs; setae of ventral surfaces mostly smooth but some setae with extremely finely ser- rulate margins (visible only under high magnification). Acrotergites with setae of two types : ( 1 ) row of ensiform setae with denticulate margins and two or three ribs between margins and inserted close to antecostal suture, (2) number of simple setae nearly as long as half length of tergite and gradually becoming extremely fine (al- most beyond resolution except with oil-immersion lens) and inserted anterior to ensiform setae. Tergites with linear to lanceolate setae with one to five ribs between margins but most commonly with four ribs between margins; lateral areas of nota and anterior tergites also with few long simple setae. Sockets of submarginal erect setae with one lateral adjacent sensory pit except that submarginal seta nearest spiracle may have two adjacent sensory pits. Sterna with linear setae only; these with two to three ribs between margins except for slender setae on anterior area of sternum; anterior area of sterna without row of longer, stouter, linear setae; first abdominal sternum with somewhat longer setae at midline. Spiracle and associated structures : Spiracle close to margin of tergum but closed by tergum and adjacent to spiracular sclerite; anterior margin more or less straight. Spiracular sclerite somewhat rhomboidal, not enclosed by tergum. Tergum forming sclerotized margin along anterior edge of sclerite but area anterior to margin not sclerotized; sclerite appearing to project laterad and somewhat posteriad of tergum. Abdomen : Eighth abdominal tergum with antecostal suture. Ninth abdominal tergum about two-sevenths as long as tergite of eighth abdominal segment. No sclerites present in sternum of ninth abdominal segment. Legs : Femur of prothoracic leg about 9/16 as wide as long; ventral margin about one-half as long as dorsal margin. Tibia with five stout setae in subdorsal row on posterior face. Setae at base of pretarsus ensiform, subequal in length; seta on posterior side a third wider than seta on anterior side. 218 Entomologica Americana [Vol. 45, No. 3 Distribution: Ranging from Independence, Inyo County, Cal- ifornia, south throughout Mojave and Rorego deserts to northern Baja California, eastward into Clark County, Nevada, Washington County, Utah, all desert areas of Arizona south of the Mogollon Rim, Hidalgo County, New Mexico, and southeast along the Rio Grande River to Brewster County, Texas. The species probably extends well into Mexico, but I have seen no collections of it from Mexico outside Baja California. Diagnosis: Males of this species have a distinctive color pattern on the elytra. Contrasting with the dark brown hairs of the elytron are white hairs which form a basal patch or short band, a diagonal band extending from the humeral angle to the median suture at about the basal two-fifths, and a number of lines extending from the diagonal band nearly to the apex of the elytron. Some females, particularly those from Pima and Santa Cruz counties in Arizona, resemble the males, but females from other Arizona areas, California, Nevada, and Utah are covered dorsally with white pubescence alone. No other Nearctic Attagenini are likely to be confused with it, except perhaps A. lobatus , which is uniformly covered with light golden- brown pubescence. However, A. lobatus has an 11 -segmented an- tenna, while N. uteana has a 10-segmented antenna. Other distin- guishing features of A. lobatus are discussed under the diagnosis of that species. The color of the setae of the larvae distinguishes them at once from known larvae of all other Nearctic species. Those on the head are dark brown, on the dorsum predominantly grayish white, on the ventral surfaces black, and on the legs light golden-brown. The larvae are likewise unique in that the hairs of the acrotergites consist of a row of ensiform setae near the antecostal margin and of numerous, extremely long and fine, simple setae. The former have denticulate margins with two or three ribs between the margins. The simple setae are tapering, the ends becoming exceedingly fine, visible only under the high powers of a compound microscope. Discussion: Casey treated the light-colored females and the forms marked with light and dark hairs as separate species. In my previous revision of Novelsis (1954) I synonymized the forms on the basis of the geographic distribution of museum specimens. Sub- sequently I have been able to obtain live adults from near Peoria, Arizona. These bred readily in the laboratory, and from them I have been able to rear several generations of the species. Observa- tions on the breeding of the forms confirmed my supposition that 1970] Beal: Taxonomic Study of Attagenini 219 the forms are no more than sexually dimorphic variants of one species. Ecology: The habitats of the larvae have not yet been dis- covered. The larvae have been reared to maturity in the laboratory on dried insects, but an effort to rear the species on dried dog food, which is suitable for most dermestids, was a failure. Adults mate and oviposit in the laboratory without the necessity of food or water. Adults are ordinarily collected out-of-doors on flowers and have been taken on mesquite [Prosopis juliflora (Swartz) DC.], catclaw ( Acacia greggii A. Gray), ironwood ( Olneya tesota A. Gray), French tamarix ( Tamarix gallica L.), sandpaper-plant ( Petalonyx thurber A. Gray), heliotrope ( Heliotropum curassavicum L.), and willow. The species was collected once on a farm near Casa Grande, Arizona, in the course of inspections for the khapra beetle. How- ever, the situation in which it was found was not recorded. Novelsis liorni (Jayne) Attagenus horni Jayne, 1882, p. 356. — Sharp, 1902, p. 645. Attagenus byturodes Jayne, 1882, p. 356. Novelsis horni : Casey, 1900, p. 148. — Rees, 1943, p. 15. — Beal, 1954, pp. 85-86. Adults : Redescribed by Beal ( 1 954 ) . Mature Larvae: Integument of head and dorsal surfaces yellowish brown. Setae of head dark golden-brown, sometimes with black hairs around occiput; setae of dorsal and ventral surfaces of body black; setae of legs golden-brown. Head : Antenna with terminal seta about one and one-half times as long as length of terminal segment; second antennal seg- ment without setae. Epipharynx with proximal series of about 24 sensory pits and middle series of six sensory pits but without usual distal sensory pits. Maxillary palp with two setae inserted on third segment. Labium with three ensiform setae on dorsal (inner) surface of each lobe of ligula; medial seta wide with lateral setae progressively narrower; palp without seta on first segment. Body setae : Margins of setae on disc of pronotum smooth; margins of setae of mesonotum, metanotum, and terga minutely serrulate (serrulations visible only under high power of compound microscope); anterior setae on each tergum with coarser serrulations than posterior setae; setae of sterna smooth or extremely minutely serrulate. Acrotergites with small, linear setae with one rib between margins and larger, linear-lanceolate setae with one, two, or rarely 220 Entomologica Americana [Vol. 45, No. 3 three ribs between margins. Tergites with most setae spindle-shaped but some linear; widest setae usually with two, sometimes three ribs distinctly visible between margins. Sockets of submarginal erect setae with one sensory pit near lateral margin. Sterna with linear setae only; widest setae usually with no more than two ribs dis- tinctly visible between margins but three ribs occasionally visible. First abdominal sternum with setae along midline somewhat longer than other setae of disc. Setae of each sternum becoming smaller toward anterior margin without anterior row of longer, stouter setae. Spiracle and associated structures : Spiracle opening on margin of tergum, not closed by tergum, and adjacent to spiracular sclerite; anterior margin curved. Spiracular sclerite subrectangular, weakly sclerotized, bearing six or fewer setae, not enclosed by tergum. Tergum forming sclerotized margin along anterior edge of sclerite but area of tergum anterior to margin not sclerotized. Sclerite ap- pearing to project laterad and somewhat posteriad of tergum. Abdomen : Eighth abdominal tergum with antecostal suture. Ninth abdominal tergum about half as long as tergite of eighth abdominal segment. No sclerites present in sternum of ninth ab- dominal segment. Legs : Femur of prothoracic leg about half as wide as long; ventral margin about 5/12 as long as length of segment. Tibia with six or seven stout setae on posterior face below dorsal margin; setae arranged more or less in two rows. Setae inserted at base of pre- tarsus subequal in length, not extending beyond basal fourth of claw; posterior seta ensiform, about twice as wide as anterior seta. Distribution: The species is recorded from southern Cali- fornia, southern Arizona, New Mexico, southwestern Texas, Wills Point, Louisiana, Mexico City and Tabasco, Mexico. New records extending the periphery of the known distribution are the following. Arizona : Cottonwood, Yavapai County, as larva March 30, 1963 (R. S. Beal); Flagstaff, Coconino County, as larva January 11, 1963 (R. S. Beal). New Mexico : Tucumcari, Quay County, May 23, 1957 (R. M. Eads). Texas : Clear Fork, Brazos River, Jones County, September 29, 1957 (R. S. Beal). Mexico: Basuchil, 6-7,000 ft. elev. (Mrs. Y. Mexia). Diagnosis: Adults of this species and N. andersoni are readily distinguished from all others in the tribe by the cross-shaped, light yellowish brown marking on the blackish elytra. The longitudinal part of the cross is formed by a narrow line that extends along the suture from the base nearly to the apex. A somewhat broader transverse band intersects this line at the basal third. In N. horni 1970] Beal: Taxonomic Study of Attagenini 221 the transverse band is somewhat expanded laterally, often into a short, posteriorly directed spur near the lateral margin. In no case does the spur extend posteriorly beyond the middle of the elytron. In N. andersoni the transverse band is also expanded laterally, but there is always a light colored line extending from the humerus along the margin of the elytron to the apical third or beyond. Females of N. horni may also be distinguished from those of N. andersoni by the 11-segmented antenna. Females of N. andersoni have a 10-segmented antenna. Males of both species have an 11- segmented antenna. Larvae of N. horni and N. andersoni are apparently indistinguish- able from each other. They may be separated from larvae of other Attagenini by the following combination of characters. The color of the setae of the head is dark golden-brown, of the dorsal and ventral surfaces black, and of the legs golden-brown. The setae of the acrotergites are linear and linear-lanceolate, those of the tergites mostly spindle-shaped with two or three ribs visible between the margins. The spiracular sclerite, which is difficult to observe be- cause of the dense, black setae, appears to be separated from the tergite, but is actually inserted within a narrow ring of the tergite that extends peninsula-like from its lateroposterior angle. The two setae inserted on the pretarsus of each leg are unequal, one being twice as wide as the other. Ecology: The species occurs in a wide variety of habitats and is obviously a general scavenger on dried protein materials. It has been collected in Sceliphron mud dauber nests in Arizona (Beal, 1951) and in Texas. I found it in Flagstaff, Arizona, on a dead bat in an attic and in large numbers in Cottonwood, Arizona, in nests of the spider Physocyclus tanneri Chamberlin. It was reared from larvae found by John LeCave in an English sparrow nest in Tucson, Arizona. H. G. Hubbard recorded in his dairy (deposited at the United States National Museum) on April 15, 1897, that he took the species at Tucson “in bunch of curled leaves of cotton- wood with aphis.” The adults reproduce readily in the laboratory without the necessity of drinking or of flying. However, the adults probably regularly fly out-of-doors to flowers. The species has been taken at Nogales, Arizona, on blossoms of mesquite ( Prosopis juliflora). The species can be considered a negligible pest of stored prod- ucts. K. S. Rohwer collected it in a feed and seed store in Willcox, Arizona. R. M. Eads found it on cotton seed in a warehouse at Tucumcari, New Mexico. Occasional specimens are found in homes. 222 Entomologica Americana [Vol. 45, No. 3 I reared it through a number of generations on dried dog food and through at least one complete generation on oatmeal. Novelsis andersoni Beal Novelsis andersoni Beal, 1954, pp. 86-88. Adults and Larvae: Adults were described by Beal (1954). No characters have been found by which the larvae differ from larvae of N. horni. Recorded Distribution: The type locality is forty miles west of Mexicali, Baja California. It has also been recorded twenty miles west of Mexicali and twenty miles south of Palacio, Baja Cali- fornia, at Fort Yuma, Imperial County, California, and at Phoenix, Arizona. New Records: Arizona : Maricopa County: Fort McDowell, May 12, 1962 (R. S. Beal); Gillespie Dam, as larva November 13, 1958 (R. S. Beal); Komatke, April 9, 1960 (R. S. Beal); Maricopa, adults, pupa, and larva November 8, 1956 (R. S. Beal); Tempe, April 16, 1959 (E. O. Johnson); same locality, April 24, 1’962 (R. K. Weaver). Yuma County: Parker, January 22, 1958 (G. H. Spider); Roll, April 18, 1958 (N. Berry and E. F. Pittman); Yuma, April 13, 1955 (Butler and Tuttle); same data (Butler and Werner); same locality, April 6, 20, 1959 (D. Muse). California : Indio (Fall). Diagnosis: Adults of this species are strikingly similar to those of N. horni and were confused with them by earlier authors. Specimens of this species stand in both the LeConte and the Fall collections at the Museum of Comparative Zoology under the name horni. Nevertheless, the species are readily distinguished by dif- ferences in the pattern of elytral maculation. In N. horni there is a subbasal band at the basal third that is expanded laterally, sometimes forming a short lateral line along the margin of the elytron in the basal fourth. Occasionally there is a lateral spur extending a short distance posteriad. In no instance does the spur extend beyond the middle of the elytron. On the other hand, N. andersoni has a lateral line on the elytron that extends to the apical third or fourth. The line may be broad or very narrow and interrupted so far as the colors of the integument are involved, but it is always plainly indicated by the pattern of white hairs. The two species are also distinguished by the number of segments in the antennae of the females. N. horni females have 11 segments; N. andersoni 10 segments. 1970] Beal: Taxonomic Study of Attagenini 223 The diagnosis of the larval stages is given under the diagnosis of N. horni. Ecology: Larvae of this species have been found in a number of different situations, suggesting that it is a general scavenger on dried protein materials, like most other members of the tribe. It was taken at Maricopa, Arizona, in a Sceliphron mud dauber nest and at Gillespie Dam, Arizona, in a swallow nest. At Roll, Arizona, a specimen was found on a dead mouse. A specimen deposited at the United States National Museum reportedly came from a Tempe, Arizona, flour mill. The writer has reason to believe that the speci- men did not come from the mill itself but from a neighboring ware- house filled with grain sacks heavily infested with Ephestia and other insects, where it was likely feeding on their dead remains. Adults have been swept from mesquite ( Prosopis juliflora ), creosotebush [Larrea tridentata (DC.) Coville], and willow. Novelsis timia Beal Novelsis timia Beal, 1954, pp. 82, 88. Adult Females: Described by Beal (1945). Males were un- known at the time of the description of the species. The following is a description of a male from Highway 74 above Palm Desert, Riverside County, California. Adult Males: Integument of head dark brown; integument of pronotum light brown with lateral carina and narrow posterior border dark brown; elytra light brown; ventral surfaces brown with abdomen dark brown; legs and antenna yellowish tan. Pubescence of head and dorsum subrecumbent, piceous and golden-white; pubescence of ventral surfaces recumbent, white. Eye emarginate over base of antenna. Antenna 1 1 -segmented, in repose extending behind anterior margin of metasternal episternum; flagellar shaft (segments 3-8) little shorter than first segment of club (segment 9); ratios of lengths of first, second, and third segments of club 11:13:21; ratio of length of terminal segment to length of pronotum and elytra combined 1:7.3. Channel below eye for reception of flagellar shaft of antenna shallowly concave; anterior margin of channel forming low carina not visibly projecting from side of head when seen from front (similar to Fig. 2); carina not curved behind base of maxilla but projecting diagonally behind base of maxilla parallel to margin of eye (relation of carina to gular suture not apparent since suture not distinct). Pattern of pubescence of pronotum and elytra as described for female. Pronotum with lateral 224 Entomologica Americana [Vol. 45, No. 3 carina continued thread-like around anterolateral angle; basal lobe feebly rounding or truncate and not produced abruptly posteriad; disc with intermingled craterform punctures about diameter of facet of eye and simple punctures about half diameter of facet of eye; punctures separated by less than half diameter of larger punctures; punctures toward side becoming slightly larger but not quite con- tiguous; setae on basal lobe and lateroposterior angles about twice length of setae of disc. Elytron with hairs of disc about one and one-half times as long as third segment of antenna. Hypomeron slightly inflated. Posternum with posterior margin of lateral lobe scarcely reflected against procoxa; prosternal process slightly ex- panded at apex, not extending posteriad as far as apex of procoxa in repose; disc of prosternum without longitudinal carina, without distinct lateral carina before anterior margin, and without median anterior denticle. Epipleuron terminating gradually behind first abdominal sternum. Ventral plate of hind coxa not forming tooth lateral to insertion of trochanter; plate extending behind metepi- sternum but not quite attaining metepimeron. Protibia not carinate on dorsal margin. Mesofemur with crural cavity in nearly horizontal plane; anteroventral carina extending from apex to about basal third; posteroventral carina indistinct. Metatibia with stout spines and setae along shaft not longer than apical spines. Length (of pronotum and elytra combined) : 2.5 mm. Recorded Distribution: The species has been known only from its type locality, Potholes, Imperial County, California, and one other locality, Palm Springs, California. New Records: Arizona : Junction of the North and South Forks of the Santa Maria River, Yavapai County, as larva November 20, 1956 (R. S. Beal). California : Highway 74, 3,000 ft. elev., above Palm Desert, Riverside County, June 21, 1961 (G. H. Nel- son); Painted Canyon (near Indio), April 18, 1926 (Low); Palm Springs, May 20, 1916 (H. C. Fall Collection). Diagnosis: The three transverse bands of light pubescence on the elytra distinguish this from most of the other species considered here. N. perplexus, N. varicolor, and N. aequalis also have three or more bands of light colored pubescence on the elytra, but in these species the pubescence is long, giving the specimens a shaggy ap- pearance, and the pubescence is tricolorous. In N. timia the dorsal hairs are quite short and are bicolorous. N. picta might be con- fused with N. timia; the two species mutually resemble each other more than they do any other species. However, N. picta has a 10- 1970] Beal: Taxonomic Study of Attagenini 225 segmented antenna, N. timia an 11 -segmented antenna. N. picta also has a number of long hairs on the hind tibia that are two or three times as long as the apical spines of the tibia. The tibial hairs of N. timia are at most no more than subequal to the tibial spines. Ecology: Almost nothing is known of the ecology of the species. Adults have been taken on catclaw ( Acacia greggii ). One larva was found in a sheltered bird’s nest in a recess in a cliff on the Santa Maria River in central-western Arizona. The nest was not positively identified, but possibly was a phoebe nest. Unnamed Forms near N. timia : Three Arizona specimens that may represent a geographic variant of N. timia or possibly a new species near N. timia have been collected by the author. They closely resemble specimens of N. timia except that the integument is darker brown and the antennae of two of the specimens are 10- segmented. Data for the specimens are as follows: McDowell Mountains, Maricopa County, May 6, 1962, on flowers of Acacia (11-segmented antenna; female); same locality, May 19, 1961, at light (10-segmented antenna; male); Komatke, Maricopa County, August 25, 1966, at light (10-segmented antenna; male). Novelsis picta Casey Novelsis picta Casey, 1900, p. 184. — Beal, 1954, pp. 81—82, 89. Adult Males : Redescribed by Beal ( 1 954 ) . Adult Females: Previously undescribed; differing from males in the following observed characteristics: Antenna in repose extend- ing to middle of lateral margin of pronotum; ratio of length of first, second, and terminal segments of club 11:10:20. Length of pro- notum and elytra combined) : 3.6 mm. to 3.8 mm. Recorded Distribution: The type locality is given as River- side, Arizona. The locality was later named Kelvin (Barnes, 1960) and is located in Pinal County seven miles northwest of Kearny. In addition the species has been recorded from the following Arizona localities: west of Tucson, San Luis (Yuma County), and Hot Springs (Castle Hot Springs, Maricopa County?). It has also been recorded from four miles east of Yermo, San Bernardino County, California. New Records: Arizona : Globe (D. K. Duncan); Sabino Canyon, Santa Catalina Mountains, July 12, 1935 (E. D. Ball); same locality, June 29, 1955 (G. D. Butler and F. W. Werner); Santa Rita Range Reserve, Pima County, May 23, 1957 (G. D. Butler and F. W. Werner); Tucson, June 25, 1937 (O. Bryant). 226 Entomologica Americana [Vol. 45, No. 3 California : Cathedral City, Riverside County, June 20, 29, August 24, 28, 1950 (L. W. Isaak); 15 miles east of Calexico, Imperial County, June 5, 1961 (G. H. Nelson and H. F. Howden); Trona, San Bernardino County, June 13, 1961 (G. H. Nelson). Diagnosis: Adults of the species are easily distinguished from most other Nearctic Attagenini by the three bands of light pubes- cence on the elytron. There is a broad subbasal band, a narrow submedian band, and a narrow subapical band. Sometimes the subbasal band occupies the entire basal area of the elytron except for a patch of darker pubescence that forms a subbasal “eye.” The subapical band may or may not extend to the elytral apex or there may be a separate apical patch of light pubescence. N. aequalis, N. varicolor, and N. perplexus also have three or more bands of light-colored pubescence on the elytra. However, in these species the darker colored hairs of the elytra are blackish rather than a golden-brown, as they are in N. picta. Only N. timia is likely to be confused with N. picta , but N. timia has an 11 -segmented rather than 10-segmented antenna. N. picta is unique among Nearctic Attagenini in having long, erect or suberect setae along the posterior face of the shaft of each tibia in addition to the usual appressed hairs and short, stout, erect, spine-like setae. On the hind tibia these hairs are up to a fourth longer than the longest apical spines of the tibia. The species is also remarkable for the length of the segments of the antennal club of the male. The antenna in repose may extend beyond the hind margin of the metepimeron. Ecology: No information on the species is available other than that adults have been swept from mesquite ( Prosopis juliflora) and Condalia sp. (Rhamnaceae). They have also been taken at light. The larval stages remain unknown. SPECIES DESCRIBED BUT UNRECOGNIZED There is no way of knowing the identity of the following species, since the types have been lost and the original descriptions are not definitive : Attagenus cylindricornis Say, 1825, p. 185. Attagenus cylindricollis Melsheimer, 1853, p. 48. ACKNOWLEDGMENTS This paper could not have been completed without the kind help of a great many workers who either donated specimens for study or loaned specimens under their care. Among them should 1970] Beal: Taxonomic Study of Attagenini 227 be mentioned the following: W. F. Barr of the University of Idaho; L. J. Bayer of the University of Wisconsin; W. E. Burkholder, Market Quality Research Division, Agricultural Research Service, Madison, Wisconsin; J. F. Gates Clarke of the Department of Ento- mology of the United States National Museum; J. B. Dimond and E. A. Osgood, Jr., of the University of Maine; the late S. A. Graham of the University of Michigan; M. H. Hatch of the University of Washington; C. L. Hogue of the Los Angeles County Museum; P. D. Hurd of the California Insect Survey of the University of California; M. T. James of Washington State University; V. Kalik of Pardubice, Czechoslovakia; V. M. Kirk of the Entomology and Crops Research Division of the Agricultural Research Service, Brookings, South Dakota; G. F. Knowlton of Utah State University; H. B. Leech of the California Academy of Sciences; Arnold Mallis of the Gulf Research Development Company; T. E. Moore of the Museum of Zoology of the University of Michigan; W. L. Nutting of the University of Arizona; J. A. Payne of the Market Quality Research Division, Agricultural Research Service, Tifton, Georgia; L. L. Pechuman of Cornell University; Jerome G. Rozen, Jr., of the Department of Entomology, American Museum of Natural His- tory; M. W. Sanderson of the Illinois Natural History Survey; W. B. Showers, Jr., of the Plant Pest Control Division of the Agricultural Research Service, Phoenix, Arizona; the late G. J. Spencer of the University of British Columbia; C. A. Triplehorn of Ohio State University; H. V. Weems, Jr., of the Florida State Plant Board; F. G. Werner of the University of Arizona; and R. D. Zhantiev of Lomonosov University, Moscow. I also wish to thank V. D. Roth of the Southwestern Research Station of the American Museum of Natural History for determination of some spider hosts as well as for contributing specimens collected in spider nests. 1 am grateful to Maciej Mroczkowski of the Institute of Zoology of the Polish Academy of Science for the privilege of examining the manuscript of his catalogue of the Dermestidae of the world in advance of its publication. J. M. Kingsolver of the Insect Identification and Parasite Introduction Branch of the Entomology Research Division deserves particular thanks for many helpful comments and suggestions during the course of my study, for checking many bibliographic references inaccessible to me, and for a critical reading of the manuscript. The study was funded in part by Grant No. G 13280 and Grant No. GU 691 from the National Science Foundation. 228 Entomologica Americana [Vol. 45, No. 3 LITERATURE CITED (All references have been verified from the original by the author or by Dr. J. M. Kingsolver except those followed by an asterisk.) Armstrong, J. W. T. 1945. On Australian Dermestidae. Part IV. Proc. Linnean Soc. N.S.W., 70: 47-52. Arrow, G. J. 1915. Notes on the Coleopterous family Dermestidae, and descrip- tions of some new forms in the British Museum. Ann. and Mag. Natur. Hist. London [8] 15: 425-451. Barnes, W. C. 1960. Arizona place names (ed. rev. by Byrd H. Granger). Tucson, University of Arizona Press, xxi +519 pp. Beal, R. S., Ir. 1951. Habitats of species of Novelsis (Coleoptera: Dermestidae). Pan-Pac. Entomol., 27: 57. 1954. A revision of the species included in the genus Novelsis (Coleoptera: Dermestidae). Trans. Amer. Entomol. Soc., 80: 73-90. Boush, G. M., F. V. Dunkel, and W. Burkholder 1968. Progeny suppression of Attagenus megatoma and Trogoderma parabile by dietary factors. J. Econ. Entomol., 61: 644-646. Brahm, N. J. 1791. Insektenkalender fur Sammler und Oekonomen. Handbuch der okonomischen Insektengeschichte in Form eines Kalenders bearbeitet. Mainz. Erster Theil, part II, pp. 129-248. Brulle, A. 1832. Expedition scientifique de Moree. Section des sciences physi- ques. Tome III. Ire Partie. Zoologie. Deuxieme Section — Des animaux articules. Paris, 395 pp., 22 pi. Casey, T. L. 1900. Review of the American Corylophidae, Cryptophagidae, Tritomidae, and Dermestidae, with other studies. J. N. Y. Entomol. Soc., 8: 51-172. 1916. Some random studies among the Clavicornia. Mem. Coleop- tera, 7: 35-292. Chao, Yung-Chang 1954. Insects in grain elevators at Pullman and Albion, Washington. Pan-Pac. Entomol., 30: 260-262. Charles, V. K. 1941. A preliminary check-list of entomogenous fungi of North America. U. S. Dep. Agr. Insect Pest Survey Bull., 21: 770- 785.* 1970] Beal: Taxonomic Study of Attagenini 229 Chevrolat, A. 1863. Coleopteres de l’ile de Cuba. Ann. Soc. Entomol., Fr., [4] 3: 489-620. Crotch, G. R. 1873. Check list of the Coleoptera of America, North of Mexico. Salem, Mass., 136 pp. Dahl, G. 1823. Coleoptera und Lepidoptera. Wien, 105 pp. von Dalla Torre, K. W. 1911. Nosodendridae, Byrrhidae, Dermestidae. In Junk, Coleopter- orum Catalogus, pars 33, pp. 1-96. Degeer, C. 1774. Characteres des genres des insectes. Memoires pour servir a l’histoire des insectes, Stockholm, vol. 4, pp. 1-456, 19 pi. Dejean, P. F. 1837. Catalogue des Coleopteres de la collection de M. le Comte Dejean. Troisieme edition, revue, corrigee et augmentee. Paris, viii + 503 pp. Dunkel, F. V., and G. M. Boush 1968a. Studies on the internal anatomy of the black carpet beetle, Attagenus megatoma. Ann. Entomol. Soc. Amer., 61: 755— 765. 1968b. Biology of the gregarine Pyxinia frenzeli in the black carpet beetle, Attagenus megatoma. J. lnvertbr. Pathol., 11: 281-288. Eichler, W. 1939. Gregarinen als Feinde von Vorratsschadlingen. Mitt. Ges. Vorratsschutz, 15: 48-50. Erichson, W. F. 1846. Naturgeschichte der Insekten Deutschlands. Erste Abtheilung. Coleoptera, vol. 3, pp. 321-480. Fabricius, J. C. 1798. Supplementum entomologiae systematicae. Hafniae, ii + 572 pp. 1801. Systema eleutheratorum secundum ordines, genera, species; adjectis synonymis, locis, observationibus, descriptionibus. Kiliae, vol. 1, pp. xxiv + 1-506. Faldermann, F. 1835. Additamenta entomologica ad faunam Rossicam in itineribus Jussu Imperatoris Augustissimi annis 1827-1831 a Cl. Mene- tries et Szovitz susceptis collecta, in lucem edita. Coleoptera Persico-Armeniaca. Nouv. Mem. Soc. Imp. Nat. Moscou, 4: 1-310.* Fauvel, A. 1903. Faune analytique des coleopteres de la Nouvelle-Caledonie. Rev. d’Entomol., 22: 203-378. 230 Entomologica Americana [Vol. 45, No. 3 Foerster, H. 1938. Beobachtungen uber das Auftreten von Gregarinen in In- sekten. Z. Parasitenk., 10: 157-209. Ganglbauer, L. 1904. Die Kafer von Mitteleuropa, Wien, vol. 4, pp. 1-286. Griswold, G. H., and M. Greenwald 1941. Studies on the biology of four common carpet beetles. Part I. The black carpet beetle ( Attagenus piceus Oliv.), the varied carpet beetle ( Anthrenus verbasci L.), and the furniture carpet beetle ( Anthrenus vorax Waterh.). Mem. Cornell Agr. Exp. Sta., 240: 3-57, 70-75. von Harold, F. E. 1868. Berichtigungen und Zusatze zum Catalogus Coleopterorum synonymicus et systematicus. Coleopterologische Hefte, 4: 104. Hatch, M. H. 1962. The beetles of the Pacific Northwest. Part III: Pselaphidae and Diversicornia I. Univ. Wash. Publ. Biol., 16: 1-503. Herbst, J. F. W. 1792. Natursystem aller bekannten in- und auslandischen Insekten als eine Fortsetzung der von Buffonschen Naturgeschichte. In Jablonsky, Der Kafer, vol .4, pp. 1-197, 8 pi. Hicks, E. A. 1959. Check-list and bibliography on the occurrence of insects in birds’ nests. Ames, Iowa State College Press, 681 pp. Hinton, H. E. 1943. Notes on two species of Attagenus (Col., Dermestidae) re- cently introduced into Britain. Entomol. Mon. Mag., 79: 224- 227. 1945. A monograph of the beetles associated with stored products. London, British Museum, vol. 1, pp. 1-443. 1946. The “gin-traps” of some beetle pupae; a protective device which appears to be unknown. Trans. Roy. Entomol. Soc. London, 97: 473-496. Hope, F. W. 1840. Coleopterists’ manual, London, vol. 3, pp. 1-191, pi. I— III. Howe, R. W. 1952. A note on the egg output of Attagenus megatoma (F.) ( = piceus (Ol.)) (Col., Dermestidae). Entomol. Mon. Mag., 88: 40-41. 1962. The influence of diapause on the status as pests of insects found in houses and warehouses. Ann. Appl. Biol., 50: 611- 614. Illiger, J. K. W. 1798. In Kugelann, Verzeichniss der Kafer Preussens. Halle, xlii -f 510 pp. 1970] Beal: Taxonomic Study of Attagenini 23 Jacquelin du Val, C. 1858. Manuel entomologique. Genera des coleopteres d’Europe, vol. 2, pp. 1-285; Catalogue, pp. 1-124, 67 pi. Jayne, H. F. 1882. Revision of the Dermestidae of the United States. Proc. Amer. Phil. Soc., 20: 343-377, pi. 1-4. Johansen, C., and J. Eves 1966. Parasites and nest destroyers of the alfalfa leafcutting bee. Wash. State Univ. Agr. Exp. Sta., cir. 469, pp. 1-12. Kalik, V. 1948. Contribution to knowledge of Czechoslovak Dermestidae. Sbornik Pnrodovedeckeho klubu v Pardubicich , pp. 1—13, pi. 1-3. 1955. Dermestidae der aethiopischen Region aus den Sammlungen des Konigl. Museums von Belgisch Kongo in Tervuren (mit Bemerkungen liber verwandte Arten). Rev. Zool. Bot. Afr., 52: 294-322. Kempers, K. J. W. B. 1923. Abbildungen von Fliigelgeader der Coleopteren. Entomol. Mitt., 12: 71-115. Kirby, W. 1837. The Insects. In Richardson, Fauna Boreali- Americana. Lon- don, vol. 4, pp. 1-325, 8 pi. Korschefsky, R. 1944. Bestimmungstabelle der bekanntesten deutschen Dermestiden- larven (Coleoptera) . Arb. Physiol. Angew. Entomol. Berlin- Dahlem, 11: 140-152. Kugelann, J. G. 1792. Verzeichniss der in einigen gegenden Preussens bis jetzt entdeckten Kafer- Arten, nebst kurzen Nachrichten von den- selben. Fortsetzung. Neu. Mag. Entomol., 1: 477-512.* Lacordaire, J. T. 1854 (1855). Paussides-Heterocerides. Histoire naturelle des in- sectes. Genera des Coleopteres, ou expose methodique et critique de tous les genres proposes jusqu’ici dans cet ordre d’insectes. Paris, vol. 2, pp. 1-548. de Laporte, F. L. 1840. Histoire naturelle des insectes Coleopteres. Histoire naturelle des animaux articules. Paris, vol. 2, pp. 1-563. Latreille, P. A. 1802. Histoire naturelle, generate et particuliere des crustaces et des insectes. Paris, vol. 3, pp. i-xii + 13-468. 1807. Genera Crustaceorum et Insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisque plurimis explicata. Paris, vol. 2, pp. 1-280. 232 Entomologica Americana [Vol. 45, No. 3 LeConte, J. L. 1854. Synopsis of the Dermestidae of the United States. Proc. Acad. Natur. Sci. Philadelphia, 7: 106-113. 1859. Catalogue of the Coleoptera of Fort Tejon, California. Proc. Acad. Natur. Sci. Philadelphia, 11: 69-90, 281-292. Lepesme, P., and R. Paulian 1939. Etude biologique et morphologique d 'Entomotrogus mega- tomoides Rtt. (Col. Dermestidae). Bull. Soc. Zool. Fr., 64: 159-168. Linnaeus, C. 1758. Systema Naturae per regna tria naturae secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Edito decima, Reformata. Stockholm: vol. 1, pp. 1-823. Linsley, E. G. 1944. Natural sources, habitats, and reservoirs of insects associated with stored food products. Hilgardia, 16: 187-224. 1946. Some ecological factors influencing the control of carpet beetles and clothes moths. Pests and Their Control, July. Lutz, K. G. 1911. Die Kafer des Deutschen Reiches. In Reitter, Fauna Ger- manica, vol. 3, pp. 1-436, pi. 81-128. Mallis, A., B. T. Burton, and A. C. Miller 1962. The attraction of salts and other nutrients to the larvae of fabric insects. J. Econ. Entomol., 55: 351-355. Mallis, A., A. C. Miller, and R. C. Hill 1958. Feeding of four species of fabric pests on natural and synthetic textiles. J. Econ. Entomol., 51: 248-249. 1959. The attraction of stains to three species of fabric pests. J. Econ. Entomol., 52: 382-384. Melsheimer, F. E. 1853. Catalogue of the described Coleoptera of the United States. Smithson. Misc. Collect. Washington, D.C., xvi +174 pp. Moore, W., and M. B. Moore 1942. Two species of black carpet beetle. J. Econ. Entomol., 35: 288. von Motschulsky, T. V. 1858. Sur les collections Coleopterologiques de Linne et de Fabricius. Etudes Entomol., Part. 7: pp. 123-152. Mroczkowski, M. 1954. Skorniki-Dermestidae. Klucze do oznaczania owadow Polski Polski Zwiazek Entomol., [2] 19: 1-47. 1958. Dermestidae (Coleoptera) collected in Bulgaria by members of the staff of the Zoological Institute of the Polish Academy of Sciences in the years 1950 and 1956. Fragmenta Faunistica (Warszawa) 8: 2-9. 1970] Beal: Taxonomic Study of Attagenini 233 1962. Anthrenus minutus Er. and faunistic notes on other Sardinian Dermestidae (Coleoptera) . Studi Sassarensi, Sassari, [3] 9: 544-549. 1964. Systematic and synonymic notes upon certain species of Dermes- tidae (Coleoptera). Ann. ZooL, 22: 179-187. 1965. Ergebnisse der Albanien-Expedition 1961 des Deutschen Ento- mologischen Institutes. 36 Beitrag. Coleoptera: Dermestidae. Beitr. EntomoL, 15: 665—671. 1968. Distribution of the Dermestidae (Coleoptera) of the world with a catalogue of all known species. Ann. ZooL, 26: 15-191. Mutchler, A. J., and H. B. Weiss 1927. The dermestid beetles of New Jersey including the carpet beetles and those which feed on animal products. N. J. Dep. Agr., cir. 108, pp. 1—31. Mulsant, E., and C. Rey 1867 (1868). Tribu des Scuticolles. Ann. Soc. Linnean. Lyon, N.S., 15: 1-188, pi. 1-3. Olivier, A. G. 1790. Entomologie, ou histoire naturelle des insectes, avec leurs caracteres generiques et specifiques, leur description, leur synonymie et leur figure enluminee. Paris, Coleopteres, vol. 2, No. 9. Dermeste. 16 pp. Panzer, G. W. F. 1795. Entomologia Germanica exhibens Insecta per Germaniam indigena secundum Classes, Ordines, Genera, species, adjectis synonymis, locis, observationibus. I. Eleutherata cum tab. aeneis. 36 + 372 pp., + 12 pi.* Pic, M. 1910. Descriptions ou diagnoses et notes diverses. Suite. Echange, 26: 17-19. Redtenbacher, L. 1843. Bemerkungen liber die in Syrien von Theodor Kotschy gesam- melten Kiifer [and] Coleopterorum Syriae genera et species novae. In Russegger, J., Reisen in Europa, Asien und Afrika. Erster Band. Reise in Griechenland, Unteregypten, im nord- lichen Syrien und Siidostlichen Kleinasien, mit besonderer Riicksicht auf die naturwissenschaftlichen Verhaltnisse der betreffenden Lander, unternommen in dem Jahre 1836. Zweiter Theil. Stuttgart, pp. 973-990.* Rees, B. E. 1943. Classification of the Dermestidae (larder, hide, and carpet beetles) based on larval characters, with a key to the North American genera. U. S. Dep. Agr. Misc. Publ. no. 511. 18 pp. 234 Entomologica Americana [Vol. 45, No. 3 Reitter, E. 1880 (1881). Die aussereuropaischen Dermestiden meiner S^mm- lung. Mit 70 Diagnosen neuer Arten. Verh. Naturf. Ver. Brunn, 19: 27-60. 1887. Bestimmungs-Tabellen der europaischen Coleopteren, III, ent- haltend die Familien Scaphidiidae, Lathridiidae und Dermesti- dae. Zweite vermehrte und verbesserte Auflage. Modling, 75 pp. 1889. Insecta, a Cl. G. N. Potanin in China et in Mongolia novissime lecta. VIII. Clavicornia. Hydrophilidae. Bruchidae. Horae Soc. Entomol. Rossicae., 23: 555-559. Rosenhauer, W. G. 1856. Die Thiere Andalusiens nach dem Resultate eine Reise zusammengestellt, nebst den Beschreibungen von 249 neuen oder bis jetzt noch unbeschreibungen Gattungen und Arten. Erlangen, viii + 429 pp., 3 plates.* Roth, I. R. 1851. Diagnosen neuer Coleoptera aus Abyssinien. Arch. Naturges., 17: 115-113.* Say, T. 1825. Descriptions of new species of coleopterous insects inhabiting the United States. J. Acad. Natur. Sci. Philadelphia, 5: 160-204. SCHONHERR, C. J. 1808. Synonymia Insectorum, oder: Versuch einer synonymie aller bisher bekannten Insecten, nach Fabricii Systema Eleuthera- torium &c. geordnet, mit Berichtigungen und Anmerkungen, wie auch Beschreibungen neuer Arten und illuminirten Kup- fern. Erste Band. Eleutherata oder Kafer. Zweiter Theil. Spercheus — Cryptocephalus. Stockholm, ix + 423 pp., 4 pi. Sharp, D. 1885. On some new species and genera of Coleoptera, Fam. Dermesti- dae. Sci. Trans. Roy. Dublin Soc., N.S., 3: 147-150. SOLODOVNIKOVA, O. 1938. [Biology of the Dermestidae Attagenus byturoides Solo, and Trogoderma versicolor Creutz. and control of them.] Acta Univ. Asiae med., (8 zool.), fasc. 41, 20 pp. Abstract (1941) in Rev. Appl. Entomol., 29: 453-454. Solsky, S. M. 1876. A second tetrad of beetles (Coleoptera) [in Russian], in A. P. Fedchenko, Expedition to Turkestan [in Russian]. Izv. Imp. Obshch. Lyub. Estest. Antrop. Entogr., Imp. Mosk. Univ. 21(1): 223-398. Spencer, G. J. 1947 (1948). Notes on some Dermestidae of British Columbia (Coleoptera). Proc. Entomol. Soc. Brit. Columbia, 44: 6-9. 1970] Beal: Taxonomic Study of Attagenini 235 Stephens, J. F. 1830. Dermestidae. Illustrations of British entomology; Mandibulata. London, vol. 3, pp. 118-128. Thunberg, C. P. 1795. Dissertatio entomologica, sistens insecta suecica. Part 9. Up- saliae, pp. 105-113.* Triplehorn, C. A. 1965. Insects found in Ohio grain elevators and feed mills. J. Econ. Entomol., 58: 578-579. Wollaston, T. V. 1864. Catalogue of the coleopterous insects of the Canaries in the Collection of the British Museum. London, Trustees of the British Museum, xiii + 648 pp. Woodroffe, G. E. 1953. An ecological study of the insects and mites in the nests of certain birds in Britain. Bull. Entomol. Res., 44: 739-772, pi. xiv-xvi. Wradatsch, G. 1914. Die Kaferausbeute von 1913 und die angewendete Fangtechnik. Entomol. Blatt., 10: 150-154. Zhantiev, R. D. 1963a. New species of dermestids belonging to the genus Attagenus Latr. (Coleoptera, Dermestidae) from Central Asia [in Rus- sian]. Zool. Zh., 42: 454-457. 1963b. Dermestid beetles of the genus Attagenus Latr. (Coleoptera, Dermestidae) from the U.S.S.R. [in Russian]. Entomol. Obozrenie (Rev. d’Entom. de l’URSS), 42: 415-424. ZlNKERNAGEL, R. 1952. Remarks on Attagenus species. Trans. Ninth Int. Congr. Entomol., No. 1, pp. 844-848. ZlNKERNAGEL, R., AND E. MULLER 1952. Kreuzungsversuche mit Attagenus piceus Oliv. (Dermestidae). Verh. Schweiz Naturf. Ges., 132: 156. Entomologica Americana A REVISION OF THE GENUS LOX ANDRUS LECONTE (COLEOPTERA: CARABIDAE) IN NORTH AMERICA Robert T. Allen PUBLISHED BY THE NEW YORK ENTOMOLOGICAL SOCIETY INCORPORATING THE BROOKLYN ENTOMOLOGICAL SOCIETY Vol. 46, No. 1 August 1, 1972 The New York Entomological Society (incorporating the Brooklyn Entomological Society) publishes two journals: Entomologica Ameri- cana (irregularly, according to the availability of acceptable papers) and the Journal of the New York Entomological Society (quarterly). Monographs and longer papers (minimum length approximately 70 manuscript pages) are published in Entomologica Americana. Shorter papers may be submitted to the Journal. Manuscripts should be sent to the Editor, Entomologica Americana, Department of Entomology, The American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. Entomologica Americana was first published by the Brooklyn Entomolog- ical Society in April, 1885. Publication was suspended after six volumes, resumed in 1926, again suspended in 1964 with Volume 44, and resumed in 1969 with Volume 45. The subscription rate to Entomologica Ameri- cana is $9.00 per volume. Correspondence regarding subscriptions and other non-editorial matters should be addressed to Publication Business Manager, New York Entomological Society, Department of Entomology, The American Museum of Natural History. EDITOR J. A. L. Cooke Mailed August 1, 1972 Entomologica Americana is published for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Vol. 45, No. 3 was published on October 26, 1970. Entomologica Americana Vol. 46, 1972, pp. 1-184 A REVISION OF THE GENUS LOXANDRUS LECONTE (COLEOPTERA: CARABIDAE) IN NORTH AMERICA1 2 3 4 5’2 3 By Robert T. Allen4 5 Table of Contents Introduction 3 Historical Summary 4 Biology and Distribution 5 Methods and Materials 5 Measurements 6 Abbreviations 7 Types 8 Morphology 1 1 The Genus Loxandrus 15 Key to the Species of Loxandrus Found in North America 18 Loxandrus infimus Bates 30 Loxandrus pactinullus new species 34 Loxandrus sculp til is Bates 38 Loxandrus rectangulus LeConte 40 Loxandrus elongatus new species 42 Loxandrus celeris (Dejean) 44 Loxandrus cubanus Tschitscherine 45 Loxandrus balli new species 48 Loxandrus elnae new species 52 Loxandrus whiteheadi new species 54 Loxandrus tropicus new species 56 Loxandrus lepidus new species 58 Loxandrus fraus new species 60 Loxandrus unispinus new species 62 Loxandrus yeariani new species 64 Loxandrus rectus (Say) 68 Loxandrus pravitubus new species 72 1 Modified from a thesis submitted to the Graduate School of the University of Illinois in partial fulfillment of the requirements for the degree of Doctor of Philosophy. 2 Partially supported by grants from the Bach Fund (National Academy of Sciences) and a Sigma Xi Grant-in-Aid. 3 Published with the approval of the Director, Arkansas Agricultural Ex- periment Station. 4 The study was begun while the author was employed by the Illinois State Natural History Survey and was completed as an Assistant Professor, Entomol- ogy Department, University of Arkansas. 5 Author’s address: Department of Entomology, University of Arkansas, Fayetteville, Arkansas 72701. [1] 2 Entomologica Americana [Vol. 46, No. 1 The L. unilobus, L. saccisecundaris, L. extendus and L. taeniatus complex 76 Loxandrus unilobus new species 76 Loxandrus saccisecundaris new species 78 Loxandrus extendus new species 82 Loxandrus taeniatus LeConte 84 Loxandrus florid anus LeConte 89 Loxandrus agilis (Dejean) 91 The L. aduncus, L. cervicalis, L. parallelus, L. taeniatus and L. spini- lunatus complex 94 Loxandrus aduncus new species 96 Loxandrus parallelus Casey 97 Loxandrus cervicalis Casey 98 Loxandrus spinilunatus new species 100 Loxandrus nocticolor Darlington 102 The L. rossi, L. algidus, L. parallelus , L. unilobus, L. velox and L. taeniatus complex 106 Loxandrus rossi new species 106 Loxandrus algidus new species 108 Loxandrus crenatus LeConte 110 The L. micans, L. vitiosus and L. duryi complex 112 Loxandrus micans Chaudoir 1, 112 Loxandrus vitiosus new species 115 Loxandrus duryi Wright 116 Loxandrus spinigrandis new species 120 Loxandrus discolor new species 122 Loxandrus saphyrinus (Chaudoir) 126 Loxandrus lucens Chaudoir 129 Loxandrus uniformis new species 130 Loxandrus minutus new species : 134 Loxandrus velocipes Casey 136 Loxandrus infer us new species 137 Loxandrus remotus new species 138 Loxandrus brevicollis (LeConte) 142 Loxandrus erraticus (Dejean) 145 Loxandrus gibbus new species 146 The L. nitidulus, L. robustus, L. cincinnatiensis, L. minor and L. cir- culus complex 148 Loxandrus nitidulus (LeConte) 150 Loxandrus robustus new species 153 Loxandrus cincinnatiensis Casey 154 Loxandrus minor (Chaudoir) 156 Loxandrus circulus new species 157 Loxandrus velox (Dejean) 158 Postulated Ancestral and Derived States of Characters in the Genus Loxandrus 159 The Phylogenetic Relationships Among the Species of Loxandrus 164 Zoogeography 174 North America 174 Loxandrus and the insect fauna of the Eastern United States 177 Acknowledgments 180 References 180 1972] Allen: Revision of Loxandrus LeConte 3 ABSTRACT: Fifty-two species in the genus Loxandrus are recog- nized as occurring in North America. Twenty-seven new species are described. A key to the species, descriptions, locality records, illus- trations of pertinent anatomical parts and a discussion of the phylo- genetic relationships and the zoogeography of the 52 species are presented. Nine additional species from Australia and three species from South America are treated in the phylogenetic and zoogeo- graphic sections but not in the taxonomic section. Based on the information now available it is postulated that the genus may have arisen in Middle or South America and then dispersed to North America, Australia and New Guinea. INTRODUCTION The genus Loxandrus , a taxon of the order Coleoptera, family Carabidae, has historically been considered as having its closest affinities with other genera comprising the tribe Pterostichini. Ball (1962), in the most recent comprehensive study of the North American beetle family Carabidae, placed the tribe Pterostichini in the subfamily Carabinae and in division B, the Caribida. Other workers (Darlington, 1962; Blackburn, 1890, 1892, 1907; Sloane, 1898, 1903; Straneo, 1949, 1951, 1953; Tschitscherine, 1898, 1900, 1901; Moore, 1965) who studied the species of Loxandrus occurring in South America, Australia and New Guinea also considered the genus to belong in the tribe Pterostichini. I concur with Ball’s hierarchical arrangement and placement of Loxandrus. The need for a generic revision of the genus Loxandrus was pointed out by Ball (1962). The complexity of the evolution and phylogenetic relationships in the tribe Pterostichini and its genera was discussed by Darlington (1962), Moore (1965) and Ball (1966). The lack of information in these two areas prompted my original investi- gation of Loxandrus. The following paper is a presentation of findings, thus far, concerning the taxonomy and phylogenetic relationships among species of the genus Loxandrus. LeConte’s original description characterized the genus Loxandrus as follows : Antennae slender, base not carinate; palpi cylindrical, slender, ultimate article not short, mentum dentate, ligula flat, apex truncate; labrum not emarginate; mandibles short; elytra with one dorsal puncture; scutellar stria not impressed; tarsi slender; outside of posterior tarsi sulcate, ante- rior tarsal segments one to three dilated, inside prolonged, outside oblique. 4 Entomologica Americana [Vol. 46, No. 1 All the characters mentioned in the original description are valid and are used in present day taxonomic works to characterize Lox- andrus. In addition to those characters mentioned by LeConte an ad- ditional character is necessary to distinguish the genus Loxandrus from the genus Oxycrepis Reiche. Members of the genus Oxycrepis have the sides of the pronotum strongly constricted towards the base. This constriction is not evident in any species of Loxandrus. The genus Loxandrus , as it is interpreted in this paper, cannot be set apart on any one single character but can be defined by utilizing the combi- nation of characters recorded by LeConte plus the additional character of the base of the pronotum not being constricted. HISTORICAL SUMMARY LeConte (1852), in his synoptic work on Pterostichus Bonelli and allied genera, “synthesized” and described Loxandrus. He in- cluded in Loxandrus three new species and eight species previously described by himself and other authors in other genera. The eight previously named species were described by LeConte (1848), Say (1823), Dejean (1828), and Chaudoir (1843) and were assigned to one of the following genera: Argutor Stephens, Feronia Latrielle, Meg- alostylus Chaudoir or Pogonus Nichol. Since 1852 several workers published on new species of Loxandrus from North America. Mots- chulsky (1865) described a new species from Mississippi and one listed from North America. Chaudoir (1868) described three new species, one each from Texas, Louisiana and South Carolina. LeConte (1878) also described three new species from Florida. Blatchley (1918) described two new species from Florida. Casey (1918) revised the genus Loxandrus but did not include all the species that had been described from North America. He also described nineteen new species at that time as well as two additional new species in 1924. Since 1924, only one worker, Wright (1939), has described a new species of Loxandrus from the United States and the group as a whole has not been treated since Casey’s work in 1918. Darlington (1938) recorded L. infimus Bates, previously known only from Mexico and Central America, from Brownsville, Texas. Darlington (1934, 1935) treated the genus as it occurrs in the West Indies. He described two new species, one from Cuba and one from Haiti and recorded the occurrence of several other previously- described species but did not provide a key to these forms. Bates (1871, 1881, 1891) described five species occurring in the Mexico-Central America area. He did not give a key to the species and only listed the general location from which the type material came. 1972] Allen: Revision of Loxandrus LeConte 5 Lindroth (1966) recorded two species from Southern Ontario, Canada. A key was presented and a type locality for a third species not occurring in Canada, L. erraticus (Dejean), was designated. BIOLOGY AND DISTRIBUTION Very little is known concerning the biology of Loxandrus species. Adults are found near water, under stones and logs along stream banks, and on flood plain areas. They can be obtained by treading vegetation in swamps and along stream banks. Some species have also been collected at black light traps. Life history studies have not been made for any of the species and no larval forms have been associated with the genus. Species of the genus occur as far north as Southern Canada in the Eastern United States and west to Illinois, Missouri, Arkansas and South-western Texas. They are also found in Mexico, Central America, the Antilles, Australia and in New Guinea. One species has been recorded from Celebes. METHODS AND MATERIALS The material for this study consisted of over 3,000 borrowed specimens of Loxandrus representing 61 species. Several hundred specimens belonging to related groups were also examined. These con- sisted of 58 species representing 29 genera and subgenera in three tribes belonging to the family Carabidae. These related species, gen- era, subgenera and tribes were studied in order to establish the an- cestral and derived states of characters in the genus Loxandrus. These specimens were preserved on insect pins or points. The external morphology and genitalia of all these species were examined. The external anatomical parts are illustrated in Figure 1 A, B. The genitalia of the males of all species, when available, were examined. The genitalia of the females were also examined but few stable taxonomic characters were found. The following method was used to remove the genitalia and pre- pare them for study. Specimens were placed in household ammonia for about 20 minutes. They were then removed and placed in boiling water for 2-3 minutes. As each specimen was removed from the boiling water, it was placed on a paper towel, which absorbed the excess water, and then dissected under a stereo-microscope. Using a pair of small scissors, an incision was made on the dorsal surface of the abdomen from the midpoint of the apex along the lateral margin for a short distance. The genitalia were removed by reaching under the “flap” with a pair of forceps and removing them. Internal membranes and muscles still attached to the genitalia were cut with the scissors. 6 Entomologica Americana [Vol. 46, No. 1 The basal ring of the males was left in the abdomen and not removed. The genitalia were then placed in individual small tubes containing 10-20 percent potassium hydroxide in a water bath and heated to just below boiling for 10-15 minutes. The genitalia were removed and placed under the microscope in 70 percent alcohol and agitated to force the potassium hydroxide and dissolved tissues from the median lobe of the males and from around the coxites of the females. The female genitalia were then placed in glycerine. After being subjected to hot potassium hydroxide, the walls of the median lobe of the aedeagus become somewhat transparent, and the scale groups on the internal sac were visible. In most instances, it was desirable to evert the sac to gain a better idea of the arrangement of the scale groups. This was accomplished by either of two methods. The first method employed the use of a minute insect pin that had been bent to form a hook at its tip. The base of the pin was inserted in a match stem that served as a handle. The hooked pin was inserted into the apex of the median lobe and the internal sac gradually pulled out. The second method, the one used most often by this writer, utilized a hypodermic syringe, a small hypodermic needle and 70 percent alcohol. The syringe was filled with 70 percent alcohol and the needle tip was placed in or against the basal foramen of the median lobe. The median lobe was held firmly but not tightly between a pair of forceps. The alcohol was injected into the basal foramen and the pressure thus created forced the internal sac out. Using this technique, the internal sac of about 98 percent of all specimens examined were everted. After washing in alcohol, whether the internal sacs were everted or not, the genitalia were placed in glycerine for 30 minutes before transferring them to genitalia vials containing glycerine. The vials were then placed on the insect pins under the specimens. Measurements. Measurements were made of the length and width of the head, thorax, and elytra. Abbreviations used for the measure- ments are as follows: TL — total length; WE — width across eyes, outside margins; WP — width of pronotum at widest point; WW — width across elytra at widest point. The total length was derived by combining the lengths of the head, thorax, and abdomen which were measured individually as follows: head — from the anterior margin of the clypeus along the midline to an imaginary line connecting the poste- rior margins of the compound eyes; thorax— from the anterior to the posterior margin along the midline; elytra — from the apex of the scutellum to the apex of the right elytron along the midline. All measurements are in millimeters. 1972] Allen: Revision of Loxandrus LeConte 7 Abbreviations. An asterisk (*) has been used in the locality sections of the descrip- tions to note localities that could not be located and mapped. Museums and Private Collectors AM BM CAS CINM CNC CNHM CU FSPB GB GN INHS MCZ MNHP MSU NCSC osu PSU PU RTB TAM TLE UA UK UM UMIN UMO USNM — American Museum of Natural History, New York — British Museum (Natural History), London — California Academy of Sciences, San Francisco — Cincinnati Museum of Natural History, Cincinnati — Canadian National Collection, Ottawa — Chicago Natural History Museum, Chicago — Cornell University, Ithaca — Florida State Collection of Arthropods, Gainesville — George Ball Collection, Edmonton — Gerald Noonan Collection, Washington, D. C. — Illinois Natural History Survey, Urbana — Museum of Comparative Zoology, Cambridge — Museum of Natural History, Paris — Michigan State University, East Lansing — North Carolina State University, Raleigh — Ohio State University, Columbus — Pennsylvania State University, University Park — Purdue University, Lafayette — Ross T. Bell Collection, Burlington — Texas A & M University, College Station — Terry L. Erwin Collection, Cambridge — University of Arkansas, Fayetteville — University of Kansas, Lawrence — University of Michigan, Ann Arbor — University of Minnesota, St. Paul — - University^ Missouri, Columbia — United States National Museum, Washington Anatomical Regions AN — Antenna AS — Ambulatory setae ATS — Anterior sulcus BI — Basal impression C — Clypeus CE — Compound eye CX — Coxae F — Femur FR — Frons G — Gula GE — Gena H — Humerus 11-19 — Intervals IS — Internal sac LM — Lobes of mentum LP — Labial palpi LPA — Left paramere MC — Mesocoxae FS — Frontal sulci ML — Median lobe MN — Mentum MS — Mesosternum MT — Metasternum MTM — Metepimeron 8 Entomologica Americana [Vol. 46, No. 1 MTP — Metepisternum S — Sclerite MXP — Maxillary palpi SA — Scale area PA — Posterior angle SC — Scutellum PC — Procoxae SG — Scale group PE — Epipleura of pronotum SR — Sclerotized ring PM — Proepimeron STI — Sternites of abdomen PP — Proepisternum T — Tibia PR — Intercoxal process TC — Tarsal claw MES — Mesepimeron TO — Tooth of mentum PS — Prosternum TR — Trochanter RPA — Right paramere TS — Tarsal segments TYPES Besides trying to follow the Rules of Zoological Nomenclature, two additional objectives influenced my work with types. First, in most instances species of Loxandrus cannot be accurately identified unless the internal sac of the male aedeagus is examined. Thus most species that are based on female types cannot be, at this time, adequately diagnosed. Secondly, I do not wish to multiply the number of names available for Loxandrus if it can be avoided. Therefore in this paper species are utilized that are based on female types but have a male that was in the original type material. I have tried to determine from the original description and from any other sources (mainly personal communication with other specialists) precisely what material was in the original series. The types of three species, Loxandrus parvulus Chaudoir, L. inquietus Casey, and L. duryi Wright, could not be located. However, a male paratype of L. duryi was studied and this species is considered in more detail below. The types of the two species described by Mots- chulsky, L. commutables and L. iris, were not available for study. The types of the latter two species are presumably in the Moscow Museum in Moscow, USSR. The following tables contain information accumulated concerning Loxandrus types for North America. TABLE I. Species in the genus Loxandrus based on male types or with a male in the original material. Type localities given in text. L. agilis Dejean 1828:244 $ (MNHP) L. brevicollis LeConte 1848:338 2 3 specimens in the series; the third specimen in the series is a S (MCZ) 1972] Allen: Revision of Loxandrus LeConte 9 TABLE I ( Continued ) L. brunneus Blatchley 1918:417 L. calathinus LeConte 1878:376 L. celeris Dejean 1828:246 L. cervicalis Casey 1918:391 L. cincinnatiensis Casey 1924:80 L. crenatus LeConte 1852:252 L. crenulatus Cbaudoir 1868:343 L. cubanus Tschit- scherine 1903:60 L. duryi Wright 1939:257 paratype L. erraticus Dejean 1828:24 L. flavilimbus Blatchley 1918:418 L. floridanus LeConte 1878:376 L. infimus Bates 1882:87 L. laticollis Chaudoir 1868:342 L. lucens Chaudoir 1868:342 L. lucidulus Dejean 1831:239 L. mandibularis Casey 1918:382 L. micans Chaudoir 1868:342 L. minor Chaudoir 1843:766 L. mutans Darlington 1935:180 L. nitidulus LeConte 1848:239 L. nocticolor Darlington 1934:91 L. parallelus Casey 1918:396 L. rapidus Chaudoir 1868:344 L. rectangulus LeConte 1878:377 L. rectus Say 1825:58 L. reflexus LeConte 1878:376 L. saphyrinus Chaudoir 1843:766 L. scitus Casey 1918:388 L. sculptilis Bates 1884: (278) L. taeniatus LeConte 1852: (252) L. tetrastigma Bates 1882:87 L. unistigma Bates 1882:87 L. velocipes Casey 1918:390 L. velox Dejean 1828:245 8 (PU) $ Type no. 5693, one additional specimen in the series, a 8 (MCZ) 8 (MNHP) 8 (USNM) 8 (USNM) 8 (MCZ) 8 (MNHP) 8 (MNHP) 8 (Unlocated) 8 (CINM) 8 (MNHP) 8 (PU) $ 7 specimens in the series, second specimen is a $ , two additional 8 8 in the series, all the same species (MCZ) $ lectotype here designated (BM) 8 (MNHP) 8 (MNHP) 8 (MNHP) 8 (USNM) 8 (MNHP) 8 (MNHP) 8 Type No. 22022 (MCZ) 9 4 specimens in the series, the second specimen is a 8 (MCZ) 8 (MCZ) 8 (USNM) 8 (MNHP) 8 (MCZ) 8 neotype designated by Lindroth & Freytag (1969) (MCZ) $ Type No. 5692, three additional specimens in the series, second specimen is a 8 (MCZ) 8 (MNHP) 8 (USNM) 8 lectotype here designated (BM) 8 (MCZ) 8 (MNHP) 8 lectotype here designated (BM) 8 (USNM) 8 (MNHP) 10 Entomologica Americana [Vol. 46, No. 1 TABLE II. Species in the genus Loxandrus based on female types without a male in the original material. Type localities given. L. accelerans Casey 1918:386 L. breviusculus Casey 1924:80 L. collucens Casey 1918:382 L. comptus Casey 1918:387 L. contumax Casey 1918:388 L. cursitans Casey 1918:387 L. concinnus Casey 1918:341 L. julgens Casey 1918:388 L. lateralis Casey 1918:381 L. limatus Casey 1918:382 L. ludovicianus Casey 1918:383 L. mondus Casey 1918:385 L. parvicollis Casey 1918:389 L. piceolus Chaudoir 1868:343 L. piciventris LeConte 1848:237 L. proximus Chaudoir 1868:344 L. pusillus LeConte 1852:252 L. rubricatus Bates 1891:250 L. suturalis Casey 1918:384 L. vulneratus Casey 1918:390 2 Type No. 47338 Texas, Galves- ton. (USNM) 2 Type No. 47350 Florida, Dune- din. (USNM) 2 Type No. 47336 Texas, Houston (USNM) 2 Type No. 47349 Florida, Dune- din (USNM) 2 Type No. 47352 Florida, Indian River (USNM) 2 Type No. 47348 Florida, Dune- din (USNM) 2 Type No. 47353 Florida, Dune- din (USNM) 2 Type No. 47344 Mississippi, Vicksburg (USNM) 2 Type No. 47347 Florida, Sara- sota (USNM) 2 Type No. 47335 District of Co- lumbia. (USNM) 2 Type No. 47337 Louisiana, Cane River. (USNM) 2 Type No. 47341 Florida, La- Belle. (USNM) 2 Type No. 47343 Texas, Galves- ton. (USNM) 2 A single specimen from Texas, C. Pilate (Coll.?). (MNHP) 2 Type No. 5698 Georgia (MCZ) 2 Chaudoir states in the original description that he was studying a $ . The specimen I found in the MNHP was a 2 . The geni- talia had been destroyed or lost. The Chaudoir determination la- bel gives Texas as the locality. This locality agrees with the original description. 2 Type No. 5699 Georgia. (MCZ) 2 (BM) 2 Type No. 47339 Florida, Dune- din (USNM) 2 Type No. 47354 Indiana. (USNM) 1972] Allen: Revision of Loxandrus LeConte 11 TABLE III. Species in the genus Loxandrus for which a type could not be located or was unavailable for study. L. blatchelyi Csiki 1930 A name proposed for L. brun- neus Blatchley now a synonym of L. cervicalis Casey L. commutablis Mots- chulsky 1865:243 L. inquietus Casey 1918:389 2 Indiana. L. iris Mots- chulsky 1865:242 L. parvulus Chaudoir 1868:342 2 Carolina MORPHOLOGY The general external morphological characters are illustrated in Figures 1 A, B. The characters used in this paper are familiar to most systematists. Thus, most of them are self-explanatory and only a few warrant discussion in this section. In most species of Loxandrus, the lobes of the mentum are evi- dently longer than the mental tooth (Figs. 141, 142, 143). However, Fig. 1. Habitus, L. unispinus n. sp. 12 Entomologica Americana [Vol. 46, No. in a few species, the length of the tooth is equal to the length of the mental lobes (Figs. 138-140). This gives the mentum a more trans- verse appearance. The external surface of all sclerites has a characteristic type of microsculpture, or the microsculpture is entirely absent. Two basic types of microsculpture with several intermediate types are present. The two basic types are referred to as isodiametric (Figs. 11 A, B; 16 A, B, C; 20 A, B, C) and transverse (Figs. 12, 15). The trans- verse type can be extremely stretched into fine parallel lines (Fig. 15) or only slightly stretched. The shape of the pronotum is characteristic for many species groups as well as individual species. The lateral margins are either evenly rounded from the posterior to the anterior margins (Fig. 35 B) or slightly oblique (Fig. 47) or sinuate (Fig. 45) in the posterior *4 to Vs. Since these variations are sometimes difficult to convey in words, they have been illustrated (Figs. 25 to 70). Figs. 3-6. Habitus, dorsal. 3. L. infimus Bates. 4. L. parallelus Casey. 5. L. remotus n. sp. 6. L. gibbus n. sp. 1972J Allen: Revision of Loxandrus LeConte 13 L. REMOTUS 5 L. GIBBUS 14 Entomologica Americana [Vol. 46, No. 1 Figs. 7-10. Male genitalia, left lateral aspect. 7. Pterostichus ( Poe - cilus) chalcites Say. 8. Pterostichus ( Hypherpes ) sp. 9. L. celeris (Dej.). 10. L. circulus n. sp. Characters associated with the genitalia, especially the males, are complex and difficult to describe. In many species of Loxandrus , only the males can be recognized, based on characters associated with the male genitalia. For these reasons, numerous illustrations of the male genitalia are provided. The male genitalia of Loxandrus are composed of a median lobe with an internal sac and a pair of parameres at the base of the median lobe (Fig. 7). The internal sac can be everted as previously described. Associated with the genitalia is a sclerotized ring process : the modified 9th abdominal sternite which appears to possess no taxonomic or phylogenetic characters. The scale groups on the internal sac are constant in position and characteristic for each species of Loxandrus. The number of scales in each group varies from specimen to specimen but the variation is not significant in species recognition. With the stereo-microscope, several different types of scales can be identified. The most common type is moderately long with a pointed apex (Fig. 10 SGI). The size range of this type of scale is variable (Fig. SG2). They are usually found in groups but occasionally a small group may be reduced to one scale 1972] Allen: Revision of Loxandrus LeConte 15 (Fig. 109 D). A second common scale type is a very small weakly- sclerotized scale with the apex usually rounded but sometimes minutely pointed (Fig. 9 SG3 ). These do not usually form well-defined groups. They are more common in the basal portion of the sac but are not uncommon over much of the surface of the sac. A third type of scale that is relatively rare is hair-like (Fig. 10 SG4). They are very slender and pointed and are usually found in small groups (Figs. 102 A; 103 B). In one species, L. rectangulus LeConte, they occur over much of the surface of the sac (Fig. 74 A, B). In L. spinigrandis (Fig. 83 A, B) large, strong spines are present on the internal sac. These different types and their significance have not been studied in detail. The function of the scales is also unknown. They may or may not be a part of a lock and key mechanism essential for fertilization of the female. Sclerotized bars, plates or sclerites are found on the internal sac of only one species of Loxandrus , L. dubius Curt. Sclerites are com- mon on the internal sacs of other Pterostichine species however (Fig. 8 S). In many species of Loxandrus a sclerotized ring occurs on the in- ternal sac towards the apex. In other species the ring is weakly sclero- tized and in a third group it is totally absent. This may be the exit of the ejaculatory duct, the gonopore. In those species in which a well- developed ring is present the ejaculatory duct was traced to the open- ing. However, an opening was not found for the ejaculatory duct in those species without the ring. I have, therefore, referred to this only as a “sclerotized ring” in this paper. THE GENUS LOXANDRUS Type Species & Locality Feronia recta Say 1823:58. “A $ with genitalia dissected from South Carolina, designated by R. T. Allen as neotype and the state as type area” (Lindroth & Freytag 1969). Description. General characteristics that prevail among all species of Loxandrus found in North America. Head. Labrum: Anterior margin straight or slightly concave, both conditions apparent among specimens of the same species; 6 setae along the anterior margin, decreasing in length inwardly; anterior lateral angles rounded; microsculpture usually distinct, isodiametric; color always of a lighter hue than rest of head. Clypeus: anterior lateral margins rounded; 1 seta each side, removed from the anterior and lateral margins (distance variable). Mentum: tooth always well developed, apex rounded or straight; a pair of setae at base of tooth. 16 Entomologica Americana [Vol. 46, No. Submentum: a slightly elevated, well-rounded ridge; microsculpture elongate. Gular sutures: distinct; microsculpture elongate or absent. Postgenae: microsculpture elongate or absent. Eyes: large and dis- tinct. Antennal segments: I, one dorsal seta near the apex, otherwise glabrous; II— III, a ring of setae at apex, otherwise glabrous; IV-XI, somewhat compressed laterally, pubescence covering each segment except a narrow, elongate area on each lateral surface. Mandibles: scrobes always distinct, without setae, microsculpture isodiametric. Anterior tentorial pits: distinct, located in the frontal sulci, approxi- mate to the epistomal suture. Thorax. Pronotum: 1 seta in or approximate to each hind angle; 1 seta along each lateral margin (position variable); anterior lateral angles rounded; lateral margin narrowly reflexed its entire length. Prosternum: intercoxal process not margined. Mesosternum: de- clivate medially; never punctate. Metasternum: area between meso- coxae margined; median sulcus reaching from posterior margin of metasternum to posterior margin of mesocoxae; lateral sulci elongate but never reaching lateral margins. Metepisternum : lateral margin always much longer than anterior margin. N. B. Microsculpture and punctation are variable on all ventral thoracic sclerites except where specifically noted above. Legs. Segment 5 of all tarsi usually with 2 ventral rows of very minute setae toward the apex; protarsal segments 1-3 of male obliquely dilated with 2 rows of large scales on the ventral surface. Elytra. 1 seta at base of stria 2; 1 seta on interval 3 approxi- mate to stria 2 (longitudinal position variable); usually 15 setiferous punctures in stria 8 along the lateral margin; 2 setae near apex of stria 7; scutellar stria absent; lateral margin near apex usually with distinct internal plica (absent in some Central American species); lateral margin near apex distinctly sinuate. Abdominal sternites. Segments 3, 4 and 5 always with a pair of ambulatory setae; segment 2 with or without ambulatory setae; segment 6, males with 1 pair of setae along apical margin, females with 2 pair of setae along apical margin. Aedeagus. Median lobe: usually an elongate tube (modified in 4 species). Parameres: left paramere short, conchoid shaped; right paramere slightly longer than left. Internal sac: everts apically; surface covered with scales which may or may not form distinct groups; a sclerotized ring near the apex may or may not be present. Figs. 11-15. Elytra, dorsal aspect. 1 1 A. L. yeariani n. sp. 11 B. Same, close-up. 12. L. nitidulus (LeC.). 13. L. agilis (Dej.). 14. L. brevicollis (LeC.). 15. L. discolor n. sp. 1972] Allen: Revision of Loxandrus LeConte 17 1 1 A L. YEARIANI L. AGILIS 12 L. NITIDULUS 14 L. BREVICOLLIS 15 L. DISCOLOR 18 Entomologica Americana [Vol. 46, No. 1 KEY TO THE SPECIES OF LOXANDRUS FOUND IN NORTH AMERICA 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 0). (2). 0). (4) . (5) . (4). (7) . (8) . (9). Microstructure on elytra distinct, isodiametric (Figs. 1 1 A, B) 2 Microsculpture on elytra present or absent; if present, composed of fine elongate, parallel lines (Figs. 12; 13; 14; 15) 4 Mentum with distinct fovae (Fig. 139) 3 Mentum without fovae (Fig. 143) 1. L. infimus Bates Large specimens, total length 7. 8-8. 6 2. L. pactinullus New Species Smaller specimens, total length 5. 8-7.3 mm. .... 3. L. sculptilis Bates Microsculpture on head and pronotum distinct, mostly isodiametric (Figs. 16 A, B, C); microsculpture on elytra distinct or not; if so, composed of elongate stretched lines on the intervals; hind angles of pronotum acute (Figs. 31, 32); lateral areas of pronotum explinate - 5 Microsculpture on head present or absent; microsculpture on pro- notum present or absent, if present never as distinct as on head (Figs. 18 A, B); microsculpture on elytra, if present, obscure and elongate; if hind angles of pronotum acute, then lateral areas not explinate , 7 Males 6 Females keyed to this couplet belong to one of the following species L. rectangulus & L. elongatus Mid portion of IS* covered with elongate, slender, hair-like scales (Figs. 74 A, B) 4. L. rectangulus New Species IS without elongate, slender, hair-like scales (Fig. 72) 5. L. elongatus New Species Metepisternum punctate (Fig. 19 A) 53 Metepisternum impunctate (Fig. 20 A) 8 Elytra bicolored (at least one central, pale-colored spot near the apex of the elytra, or two spots basally and two spots apically on the elytra) 9 Elytra unicolorous 22 Only one centrally located pale area towards the apex of the elytra 10 Pale areas on each elytron located at the humeri and toward the apex; area may be oval or elongate 14 Males 1 1 Females 12 * IS Internal sac of male aedcagus. Figs. 16-18. 16. Microsculpture distinct, isodiametric, L. rectangulus LeC. 16 A. Prothorax, dorsal aspect. 16 B. Head, dorsal aspect. 16 C. Head and prothorax, dorsal aspect. 17. Microsculpture slightly stretched, weakly present, L. floridanus LeC. 18. Microsculpture ab- sent, L. brevicollis (LeC.). 18 A. Head, dorsal aspect. 18 B. Pro- thorax, dorsal aspect. 1972] Allen: Revision of Loxandrus LeConte 19 I6A L. RECTANGULUS 17 L. FLORIDANUS I8A L. BREVICOLLIS I8B 20 Entomologica Americana [Vol. 46, No. 1 11. (10). 12. (10). 13. (12). 14. (9). 15. (14). 16. (15). 17. (16). 18. (17). 19. (16). 20. (19). 21. (14). Internal sac of the aedeagus with an elongate group of scales on the ventral surface at the base of the sac (Figs. 9; 75); usually larger in size, total length 6. 1-7.2 mm. (in part) 6. L. celeris (Dejean) Internal sac without any distinct scale groups (Fig. 76); usually smaller in size, total length 5. 8-6. 4 mm. (in part) 7. L. cubanus Tschitscherine Females from the United States (in part) L. celeris (Dejean) Females from Mexico, Central America or the West Indies 13 Females from Costa Rica, or Mexican states of Jalisco, Oaxaca, Nayarit, Sinaloa (in part) L. cubanus Tschitscherine Females from Cuba or Mexican states of Campeche, Chiapas, Colima, Veracruz, San Luis Potosi (in part) __ L. celeris (Dejean) Males 15 Females 21 IS without a secondary sac 16 IS with a secondary sac on the dorsal surface in the basal portion (Figs. 90 A, B) 9. L. eliiae New Species IS with scales but without a distinct scale group 17 IS with at least one distinct scale group 19 IS very elongate, almost cylindrical 18 IS shorter, sac-like, not cylindrical in shape (Fig. 85) 12. L. lepidus New Species IS with a small bulbous projection at the base on the right dorso- lateral surface (Figs. 86 A, B); smaller (in part) 10. L. whiteheadi New Species IS without a bulbous projection at the base on the right dorso-lateral surface (Fig. 84); larger 11. L. tropicus New Species IS with only 1 distinct scale group 20 IS with 2 distinct scale groups (Fig. 89) 13. L. fraus New Species Scale group on the right lateral side of the sac, elongate, length % or more than % the entire length of the sac (Fig. 88) 8. L. balli New Species Scale group on the right lateral side of the sac, short, oblong, length % or less than 34 the entire length of the sac (Figs. 87 A, B, C) 14. L. unispinus New Species Area between basal fovae of pronotum and midline usually im- punctate or with only 1 puncture on each side, rarely with 2-3 punctures L. balli New Species Figs. 19-20. 19. Thorax, ventral aspect, L. saphyrinus (Chaud.) . 19 A. Metepisternum, microsculpture absent, punctate. 19 B. Mesepister- num, microsculpture absent, punctate. 19 C. Metasternum, micro- sculpture absent, punctate laterally. 20. Thorax, ventral aspect, L. agilis (Dej.). 20 A. Metepisternum, microsculpture isodiametric, impunctate. 20 B. Mesepisternum, microsculpture isodiametric, im- punctate. 20 C. Metepisternum, microsculpture isodiametric, impunc- tate. 1972] Allen: Revision of Loxandrus LeConte 21 I9A L. SAPHYRINUS 20A L. AGILIS 20 B 20 C 22 Entomologica Americana [Vol. 46, No. 22. (8). 23. (22). 24. (23). 25. (22). 26. (25). 27. (26). 28. (27). 29. (28). 30. (25). Area between basal fovae of pronotum and midline usually with at least 4 punctures Females keyed to this couplet belong to one of the following species: L. unispinus, L. elnae, L. fraus, L. lepidus, L. tropicus, L. whiteheadi (in part). Mesepisternum punctured anteriorly (Fig. 19 B); abdominal seg- ment 2 without ambulatory setae; posterior angles of pronotum broadly rounded and distinctly explinate (Figs. 43 A, B; 44 A, B ); large species 23 Mesepisternum not punctured anteriorly (Fig. 20 B); abdominal segment 2 with or without ambulatory setae, posterior angles or pronotum broadly rounded or not, not distinctly explinate; usu- ally smaller species 25 Specimens from United States 24 Specimens from Mexico (in part) 15. L. yeariani New Species Larger, total length not less than 10.2 mm.; $ & $ genitalia char- acteristic (in part) 16. L. rectus (Say) Smaller, total length not greater than 9.3 mm.; $ & 9 genitalia characteristic (in part) 17. L. pravitubus New Species Metasternum punctured toward lateral margin (Fig. 19 C) ____ 26 Metasternum not punctured toward lateral margin (Fig. 20 C) 30 Males 27 Females keyed to this couplet belong to one of the following species: L. unilobus, L. saccisecundaris, L. taeniatus, L. extendus. 1 or 2 scale groups present on the left lateral surface at the base of the IS approximate to the median lobe (Figs. 121 A; 131 B); IS without a small secondary sac near the apex on the right lateral surface - 28 Scale groups absent on the left lateral surface at the base of the IS (Fig. 123 A); a small secondary sac near the apex on the right lateral surface (Fig. 123 B) 18. L. unilobus New Species No secondary sacs protruding from the main IS on the left lateral surface 29 A distinct secondary sac protruding from the left side of the main IS (Fig. 121 A) 19. L. saccisecundarius New Species Apex of IS elongate and not or only sparsely covered with weak small scales (Figs. 131 A, B) 20. L. extendus New Species Apex of IS truncate, rounded, almost entirely covered with small weakly sclerotized scales (Figs. 127 A, B) 21. L. taeniatus LeConte Pronotum with hind angles obtusely pointed, not rounded and usu- ally weakly turned to the outside (Figs. 30 A, B; 68 A, B) ____ 31 Figs. 21-24. Hind coxae, inner surface. 21. L. agilis (Dej.), micro- sculpture isodiametric. 22. L. saphyrinus (Chaud.), microsculpture absent. Figs. 23-34. Prothorax, dorsal aspect. 23 A. L. fraus n. sp. 23 B. Same, close-up of hind angle. 24 A. L. unispinus n. sp. 24 B. Same, close-up of hind angle. 1972] Allen: Revision of Loxandrus LeConte 23 21 L. AGILIS 22 L. SAPHYRINUS 23 B 24 B 24 A L UN1SPINUS 24 Entomologica Americana [Vol. 46, No. 1 31. (30). 32. (31). 33. (31). 34. (33). 35. (30). 36. (35). 37. (36). 38. (37). 39. (38). 40. (39). 41. (40). 42. (41). Pronotum with hind angles rounded or if obtusely pointed then never turned to the outside 35 Males 32 Females 33 IS of the aedeagus with an elongate group of scales on the ventral surface at the base of the sac (Figs. 9; 75); usually larger in size, total length 6. 1-7.2 mm. (in part) L. celeris Dejean IS of the aedeagus without any distinct scale groups (Fig. 76) usu- ally smaller in size, total length 5. 8-6.4 mm. (in part) L. cubahus Tschitcherine Females from the United States (in part) L. celeris Dejean Females from Mexico, Central America or the West Indies 34 Females from Costa Rica or Mexican States of Jalisco, Oaxaca, Nayarit, Sinaloa (in part) L. cubanus Tschitscherine Females from Cuba or Mexican states of Campeche, Chiapas, Colima, Veracruz, San Luis Potosi (in part) _ L. celeris Dejean Pronotum with distinct microsculpture on the dorsal discal area 36 Pronotum with the microsculpture on the dorsal discal areas very fine and obscure or absent 43 Striae narrow, microsculpture in trough usually only 1 (sometimes 2) isodiametric units wide (Fig. 13) 37 Striae wide, microsculpture in trough at least 2 and usually 3-4 isodiametric units wide (Figs. 12; 14; 15) 22. L. floridanus LeConte Inner surface of hind coxae with microsculpture usually weak very fine elongate parallel lines (Fig. 22); pronotum never explinate 38 Inner surface of hind coxae with distinct isodiametric microsculp- ture (Fig. 21); basal and lateral margin of pronotum explinate (in part) 23. L. agilis (Dejean) Males 39 Females keyed to this couplet belong to one of the following species: L. aduncus, L. parallelus, L. cervicalis, L. taeniatus, L. spinilunatus. Apex of median lobe not turned apically 40 Apex of median lobe turned apically (Fig. 124 A) 24. L. aduncus New Species IS without a secondary sac protruding from the left lateral side 41 IS with a secondary sac protruding from the left lateral side (Fig. 120 A) (in part) 25. L. parallelus New Species IS with one or two distinct scale groups on the ventral surface 42 IS without distinct scale groups on the ventral surface, although some groups may partially touch the ventral surface (Figs. 127 A, B) L. taeniatus LeConte IS with two distinct scale groups on ventral surface basal group oriented obliquely, apical group oriented transversely (Figs. 130 A, B, C) 26. L. cerivicalis Casey IS with one distinct scale group on ventral surface (Figs. 132 A, B) 27. L. spinilunatus New Species 1972] Allen: Revision of Loxandrus LeConte 25 43. (35). 44. (43.) 45. (43). 46. (45). 47. (46). 48. (47). 49. (48). 50. (49). 51. (50). 52. (50). 53. (7). 54. (53). 55. (54). Specimens from Mexico or West Indies 44 Specimens from U. S. 45 Specimens from Mexico 15. L. yeariani New Species Specimens from Cuba 28. L. nocticolor Darlington Striae narrow, microsculpture in trough usually 1 (sometimes 2) isodiametric units wide (Fig. 13) 46 Striae wider, microsculpture in trough at least 2 and usually 3-4 or more isodiametric units wide (Figs. 12; 14; 15) (in part) „ L. rectus (Say) Microsculpture on inner surface of hind coxae if present, fine elongate parallel lines (Fig. 22); median lobe not compressed dorso-ventrally 47 Microsculpture on inner surface of hind coxae isodiametric; (Fig. 21); median lobe compressed dorso-ventrally, sinuate on the right lateral margin near the apex (Fig. 125) 23. L. agilis (Dejean) Males 48 Females keyed to this couplet belong to one of the following species: L. rossi, L. algidus, L. parallelus, L. unilobus, L. taeniatus, L. velox. Gonopore opening on the left lateral surface of the IS 49 Gonopore opening on the right lateral surface of the IS at or near the apex (Figs. 126 A, B) 29. L. rossi New Species Apex of median lobe not modified 50 Apex of median lobe indented (Figs. 134 A, B) 52. L. velox (Dejean) IS with a secondary sac everting on the left lateral surface (Figs. 120 A; 133 A) 51 IS without a secondary sac everting on the left lateral surface (Figs. 123 A; 127 B) __ 52 5 distinct scale groups present on the IS (Figs. 133 A, B, C) 30. L. algiclus New Species 7 distinct scale groups present on the IS (Figs. 120 A, B) (in part) L. parallelus New Species IS with a rather large distinct scale group covering most of the apex (Figs. 127 A, B) (in part) L. taeniatus LeConte IS without a distinct scale group at the apex although very small, weakly sclerotized scales may cover the sac’s surface (Figs. 123 A, B) (in part) L. unilobus New Species Mentum transverse and with deep, distinct fovae (Fig. 138) 54 Mentum not transverse, with or without fovae; if fovae are present, small and shallow (Fig. 141) 58 Dorsal color black (sometimes with a small pale area towards the apex of the elytra); posterior angles of pronotum narrowly rounded, never turned to the outside 55 Dorsal color brown; posterior angles obtusely pointed, usually turned to the outside (Fig. 40) 31. L. crenatus LeConte Males 56 Females keyed to this couplet belong to one of the following spe- cies: L. micans, L. vitiosus, L. duryi. 26 Entomologica Americana [Vol. 46, No. 1 56. (55). 57. (56). 58. (53). 59. (58). 60. (59). 61. (58). 62. (61). 63. (62). 64. (63). IS with a group of scales on the ventral surface in the basal V2 on the midline, more or less lying between two lateral groups of scales (Figs. 109 E; 110 C) 57 IS without a distinct scale group on the ventral surface (Figs. 108 A, B, C) 32. L. micans Chaudoir IS with a secondary sac everting near the apex on the left lateral surface, the apex of which is covered with a distinct scale group (Figs. 110 A, B, C) 33. L. vitiosus New Species IS without a secondary sac, although a small lobe may appear to be present: if so, the apex of the lobe is never covered with a dis- tinct scale group (Figs. 109 A, B, C, D, E, F) 34. L. duryi Wright Elytra bicolored (discolored areas on the humeri and sometimes near the apex) 59 Elytra unicolorous 61 Lateral margins of pronotum broadly explinate posteriorly, poste- rior angles narrowly rounded but not acute (Fig. 26); IS with 2 parallel rows of very large spines extending the entire length of the sac (Figs. 83 A, B) 35. L. spinigrandis New Species Lateral margin of pronotum not explinate posteriorly, posterior angles acute; IS with or without distinct scale groups but never with spines 60 Lateral and basal margins of pronotum coarsely and profusely punc- tate (Figs. 29 A, B); IS of males elongate, more or less an evenly rounded tube, entirely covered with small scales (Fig. 82 A) — _ 36. L. discolor New Species Lateral and basal margins of pronotum punctate but not as coarse or as numerous as in L. discolor (Figs. 27 A, B); IS an elongate tube convoluted towards the apex and with a small bulbous pro- jection on the right side at the midpoint, some surface areas lack- ing scales (Figs. 86 A, B) (in part) L. whiteheadi New Species Pronotum with the hind angles broadly rounded (Figs. 36 A, B; 37 A, B; 38 A, B; 43 A, B; 44 A, B) 62 Pronotum with the hind angles rounded or not but not broadly rounded 66 Metasternum coarsely and distinctly punctate over a large area toward the lateral margin 63 Metasternum punctate or not, if so then only a few shallow punc- tures, never coarse punctures (in part) __ L. pravitubus New Species Lateral margins of pronotum near basal angles distinctly explinate 37. L. saphyrinus (Chaudoir) Lateral margins of pronotum near basal angles not explinate 64 Smaller, total length not greater than 10.0 mm. 65 Larger, total length not less than 10.2 mm. (in part) L. rectus (Say) Figs. 25-28. Prothorax, dorsal aspect. 25. L. balli n. sp. 26. L. spini- grandis n. sp. 27 A. L. whiteheadi n. sp. 27 B. Same, close-up of hind angle. 28 A. L. elnae n. sp. 28 B. Same, close-up of hind angle. 1972] Allen: Revision of Loxandrus LeConte 27 25 L. BALLI 27A L. WHITEHEADI 26 L. SPINIGRANDIS 27 B 28B 28A L. ELNAE 28 Entomologica Americana [Vol. 46, No. 65. (64). 66. (61). 67. (66). 68. (67). 69. (68). 70. (67). 71. (70). 72. (71). Lateral margins of pronotum evenly rounded giving the pronotum a more rotund appearance; basal angles broadly rounded (Figs. 37 A. B); IS distinctly turned to the left side, without scale groups on the ventro-lateral margins at the base of the sac (Fig. 105) L. lucens (Chaudoir) Lateral margins of pronotum less rounded basally, pronotum less rotund, posterior angles more distinct than in L. lucens (Figs. 38 A, B); IS elongate, not turned to the left side with 1 elongate scale group on the ventro-lateral margin on each side at the base of the sac (Fig. 107) 39. L. uniformis New Species Pronotum with hind angles more or less acute; species with a total length not less than 8 mm. 67 Pronotum with hind angles rounded (Figs. 65 A, B); species with a total length not greater than 7 mm. __ 40. L. minutus New Species Pronotum with distinct microsculpture AND/OR the IS with a large distinct secondary sac everting on the right lateral surface near the apex, sclerotized ring on the secondary sac (Figs. 99 A, B) ____ 68 Pronotum without distinct microsculpture (sometimes vaguely present); IS without a large secondary sac on the right lateral surface near the apex 70 Males 69 Females keyed to this couplet belong to one of the following species: L. velocipes, L. injerus. Five distinct scale groups on the IS (Figs. 99 A, B ) 41. L. velocipes Casey Four distinct scale groups on the IS 42. L. inferus New Species Lateral margin of pronotum sinuate posteriorly (Figs. 45; 46; 48; 70 A, B); IS without parallel scale groups at the base on the dorsal or ventral surface or if with parallel scale groups then an elongate secondary sac everting on the left lateral surface (Fig. 101 A) 71 Lateral margin of pronotum not at all sinuate posteriorly, evenly rounded; IS with parallel scale groups at the base on the dorsal or ventral surface without an elongate secondary sac (Fig. 102 D) 74 Basal impressions of pronotum continuous with a secondary poste- rior marginal sulcus (Figs. 5; 70 A, B); specimens from Central America 43. L. remotus New Species Basal impressions of pronotum not continuous with a secondary posterior marginal sulcus; specimens from the Continental United States 72 IS with an elongate secondary sac everting on the left lateral surface (Fig. 101 A); width of pronotum 3.4 mm. or more 44. L. brevicollis (LeConte) -» Figs. 29-32. Prothorax, dorsal aspect. 29 A. L. discolor n. sp. 29 B. Same, close-up of hind angle. 30 A. L. celeris (Dej.). 30 B. Same, close-up of hind angle. 31. L. rectangulus LeC. 32. L. elongatus n. sp. 1972] Allen: Revision of Loxandrus LeConte 29 29A L. DISCOLOR 30A L. CELERIS 3! L. RECTANGULUS 29B 30B 32 L. ELONGATUS 30 Entomologica Americana [Vol. 46, No. 1 73. (72). 74. (70). 75. (74). 76. (75). 77. (76). 78. (75). IS without an elongate secondary sac everting on the left lateral sur- face; width of pronotum 3.3 mm. or less 73 Specimen usually larger; TL— 9.6, WH— 1.9, WP— 3.0, WW— 3.9; IS elongate without a secondary sac (Figs. 95 A, B) 45. L. er rations (Dejean) Specimens usually smaller, TL — 8.6, WH — 1.5, WP — 2.6, WW — 3.0; IS not elongate, with a bulbous secondary sac everting on the left lateral surface (Figs. 96 A, B; 97) . 46. L. gibbus New Species Males 75 Females keyed to this couplet belong to one of the following species: L. nitidulus, L. minor, L. cincinnatiensis, L. robustus, L. circulus. IS with two elongate groups of scales lying parallel to one another on the dorsal surface in the basal % 76 IS without two parallel groups of scales on the dorsal surface in the basal % (one elongate group may be present on the left side but never a matching group on the right side) 78 IS with 5 or 6 distinct scale groups 77 IS with 4 distinct scale groups (Figs. 118 A, B) 48. L. robustus New Species IS with 5 distinct scale groups (Fig. 116) L 49. L. cincinnatiensis Casey IS with 6 distinct scale groups (Fig. 117) 50. L. minor Chaudoir IS with 5 distinct scale groups excluding a group of very elongate, slender scales near the sclerotized ring .. 47. L. nitidulus LeConte IS with 6 or 7 distinct scale groups excluding a group of very elon- gate scales near the sclerotized ring which may or may not be present 51. L. circulus New Species 1. Loxantlrus infimus Bates Figs. 3; 33; 71 A, B; 144 l,oxandrus infimus Bates 1881:87. Lectotype (here selected). $. The data label lists the specimen from Cubilguitz, Vera Paz. Collected by Champion. One additional specimen is in the syntypic series (BM). This does not agree with Bates’ description in which he stated there was a male and a female. L. mutans Darlington 1935: 180. Holotype. $. Type No. 22022. Haiti, Etang Lachau. Description: Color. Dorsal: black, dull, not shining, apex of elytra in some specimens orange or light brown. Ventral: black to Figs. 33-36. Prothorax, dorsal aspect. 33. L. infimus Bates. 34. L. sculptilis Bates. 35 A. L. pactinullus n. sp., close-up of hind angle. 35 B. Same. 36 A. L. saphyrinus (Chaud.), close-up of hind angle. 36 B. Same. 1972] Allen: Revision of Loxandrus LeConte 3 33 L. INFIMUS 35A L. PACTINULLUS 34 L. SCULPTILIS 36A L. SAPHYRINUS 36B 32 Entomologica Americana [Vol. 46, No. 1 dark brown; last abdominal segment sometimes light brown. Ap- pendages: dark to light brown, occasionally black. Head. Clypeus: anterior margin straight. Epistomal suture: distinct, complete. Sub- genal suture: obscure or absent. Frontal sulci: obscure or absent. Microsculpture: clypeus, frons and vertex absent. Mentum: apex of lobes rounded; tooth distinct, slightly shorter than lobes; furrows shallow; fovae absent; microsculpture isodiametric. Thorax. Pro- notum: microsculpture elongate on disk, isodiametric along the lateral margins and posteriorly; lateral margins sinuate posteriorly; posterior angles subacute; anterior marginal sulcus complete; posterior marginal sulcus incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions short, broad, punctate, area between basal impressions punctate; area between basal impres- sions and lateral margins sparsely punctate posteriorly. Scutellum: apex pointed; microsculpture isodiametric. Prosternum: microsculp- ture elongate medially, isodiametric laterally; impunctate. Proepi- sternum and Proepimeron: microsculpture isodiametric; impunctate. Mesosternum: microsculpture elongate; impunctate. Mesepisternum : Microsculpture isodiametric; anterior quarter punctate. Metasternum: microsculpture elongate medially, isodiametric laterally; impunctate. Metepisternum and Metepimeron: microsculpture isodiametric; im- punctate. Legs. Hind coxae: microsculpture on inner surface isodia- metric. Tarsal segments: 1-3 of middle and hind tarsi grooved dorso- laterally. Elytra. Microsculpture on intervals distinct, isodiametric; all striae distinct, punctate, trough 1-2 isodiametric units wide; internal plica well developed, weakly developed, or absent; humeri rounded; microsculpture of epipleura isodiametric. Abdominal sternites. Micro- sculpture elongate medially, isodiametric laterally, impunctate; seg- ments 1-5 rugose laterally. Aedeagus. Median lobe: apical half to quarter enlarged. Internal sac: everts medially; a scale group devel- oped at apex of sac; most of the remainder of surface covered with small scales; sclerites absent; sclerotized ring absent. Measurements. (18 specimens). TL— 7.4 (6.3-8.0) ; WH— 1.5 ( 1. 3-1.7) ; WP— 2.2 (1. 8-2.3); WW— 2.9 (2.4-3.3). Distribution. Central and South America. Localities. Mexico. Colima: 16.5 mi. E. Colima, Rte. 110, 1900 ft. (GB); Colima, Rte. 110, 1500 ft. (GB); 8 mi. S. W. Colima (GB). Guer- rero: 30.8 mi. N. Zumgango del Rio, Rte. 95, 1900 ft. (GB). Michoacan: Figs. 37-40. Prothorax, dorsal aspect. 37 A. L. lucens Chaud. 37 B. Same, close-up of hind angle. 38 A. L. uniformis n. sp. 38 B. Same, close-up of hind angle. 39. L. micans Chaud. 40. L. crenatus LeC. 1972] Allen: Revision of Loxandrus LeConte 33 37A L LUCENS 38A L. UNIFORMIS 39 L. MICANS 37B 38 B 34 Entomologica Americana [Vol. 46, No. 1 15 mi. W. La Piedad (GB). Morelos: Tejalpa (GB). Quintana Roo: 15.3 mi. W. Chetumal, 100 ft. (GB). Sinaloa: Mazatlan (GB). Sonora: El Oasis, Rte. 15 (GB). Tabasco: San Juan Bautista (BM). Tamaulipas: 73.1 mi. N. Manuel, 200 ft. (GB). united states. Texas. Cameron; Brownsville (Darlington, 1935). west indies. Haiti. Etang Lachaux; swamps north of Dessalines; Damien (near Port-au-Prince) (MCZ). *Cubilguitz, Vera Paz (BM). Comments. This species resembles L. sculptilis and L. perplexus in external appearance. It can readily be separated from them by the absence of fovae on the mentum, by the sinuate lateral margins of the pronotum, and by the more acute posterior angles of the pronotum. I have seen one specimen of this species from Brazil. Two specimens I examined were collected in New Orleans, Louisiana (USNM) but were intercepted on banana debris from Mexico. Darlington (1938) recorded this species from Brownsville, Texas. Darlington (1935) described L. mutans, a synonym of L. infimus , from Haiti. There were 19 paratypes in the material he studied from three different localities. He states that “My specimens were taken under trash and in loose soil a little above water level beside standing water.” The remainder of the specimens I have examined were collected in Mexico. Csiki’s Catalogue also records this species from Guatemala. Collecting data shows that the species has been collected in the months of April, June, July, August, September, October, and November. It has oc- occasionally been taken at lights. 2. Loxandrus paetinullus New Species Figs. 35 A, B; 93 A, B, C; 154 Holotype. S. Mexico. Michoacan: 8.1 mi. E. Villamar, Rte. 15, 5500 ft., July 31, 1966. George E. Ball and D. R. Whitehead, Collectors (MCZ). Paratypes. 12$ $ 8 $ $ (GB); 1 $ 1$ (UA); 1 $ (BM). Description. Color. Dorsal: black to dark brown. Ventral: dark to light brown. Appendages: usually light brown, sometimes dark. Head. Clypeus: anterior margin straight. Epistomal suture: distinct, complete. Subgenal suture: present or absent. Frontal sulci: obscure or absent. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes narrowly rounded; tooth dis- tinct, as long as lobes; furrows present, shallow; fovae deep; micro- sculpture isodiametric. Thorax. Pronotum: microsculpture elongate Figs. 41-44. Prothorax, dorsal aspect. 41. L. vitiosus n. sp. 42. L. duryi Wright. 43 A. L. rectus (Say). 43 B. Same, close-up of hind angle. 44 A. L. pravitubus n. sp. 44 B. Same, close-up of hind angle. 1972] Allen: Revision of Loxandrus LeConte 35 41 L. VITIOSUS L t lllllll , >. 42 L. DURYI 43A L. RECTUS 43B 44A L. PRAVITUBUS 44B 36 Entomologica Americana [Vol. 46, No. 1 on the disk, isodiametric along the lateral margins and posteriorly; lateral margins punctate, not sinuate posteriorly; posterior angles weakly pointed; anterior marginal sulcus complete; posterior marginal sulcus present, never complete; median sulcus may or may not reach the anterior and/or posterior sulci; basal impressions shallow, short, punctate; area between basal impressions punctate (more so than in L. sculptilis Bates) ; area between basal impressions and lateral margins punctate. Scutellum : apex narrowly rounded; microsculpture distinct, isodiametric. Prosternum: microsculpture elongate medially, iso- diametric laterally; impunctate. Proepisternum and Proepimeron: microsculpture isodiametric; impunctate. Mesosternum: microsculp- ture elongate; impunctate. Mesepisternum: microsculpture isodia- metric; impunctate. Metasternum: microsculpture elongate medially, isodiametric laterally; impunctate. Metepisternum: microsculpture isodiametric; impunctate. Metepimeron: microsculpture elongate; impunctate. Legs. Hind coxae: microsculpture on inner surface iso- diametric. Tarsal segments: 1-3, sometimes 4, of middle and hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals dis- tinct, isodiametric; all striae distinct, trough 1-2 isodiametric units wide; internal plica weakly developed or absent; humeri rounded; microsculpture of epipleura isodiametric. Abdominal sternites. Micro- sculpture elongate medially, isodiametric laterally; impunctate; seg- ments 1-4 rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts medially; scales present but no distinct scale groups; sclerites absent; the ring is a very elastic process but not sclerotized. Measurements. (19 specimens). TL — 8.0 (7. 8-8. 6); WH — 1.7 ( 1. 6-1.9 ); WP— 2.7 (2.6-2.9) ; WW— 3.3 (2.9-3.5). Distribution. Mexico. Localities. Mexico. Colima: 16.5 mi. E. Colima, Rte. 110, 1900 ft. (GB). Michoacan: 8.1 mi. E. Villamar, Rte. 15, 5500 ft. (GB). Morelos: Cuatla (GB); Cuernavaca (GB); Tejalpa (GB). Sonora: El Oasis, Rte. 15 (GB). Discussion. This species is similar to L. infimus but can be readily separated from that species by the absence of fovae on the mentum. L. pactinullus agrees in most external characters with L. sculptilis but the latter species is considerably smaller. Bates confused this species with L. sculptilis and it is probably misidentified in most collections. -> Figs. 45-50. Prothorax, dorsal aspect. 45. L. erraticus (Dej.). 46. L. gibbus n. sp. 47. L. velocipes Casey. 48. L. brevicollis (LeC.). 49. L. nitidulus (LeC.). 50. L. circulus n. sp. 49 L. NITIDULUS 50 L. CIRCULUS 1972] Allen: Revision of Loxandrus LeConte 45 L. ERRATICUS 46 L. GIBBUS 47 L. VELOCIPES 48 L. BREVICOLLIS 38 Entomologica Americana [Vol. 46, No. 1 It also resembles some Agonini due to the absence on the elytra of the internal plica in some specimens. For further comments see the de- scription of L. sculptilis and L. pactinullus. It is known only from Mexico where it has been collected during the months of June, July, and August. 3. Loxandrus sculptilis Bates Figs. 34; 91 A, B; 153 Loxandrus sculptilis Bates, 1881:278. Lectotype (here designated). $ . The data label gives the following information: Tole, Panama, Champion (Collector) (BM). Four additional specimens are marked as syntypes in the BM collection. This does not agree with Bates’ description which listed only a male and female. Description. This species resembles L. pactinullus in all char- acters except the following: (1) furrows on the mentum are absent; (2) the posterior areas of the pronotum are not as densely punctate; ( 3 ) the posterior marginal sulcus of the pronotum is sometimes com- plete; (4) the species is smaller in almost all respects. A comparison of size ranges follows : TL WH WP WW L. sculptilis 5. 8-7.3 1.3-1. 5 1. 9-2.2 2.5-2.9 L. pactinullus 7.8-S.6 1.6-1. 9 2.6-2.9 2.9-3.5 The aedeagus is also smaller in L. sculptilis. Distribution. Central America. Localities. Mexico. Morelos: Santa Rosa, 3.2 mi. N. Zacatepec, 3100 ft. (GB). Sinaloa: Presidio River vs. Union Sin (USNM) . Tamau- lipas: 27.5 mi. W. Soto la Marina, 1600 ft. (GB); *1.4 mi. SE. Chamal, 500 ft. (GB); 73.1 mi. N. Manuel, 200 ft. (GB). Chiapas: 7.7 mi. N. Frontera Comalpa, 2600 ft. (GB). Jalisco: 23.2 mi. S. Autlan, 800 ft., Rte. 80 (GB). Colima: Colima, 1500 ft., Rte. 110, at light (GB). Oaxaca: Tehuantepec (RTB). panama. Riqui: Tole (BM). Discussion. This is one of several species of Loxandrus that closely resemble some species in the tribe Agonini. The presence of the internal plica is variable and the presence of distinct isodiametric microsculpture on the elytra is unlike most species of Loxandrus. It Figs. 51-56. Prothorax, dorsal aspect. 51. L. minor (Chaud.). 52. L. cincinnatiensis Casey. 53. L. robustus n. sp. 54. L. saccisecundaris n. sp. 55. L. parallelus Casey. 56. L. agilis (Dej.). 1972] Allen: Revision of Loxandrus LeConte 39 51 L. MINOR 53 L. ROBUSTUS 52 L. CINCINNATIENSIS 54 L. SACCISECUNDARIS 55 L. PARALLELUS 56 L AGILIS 40 Entomologica Americana [Vol. 46, No. shares most characters in common with L. pactinullus but can be dis- tinguished from this species by its smaller size. This species has been collected in March, April, June, July, August, September, and October. 4. Loxandrus rectangulus LeConte Figs. 31; 74 A, B; 146 Loxandrus rectangulus LeConte 1878:377. Holotype. $. Enterprise, Florida, May 19 (MCZ). One additional specimen in the series, a female. This agrees with LeConte’s original description. Description. Color. Dorsal and ventral: usually black, some- times dark brown,, shining but not iridescent. Appendages: dark to light brown. Head. Clypeus: anterior margin straight. Epistomal and subgenal sutures distinct, complete. Frontal sulci: moderately impressed; short, broad. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes narrowly rounded; tooth distinct, almost as long as lobes; furrows and fovae absent; mi- crosculpture isodiametric. Thorax. Pronotum: microsculpture distinct, isodiametric anteriorly, sometimes slightly elongate posteriorly and lat- erally; lateral margins punctate in the posterior half to quarter, sinuate posteriorly; posterior angles acute; anterior marginal sulcus deep, complete; posterior marginal sulcus reaching past the basal fovae but incomplete; median sulcus distinct, reaching anterior sulcus but not posterior margin; basal impressions moderately deep, elongate, punc- tate; area between impressions punctate; area between impressions and lateral margins sparsely punctate or not. Scutellum: apex narrowly rounded; microsculpture isodiametric. Ventral surface entirely im- punctate, microsculpture elongate, more or less present on all ventral sclerites. Legs. Hind coxae: Microsculpture on inner surface elon- gate. Tarsi: segments 1-4 of hind tarsi, 1-3 and sometimes 4 of middle tarsi grooved dorso-laterally. Elytra. Microsculpture present or absent, if present composed of very elongate parallel lines; all striae distinct, punctate, trough 2-4 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura isodia- metric along the margin, very elongate or absent elsewhere. Abdom- inal sternites. Microsculpture isodiametric laterally, slightly elongate medially; segment 1 sometimes punctate at the base, laterally, other segments impunctate; segments 1-4 sometimes weakly rugose laterally. Figs. 57-62. Prothorax, dorsal aspect. 57. L. aduncus n. sp. 58. L. floridanus LeC. 59. L. unilobus n. sp. 60. L. taeniatus LeC. 61. L. rossi n. sp. 62. L. extendus n. sp. 1972] Allen: Revision of Loxandrus LeConte 41 57 L. ADUNCUS 58 L. FLORIDANUS 60 L. TAENIATUS 59 L. UNILOBUS 61 L. ROSSI 62 L. EXTENDUS 42 Entomologica Americana [Vol. 46, No. 1 Aedeagus. Median lobe: not modified. Internal sac: everts in a median position, very elongate; one distinct scale group present; much of the mid-portion covered by very elongate scales; other areas covered with small, weakly sclerotized scales. Sclerotized ring absent. Mea- surements. (9 S specimens). TL — 7.3 (6. 7-7. 6); WH — 1.5 (1.3- 1.5); WP— 2.1 ( 1. 9-2.2 ); WW— 3.1 (2.8— 3.4). Distribution. United States (Southern Florida) and Southern Mexico. Localities. Mexico. Campeche: 14.9 mi. S. Champoton, Rte. 180, 20 ft. (GB). Chiapas: San Quintin, 91°20'-16°24/, 700 ft. (GB). Vera- cruz: .8 mi. W. Sontecomapan, sea level (GB). united states. Florida. Highlands Co.: Archbold Bio. Sta., Lake Placid (CU). Manatee Co.: Oneco (GB). Volusia Co. : Enterprise (MCZ). *Texas. An unrecorded locality (MCZ, LeConte Coll.), west indies. Grand Cayman Island. South Sound (MCZ). Discussion. This species and L. elongatus appear to be alike in all external characters. They can be separated only on characters associated with the male genitalia. L. rectangulus has been collected at two localities in Florida while L. elongatus is known only from Mexico. Collecting data indicate that L. rectangulus has been collected in swamp and marsh habitats. It has been found in February, April and September. 5. Loxandrus elongatus New Species Figs. 32; 72; 145 Holotype. $. Mexico. Veracruz: 3.6 mi. N. Catemaco, 1000 ft., June 5, 1966. George E. Ball and D. R. Whitehead, Collectors (MCZ). Paratypes. 12 S $ (GB) 1 $ (UA). Description. This species resembles L. rectangulus in most external characters. The only distinguishable external difference appears to be in the microsculpture on the intervals of the elytra. In this species the microsculpture is usually evident as fine elongate parallel lines whereas in L. rectangulus the microsculpture is always absent. Other distinguishing differences are found in the male geni- talia. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position and is directed to the left side; a small bulbous secondary sac on the basal quarter of the main sac protruding from Figs. 63-67. Prothorax, dorsal aspect. 63. L. cervicalis Casey. 64. L. spinilunatus n. sp. 65 A. L. minutus n. sp. 65 B. Same, close-up of hind angle. 66. L. algidus n. sp. 67. L. lepidus n. sp. 1972] Allen: Revision of Loxandrus LeConte 43 63 L. CERVICALIS 65A L. MINUTUS 66 L. ALGIDUS 64 L. SPINILUNATUS 65B 67 L. LEPIDUS 44 Entomologica Americana [Vol. 46, No. 1 the right lateral surface, a small scale group at its apex; no distinct scale groups present on the main sac although the surface is covered with very small weakly sclerotized scales; sclerotized ring absent. Measurements. (11 $ specimens). TL — 7.6 (6.7-8.1); WH — 1.5 ( 1. 4-1.6 ); WP— 2.1 (1. 8-2.3); WW— 3.3 (3.0-3. 5). Distribution. Southern Mexico. Localities. Mexico. Chiapas: San Quintin, 91°20'-16°24' (GB); 700 ft. Veracruz: 3.7 mi. N. Catemaco 1000 ft. (GB); 2.5 mi. W. Sonte- comapan 100 ft. (GB); .8 mi. W. Sontecomapan (GB). Discussion. This species has been collected in February, June and September. It closely resembles L. rectangulus in external appear- ances. Females of the two species cannot be separated. 6. Loxandrus celeris (Dejean) Figs. 9; 30 A, B; 75; 147 Feronia celeris Dejean 1828:246. Lectotype (here selected) . $. The first specimen in a series of four specimens in the Paris Museum. The locality label lists it from Boreal America. Loxandrus rapidus Chaudoir 1868:344. Lectotype (here selected). $. The first specimen in a series of three specimens in the Paris Museum. The specimen bears no data labels on the pin but the Chaudoir determination label lists it from Louisiana. Description. This species is like L. cubanus Tschitscherine in all external characters except the following: (1) the microsculpture on the pronotum is not as distinct; (2) the microsculpture on the mesepi- sternum is isodiametric in the anterior half and elongate or absent in the posterior half. Aedeagus. Median lobe: not modified. Internal sac: everts medially; slightly directed to the right side; one elongate, well developed scale group on the ventral surface at the base of the sac; remainder of the sac’s surface covered with small weakly sclero- tized scales; sclerotized ring absent. Measurements. (20 specimens). TL— 6.8 (6. 1-7.2); WH— 1.4 (1.3-1. 5); WP— 2.1 (1.9-2.2); WW —2.8 (2.6-3. 1). Distribution. Southern United States and Southern Mexico. Localities. Mexico. Campeche: 14.9 mi. W. Champaton, Rte. 180, 20 ft. (GB); 31.8 mi. E. Escarcega, 200 ft. (GB). Chiapas: 6 mi. W. Cintalapa, Rte. 190, 1950 ft. (GB); San Quintin, 90°20,-16°24', 700 ft. (GB). Colima: 16.5 mi. E. Colima, Rte. 110, 1900 ft. (GB). San Luis Potosi: 1.8 mi. N. El Narajo, 1000 ft. (GB). Veracruz: E. Minatilan, Rte. 180, 60 ft. (GB); 2.5 mi. W. Sontecompan 100 ft. (GB). united states. Alabama. Mobile Co.: Mobile (MCZ). Winston Co.: Winston Co. National Forest (CU). Florida. Alachua Co.: Gainesville (FSPB). 1972] Allen: Revision of Loxandrus LeConte 45 Dade Co.: Homestead (FSPB). Duval Co.: Jacksonville (CAS). High- lands Co.: Archbold Biological Station, Lake Placid (PSU, CU). Hills- borough Co.: Tampa (MSU). Jefferson Co.: Monticello (DW). Lucie Co.: Capron (Ft. Capron on an 1861 map) (USNM). Manatee Co.: Oneco (GB). Okeechobee Co.: Shore Lake Okeechobee, 6 mi. S. Okeechobee (GB, CU); Basinger (PU). Orange Co.: Titusville (CNC). Pinellas Co.: Clearwater (CMNH); Dunedin (CU, PU); Royal Palm Park (PU). Sarasota Co.: Myakka River State Park (TLE, GB). St. Johns Co.: St. Augustine (MCZ). Volusia Co.: Enterprise (USNM); New Smyrna (OSU). Unrecorded localities in Florida (OSU, USNM, CU). Georgia. Baker Co.: Emory Univ. Field St., Newton (CU?). Bibb Co.: Macon (TLE); Glynn Co.: St. Simons Island (USNM). Lee Co.: Smithville (CU). Louisiana. Calcasieu Co.: Sabine River Ferry Landing (CU, CAS). Franklin Co.: Chase (UA). Orleans Co.: New Orleans (MCZ, USNM, CAS). Plaquemines Co.: Myrtle Grove (INHS). St. Mary Co.: Franklin (DW). Vermilion Co.: Gueydan (USNM). Un- recorded localities in Louisiana (UM, CAS, AM, USNM). Mississippi. Jackson Co.: Horn Island (CU) ; Lucedale (CU). South Carolina. Beau- fort Co.: no locality (CNC). Texas. Brazos Co.: College Station (TAM). Dallas Co.: Dallas (MCZ). Galveston Co.: Galveston (UK). west indies. Bahamas. Cat Island: Arthur’s Town (MCZ). Grand Cayman: South Sound (MCZ). Inayua (Island?) (MCZ). Cuba. Soledad (MCZ). Discussion. L. celeris resembles L. cubanus in most external char- acters. Identification of females from outside the United States can be made only on the basis of their geographical distribution. Identifica- tion of females and males from the United States is based on the shape of the posterior angles of the pronotum and on the aedeagii of males. Specimens from the eastern part of the United States usually have a distinct orange spot near the apex of the elytra while those from farther west usually lack this spot. This species has been collected in all months of the year. It is commonly found in very moist areas and can be collected under leaf debris and by treading. It has also been collected at lights. 7. Loxandrus cubanus Tschitscherine Figs. 68 A, B; 76; 148 Loxandrus cubanus Tschitscherine 1903 : 60. Holotype. $. Brussels Mu- seum of Natural History. Paratypes. 1 9 . Brussels Museum of Natural History. Description. Color. Dorsal: black or dark brown; shining; elytra iridescent; distal area of elytra sometimes with a centrally located orange spot. Ventral: black or dark brown; apex of last abdominal segment usually light brown. Appendages: dark to light brown or straw yellow. Head. Clypeus: anterior margin straight or very 46 Entomologica Americana [Vol. 46, No. 1 slightly concave. Epistomal suture: distinct, complete. Subgenal suture: present or absent. Frontal sulci: obscure, broad, shallow. Microsculpture: clypeus, frons and vertex isodiametric. Mentum: apex of lobes narrowly rounded; tooth distinct, almost as long as lobes (mentum more or less transverse); furrows shallow; fovae, if present, small, shallow; microsculpture isodiametric or slightly elongate. Thorax. Pronotum: microsculpture elongate on the disk, isodiametric along the lateral and basal margins and in the basal impressions; lateral margins rounded in the anterior half, more or less taper- ing in the posterior half, not sinuate, impunctate, posterior angles acute; anterior marginal sulcus present or absent, not complete if present; posterior marginal sulcus reaching well past the basal im- pressions but incomplete; median sulcus usually not reaching the ante- rior marginal sulcus, never reaching the posterior margin; basal impressions elongate, moderately deep, sparsely punctate; area be- tween the basal impressions punctate; area between the basal impres- sions and the lateral margins, if punctate, with only 1 or 2 punctures. Scutellum: microsculpture isodiametric; apex pointed. N. B. Entire ventral surface impunctate. Prosternum : microsculpture elongate me- dially, isodiametric laterally. Proepisternum and proepimeron: micro- sculpture slightly to very elongate. Mesosternum: microsculpture elongate. Mesepisternum: microsculpture isodiametric in anterior three quarters, elongate in posterior quarter. Metasternum: micro- sculpture elongate medially, isodiametric laterally. Metepisternum : microsculpture isodiametric. Metepimeron: microsculpture, if pres- ent, very elongate. Legs. Hind coxae: microsculpture on inner surface absent or very elongate. Tarsi: segments 1-3 of middle and hind tarsi and sometimes 4 of hind tarsi grooved dorsolaterally. Ely- tra. Microsculpture on intervals absent; all striae distinct, punctate, trough 1-3 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture distinct, isodiametric laterally, elongate medially; impunctate; segments 1-5 sparsely rugate laterally. Aedea- gus. Median lobe: not modified. Internal sac: everts medially, strongly turned to the right side; district scale groups absent; small, weakly sclerotized scales covering most of the sac’s surface; sclerotized ring absent. Measurements. (16 specimens). TL — 6.0 (5. 8-6.4); WH— 1.3 (1.2-1.4); WP— 1.8 ( 1. 7-2.0) ; WW— 2.5 (2.3-3. 1). Figs. 68-70. Prothorax, dorsal aspect. 68. A. L. cubanus Tschit. 68 B. same, close-up of hind angle. 69 A. L. tropicus n. sp. 69 B. Same, close-up of hind angle. 70 A. L. remotus n. sp. 70 B. Same, close-up of hind angle. 1972] Allen: Revision of Loxandrus LeConte 47 68A L. CUBANUS 68 B 69A L. TROPICUS 70 A L. REMOTUS 69B 70B 48 Entomologica Americana [Vol. 46, No. 1 Distribution. Mexico, Central America and West Indies. Localities, costa rica. Guanacaste: 5 km. N. Canas (GN). Mex- ico. Jalisco: 10.2 mi. NE. La Huerta, Rte. 80, 1200 ft. (GB). Sinaloa: Los Machis (GB). Nayarit: 5.7 mi. E. San Bias, Rte. 46, 140 ft. (GB). Oaxaca: 13.8 mi. W. Zanatepec, Rte. 190, 475 ft. (GB); 1.7 mi. W. Zana- tepec, Rte. 190, 300 ft. (GB). west indies. Bahamas. Arthur’s Town (Cat I.) (MCZ); Inagua (MCZ). Cuba. Soledad (Cienfuegos) (MCZ). Oriente: *Aguadores (MCZ). Dominican Republic. Puerto Plata (MCZ), *Sanchez (MCZ). Haiti. Swamps N. of Dessalines (MCZ). Puerto Rico. Guanica Lagoon (MCZ). Discussion. This species closely resembles L. ceteris. Females of these two species can only be identified by using geographical data. Specimens from Oaxaca have an orange colored spot near the apex of the elytra. Specimens from other areas in Mexico and Central America lack this spot. Darlington (1934) synonymized this species with L. ceteris. As I have stated, the external characters of the two species are very similar. The genitalia of the males, however, are distinct. L. cubanus has been found in marsh habitats. It has been collected in July and August. 8. Loxandrus belli New Species Figs. 25; 88; 151 Holotype. $. Mexico. Veracruz: 2.5 mi. W. Sontecomapan, 100 ft., September 20-26, 1965. George E. Ball and D. R. Whitehead, Collectors (MCZ). Paratypes: 65 S S 66 $ $ (GB); 1 <3 1 $ (GN); 1 52 5$ (UA); 1 $ (BM). Description. Color. Dorsal: black to dark brown, shining; elytron with 1 yellow or orange spot near the humeri and 1 near the apex. Ventral: black to dark brown, apex of last abdominal segment yellow. Appendages: usually pale yellow, sometimes brown or orange. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: obscure, ill defined. Frontal sulci: obscure; shal- low, broad, short. Microsculpture: clypeus, frons and vertex isodia- metric. Mentum: apex of lobes narrowly rounded; tooth distinct, Figs. 71-74. Male genitalia. 71 A. L. infimus Bates, left lateral aspect. 71 B. Same, right lateral aspect, IS everted. 72. L. elongatus n. sp., dorsal aspect, IS everted. 73 A. L. dubius (Curtis), left lateral as- pect. 73 B. Same, left lateral aspect, IS everted. 74 A. L. rectangu- lus LeC., left lateral aspect, IS everted. 74 B. Same, dorsal aspect, IS everted. 1972] Allen: Revision of Loxandrus LeConte 49 50 Entomologica Americana [Vol. 46, No. 1 not as long as lobes; furrows shallow; fovae small; microsculpture elongate. Thorax. Pronotum: microsculpture absent; lateral margins very slightly sinuate posteriorly, punctate in posterior quarter to half; posterior angles acute; anterior marginal sulcus complete or not; posterior marginal sulcus reaching past the basal impressions but incomplete; median sulcus not reaching anterior marginal sulcus or posterior margin; basal impressions elongate, moderately deep, usually impunctate; area between basal impressions with only 1-2 punctures if punctate; area between basal impressions and lateral margins usu- ally impunctate. Scutellum: microsculpture isodiametric and elon- gate; apex pointed. N. B. Microsculpture on ventral surface absent ex- cept around hind coxae. Prosternum, proepisternum, proepimeron: impunctate. Mesosternum: impunctate. Mesepisternum: weakly punctate in anterior quarter. Metasternum : punctate toward the lateral margin. Metepisternum, metepimeron: impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi grooved dorso-laterally. Elytra. Microsculp- ture on intervals absent; all striae distinct, punctate, trough 1 isodia- metric unit wide; internal plica well developed; humeri rounded; micro- sculpture on epiplura absent. Abdominal sternites. Microsculpture vaguely present around metacoxae; segment 1 punctate laterally at the base; segments 1-3 rugate laterally at the base. Aedeagus. Median lobe: not modified. Internal sac: everts medially and directed dorsally; 1 elongate scale group on the right lateral surface; most of the sac’s surface covered with small weakly sclerotized scales; sclero- tized ring absent. Measurements. (20 specimens). TL — 5.3 (5.1— 5.3); WH— 1.1 (1. 0-1.2); WP— 1.6 (1.5— 1.7); WW— 2.4 (2.5- 2.2). Distribution. Mexico and Central America. Localities, costa rica. Cartago: Turrialba, IICA Experiment Sta- tion (UA). Guanacaste: 5 Km. N. Canas (GN). Mexico. Chiapas: 11.6 mi. N. Ocozocuautla, 3200 ft. (GB); San Quintin, 90°20/-16°24/, 700 ft. (GB); San Quintin, Sierra de la Colmena ca. 300-700 ft. (GB); Palenque Ruins, 400 ft. (GB). Veracruz: 2.5 mi. W. Sontecomapan, 100 ft. (GB); 6.7 mi. NW. Sontecomapan, 200 ft. Coyame, Lake Catemaco (GB). Figs. 75-78. Male genitalia, IS everted. 75. L. ceteris (Dej.), ventral aspect. 76. L. cubanus Tschit., left lateral aspect. 77 A. L. remotus n. sp., left lateral aspect. 77 B. Same, left lateral aspect. 78 A. L. subiridescens M’Leay, ventral aspect. 78 B. Same, dorsal aspect. 78 C. Same, left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 51 52 Entomologica Americana [Vol. 46, No. 1 Discussion. More specimens of this species were collected by Ball and company in Mexico than any other Loxandrus. It is a rather small species. The males can be readily identified utilizing the characters associated with the genitalia. The females can usually be identified on the basis of external characters set forth in the key. It has been collected in the months of Februrary, June, July and Sep- tember in leaf litter and at black lights. It is named in honor of my friend and colleague Dr. George E. Ball whose contributions to the study of Carabidae are without equal. 9. Loxandrus elnae New Species Figs. 28 A, B; 90 A, B; 152 Holotype. $. MEXICO. Chiapas: 32.5 mi. E. Comitan, Rte. 190, 2200 ft., September 3, 1965. George E. Ball and D. R. Whitehead, Collectors (MCZ) . Paratypes. 19 8 $ (GB); 1 2 (UA); 1 $ (GN). Description. Color. Dorsal: head and pronotum and back- ground of elytra black to dark brown; elytra various, usually with at least two orange to yellow spots at the humeri and two apically, some- times the humeral spots greatly elongate reaching or almost reaching the apical spots; labrum and apex of clypeus usually orange or yellow. Ventral: black to dark brown, apex of last abdominal segment usually light yellow. Appendages : brown, apex yellow or light yellow. Head. Clypeus: anterior margin straight or slightly concave. Epistomal suture: distinct, complete. Subgenal suture: obscure if present. Frontal sulci: obscure, broad, short, shallow. Microsculpture: clyp- eus isodiametric. Mentum: apex of lobes acute; tooth distinct, not as long as lobes; furrows shallow; fovae distinct, small; microsculpture elongate. Thorax. Pronotum: microsculpture entirely absent; lateral margins punctate in the posterior half, evenly rounded their entire length; posterior angles narrowly rounded: anterior marginal sulcus complete; posterior marginal sulcus complete or not; median sulcus not reaching anterior or posterior marginal sulci; basal impressions elongate, moderately deep, punctate; area between basal impressions sparsely punctate; area between impressions and lateral margins with or without 1-2 punctures. Scutellum: microsculpture isodiametric; apex pointed. N. B. Microsculpture evident only near the pro- and Figs. 79-81. Male genitalia. 79 A. L. latus Dari., right lateral aspect, sac everted. 79 B. Same, left lateral aspect, sac everted. 80 A. L. subrectus Dari., left lateral aspect, sac everted. 80 B. Same, right lateral aspect, sac everted. 81. L. discolor n. sp., left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 53 81 L. DISCOLOR 54 Entomologica Americana [Vol. 46, No. 1 metacoxae. Prosternum: impunctate. Proepisternum: sparsely punc- tate. Proepimeron: impunctate. Mesosternum: impunctate. Mesepi- sternum: punctate in anterior half to three quarters. Metasternum: punctate toward lateral margins. Metepisternum sparsely punctate or impunctate. Metepimeron: impunctate. Legs. Hind coxae: micro- sculpture on inner surface absent. Tarsal segments: 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Mi- crosculpture absent on the intervals; all striae distinct, punctate, trough 1-2 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Mi- crosculpture absent; segment 1 punctate laterally at the base; segments 1—5 rugate laterally. Adeagus. Median lobe: not modified. Internal sac: everts in a median position, distinctly turned dorsally; three distinct scale groups on the main sac; secondary sac everts on the dorsal mem- braneous portion of the median lobe, scale group at the apex; much of the remainder of sac’s surface covered with small weakly sclerotized scales; sclerotized ring absent. Measurements. (25 specimens). TL — 6.8 (5. 6-7. 4) ; WH— 1.3 (1.2— 1.4); WP— 2.1 (1.7-2.4); WW— 3.0 (2.5-3. 3). Distribution. Mexico and Central America. Localities. Costarica. Guanacaste: 5 Km. N. Canas (GN). Mex- ico. Chiapas: 32.5 mi. E. Comitan, Rte. 190, 2200 ft. (GB); 37.7 mi. E. Comitan, Rte. 190, 2200 ft. (GB); 37.7 mi. E. Comitan, Rte. 190, Rio Chiapa, 2200 ft. (GB). Morelos: Santa Rosa, 3.2 mi. N. Zacatepec, 3100 ft. (GB). Oaxaca: 5 mi. E. Tapanatepec, Rte. 190, 800 ft. (GB). Discussion. This species closely resembles several other Lox- andrus species from Mexico and Central America. It can be separated from them only by utilizing characters associated with the aedeagii of the males. It has been collected in the months of April, June, July and September. This species is named in honor of my wife Mrs. Elna Allen. 10. Loxandrus whiteheadi New Species Figs. 27 A, B; 86 A, B; 150 Holotype. $. Mexico. Chiapas: 32.5 mi. E. Comitan, Rte. 190, 2200 ft., September 4, 1965. G. E. Ball and D. R. Whitehead, Collectors (MCZ) . Paratypes: 24 $ $ (GB);2 $$ (RTB); 1 S (UA). Description. Color. Dorsal: head, labrum and clypeus light brown to straw yellow, remainder of head usually black, sometimes dark brown; pronotum, dark to light brown, lateral areas usually lighter in color than disk; elytra, dark brown, an orange or yellow spot near the humerus and toward the apex. Ventral: dark brown, occa- 1972J Allen: Revision of Loxandrus LeConte 55 sionally black, apex of last abdominal segment sometimes yellow. Appendages: dark brown to straw yellow. Head. Clypeus: anterior margin slightly concave or straight. Epistomal and subgenal sutures: complete. Frontal sulci: broad, moderately impressed. Microsculp- ture: clypeus, frons and vertex isodiametric. Mentum: apex of lobes narrowly rounded; tooth distinct, broadly rounded at apex; furrows shallow or absent; fovae present or absent, if present small and punc- ture-like; microsculpture elongate. Thorax. Pronotum: microsculp- ture absent except along narrowly reflexed lateral margins; lateral margins weakly sinuate posteriorly, punctate in posterior half; poste- rior angles distinct, acute; anterior marginal sulcus complete; posterior marginal sulcus reaching only to basal impressions; median sulcus not reaching anterior marginal sulcus or posterior margin; basal impres- sions elongate, moderately deep, punctate; area between impressions and between impressions and lateral margins sparsely punctate. Scu- tellum: microsculpture isodiametric; apex pointed. N. B. Entire ven- tral surface without microsculpture. Prosternum: impunctate. Proepi- sternum: punctate. Proepimeron. impunctate. Mesosternum: im- punctate. Mesepisternum : punctate in anterior quarter to half. Metasternum: punctate toward lateral margin. Metepistemum : punctate or not. Metepimeron: impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi grooved dorso-laterally. Elytra. Microsculp- ture on intervals absent; all striae distinct, punctate, trough 2-3 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture on epipleura absent. Abdominal sternites. Microsculp- ture absent; segment 1 punctate and rugose basally towards the lateral margin; segments 2 and 3 weakly rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position, directed dorsally; no distinct scale groups present; most of the sac’s surface covered with small weakly sclerotized scales; a small bulbous projec- tion on the right dorso-lateral margin of the sac; sclerotized ring absent. Measurements. (20 specimens). TL — 6.5 (6. 0-7. 2); WH — 1.4 (1.2-1. 5); WP— 2.0 (1. 8-2.2); WW— 2.9 (2.6-3. 1). Distribution. Central America. Localities. Guatemala. Escointla: Zapote. Mexico. Chiapas: 8 mi. SE. Arriaga (RTB); 32.5 mi. E. Comitan, Rte. 190, 2200 ft. (GB); 0.9 mi. N. Frontera Comalapa, 2100 ft. (GB); 11.6 mi. N. Ocozocuautla, 3200 ft. (GB). Oaxaca: 5 mi. E. Tapanatepec, Rte. 190, 800 ft. (GB). Veracruz: 2.5 mi. W. Sontecomapan, 100 ft. (GB). Discussion. L. whiteheadi is named in honor of Mr. Don White- head. Specimens having a punctate metepistemum can readily be 56 Entomologica Americana [Vol. 46, No. 1 distinguished on the basis of external characters. Those specimens in which the metepisternum is impunctate cannot be separated from L. unispinus , L. elnae , L. fraus , L. lepidus, or L. tropicus using ex- ternal characters. For this reason females have been excluded from the paratype material. L. whiteheadi has been collected in the months of June, August, and September. 1 1 . Loxandrus tropicus New Species Figs. 69 A, B; 84; 149 Holotype. S. Costarica. Cartago Province; Turrialba. July 11, 1965; R. J. Hampton, Collector (MCZ). Paratypes: 1 S (GN); 1 $ (UA). Description. Color. Dorsal surface: shining; head, disk of pronotum and elytra black to dark brown; elytra with two apical and two basal orange to yellow spots; pronotum with lateral areas light brown to straw yellow. Ventral surface: black to dark brown, ab- dominal segments tinged with yellow. Appendages: light brown to straw yellow. Head. Clypeus: anterior margin straight or slightly concave. Epistomal suture: distinct. Subgenal suture: distinct or absent. Frontal sulci: distinct, shallow, short. Microsculpture: clypeus, frons and vertex, distinct, isodiametric. Mentum: apex of lobes rounded; tooth distinct, not as long as lobes; furrows present; fovae present or absent; microsculpture elongate or absent. Thorax. Pronotum: microsculpture absent or very elongate, vaguely present only along the lateral margins in the anterior half; lateral margins coarsely punctate in the posterior half to three quarters, tapering evenly to the posterior angles, not sinuate posteriorly; posterior angles pointed; anterior marginal sulcus complete; posterior marginal sulcus complete or not; median sulcus distinct, reaching the anterior and posterior sulci or not; basal impressions distinct, elongate, punctate; basal quarter coarsely punctate. Scutellum: apex pointed; microsculp- ture distinct, isodiametric. N. B. Entire ventral surface without micro- sculpture. Prosternum; impunctate. Proepistemum : coarsely but sparsely punctate. Mesosternum: impunctate. Mesepisternum : coarsely punctate anteriorly. Metasternum: coarsely punctate later- ally. Metepisternum: punctate or not. Metepimeron: impunctate. Figs. 82-84. Male genitalia. 82 A. L. discolor n. sp., left lateral aspect, apex of median lobe with IS everted. 82 B. Same, dorsal aspect of median lobe. 83 A. L. spinigrandis n. sp., left lateral aspect. 83 B. Same, right lateral aspect, IS everted. 84. L. tropicus n. sp., left lateral aspect, IS everted. 84 L. TROPICUS 58 Entomologica Americana [Vol. 46, No. 1 Legs. Hind coxae: microsculpture on inner surface absent; tarsal segments 1-4 of middle and hind tarsi grooved dorso-laterally. Elytra. Microsculpture absent; all striae distinct, trough 2-3 isodiametric units wide; internal plica well-developed; humeri rounded; micro- sculpture of epipleura absent. Abdominal sternites. Microsculpture absent; segment 1 coarsely punctate near the lateral anterior surface; segments 2 and 3 sometimes punctate in the central area; other seg- ments impunctate. Aedeagus. Median lobe: apex not modified. Internal sac : everts medially but is turned dorsally and then ventrally and to the left side; no distinct scale groups; most of the surface of the sac is covered by small lightly sclerotized scales; sclerotized ring absent. Measurements. (2 $ specimens). TL — 7.3 (7. 1-7. 6); WE —1.5 ( 1. 4-1.5 ); WP— 2.3 (None) ; WW— 4.7 (4.6-4.8). Distribution. Central America. Localities, costa rica. Cartago: Turrialba (MCZ) (GN). Discussion. This species is known from Costa Rica. Two males and two females were collected by Mr. R. J. Hampton and donated to Mr. Gary Noonan. The specimens were probably collected on or near the IICA Experiment Station just outside of Turrialba. The external morphological characters are similar to the characters of the other species in this complex. The females can not be separated with any accuracy and are not marked as paratypes. 12. Loxandrus lepidus New Species Figs. 67; 85; 149 Holotype. $. Mexico. Veracruz: Fortin de las Flores, June 26-30, 1963, leaf litter. D. R. Whitehead, Collector (MCZ). Description. Color. Dorsal surface: shining, background color black, lateral areas of pronotum and lateral and apical areas of elytra brown; elytra with two apical and two basal orange to yellow spots. Ventral surface: shining, background color black, lateral and apical areas of segments 4 and 5 light brown. Appendages: brown to light brown. Head. Clypeus: anterior margin straight. Epistomal and subgenal sutures: distinct, complete. Frontal sulci: distinct, broad, Figs. 85-88. Male genitalia. 85. L. lepidus n. sp., left lateral aspect. IS everted. 86 A. L. whiteheadi n. sp., left lateral aspect, IS everted. 86 B. Same, dorsal aspect, apex of median lobe with IS everted. 87 A. L. unispinus n. sp., right lateral aspect, IS everted. 87 B. Same, left lateral aspect. 87 C. Same, dorsal aspect. 88. L. balli n. sp., left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 59 88 L. BALLI 60 Entomologica Americana [Vol. 46, No. 1 shallow, short. Microsculpture: clypeus, frons and vertex, distinct, isodiametric. Mentum: apex of lobes rounded; tooth distinct, not as long as lobes; furrows present; fovae absent; microsculpture elongate. Thorax. Pronotum: microsculpture absent on most of the surface, present only along the reflexed lateral margins; lateral margins weakly punctate, slightly sinuate near the basal angle; posterior angles pointed; anterior marginal sulcus complete; posterior marginal sulcus incom- plete; median sulcus present, not reaching anterior or posterior sulci; basal quarter sparsely punctate. Scutellum: apex pointed; microsculp- ture distinct, isodiametric. N. B. Entire ventral surface without micro- sculpture. Prosternum: impunctate. Proepisternum : weakly and sparsely punctate. Proepimeron : impunctate. Mesosternum : impunc- tate. Mesepisternum : punctate in the anterior half to three quarters. Metasternum: punctate toward lateral margin. Metepisternum and Metepimeron: impunctate. Legs. Hind coxae: microsculpture on inner surface absent; segments 1-3 of middle and hind tarsi grooved dorso-laterally. Elytra. Microsculpture absent; all striae distinct, trough 1 isodiametric unit wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture absent; segment 1 coarsely punctate along the lateral basal margin; segment 2 sparsely punctate laterally; other segments impunctate. Aedeagus. Median lobe: apex not modified. Internal sac: everts medially, deflected dorsally. Internal sac: no distinct scale groups; small lightly sclerotized scales cover most of the surface of the sac, somewhat more dense towards apex and in the mid region of the right side; sclerotized ring absent. Measurements. (1 speci- men). TL— 7.3; WE— 1.5; WP— 2.1; WW— 3.3. Distribution. Mexico. Localities. Mexico. Veracruz: Fortin de las Flores (MCZ). Discussion. This species is known from a single male specimen collected in leaf litter between June 26-30, 1963, by D. R. Whitehead. Its nearest relative appears to be L. tropicus. It differs from L. tropicus only in the shape of the internal sac of the aedeagus as characterized in the key. 13. Loxandrus f raus New Species Figs. 23 A, B; 89; 152 Holotype. $. Mexico. Morelos : Santa Rosa, 3.2 mi. N. Zacatepec, 3100 ft. elevation, March 31, 1966. McFadden, Whitehead, Collectors (MCZ). Paratypes: 44 S $ (GB); 1 $ (UA). Description. Color. Dorsal: shining, head and elytra black to dark brown, pronotum dark to light brown, elytra with two apical and 1972] Allen: Revision of Loxandrus LeConte 61 two basal orange to yellow spots, spots may sometimes be very elon- gate. Ventral: usually black to dark brown, sometimes light brown, abdominal segments sometimes tinged with yellow. Appendages: dark brown to straw yellow. Head. Clypeus : anterior margin straight or slightly concave. Epistomal and subgenal sutures: distinct, com- plete. Frontal sulci: obscure, broad, shallow, short. Microsculpture: clypeus, frons and vertex isodiametric. Mentum: apex of lobes nar- rowly rounded; tooth distinct, not as long as lobes; furrows shallow, broad; fovae small but distinct; microsculpture absent or very elongate. Thorax. Pronotum: microsculpture absent even along the lateral margin and in the basal furrows; lateral margins evenly rounded their entire length, not sinuate, punctate in the posterior half to three quarters; posterior angles narrowly rounded; anterior marginal sulcus complete; posterior marginal sulcus complete or not; median sulcus reaching anterior marginal sulcus or not, never reaching posterior marginal sulcus; basal impressions elongate, moderately deep, punc- tate; area between basal impressions punctate; area between basal impressions and lateral margins sparsely punctate. Scutellum : micro- sculpture isodiametric and/or elongate; apex pointed. N. B. Entire ven- tral surface of the pronotum without microsculpture. Prosternum: im- punctate. Proepisternum : sparsely and weakly punctate. Proepi- meron: impunctate. Mesosternum: impunctate. Mesepisternum : anterior quarter to half punctate. Metasternum: punctate laterally. Metepisternum : usually impunctate, sometimes with 1-3 obscure punctures. Metepimeron: impunctate. Legs. Hind coxae: micro- sculpture on inner surface absent. Tarsal segments: 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct, punctate, trough 1-2 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture on epipleura absent. Abdominal sternites. Microsculpture absent except around hind coxae; segment 1 punctate laterally at the base, other segments impunctate; segments 1-5 sparsely rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts medially and turned dorsally; two distinct scale groups on the right side; remainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring absent. Measurements. (25 speci- mens). TL— 6.0 (5.6— 6.5); WH— 1.2 (1.1-1. 3); WP— 1.8 (1.7- 2.0); WW— 2.6 (2.5-2.8). Distribution. Mexico. Localities. Mexico. Chiapas: .9 mi. N. Frontera Comalapa, 2100 ft. (GB) ; 2.9 mi. N. Frontera Comalapa, 2200 ft. (GB) ; San Quintin, 700 ft., 91°2T-16°24/ (GB). Morelos: Santa Rosa, 3.2 mi. N. Zacatepec, 3100 ft. (GB). Nayarit: 5.7 mi. E. San Bias, Rte. 46, 140 ft. (GB); 13.8 mi. 62 Entomologica Americana [Vol. 46, No. 1 E. San Bias, Rte. 46, 200 ft. (GB). Oaxaca: 5 mi. E. Tapanatepec, Rte. 190, 800 ft. (GB). Discussion. In external characters this species is most like L. elnae. It also resembles L. tropicus , L. balli and L. unispinus and can- not be separated from them using external characters. Determination of females has been based on geographical distribution. L. fraus has been collected in the months of February, March, April, June, July, and August. 14. Loxandrus unispinus New Species Figs. 1 A, B; 2 A, B; 24 A, B; 87 A, B, C; 150 Holotype. $. canal zone. Barrro Colorado Island : April, 1941 . James Zetek, Collector; No. 4781, Lot No. 41-8655 (USNM). Paratypes: 11 $ $ (USNM); 2 $ S (UA). Description. Color. Dorsal surface: shining, background color dark to light brown; head usually darker than other parts; disk of pronotum may or may not be darker than lateral areas; elytra with two apical and two basal orange to yellow spots; ventral surface: shining, background color dark to light brown, occasionally tinged with yellow; appendages generally light to dark brown, sometimes straw yellow. Head. Clypeus: anterior margin straight. Epistomal and subgenal sutures: distinct, complete. Frontal sulci: distinct, broad, shallow, short. Microsculpture: clypeus, frons and vertex, distinct isodia- rnetric. Mentum: apex of lobes narrowly rounded; tooth distinct, not as long as lobes; furrows present; fovae present or absent; microsculp- ture absent or very elongate, vaguely present only along the lateral margin in the anterior fifth; lateral margins punctate, slightly sinuate posteriorly; posterior angles pointed; anterior marginal sulcus complete; posterior marginal sulcus complete or not; median sulcus present, reaching the anterior and posterior sulci or not; basal impressions distinct, elongate, sparsely punctate; basal quarter punctate. Scutel- lum: apex pointed; microsculpture distinct, isodiametric. N. B. Entire ventral surface without microsculpture. Prosternum: impunctate. Proepisternum : usually impunctate, sometimes with a few weak punc- tures. Proepimeron: impunctate. Mesosternum: impunctate. Mes- Figs. 89-92. Male genitalia. 89. L. fraus n. sp., right lateral aspect, IS everted. 90 A. L. elnae n. sp., left lateral aspect, IS everted. 90 B. Same, right lateral aspect, IS everted. 91 A. L. sculptilis Bates, left lateral aspect. 91 B. Same, left lateral aspect, IS everted. 92. L. negrei Straneo, left lateral aspect, IS everted. 1972] Allen: Revision of Loxandrus LeConte 63 92 L. NEGREI 64 Entomologica Americana [Vol. 46, No. 1 episternum: coarsely punctate anteriorly. Metasternum: coarsely punctate laterally. Metepistemum : usually impunctate, sometimes with a few weak punctures. Metepimeron: impunctate. Legs. Hind coxae: microsculpture on inner surface absent; tarsal segments 1-3 and sometimes 4 of middle and hind tarsi grooved dorso-laterally. Elytra. Microsculpture absent; all striae distinct, trough 1 isodia- metric unit wide; internal plica well-developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Micro- sculpture absent; segment 1 coarsely punctate near the lateral anterior surface; other segments impunctate. Aedeagus. Median lobe: not modified. Internal sac: everts medially, deflected dorsally; one well- developed, distinct, small scale group on right side of sac near the mid- line: other small scales covering much of the sac’s surface; sclerotized ring absent. Measurements. (14 specimens). TL — 5.7 (5.0-6.1); WE— 1.2 (1. 1-1.2); WP— 1.8 (1. 6-1.9); WW- 2.5 (2.2-2.8). Distribution. Southern Central America. Localities, canal zone. Albrook Forest, Fort Clayton (UA); Barro Colorado Island (USNM); Corozal (USNM). panama. Pearl Island, San Jose (USNM). Discussion. This species is presently known only from Panama where it has been collected at light traps in the months of April, May, June and July. The external characters of this species are very similar to those of L. f raus, L. elnae , L. tropicus and L. lepidus. No one ex- ternal character sets it apart from these species though it is generally smaller than any of them. Determination of females has been made by associating them with males having the same collecting data. 15. Loxandrus yeariani New Species Figs. 119; 169 Holotype. $. Mexico. Colima: 16.5 mi. E. Colima, 1900 ft., Rte. 110, August 2-3, 1966. G. E. Ball and D. R. Whitehead, Collectors ( MCZ) . Paratypes : 3 $ S 4 9 $ (GB). Description. Color. Dorsal and Ventral: black, shining. Ap- pendages: usually black, sometimes tinged with brown. Head. Clyp- eus: anterior margin concave. Epistomal and subgenal sutures: distinct, complete. Frontal sulci: obscure, shallow, broad, short. Figs. 93-94. Male genitalia. 93 A. L. pactinullus n. sp., left lateral aspect, IS everted. 93 B. Same, dorsal aspect, 93 C. Same, left lateral aspect. 94 A. L. tucumanus (Dej.), right lateral aspect, IS everted. 94 B. Same, left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 65 94 B 66 Entomologica Americana [Vol. 46, No. 1 Microsculpture: clypeus, frons and vertex weak, isodiametric. Men- tum: apex of lobes acute; tooth distinct, broadly rounded; furrows present; fovae present, small; microsculpture distinct, isodiametric. Thorax. Pronotum: microsculpture if present, elongate; lateral mar- gins impunctate, broadly rounded; posterior angles absent; anterior marginal sulcus complete; posterior marginal sulcus incomplete; me- dian sulcus barely reaching anterior marginal sulcus, not reaching posterior margin; basal impressions short, narrow, deep; area between impressions punctate; area between impressions and lateral margins sparsely punctate. Scutellum: microsculpture weak, isodiametric; apex pointed. N. B. Entire ventral surface impunctate. Prosternum: micro- sculpture elongate medially, isodiametric laterally. Proepisternum : microsculpture where present, elongate. Proepimeron: microsculpture isodiametric. Mesosternum: microsculpture isodiametric. Mesepi- sternum: microsculpture isodiametric anteriorly, elongate posteriorly. Metasternum: microsculpture isodiametric laterally, absent medially. Metepisternum: microsculpture isodiametric. Metepimeron: micro- sculpture absent. Legs. Hind coxae: microsculpture on inner surface isodiametric. Tarsal segments: 1-3 of middle and hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct; trough 3-4 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Ab- dominal sternites. Microsculpture mostly isodiametric, obscure and elongate medially on segments 2 and 3; impunctate; segments 1 and 2 rugose along the base laterally. Aedeagus. Median lobe: not modi- fied. Internal sac: everts medially and is directed dorsally; 3 distinct scale groups present; 1 large group at the apex of each of 2 secondary sacs, 1 small group on the right lateral surface; most of the remainder of the sac’s surface without small weakly sclerotized scales; sclerotized ring present. Measurements. (8 specimens). TL — 10.5(9.9-11.3); WH— 2.2 (2.1-2.4); WP— 3.3 (3. 1-3.6); WW- 4.1 (3. 8-4.4). Distribution. Mexico. Localities. Mexico. Colima: 16.5 mi. E. Colima, Rte. 110, 1900 ft. (MCZ, GB). Discussion. This is the largest species in the Mexican Loxandrus fauna. It is readily recognizable by the broadly rounded lateral mar- gins of the pronotum resulting in the absence of distinct posterior angles. The aedeagus is also characteristic. It is known from only one -> Figs. 95-96. Male genitalia, IS everted. 95 A. L. erraticus (Dej.), left lateral aspect. 95 B. Same, right lateral aspect. 96 A. L. gibbus n. sp., right lateral aspect. 96 B. Same, left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 67 68 Entomologica Americana [Vol. 46, No. 1 locality near Colima, Mexico, and has been collected only in August. It is named in honor of Dr. W. C. Yearian. 16. Loxandrus rectus (Say) Figs. 43 A, B; 98 A, B, C; 157 Feronia recta Say 1823:58. Neotype. From Lindroth and Freitag ( 1969) : “No type area given. The neotype agrees with Casey’s description (1918, p. 381) and also, except it is larger, with the 7 ex. in coll. Lee. (MCZ). A $ with genitalia dissected, from South Carolina, designated by R. T. Allen as neotype and the state as type area.” Feronia lucidula Dejean, 1828:239. Lectotype (here selected). A male found mixed with L. rectus material but marked with the green Dejean determination label as Lucidulus from Boreal America (Paris Museum Nat. Hist.). Megalostylus laticollis Chaudoir, 1843:96. Lectotype (here selected). A male and a female found mixed with L. rectus material belonging to the Chaudoir collection. The male bears a Laticollis Chaudoir determination label from Louisiana. The female bore no labels (Paris Museum Nat. Hist.). Loxandrus rectus sub. sp. mandibularis Casey, 1918:382. Type. Male from the District of Columbia. Type No. 47334 (USNM). Description. Color. Body entirely black, iridescent; appendages, dark brown to black. Head. Clypeus: anterior margin slightly con- cave (distinctly notched in one specimen). Epistomal and subgenal sutures: distinct, complete. Frontal sulci: shallow, broad, obscure. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes pointed; tooth distinct, shorter than lobes, rounded at apex; furrows weak; fovae absent; microsculpture elongate lines if present. Thorax. Pronotum: microsculpture absent on the disk, isodiametric in the reflexed lateral margins, sulci, and basal furrows; lateral margins impunctate, evenly rounded their entire length, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus usually present and complete; posterior marginal sulcus in- complete; median sulcus present, reaching anterior marginal sulcus but not posterior margin; basal impressions distinct, elongate and broad, punctate; area between basal impressions sometimes Figs. 97-98. Male genitalia, IS everted. 97. L. gibbus n. sp., left ventro- lateral aspect. 98 A. L. rectus (Say), right lateral aspect. 98 B. Same, left lateral aspect. 98 C. Same, dorsal aspect. 1972] Allen: Revision of Loxandrus LeConte 69 70 Entomologica Americana [Vol. 46, No. 1 sparsely punctate. Scutellum: apex pointed; microsculpture isodia- metric. Prosternum: impunctate; microsculpture absent medially, elongate towards lateral margin. Proepisternum : impunctate; microsculpture usually absent, if present, elongate. Proepimeron: impunctate; microsculpture elongate. Mesosternum: impunctate; microsculpture elongate. Mesepisternum : punctate anteriorly; micro- sculpture, if present, elongate. Metasternum: impunctate; microsculp- ture, if present, elongate. Metepisternum : impunctate; microsculpture, if present, elongate. Metepimeron: impunctate; microsculpture absent. Legs. Hind coxae: microsculpture on inner surface isodiametric; tarsal segments 1-4 of middle and hind tarsi grooved dorsolaterally. Elytra. Microsculpture absent except in striae, trough 3-4 isodiametric units wide; striae 7 not as distinct basally as 1-6; internal plica well- developed; humeri rounded; microsculpture of epipleura absent. Ab- dominal sternites. Microsculpture isodiametric; segment 1 punctate along the basal margin; other segments impunctate. Aedeagus. Me- dian lobe: not modified. Internal sac: everts medially, slightly turned dorsally; one large scale group covers much of the surface of the sac, the scales are distinct but vary in length and shape over the surface; a second distinct but smaller group of very elongate, slender scales is located apically near the sclerotized ring; sclerites not present; sclero- tized ring present. Measurements. (20 specimens). TL — 11.2 (10.2-11.8); WH— 2.4 (2.1-2.5); WP— 3.7 (3.2-4.0); WE— 4.5 (4.1-4.9). Distribution. Southeastern United States north to Washington, D. C. and Illinois, and west to Arkansas and West Texas. Localities. Alabama. Mobile Co.: Calvert (USNM, CAS); Mt. Vernon (OSU); Mobile (MCZ, TAM). Tuscaloosa Co.: Tuscaloosa (GB). Arkansas. Arkansas Co.: Bayou Meto, Hy. 11, East of Reydel (UA); Ashley Co.: Ouachita River, Hy. 84, 6 mi. W. Crossett (UA); Washington Co.: 5 mi. S. Fayetteville (INHS, MSU); White Co.: Cy- prus Bayou, 10 mi. W. Beebe (UA). District of Columbia. Washing- ton (USNM, UMIN, CU, MCZ, INHS, CNC, OSU). Florida. Marion Co. (AM). Illinois. Hardin Co. : Elizabethtown (INHS). Richland and Lawerence Co.’s: Wabash Valley (MCZ). Kentucky. Undetermined locality (CNHM). Louisiana. Calcasiew Co.: 8 mi. W. Vinton, Sabine River Relief (GB); Franklin Co.: Chase (UA); Madison Co.: Tallulah (MCZ); Orleans Co.: New Orleans (MCZ, GB, CAS, USNM, CU, KU); Ouachita Co.: Monroe (USNM). St. Martins Co. (UA); Union Co. (UA); Vernon Co.: Rosepine (UA). Undetermined localities in Louisi- ana (CAS, CU, INHS, MCZ, OSU, UM, UMIN, USNM). Maryland. Undetermined locality (UMIN). Mississippi. George Co.: Lucedale (CU); Green Co.: Leakville (CU); Harrison Co.: Gulfport (CU); Jackson Co.: Ocean Springs (CU). Missouri. St. Louis Co.: St. Charles 1972] Allen: Revision of Loxandrus LeConte 71 Figs. 99-100. Male genitalia, IS everted. 99 A. L. velocipes Casey, right lateral aspect. 99 B. Same, left lateral aspect. 100 A. L. pravi- tubus n. sp., dorsal aspect. 100 B. Same, right lateral aspect. 72 Entomologica Americana [Vol. 46, No. 1 (MCZ). North Carolina. Chowan Co.: Edenton (MCZ). South Caro- lina. Undetermined locality (MCZ). Tennessee. Lake Co.: Grays Ldg. (RTB) ; Shelby Co.: Memphis (USNM, CAS). Texas. Brazos Co.: College Station (TAM); Calhoun Co.: Alligator Head (USNM); Dallas Co.: Dallas (MCZ, USNM, MSU); Jackson Co.: Edna (USNM); Vic- toria Co.: Victoria (USNM). *Mitten (MCZ). Undetermined localities in Texas (MCZ, USNM). Virginia. Nelson Co.: Rosslyn (MCZ, USNM). Undetermined locality in Virginia (USNM). Discussion. This is one of the larger species of Loxandrus found in North America. The large size and broadly rounded and explinate posterior angles of the pronotum serve to distinguish this species and L. pravitubus from all other Loxandrus species. L. rectus can be dis- tinguished from L. pravitubus by its larger size as well as the character- istic genitalia of both the males and females. It has been collected in all months of the year except September. 17. Loxandrus pravitubus New Species Figs. 44 A, B; 100 A, B; 159 Holotype. S. u.s.a. Texas: Brownsville, June 1 1-16, 1933. Darlington, Collector. (MCZ). Paratypes. 17 $ $ 19 9 $ (MCZ); 13 S $ 13$$ (USNM); 1 $ 4 $ $ (TAM); 1 S (GB); 2 2 2 (OSU); 3 $ $ (CAS); 13 19 (UA) ; 3 2 2 (CNC); 1 2 (CU). Description. Color. Dorsal: black, iridescent. Ventral: usually black, some areas sometimes dark brown, iridescent. Appendages: dark to light brown. Head. Clypeus: anterior margin slightly con- cave. Epistomal and subgenal sutures: distinct, complete. Frontal sulci: shallow, broad, obscure. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes pointed; tooth distinct, shorter than lobes, rounded at apex; furrows weak; fovae absent; microsculpture elongate lines if present. Thorax. Pronotum: microsculpture absent on the disk, isodiametric in the reflexed lateral margins, in the sulci and basal furrows; lateral margins impunctate, evenly rounded their entire length, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus usually present and complete; posterior marginal sulcus incomplete; median sulcus present, reaching anterior marginal sulcus but not posterior margin; basal impressions distinct, elongate and broad, punctate; area between basal impressions -> Fig. 101. Male genitalia, IS everted. 101 A. L. brevicollis (LeC.), left lateral aspect. 101 B. Same, right lateral aspect. 101 C. Same, ventral aspect. 101 D. Same, apex of median lobe, left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 73 74 Entomologica Americana [Vol. 46, No. 1 Fig. 102. Male genitalia, IS everted. 102 A. L. circulus n. sp., left lateral aspect. 102 B. Same, right lateral aspect. 102 C. Same, left lateral aspect. 102 D. Same, ventral aspect. 102 E. Same, right lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 75 sometimes sparsely punctate. Scutellum: apex pointed; microsculpture variable, if present usually isodiametric or only slightly elongate. Pro- sternum: impunctate; microsculpture absent medially, elongate towards lateral margin. Proepisternum : sometimes weakly punctate; microsculpture distinctly elongate. Proepimeron: impunctate; micro- sculpture elongate. Mesosternum: impunctate; microsculpture elongate. Mesepisternum: punctate anteriorly (punctures usually more numerous than in L. rectus and covering more of the sclerite). Meta- sternum: weakly punctate towards lateral margins; microsculpture distinct laterally, elongate and isodiametric. Metepisternum : punctate, shallow but usually distinct; microsculpture distinct, elongate. Metepi- meron: impunctate; microsculpture absent. Legs. Hind coxae: mi- crosculpture on inner surface isodiametric; tarsal segments 1-4 of hind tarsi and 1-3 of middle tarsi grooved dorso-laterally. Elytra. Micro- sculpture absent except in striae, trough 4-5 isodiametric units wide; striae 7 usually as distinct as 1-6; internal plica well-developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture isodiametric; segment 1 punctate along the basal margin; other segments impunctate. Aedeagus. Median lobe: modi- fied, expanded and turned to the left side in the apical two thirds. Internal sac: everts medially and turns dorsally, several small bulbous projections on the sac; one distinct scale group present; the remainder of surface of the sac covered with very small, weakly sclerotized scales; sclerotized ring present. Measurements. (20 specimens). TL —9.3 (8.7— 9.3); WH— 1.9 (1. 8-2.1); WP— 3.0 (2.8-3. 3); WW— 3.7 (3. 3-4.1). Distribution. Southeastern United States, west through South Texas to Mexican border. Localities. Alabama. Mobile Co.: Calvert (USNM). Mobile (MCZ). Louisiana. Madison Co.: Tallulah (MCZ). North Carolina. Robeson Co. : Lumberton (USNM). Texas. Brazos Co. : College Station (TAM); Cameron Co.: Brownsville (CNC, MCZ, OSU, CAS, USNM); Colorado Co.: Columbus (USNM, MCZ, MSU); Coma Co.: New Brun- fels (USNM, MCZ); Dallas Co.: Dallas (MSU, MCZ, USNM); Demmit Co.: Carrizo Springs (USNM); Galveston Co.: Dickinson (TAM); Gil- lespie Co.: (OSU) ; Harris Co.: Houston (CAS) ; Hildago Co.: Weslaco (TAM); Victoria Co.: Victoria (USNM, GB, UM). Wharton Co.: Wharton (CU). * Mitten (MCZ). Unrecorded localities in Texas (MCZ, USNM). Discussion. This species closely resembles L. rectus in most ex- ternal characters but it is smaller. It has probably been confused with L. rectus in most collections. L. pravitubus can be distinguished from L. rectus by its smaller size and the differences in both the male and 76 Entomologica Americana [Vol. 46, No. 1 female genitalia. It has been collected in April, May, June, July, October, and December. The L. unilobus , L. saccisecundaris , L. extendus and L. taeniatus Complex. Discussion. The four species in this complex cannot be distin- guished from each other using external characters. Their determination depends on the examination of characters associated with the male genitalia. L. saccisecundaris is known from a single specimen collected from an undetermined locality in Georgia. L. unilobus has been col- lected at Carlisle, Arkansas, at Jacksonville, Florida, and at Lake Chicot State Park in Louisiana. It is also recorded from one or two undetermined localities in Texas. It was collected in June in Florida, L. taeniatus is known from eight localities. It has been collected in Alabama, Louisiana, Mississippi and South Carolina in the months of March, April, June and December. L. extendus has been collected from an undetermined locality in Arkansas and from two counties in Indiana. The Indiana specimens were collected in January and May. 18. Loxanclrus unilobus New Species Figs. 59; 123 A,B;172 Holotype. $. Arkansas. Lonoke Co.: Carlisle (MCZ). Paraypes. 1 $ (MCZ) ; 2 $ 2 (CU); 1 2 (CAS); 1 2 (UA); 1 $ (TLE). Description. Color. Dorsal: black, shining. Ventral: black to dark brown. Appendages: black to dark brown except palpi which are reddish brown. Head. Clypeus: anterior margin straight or slightly concave. Epistomal and subgenal sutures: obscure, complete. Frontal sulci: obscure, shallow, broad. Microsculpture: absent on the ante- rior part of the clypeus; isodiametric on the frons and vertex. Men turn: apex of lobes narrowly rounded; tooth not as long as lobes; furrows shallow; fovae absent; microsculpture isodiametric. Thorax. Pro- notum: microsculpture on disk weak, elongate; lateral margins evenly rounded, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus complete or not; posterior marginal sulcus reaching past basal fovae but incomplete; median sulcus reaching anterior -> Figs. 103-106. Male genitalia, IS everted. 103 A. L. nitidulus (LeC.), right lateral aspect. 103 B. Same, left lateral aspect. 103 C. Same, ventral aspect. 104. L. crenatus Chaud., dorsal aspect. 105. L.lucens Chaud., dorsal aspect. 106. L. saphyrinus (Chaud.), left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 77 106 L. SAPHYRINUS 78 Entomologica Americana [Vol. 46, No. 1 marginal sulcus or not, not reaching posterior margin; basal impres- sions elongate, moderately deep, punctate or not; area between im- pressions sparsely punctate or not; area between impressions and lateral margins if punctate only one or two punctures. Scutellum: microsculpture isodiametric; apex pointed. Prosternum: microsculp- ture elongate, distinct laterally, less so medially; impunctate. Proepi- sternum: microsculpture isodiametric; impunctate. Proepimeron: mi- crosculpture isodiametric and elongate; impunctate. Mesosternum: microsculpture slightly elongate; impunctate. Mesepisternum: micro- sculpture isodiametric; impunctate but a few weak, small ridge-like structures anteriorly. Metasternum: microsculpture elongate, more distinct laterally; weakly punctate laterally in some specimens. Met- episternum: microsculpture isodiametric; weakly and sparsely punc- tate in some specimens. Metepimeron: microsculpture elongate; impunctate. Legs. Hind coxae: microsculpture on inner surface elongate. Tarsal segments: 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture absent on intervals; all striae distinct, punctate, trough 1 isodiametric unit wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture elongate medially, isodiametric laterally; impunctate; segments 1-3 rugose laterally towards the base; ambulatory setae on segment 2. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position, slightly turned to the left side and directed dorsally; six distinct scale groups present; most of the remainder of the sac’s surface coverd with small weakly sclerotized scales; sclerotized ring present, opening in the left side. Measurements. (4 specimens). TL — 4.9 (5. 5-6. 4); WH— 1.2 (1. 1-1.2); WP— 1.8 ( 1. 7-1.9 ); WW— 2.3 (2.2-2.5). Distribution. Central Southern United States. Localities. Arkansas. Calhoun Co.: Bayou Moro, Hy. 160 (UA); Lonoke Co.: Carlisle (MCZ). Unrecorded localities in Arkansas (CU). Florida. Duval Co.: Jacksonville (CAS). Louisiana. Evangeline Co.: Lake Chicot State Park (TLE). Texas. Unrecorded localities (MCZ). Discussion. See above. 19. Loxandrus saccisecundaris New Species Figs. 54; 121 A, B, C Holotype. $. Georgia. Frederick Blanchard Collection (MCZ). Description. Color. Dorsal surface: black, shining; ventral sur- face: black except apex of segment 5 of abdomen which is dark brown. Allen: Revision of Loxandrus LeConte 79 1972] Figs. 107-108. Male genitalia, IS everted. 107. L. uniformis n. sp., left lateral aspect. 108 A. L. micans Chaud., dorsal aspect. 108 B. Same, left lateral aspect. 108 C. Same, right lateral aspect. 80 Entomologica Americana [Vol. 46, No. Fig. 109. Male genitalia, IS everted. 109 A. L. duryi Wright, left lat- eral aspect. 109 B. Same, dorsal aspect. 109 C. Same, right lateral aspect. 109 D. Same, apex of median lobe, left lateral aspect. 109 E. Same, apex of median lobe, ventral aspect. 109 F. Same, left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 81 Appendages: dark to light brown. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: complete. Frontal sulci: obscure, short, broad, shallow. Microsculpture: clypeus, frons and vertex isodiametric (sometimes weak). Mentum: apex of lobes narrowly rounded; tooth not as long as lobes; furrows and fovae absent; microsculpture slightly elongate. Thorax. Pronotum: micro- sculpture on disk present but weak, elongate, less elongate along lateral margins; lateral margins reflexed and evenly rounded their entire distance, not sinuate posteriorly, impunctate; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impressions but incomplete; median sulcus just reaching anterior marginal sulcus but not posterior margin; basal impressions moderately deep, elongate, punctate; area between im- pressions sparsely punctate; area between impressions and lateral margins impunctate. Scutellum: apex pointed; microsculpture iso- diametric. Prosternum: impunctate; microsculpture elongate, more distinct laterally. Proepisternum : impunctate; microsculpture iso- diametric. Proepimeron: impunctate; microsculpture isodiametric and slightly elongate. Mesosternum: impunctate; microsculpture slightly elongate. Mesepisternum : weakly punctate in anterior half; micro- sculpture elongate. Metasternum: punctate towards the lateral mar- gins; microsculpture elongate, more distinct laterally. Metepisternum : weakly punctate; microsculpture elongate. Metepimeron: impunctate; microsculpture elongate, weak. Legs. Hind coxae : microsculpture on inner surface elongate: tarsal segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture absent; all striae distinct, trough 1-2 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture isodiametric; segment 1 sparsely punctate laterally near the base; segments 1-5 rugose along the basal margin; ambulatory setae present on second abdominal sternite. Aedeagus. Median lobe: not modified. Internal sac: everts medially, slightly turned dorsally: 9 distinct scale groups are present; basal quarter of surface covered with small weakly sclerotized scales; sclerotized ring present, opening on the left side; a secondary sac everting near the midline on the left side. Measurements. (1 speci- men). TL— 7.4; WH— 1.4; WP— 2.4; WW- 3.1. Distribution and Locality. Known from a single specimen from an unidentified locality in Georgia (MCZ) . Discussion. See above. 82 Entomologica Americana fVol. 46, No. 1 20. Loxandrus extendus New Species Figs. 62; 131 A, B; 176 Holotype. $. Arkansas, undetermined locality. Strom. Frederick Blanchard Collection (MCZ). Paratypes. 3 $ S (PU). Description. Color. Dorsal: shining black. Ventral: mostly black, some areas dark brown. Appendages: dark brown. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: weak but complete. Frontal sulci: almost absent, short, broad, shallow. Microsculpture: clypeus, frons and vertex weak, isodiametric. Mentum: apex of lobes narrowly rounded; tooth not as long as lobes, apex rounded; furrows present; fovae obscure; micro- sculpture isodiametric, sometimes slightly elongate. Thorax. Pro- notum: microsculpture vaguely present on disk, elongate, isodiametric along lateral margins; lateral margins evenly rounded their entire length, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impressions but incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions moderately deep, elongate; area between basal impressions with only one puncture; area between basal impressions and lateral margins impunctate. Scu- tellum: apex pointed; microsculpture isodiametric. Prosternum: impunctate; microsculpture isodiametric and elongate laterally becom- ing very elongate or absent medially. Proepisternum: impunctate; mi- crosculpture isodiametric. Proepimeron: impunctate; microsculpture slightly elongate. Mesosternum: impunctate; microsculpture slightly elongate. Mesepisternum: weakly and sparsely punctate in the anterior half; microsculpture isodiametric to slightly elongate. Metasternum: weakly punctate towards lateral margin; microsculpture isodiametric to weakly elongate, more distinct toward lateral margin. Metepister- num: impunctate; microsculpture slightly elongate. Metepimeron: impunctate; microsculpture elongate. Legs. Hind coxae: microsculp- ture on inner surface elongate; tarsal segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculp- ture absent; all striae distinct, trough 1-2 isodiametric units wide; Figs. 110-111. Male genitalia. 110 A. L. vitiosus n. sp., left lateral aspect, IS everted. 110B. Same, right lateral view, IS everted. 110C. Same, ventral aspect. 1 1 1 A. L. major Dari., dorsal aspect. Ill B. Same, left lateral aspect, IS everted. 1972] Allen: Revision of Loxandrus LeConte 83 84 Entomologica Americana [Vol. 46, No. 1 internal plica well-developed; humeri rounded; microsculpture of epi- pleura absent. Abdominal sternites. Microsculpture distinct, isodia- metric laterally, becoming elongate to absent medially; segment 1 sparsely punctate basally and laterally; segments 1-4 sparsely rugose laterally; ambulatory setae present on second abdominal sternite. Aedeagus. Median lobe: not modified. Internal sac: everts medially, directed slightly dorsally and turned at the apex towards the left side; 7 scale groups present; sclerotized ring present; a large extension of the main sac at the apex; most of surface of the sac covered with small weakly sclerotized scales. Measurements. (1 specimen). TL — 7.5; WH— 1.4; WP— 2.3; WW— 3.0. Distribution. Central United States. Localities. Arkansas. Unrecorded locality (MCZ). Indiana. Dubois Co. (PU); Virgo Co. (PU). Discussion. See above. 21. Loxandrus taeniatus LeConte Figs. 60; 127 A, B; 175 Loxandrus taeniatus LeConte 1852:252. $ . Marked with a red type label in the MCZ collection. Type No. 5697. No data labels on this specimen but in the original description LeConte states that it is from Louisiana and was presented to him by Mr. Schaum. Description. Color. Dorsal: black, iridescent. Ventral: dark brown to black. Appendages : dark or light to reddish brown. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: fine but complete. Frontal sulci: feebly present, shallow, broad. Microsculpture: clypeus, frons and vertex, isodiametric. Men- tum: apex of lobes broadly rounded; tooth distinct, shorter than lobes, rounded at apex; furrows and fovae absent; microsculpture, isodia- metric but slightly elongate. Thorax. Pronotum: elongate microsculp- ture present on disk (distinct in most specimens), isodiametric along lateral margins; lateral margin evenly rounded, not sinuate posteriorly, impunctate except (sometimes) near basal angles; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus Figs. 112-114. Male genitalia. 112. L. rufilabris Cast., left lateral aspect, IS everted. 113 A. L. australiensis Sloane, left lateral aspect, IS everted. 113 B. Same, dorsal aspect, IS everted. 114 A. L. medius Dari., left lateral aspect. 114 B. Same, left lateral aspect, IS everted. 114 C. Same, dorsal aspect, sac everted. 1972] Allen: Revision of Loxandrus LeConte 85 86 Entomologica Americana [Vol. 46, No. 1 Figs. 115-119. Male genitalia, IS everted. 115. L. minor Chaud., right lateral aspect. 116. L. cincinnatiensis Casey, dorsal aspect, apex of median lobe. 117. L. minor Chaud., dorsal aspect, apex of median 1972] Allen: Revision of Loxandrus LeConte 87 reaching past the basal fovae but incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions elongate, moderately deep, punctate or not; area between basal im- pressions sparsely punctate; area between basal impressions and lateral margins usually impunctate. Scutellum: apex pointed; microsculpture isodiametric. Prosternum: impunctate; microsculpture where present, elongate. Proepisternum : impunctate; microsculpture isodiametric. Proepimeron: impunctate; microsculpture isodiametric and/or slightly elongate. Mesosternum: impunctate; microsculpture elongate. Mes- episternum: impunctate; microsculpture elongate. Mesepisternum : weakly punctate in groove only; microsculpture elongate. Metaster- num: sometimes sparsely and weakly punctate laterally; microsculp- ture more distinct laterally and anteriorly, elongate. Metepisternum : punctate (sometimes sparse and weak); microsculpture isodiametric. Metepimeron: impunctate; microsculpture absent. Legs. Hind coxae: microsculpture on inner surface obscure, elongate; tarsal segments 1-3 of middle and hind tarsi grooved dorso-laterally. Elytra. Micro- sculpture absent; all striae distinct, trough 1-2 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture isodia- metric laterally, elongate medially; impunctate; rugose on lateral-basal areas of segments 1-3. Aedeagus. Median lobe: not modified. In- ternal sac: everts medially; 5 distinct scale groups; most of the apical quarter of the sac covered by moderately large scales but not forming a distinct group; remainder of the surface covered with small weakly sclerotized scales; one area on the right side distinctly free of scales; sclerotized ring present. Measurements. (16 specimens). TL — 6.7 (6.2-7.5); WH— 1.3 (1.2— 1.4); WW— 2.7 (2.5-2.8). Distribution. Southern United States. Localities. Alabama. Mobile Co.: Calvert (CAS). Arkansas. Ar- kansas Co.: E. of Reydel, Hy. 11, Bayou Meto (UA). Georgia. Unre- corded localities (MCZ, UM, INHS) . Louisiana. Beauregard Co. : Rose- pine (UA); Evangeline Co.: Lake Chicot State Park (TLE); Iberia Co.: New Iberia (USNM); St. Martin Co. (UA). Mississippi. Hancock Co. (USNM). North Carolina. *Broadman (?). South Carolina. Sumpter Co.: Sumpter (GB). Discussion. See above. <- lobe. 118 A. L. robustus n. sp., left lateral aspect. 118 B. Same, right lateral aspect. 119. L. yeariani n. sp., right lateral aspect, apex of median lobe. 88 Entomologica Americana [Vol. 46, No. 1 Figs. 120-121. Male genitalia, IS everted. 120 A. L. parallelus Casey, left lateral aspect. 120 B. Same, right lateral aspect. 121 A. L. sac- cisecundaris n. sp., left lateral aspect. 121 B. Same, dorsal aspect. 121 C. Same, right lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 89 22. Loxandrus floridanus LeConte Figs. 17; 58; 122 A, B; 171 Loxandrus floridanus LeConte, 1878:376. $. Florida, Capron; 24-4; Type No. 5694 (MCZ). There are 7 additional specimens in the series, 3 males and 4 females. See discussion. Loxandrus scitus Casey, 1918:388. $ . Florida, Dunedin. Type No. 47351 (USNM) . One $ paratype. Description. Color. Dorsal: black, shining. Ventral: usually black, sometimes dark brown. Appendages: femur and tibia usually black, sometimes dark brown; tarsi, palpi and antennae dark to reddish brown. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: distinct, complete. Frontal sulci: obscure, shallow, broad. Microsculpture: clypeus, frons and vertex isodiametric. Mentum: apex of lobes narrowly rounded; tooth not as long as lobes; furrows shallow; fovae absent; microsculpture isodia- metric or slightly elongate. Thorax. Pronotum: microsculpture on disk elongate, distinct; lateral margins evenly rounded their entire distance, not sinuate or oblique posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impressions but incomplete; median sulcus reaching ante- rior marginal sulcus but not posterior margin; basal impressions elon- gate, shallow, punctate; area between basal impressions punctate; area between basal impressions and lateral margins punctate or not, when punctate sparsely so. Scutellum: microsculpture isodiametric; apex pointed. N. B. Entire ventral surface impunctate. Prosternum: micro- sculpture elongate, more distinct laterally. Proepisternum : micro- sculpture isodiametric. Proepimeron : microsculpture isodiametric and elongate. Mesosternum: microsculpture slightly elongate. Mes- episternum: microsculpture isodiametric. Metasternum: micro- sculpture elongate medially, isodiametric laterally. Metepistemum : microsculpture isodiametric. Metepimeron: microsculpture elongate, weak. Legs. Hind coxae: microsculpture on inner surface isodia- metric. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae except basal portion of 7 distinct, trough 2-3 some- times 4 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture elongate medially, isodiametric laterally; segment 1 weakly punctate basally or not, other segments impunctate; segments 1-4 weakly rugose laterally tov/ard the base; ambulatory setae present on segment 2. Aedeagus. Median lobe: not modified. Internal sac: everts medially and is directed dorsally; seven more or less distinct 90 Entomologica Americana [Vol. 46, No. 1 Figs. 122-123. Male genitalia, IS everted. 122 A. L. floridanus LeC., left lateral aspect. 122 B. Same, right lateral aspect. 123 A. L. unilobus n. sp., left lateral aspect. 123 B. Same, right lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 91 scale groups including those on the two secondary sacs present; two secondary sacs, one at the base on the dorsal surface and one toward the apex on the left lateral surface; sclerotized ring present, partially on the main sac and on the apical secondary sac. Measurements. (20 specimens). TL— 8.0 (7.3-9.4); WH— 1.6 (1. 5-1.9); WP— 2.5 (2.3— 2.8); WW— 3.2 (2.9-3.7). Distribution. Southern and Southeastern United States and West Indies. Localities, united states. Alabama. Mobile Co.: Delchamps (OSU, MCZ, USNM, CAS, GB); unrecorded locality (CU). Florida. Alachua Co.: Gainesville (CU); Charlotte Co.: Punta Gorda (CAS); Collier Co.: Everglade (USNM) ; Dade Co.: Biscayne (USNM), Biscayne Bay (AM), Hialeah (TLE), Homestead (MCZ); Duval Co.: Jackson- ville (CAS, OSU) ; Hillsborough Co.: Tampa (MCZ) ; Lucie Co.: Capron (old Fort on 1861 map) (USNM); Manatee Co.: Oneco (GB); Osceola Co.: Kissimmee (AM); Palm Co.: Lake Worth (AM); Pinellas Co.: Dunedin (PU); Sarasota Co.: Myakka River State Park (GB). ^Indian River (USNM). Unrecorded locality in Florida (MCZ). Louisiana. Calcasieu Co.: Sabine River Ferry (CU); Madison Co.: Tal- lulah (MCZ); Orleans Co.: New Orleans (MCZ, USNM, CAS); Pla- quemines Co.: Myrtle Grove (INHS). Mississippi. Hancock Co.: Waveland (USNM); Jackson Co.: Ocean Springs (CU), Horn Island (CU). *Gulf View (USNM). Texas. Comal Co.: New Braunfels (CAS); Orange Co. (TAM). Unrecorded locality in Texas (MCZ). WEST INDIES. South Bimini Island (AM). Discussion. This species was described by LeConte (1878) from Capron and Enterprise, Florida. The material in the LeConte col- lection (MCZ) consists of the female type and seven additional speci- mens. The last specimen in the series is a female from Louisiana and was probably not part of the original type material. The three males in the series are all the same species. For this reason I am considering this a valid species. The males and females appear to be distinct and can be identified by the characters given in the key. Too much reliance should not be placed on external characters in the females, however, and exact deter- minations should be based on examination of the male aedeagus. Specimens have been collected in February, March, April, May, June, July, October, November and December. 23. Loxandrus agilis (Dejean 1828) Figs. 13; 20 A, B, C; 21; 56; 125; 173 Feronia agilis Dejean 1828:244. Lectotype (here selected) . $. The first in a series of 5 specimens in the Chaudoir Collection. It is the only one 92 Entomologica Americana [Vol. 46, No. 1 marked with a green Dejean determination label and is recorded from Boreal America (PMNH) . Loxandrus flavilimbus Blatchley, 1918:418. Type. $. Collected north of Dunedin, Florida, under a board near the border of a pond on January 1 (PU). Description. Color. Dorsal: black, shining. Ventral: black to dark brown. Appendages: dark to reddish brown. Head. Clypeus: anterior margin straight. Epistomal and subgenal sutures: distinct, complete. Frontal sulci: obscure, broad, shallow. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes rounded; tooth distinct, not as long as lobes; furrows shallow; fovae, if present, shallow, obscure; microsculpture isodiametric. Thorax. Pronotum: microsculpture distinct, usually isodiametric, sometimes slightly elongate; lateral margins evenly rounded, not sinu- ate posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impressions but incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions shallow, elongate, punctate or not; area between basal impressions punctate; area between impres- sions and lateral margins punctate or not. Scutellum: apex pointed; microsculpture isodiametric. Prosternum: impunctate; microsculpture elongate medially, isodiametric laterally. Proepisternum: impunctate, microsculpture distinct, isodiametric. Proepimeron: impunctate; microsculpture isodiametric and elongate. Mesosternum: impunctate; microsculpture feebly elongate. Mesepisternum : impunctate; micro- sculpture isodiametric. Metasternum: impunctate; microsculpture elongate medially, isodiametric laterally. Metepisternum : impunctate; microsculpture isodiametric. Metepimeron: impunctate; microsculp- ture distinct, isodiametric or feebly elongate. Legs. Hind coxae: microsculpture on inner surface isodiametric. Tarsi: segments 1-4 of hind tarsi and 1-3 of middle tarsi grooved dorso-laterally. Elytra. Microsculpture sometimes weakly present on the intervals, if so then it is very elongate; all striae distinct, trough 1 sometimes 2 isodiametric units wide; lateral margin distinctly sinuate apically; internal plica distinct; humeri rounded; microsculpture of epipleura usually present, elongate. Abdominal sternites. Microsculpture distinct, elongate me- dially, isodiametric laterally; impunctate; segments 1-3 rugose on the lateral and basal surfaces; ambulatory setae sometimes present on seg- ment 2. Aedeagus. Median lobe: compressed dorso-ventrally; right lateral margin strongly sinuate apically; apex turned toward the right side, pointed. Internal sac: four distinct scale groups present; entire right side and most of the ventral surface of the sac covered with 1972] Allen: Revision of Loxandrus LeConte 93 dorsal aspect. 94 Entomologica Americana [Vol. 46, No. 1 elongate scales that do not form a well defined group; remainder of the surface covered with small, weakly sclerotized scales; secondary sac everts from the dorsal surface of the membranous portion of the main lobe, scale group at its apex; sclerotized ring present. Measurements. TL— 9.1 (8.0-9. 8); WH— 1.8 ( 1.6-1. 8) ; WP— 2.9 (2.5-3. 1);WW —3.6 (3. 2-3. 8). Distribution. Southeastern United States west to Texas. Localities. Florida. Alachua Co. (FSPB); Charlotte Co.: Punta Gorda (CAS); Desoto Co.: Arcadia (GB); Duval Co.: Mayport (FSPB), Jacksonville (CAS, OSU); Highlands Co.: Archbold Biological Station (CU, PSU), Brighton (MCZ), Highlands Hammock (TLE); Hills- borough Co.: Tampa (MSU, USNM, CU); Manatee Co.: Oneco (PU, GB); Orange Co.: Winter Park (MCZ), Titusville (CNC); Pinellas Co.: Dunedin (PU); Putnam Co.; Crescent City (USNM); Volusia Co.: Enterprise (MSU), Sanford (MCZ), Lake Helen (CAS). ^Indian River (USNM). Unrecorded locality in Florida (CAS). Texas. *Mitten (MCZ). Discussion. The males and females of this species can be identified using external characters. The male aedeagus should be examined, however, when possible. This species has been collected in the months of February, March, April, May, June, August, and December. The L. aduncus , L. cervicalis , L. parallelus , L. taeniatus and L. spinilunatus Complex Discussion. These five species closely resemble each other in all external characters. They cannot be separated from one another except on the basis of characters associated with the male aedeagii. L. paral- lelus, described by Casey from Monroe, Louisiana, appears to have a more western distribution than the other four species under discussion. It has been collected in the months of April, June, July and August. L. cervicalis is known from several localities in Florida and one locality in Arkansas. It has been collected in January, March, August and October. L. aduncus is presently known from four localities in the Southeastern United States. Collecting data indicate this species being collected in January, February, May and June. The external charac- ters of L. taeniatus are somewhat variable and the species is keyed out in several different places in the key. It is apparently a southern form although not reaching into Florida, and has been collected in the months of March, April, June and December. L. spinilunatus is known from only two male specimens from undetermined localities in Arkansas and Louisiana. There were no collecting data with the speci- mens indicating at what time of year they were taken. 1972] Allen: Revision of Loxandrus LeConte 95 Figs. 126-128. Male genitalia, IS everted. 126 A. L. rossi n. sp., right lateral surface. 126 B. Same, left lateral surface. 127 A. L. taeni- atus LeC., right lateral aspect. 127 B. Same, left lateral aspect. 128. L. minutus n. sp., left lateral aspect. 96 Entomologica Americana [Vol. 46, No. 1 24. Loxandrus aduncus New Species Figs. 57; 124 A, B; 174 Holotype. $. Florida. Winter Park; John George Gehring Collection (MCZ) . Paratypes. 2 $ $ (MCZ); 1 $ (TLE) ; 3 S $ (CU). Description. Color. Dorsal: black, shining. Ventral: black to dark brown. Appendages: dark to reddish brown. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal suture: distinct, complete. Frontal sulci: obscure, short, broad, shallow. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes narrowly rounded; tooth distinct, slightly shorter than lobes; furrows shallow; fovae absent; microsculpture iso- diametric. Thorax. Pronotum: microsculpture distinct, usually isodia- metric but sometimes weakly elongate; lateral margins sparsely punctate in posterior quarter or less, evenly rounded their entire length, not sin- uate; posterior angles broadly rounded; anterior marginal sulcus com- plete; posterior marginal sulcus reaching past basal impressions but incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions moderately deep, elongate, punc- tate; area between basal impressions punctate; area between basal impressions and lateral margin sparsely punctate, most punctures confined to the posterior margin. Scutellum: apex pointed; mi- crosculpture isodiametric. Prosternum: microsculpture elongate; impunctate. Proepisternum and Proepimeron: microsculpture iso- diametric; impunctate. Mesosternum: microsculpture elongate; impunctate. Mesepisternum : microsculpture isodiametric; impunctate. Metasternum : microsculpture elongate medially, isodiametric laterally; impunctate. Metepisternum : microsculpture isodiametric; impunc- tate. Metepimeron: microsculpture elongate; impunctate. Legs. Hind coxae: microsculpture on inner surface elongate. Tarsi: seg- ments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso- laterally. Elytra. Microsculpture absent; all striae distinct, trough 1-2 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture distinct, elongate medially, isodiametric laterally; im- punctate; base of segments 1-3 rugose, ambulatory setae sometimes present on second abdominal sternite. Aedeagus. Median lobe: apex forming a hook that is turned dorsally. Internal sac: three distinct scale groups are present on the main body of the sac; most of the remainder of the sac’s surface is covered with small weakly sclerotized scales; a secondary sac everts on the dorsal membranous portion of the median lobe, the apex is covered with a scale group; sclerotized 1972] Allen: Revision of Loxandrus LeConte 97 ring present. Measurements. (7 S specmens). TL — 8.2 (7. 8-8. 4); WH— 1.8 (1.7-1. 8); WP— 2.4 (2.2-2.6); WW— 3.2 (3.0-3. 3). Distribution. Extreme Southeastern United States. Localities. Alabama. Mobile Co.: Mobile (MCZ). Florida. High- lands Co.: Archbold Bio. Sta. (CU), Highlands Hammock (TLE); Orange Co. : Winter Park ( MCZ ) . Discussion. See above. 25. Loxandrus parallelus Casey Figs. 4; 55; 120 A, B; 170 Loxandrus parallelus Casey 1918:386. Holotype. S. Type No. 47342. USNM. Casey states in the original description that he collected “a single example” at Monroe, Louisiana. Description. Color. Dorsal: black, shining; elytra iridescent. Ventral: dark brown to black, shining. Appendages: dark to light brown or straw yellow. Head. Clypeus : anterior margin slightly con- cave. Epistomal and subgenal sutures distinct, complete. Frontal sulci: obscure, short, shallow, broad. Microsculpture: clypeus, frons, and vertex isodiametric. Mentum: apex of lobes rounded; tooth dis- tinct, shorter than lobes; furrows shallow; fovae absent; microsculp- ture slightly elongate. Thorax. Pronotum: microsculpture distinct, elongate; lateral margins evenly rounded their entire length, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus com- plete; posterior marginal sulcus reaching past basal impressions but not complete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions moderately deep, elongate, punc- tate; area between impressions punctate; area between impressions and lateral margins punctate or not. Scutellum: apex pointed; micro- sculpture isodiametric. Prosternum: microsculpture elongate, becom- ing more distinct laterally; impunctate. Proepisternum, proepimeron, mesosternum and mesepimeron: microsculpture isodiametric; im- punctate. Metasternum : microsculpture elongate laterally and around middle coxae, isodiametric in anterior median portion; impunctate. Metepisternum : microsculpture isodiametric; impunctate. Metepi- meron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface distinct, elongate. Tarsal segments: 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-later- ally. Elytra. Microsculpture absent; all striae distinct, trough 1-2 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculp- ture elongate medially, isodiametric laterally; impunctate; segments 1-3 rugose at the base and 1-4 rugose laterally. Aedeagus. Median 98 Entomologica Americana [Vol. 46, No. 1 lobe: not modified. Internal sac: everts in a median position and deflected downward; six distinct scale groups present; remainder of the surface of the primary sac is covered with very small, lightly sclerotized scales; a secondary sac everts on the left side; sclerotized ring present. Measurements. (20 specimens). TL — 6.9 (6. 5-7. 4); WH — 1.3 ( 1. 2-1.4 ); WP— 2.1 (1. 9-2.3); WW— 2.8 (2.5-3.0). Distribution. Mid-southern United States west to Texas. Localities. Arkansas. Greene Co. : Paragould (MSU); Hempstead Co.: Hope (MCZ). Kentucky. No locality (AM). Louisiana. Caddo Co.: Shreveport (UA); East Baton Rouge Co.: Baton Rouge (UA); Evangeline Co.: Lake Chicot State Park (TLE). Texas. Brazos Co.: College Station (TAM). Navarra Co.: Corsicana (USNM). Discussion. See above. 26. Loxandrus cervicalis Casey Figs. 63; 130 A, B, C; 176 Loxandrus cervicalis Casey, 1918:385. Holotype. $. One male specimen from Sarasota, Florida. Type No. 47340. (USNM). Loxandrus brunneus Blatchley, 1918:417. Holoype. S. Type locality, Lake Okeechobee, four miles southeast of Moore Haven, March 1-3 (PU). Description. Color. Dorsal: black, shining, elytra iridescent. Ventral: black to dark brown. Appendages: mostly black to dark brown but sometimes straw yellow. Head. Clypeus: anterior margin usually straight, sometimes very slightly concave. Epistomal and sub- genal sutures distinct, complete. Frontal sulci: short, broad, shallow. Microsculpture: clypeus, frons and vertex isodiametric. Mentum: apex of lobes narrowly rounded; tooth not as long as lobes; furrows and fovae absent; microsculpture isodiametric. Thorax. Pronotum: microsculpture distinct, elongate; lateral margins evenly rounded their entire length, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching just past basal fovae but incomplete; median sulcus present, reaching anterior sulcus but not posterior margin; basal impressions elongate, sparsely punctate; area between basal impressions sparsely punctate; area between basal impressions and lateral margins impunctate. Scutel- lum: apex pointed; microsculpture isodiametric. N. B. Ventral surface of thorax impunctate. Prosternum: microsculpture elongate medially, becoming isodiametric towards the lateral margins. Proepisternum : microsculpture isodiametric. Proepimeron: microsculpture elongate. Mesosternum: microsculpture elongate. Mesepisternum : microsculp- ture elongate and isodiametric. Metasternum: microsculpture elongate or absent in median areas, isodiametric towards the lateral margins. 1972] Allen: Revision of Loxandrus LeConte 99 right lateral aspect. 129 B. Same, dorsal aspect. 129 C. Same, ven- tral aspect. 130 A. L. cervicalis Casey, right lateral aspect. 130 B. Same, left lateral aspect. 130 C. Same, ventral aspect. 100 Entomologica Americana [Vol. 46, No. Metepisternum : microsculpture isodiametric. Metepimeron: micro- sculpture if present elongate. Legs. Hind coxae: microsculpture on inner surface elongate. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture ab- sent; striae 7 weakly impressed basally, other striae all distinct, trough 1 isodiametric unit wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculp- ture elongate medially, becoming isodiametric laterally; impunctate; segments 1 and 2 weakly rugose at base; ambulatory setae sometimes present on segment 2. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position; six distinct scale groups present; re- mainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present. Measurements. (17 specimens). TL— 6.5 (6. 2-7. 2) ; WH-1.3 (1.2— 1.5); WP— 2.0 (1. 9-2.2); WW —2.6 (2. 5-2. 9). Distribution. Southeastern United States, Florida to Arkansas. Localities. Arkansas. Lonoke Co.: Carlisle (CAS). Florida. De- Soto Co.: Arcadia (GB); Duval Co.: Jacksonville (OSU); Hernando Co.: Brooksville (CAS); Highland Co.: Archbold Biological Station, Lake Placid (CU, PSU); Hillsborough Co.: Tampa (USNM); Indian River Co.: Sebastian River (USNM); Levy Co.: No locality (FSPB); Pinellas Co.: Bellair (AM, USNM); Dunedin (PU); Sarasota Co.: Sarasota (CAS); Volusia Co.: New Smyrna (CAS). *Sand Pt. (USNM). Discussion. See above. 27. Loxandrus spinilunatus New Species Figs. 64; 132 A, B Holotype. $ . Arkansas. Collector Stromb?; Frederick Blanchard Collec- tion (MCZ). Paratype. 1 $ (UM). Description. Color. Dorsal surface: black, iridescent; ventral surface: dark brown to black; appendages: dark to reddish brown. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: obscure but complete. Frontal sulci: obscure, short, broad, shallow. Microsculpture: clypeus, frons and vertex, sometimes weak, isodiametric. Mentum: apex of lobes narrowly rounded; tooth almost as long as lobes (mentum transverse); furrows absent; deep fovae present; microsculpture, where present, elongate. Thorax. Pronotum: microsculpture on disk distinct, slightly elongate; lateral margins evenly rounded, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal fovae but not complete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal 19721 Allen: Revision of Loxandrus LeConte 101 Figs. 131-132. Male genitalia, IS everted. 131 A. L. extendus n. sp., right lateral aspect. 131 B. Same, left lateral aspect. 132 A. L. spi- nilunatus n. sp., right lateral aspect. 132 B. Same, left lateral aspect. 102 Entomologica Americana [Vol. 46, No. 1 impressions elongate, moderately deep, punctate; area between impres- sions sparsely punctate; area between impressions and lateral margins impunctate. Scutellum: microsculpture isodiametric; apex pointed. N. B. Entire ventral surface impunctate. Prosternum: microsculpture elongate, more distinct laterally. Proepisternum : microsculpture iso- diametric. Proepimeron: microsculpture isodiametric and elongate. Mesosternum: microsculpture elongate. Mesepisternum : microsculp- ture isodiametric. Metasternum: microsculpture isodiametric laterally, elongate medially. Metepisternum : microsculpture isodiametric. Metepimeron: microsculpture, elongate. Legs. Hind coxae: micro- sculpture on inner surface elongate. Tarsal segments: 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae (except 7 toward base) distinct, punctate, trough 1 (sometimes 2) isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epi- pleura absent. Abdominal sternites. Microsculpture elongate me- dially, isodiametric laterally; impunctate; segments 1-4 rugose laterally near the base; ambulatory setae present on segment 2 (in one speci- men). Aedeagus. Median lobe: not modified. Internal sac: six distinct scale groups arranged in complex manner; most of the remain- der of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present. Measurements. (2 specimens). TL — 7.5 (7.1-7.8); WH— 1.4 (1.3— 1.5); WP— 2.4 (2.2-2.5); WW— 2.9 (2. 7-3.0). Distribution. Central Southern United States. Localities. Arkansas. Undetermined locality. (MCZ). Louisiana. Unrecorded locality (UM). Discussion. See above. 28. Loxandrus nocticolor Darlington Fig. 135; 144 Loxandrus nocticolor Darlington 1934:91. Holotype. $. Cuba, Soledad (near Cienfuegos), October 21, 1926. P. J. Darlington, Jr., Coll. Type No. 19512 (MCZ). Description. Color. Dorsal: black, shining. Ventral: reddish- brown to black, shining. Appendages: black to reddish brown, some- Figs. 133-134. Male genitalia, IS everted, basal 34 to % of median lobe not illustrated. 133 A. L. algidus n. sp., left lateral aspect. 133 B. Same, right lateral aspect. 133 C. Same, ventral aspect. 134 A. L. velox (Dej.), right lateral aspect. 134 B. Same, left lateral aspect. 1972] Allen: Revision of Loxandrus LeConte 103 I34B 104 Entomologica Americana [Vol. 46, No. 1 times tinged with yellow. Head. Clypeus: anterior margin, almost straight. Epistomal and subgenal suture: complete, weak laterally. Frontal sulci: obscure, broad, shallow. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes rounded; tooth distinct, apex rounded; furrows and fovae absent; micro- sculpture distinct, slightly elongate. Thorax. Pronotum: microsculp- ture present and isodiametric along the reflexed lateral margins and in the basal impressions, otherwise absent; lateral margins evenly rounded their entire distance, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus incomplete; posterior marginal sulcus reaching only to basal impressions; median sulcus not reaching anterior or pos- terior margins; basal impressions moderately deep, broad, elongate, punctate; area between impressions sparsely punctate; area between impressions and lateral margins with a few punctures at most. Scutel- lum: microsculpture weak, isodiametric; apex pointed. N. B. Entire ventral surface impunetate. Prosternum: microsculpture elongate me- dially, isodiametric laterally. Proepisternum and proepimeron: micro- sculpture isodiametric and elongate. Mesosternum: microsculpture slightly elongate. Mesepisternum : microsculpture isodiametric ante- riorly becoming elongate posteriorly and absent in posterior half to quarter. Metasternum and Metepisternum : microsculpture absent or very elongate. Metepimeron: microsculpture absent. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-4 of middle and hind tarsi grooved on outside lateral margin only. Elytra. Microsculpture on intervals absent; all striae distinct, impunetate, trough 3-4 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture on epipleura absent. Abdominal s termites. Microsculpture elongate medially, isodiametric laterally; im- punctate; segments 1-4 sparsely rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: no distinct scale groups present, much of the sac’s surface covered with small, weakly sclerotized scales; sclerotized ring absent. Measurements. (12 specimen). TL — 10.6; WH— 2.0; WP— 3.0; WW— 4.1. Distribution. West Indies. Localities, west indies. Cuba. Soledad (near Cienfuegos). Discussion. This species was originally described by Darlington in 1934. I have examined the holotype and found it to be a valid -> Figs. 135-137. 135. Male genitalia, IS everted, dorsal aspect, L. nocti- color Dari. Figs. 136-137. Female genitalia, ventral aspect. 136. L. rectus (Say). 137. L. pravitubus n. sp. 1972] Allen: Revision of Loxandrus LeConte 105 137 L. PRAVITUBUS 106 Entomologica Americana [Vol. 46, No. 1 species. The above description is based on a female given to me by Dr. Darlington. The external characters of this species appear to be rather distinct. The L. rossi, L. algidus, L. parallelus , L. unilobus , L. velox and L. taeniatus Complex These six species closely resemble one another and canot be dis- tinguished using external characters. Identification is based on charac- ters associated with the male genitalia. Loxandrus parallelus , L. unilobus and L. taeniatus have been keyed out in other places in the key and discussed previously. The description of L. algidus is based on two specimens collected in Maryland from an undetermined local- ity. There were no data with the specimens indicating at what time of the year they were collected. The description of L. rossi is based on seven specimens from Arkansas, Georgia, and Alabama. It has been collected in June at all three localities. L. velox was described by Dejean in 1828 from specimens sent to him by LeConte. The type locality was given only as “Amer. Sept.” The specimens in the Paris Museum are the only ones belonging to this species I have seen. There are 2 males and 2 females in the series. One of the females is recorded from Louisiana. The description used in this paper is based on notes taken on the Paris Museum type. 29. Loxaiulrus rossi New Species Figs. 61; 126 A, B; 174 Holotype. $ . Arkansas. Greene County, Paragould, J. Eichemeri, Col- lector, June 14, 1958, (MSU). Paratypes. 2 $ S (CAS); 1 $ (USNM); 1 $ (MCZ) ; 1 $ (TLE) ; 1 S (UA). Description. Color. Dorsal: black, shining. Ventral: dark to reddish brown. Appendages : dark to reddish brown. Head. Clypeus : anterior margin slightly concave. Epistomal and subgenal sutures: obscure but complete. Frontal sulci: almost absent, broad, shallow. Microsculpture: clypeus, fovae and vertex isodiametric. Mentum: apex of lobes rounded; tooth not as long as lobes; furrows and fovae ab- sent; microsculpture slightly elongate. Thorax. Pronotum: microsculp- ture on disk elongate; lateral margin evenly reflexed the entire distance, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impres- sions but incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions elongate, moderately deep, punctate; area between basal impressions sparsely punctate; area between basal impressions and lateral margins usually impunctate. 1972] Allen: Revision of Loxandrus LeConte 107 138 L. MICANS 140 L. DURYI 139 L. PACTINULLUS Figs. 138-143. Mentum. 138. L. micans Chaud. 139. L. pactinullus n. sp. 140. L. duryi Wright. 141. L. infimus Bates. 142. L. saphy- rinus (Chaud.). 143. L. minutus n. sp. 108 Entomologica Americana [Vol. 46, No. Scutellum: microsculpture isodiametric; apex pointed. N. B. Entire ventral surface impunctate. Prosternum : microsculpture where present elongate. Proepisternum: microsculpture elongate and isodiametric. Mesosternum: microsculpture slightly elongate. Mesepisternum : microsculpture elongate. Metasternum, Metepisternum and Metepi- meron: microsculpture elongate. Legs. Hind coxae: microsculpture on inner surface elongate. Tarsal segments: 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculp- ture absent on intervals; all striae distinct, punctate, trough 1 isodia- metric unit wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculp- ture elongate medially, isodiametric laterally; impunctate; segments 1-4 rugose laterally near the base; ambulatory setae present on segment 2. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position; directed dorsally; four distinct scale groups present; most of the remainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present, opens on the right side. Measurements. (8 specimens). TL — 6.9 (6. 7-7.2); WH — 1.3 (1.2— 1.4); WP— 2.2 (1.7-1.9); WW— 2.7 (2.6-2.9). Distribution. Southern United States. Localities. Alabama. *Tumblin Gap (CAS, MCZ, USNM). Ar- kansas. Greene Co.: Paragould (MSU). Georgia. Bibb Co.: 5 mi. SE. Macon (TLE). Discussion. See above. 30. Loxandrus algidus New Species Figs. 66; 133 A, B, C; 176 Holotype. $. Maryland. Otto Lugger Collection (UM). Paratypes. 1 S (UM). Description. Color. Dorsal: black, shining, elytra iridescent. Ventral: black to dark brown. Appendages: dark to light brown, sparsely tinged with black. Head. Clypeus: anterior margin slightly concave. Epistomal suture: distinct, complete. Subgenal suture: obscure but complete. Frontal sulci: obscure, shallow, broad, short. Microsculpture: clypeus, frons and vertex isodiametric. Mentum: apex of lobes narrowly rounded; tooth distinct, not as long as lobes; furrows more or less absent; fovae absent; microsculpture slightly elongate. Thorax. Pronotum: microsculpture absent or very weak, -> Figs. 144-146. Geographical distribution maps. 144. L. infimus Bates, L. nocticolor Dari. 145. L. elongatus n. sp. 146. L. rectangulus LeC. 1972] Allen: Revision of Loxandrus LeConte 109 110 Entomologica Americana [Vol. 46, No. 1 slightly elongate if visible; lateral margins evenly rounded their entire length, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impressions but incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions elongate, moderately deep, broad, punctate; area between basal impressions sparsely punc- tate; area between impressions and lateral margins impunctate. Scutel- lum: microsculpture isodiametric; apex pointed. N. B. Entire ventral surface impunctate. Prosternum: microsculpture absent medially, elongate laterally. Proepisternum and proepimeron: microsculpture elongate. Mesosternum: microsculpture absent or very elongate. Mesepisternum : microsculpture isodiametric anteriorly, elongate me- dially. Metasternum : microsculpture absent medially, weakly present, elongate, laterally. Metepisternum: microsculpture absent or very elongate. Metepimeron: microsculpture absent. Legs. Hind coxae: microsculpture on inner surface absent or very elongate. Tarsi: segments 1-3 of middle and hind tarsi and sometimes 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae, except basal quarter of 7, distinct, punctate, trough 1 (some- times 2) isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal s termites . Microsculpture elongate medially, becoming more isodiametric later- ally; impunctate; segments 1-4 weakly rugose laterally towards the base; segment 2 with ambulatory setae. Aedeagus. Median lobe: not modified. Internal sac: everts medially, only slightly directed dorsally; five distinct scale groups present; most of the remainder of the sac’s surface covered with small weakly sclerotized scales; a secondary sac everts from the left lateral side; sclerotized ring present. Measure- ments. (2 specimens). TL — 6.2 (6. 0-6. 4); WH — 1.3 (None); WP— 2.0 (1.9-2.0); WW— 2.5 (none). Distribution. Eastern United States. Localities. Known from one undetermined locality in Maryland (UMIN) . Discussion. See above. 31. Loxandrus crenatus LeConte Figs. 40; 104; 162 Loxandrus crenatus LeConte 1852:252. S. No locality label. MCZ. Type No. 5701 (MCZ). Four additional specimens in the MCZ series from Tampa, Capron and Enterprise, Florida. Description. Color. Dorsal: reddish-brown, shining. Ventral: reddish-brown, shining. Appendages : reddish-brown to straw yellow. 1972] Allen: Revision of Loxandrus LeConte 111 Head. Clypeus: anterior margin straight. Epistomal suture: com- plete. Subgenal suture: usually present, sometimes obscure or absent. Frontal sulci: obscure or absent, broad, shallow; anterior tentorial pits in this species deep. Microsculpture: clypeus, frons and vertex weak, isodiametric. Mentum: apex of lobes pointed or narrowly rounded; tooth distinct, almost as long as lobes; furrows shallow; fovae deep, distinct; microsculpture absent or very elongate. Thorax. Pro- notum: microsculpture absent; lateral margins weakly sinuate poste- riorly; posterior angles acute, slightly turned outward; anterior mar- ginal sulcus absent; posterior marginal sulcus reaching only to basal impressions; median sulcus not reaching anterior or posterior margins; basal impressions moderately deep, elongate, punctate; area between basal impressions with only 1-2 punctures if any; area between basal impressions and lateral margin impunctate. Scutellum: micro- sculpture absent; apex pointed. N. B. Entire ventral surface without microsculpture. Prosternum, proepistemum, proepimeron: impunc- tate. Mesosternum: impunctate. Mesepisternum: coarsely punctate in anterior quarter. Metasternum: coarsely punctate laterally. Met- episternum: coarsely punctate. Metepimeron: impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi grooved dorso-laterally. Elytra. Micro- sculpture on intervals absent; all striae distinct, distinctly punctate, microsculpture in trough usually absent, if present 1-2 units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture absent; basal margin of segment 1 punctate; segments 1-4 rugose laterally or not. Aedeagus. Median lobe: not modified. Internal sac: everts medially, turned to the left side and coiled at the apex; 1 small distinct scale group present; sclerotized ring is questionable, much of the apex appears to be sclerotized, the sclerotized ring seems to be in this area; most of the remainder of the sac’s surface covered with small weakly sclerotized scales. Measurements. (15 specimens). TL — 5.8 (4.9— 6.2); WH— 1.2 (1.0-1.3); WP— 1.8 (1. 5-1.9); WW— 2.3 (2.0- 2.5). Distribution. Southern United States. Localities. Alabama. Mobile Co.: Mobile (CAS, USNM), Spring Hill (CAS). Florida. Duval Co.: Jacksonville (CAS); Highlands Co.: Archbold Biological Station at Lake Placid (CU, PSU), Highlands Ham- mock (TLE); Hillsborough Co.: Tampa (USNM); Lucie Co.: Capron (Ft. Capron on old maps?) (USNM); Manatee Co.: Oneco (GB); Pinellas Co.: Dunedin (PU); Polk Co.: Lakeland (AM); Sarasota Co.: Myakka River State Park (TLE); Taylor Co.: Steinhatchee (USNM); Volusia Co.: Coronado Bch. (CU), Enterprise (MSU). *Sand Point 112 Entomologica Americana [Vol. 46, No. 1 (USNM) . *HawCrk. (USNM). * Royal Palm (PU). Unrecorded local- ities (MCZ, MSU, USNM). Georgia. Decatur Co.: Brawbridge (CAS). Louisiana. Vernon Co.: Rosepine (UA). Unrecorded locality (UM). Mississippi. Perry Co.: Richton (CU). South Carolina. Florence Co.: Rainwater (USNM). No state, no locality (MCZ). Discussion. This is one of the more distinct species of smaller Loxandrus. Once seen it is difficult to confuse it with other species in the genus. Its highly shining surface, distinct punctate striae and brown color are characteristic. It has been collected in the months of February, March, April, May, June, August, September and Novem- ber. The specimens from Rosepine, Louisiana, were collected in a black light trap. The L. micans , L. vitiosus and L. duryi Complex This is another group of closely related Loxandrus species. The external characters are very similar and identification must be based on characters associated with the aedeagi of the males. Even these char- acters show a great deal of resemblance. Loxandrus micans was described by Chaudoir from specimens from Opelousas, Louisiana. Two additional localities, Memphis, Tennessee, and the White River- Interstate 40 highway intersection, Arkansas, are known for this species. It has been collected in March, May and December. L. vitiosus is known from a number of localities throughout the Eastern United States. It has been collected in the months of January, March, April, June, July, September and December. L. duryi was described by Wright from Clermont Co., Ohio. Present data indicate that there is also a distinct population in Louisiana and Texas. It has been col- lected in the months of January, February, March, May, June, July and August. 32. Loxandrus micans Chaudoir Figs. 39; 108 A,B,C; 165 Loxandrus micans Chaudoir 1868:342. Holotype. $. The first specimen in the series of three bears a red type label plus a small data label. The specimen is from Opelousas, Louisiana, taken here to be the type locality. Description. Color. Dorsal surface: black or dark brown, shining; sometimes with central orange spot apically on the elytra; Figs. 147-148. Geographical distribution maps. 147. L. celeris (Dej.). 148. L. cubanus Tschit., L. remotus n. sp. 1972] Allen: Revision of Loxandrus LeConte 113 14 Entomologica Americana [Vol. 46, No. 1 elytra iridescent. Ventral surface: black or dark brown. Appendages: dark to light brown. Head. Clypeus: anterior margin slightly con- cave. Epistomal suture : obscure medially, absent laterally. Subgenal suture: obscure. Frontal sulci: obscure, shallow, broad. Microsculp- ture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes rounded; tooth large, broadly rounded at apex, almost as long as lobes; furrows absent; fovae distinct, deep; microsculpture isodia- metric and elongate. Thorax. Pronotum: microsculpture absent on the disk, weakly present laterally; lateral margins evenly rounded their entire length, not sinuate posteriorly, weakly punctate in posterior quarter to half; posterior angles rounded; anterior marginal sulcus incomplete; posterior marginal sulcus reaching past basal im- pressions but incomplete; median sulcus present but not reaching anterior or posterior margins; basal impressions moderately deep, elongate, narrow, punctate; area between basal impressions sparsely punctate; area between basal impressions and lateral margins with only 1-2 punctures. Scutellum: apex pointed; microsculpture isodiametric. Prosternum: microsculpture absent medially, weakly present laterally; impunctate. Proepisternum and proepimeron: microsculpture absent; impunctate. Mesosternum: microsculpture isodiametric; impunctate. Mesepisternum: microsculpture absent; anterior three quarters punc- tate. Metastemum: microsculpture absent except around middle coxae; punctate toward lateral margin. Metepisternum : microsculp- ture absent; punctate. Metepimeron: microsculpture absent; impunc- tate. Legs, Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi grooved dor so-later ally. Elytra. Microsculpture on intervals absent; all striae distinct, punctate, trough 1-2 isodiametric units wide or totally absent; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture weak, elongate; segment 1 punctate along basal margin; segments 1-5 rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts medially, slightly directed dorsally: 3 distinct scale groups and possibly a fourth on the main sac; a secondary sac protrudes from the left side of the main sac; secondary sac with a scale group at its apex; sclerotized ring present on the left dorso-lateral surface. Measurements. (3 specimens). TL —6.3 (6.1-6.5); WH— 1.3 (none); WP— 2.1 (2.0-2.1); WW- 2.6 (none). Distribution. Eastern United States. Localities. Arkansas. Prairie Co.: Rest stop on Interstate 40 at White River bridge (UA). Tennessee. Shelby Co. : Memphis (USNM). Discussion. See above. 1972] Allen: Revision of Loxandrus LeConte 115 33. Loxandrus vitiosus New Species Figs. 41; 110 A, B,C; 167 Holotype. $ . Arkansas. Howard Co., 4 mi. W. Mineral Springs, Rte. 27, July 12, 1968. River bottom debris, R. T. Allen and J. W. Stewart (UA). Paratypes. 5 S S (CAS); 10 $ $ (CU); 1 $ (GB); 12 $ $ (MCZ); 1 S (NCSC) ; 2 S $ (OSU) ; 4 $ $ (UA) ; 6 $ $ (USNM) ; 1 $ (PU) ; 2 S $ (CU). Description. Color. Dorsal: black shining; apical area of elytra sometimes with an orange spot. Ventral: black to dark brown. Appendages: black to dark brown or dark to reddish brown. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: distinct, complete. Frontal sulci: vaguely present. Micro- sculpture: clypeus, frons and vertex isodiametric. Mentum: trans- verse; lobes narrowly rounded; tooth distinct, blunt at apex, almost as long as lobes; furrows absent; fovae distinct, deep; microsculpture elongate. Thorax. Pronotum: microsculpture usually absent; if pres- ent, elongate; lateral margins sparsely punctate posteriorly, evenly rounded their entire length, not sinuate posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal fovae but incomplete; median sulcus reaching anterior sulcus or not, never reaching posterior margin; basal impres- sions elongate, moderately deep, punctate; area between basal im- pressions sparsely punctate; area between basal impression and lateral margins punctate or not; if punctate, punctures near lateral or posterior margins. Scutellum: microsculpture isodiametric; apex pointed. Pro- sternum: microsculpture absent medially, elongate toward the lateral margin; impunctate. Proepisternum and proepimeron: microsculpture weak, elongate; impunctate. Mesosternum: microsculpture elongate; impunctate. Mesepisternum : microsculpture usually absent; if pres- ent, elongate; punctate in anterior half. Metastemum: microsculpture present only laterally and around mesocoxae, elongate; punctate later- ally. Metepisternum : microsculpture, if present, elongate; punctate. Metepimeron: microsculpture usually absent or very elongate; im- punctate. Legs. Hind coxae: microsculpture on inner surface absent or weak and elongate. Tarsi: segments 1-3 and sometimes 4 (ob- scure) grooved dorso-laterally. Elytra. Microsculpture on intervals absent; striae 1-6 and 8 distinct, punctate, basal quarter of striae 7 not as distinct as others, trough 1-2 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture on epipleura absent. Abdominal sternites. Microsculpture isodiametric laterally, becoming elongate and then absent medially; basal and lateral areas of segment 1 116 Entomologica Americana [Vol. 46, No. 1 and sometimes 2 punctate; segments 1-4 with basal and lateral areas rugose. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position, slightly directed dorsally; 5 distinct scale groups on main sac; secondary sac everts toward the apex on the left side; secondary sac with a group of small scales at its apex; most of the remainder of the sac’s surface covered with small, weakly sclerotized scales; sclerotized ring present, on left lateral surface. Measurements. (20 specimens). TL— 6.4 (6.0-7.0); WH— 1.4 (1.2— 1.4); WP— 2.2 (1.9-2.4); WW— 2.7 (2. 5-3.0). Distribution. Eastern United States west to Missouri, Arkansas, and Louisiana. Localities. Alabama. Mobile Co.: Delchamps (CAS), Mobile (MCZ, OSU), Mt. Vernon (CU); Tuscaloosa Co.: Tuscaloosa (GB). Arkansas. Howard Co.: 4 mi. W. Mineral Springs, Rte. #27 (UA). District of Columbia. Washington (MCZ, USNM) . Florida. Unrecorded locality (OSU). Louisiana. East Baton Rouge Co.: Baton Rouge (UA). Mississippi. George Co. : Lucedale (CU); Green Co. : Leaksville (CU). Missouri. St. Louis Co.: St. Louis (MCZ). North Carolina. Hoke Co. (NCSC). Pennsylvania. Dauphin Co.; Harrisburg (CAS, MCZ, USNM). * Philadelphia Neck (OSU). Unrecorded localities in Pennsylvania (MCZ, USNM). Ohio. Hocking Co. (PU). Tennessee. Smith Co.: Elmwood (CAS). Virginia. Southampton Co.: Boykins (MCZ). *Blackpond (MCZ). Discussion. See above. 34. Loxandrus duryi Wright Figs. 42; 109 A, B, C, D, E, F; 166 Loxandrus duryi Wright 1939: 1. Holotype. Not examined. Wright stated in his original description “Holotype male and female are in my collec- tion.” Mr. Charles Oehler (personal communication) of the Cincinnati Museum of Natural History informs me that he has been unable to locate the type specimen of this species. I have, however, examined paratypes and numerous specimens from the type locality. Based on a study of this material, I am tentatively assuming the holotype to be the same as the paratypes and other specimens collected at the type locality. The type locality is near Goshen, Ohio, in Clermont County. Figs. 149-151. Geographical distribution maps. 149. L. discolor n. sp., L. spinigrandis n. sp., L. tropicus n. sp., L. lepidus n. sp. 150. L. whiteheadi n. sp., L. unispinus n. sp. 151. L. balli n. sp. 1972] Allen: Revision of Loxandrus LeConte 117 118 Entomologica Americana [Vol. 46, No. 1 Description. Color. Dorsal: black, shining, elytra sometimes with central orange spot near the apex. Ventral: usually black, some- times dark to light brown. Appendages : dark to light brown to straw yellow. Head. Clypeus: anterior margin feebly concave. Epistomal and subgenal suture: finely produced but complete. Frontal sulci: obscure, short, shallow. Microsculpture; clypeus, frons and vertex weak, isodiametric. Mentum: apex of lobes rounded; tooth as long as or almost as long as lobes, truncate at apex; furrows absent; fovae distinct, deep; microsculpture isodiametric and elongate. Thorax. Pronotum: microsculpture absent; lateral margins evenly rounded, not sinuate posteriorly; sometimes weakly punctate posteriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal fovae but incomplete; median sulcus present as a fine line, reaching anterior marginal sulcus but not posterior margin; basal impressions elongate, moderately deep, punctate; area between basal impressions sparsely punctate; area between basal impressions and lateral margins usually impunctate. Scutellum: microsculpture isodiametric; apex pointed. Prosternum: microsculpture present only along lateral margins and procoxae; impunctate. Proepisternum and Proepimeron: microsculpture elon- gate; impunctate. Mesosternum: microsculpture isodiametric to elongate; impunctate. Mesepisternum : microsculpture elongate; punctate in anterior half. Metasternum: microsculpture more evident laterally, elongate; punctate laterally. Metepisternum : microsculpture weak, elongate; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface weak, elongate. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on inter- vals absent; all striae distinct, punctate, trough 1 isodiametric unit wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture elongate medially, isodiametric laterally; impunctate; segments 1-3 rugose laterally and basally. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position, slightly directed dorsally; 5 distinct scale groups present; most of the remainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present, opening on the left side. Measurements. (17 specimens). TL — 7.0 (6. 8-7. 3); WH— 1.4 (1.3-1. 5); WP— 2.3 (2.1-2.3); WW— 2.8 (2.6-2.9). Distribution. Eastern United States west to Texas. Figs. 152-154. Geographical distribution maps. 152. L. elnae n. sp., L. fraus n. sp. 153. L. sculptilis Bates. 154. L. pactinullus n. sp. 1972] Allen: Revision of Loxandrus LeConte 119 120 Entomologica Americana [Vol. 46, No. 1 Localities. Indiana. Marion Co. (PU); St. Joseph Co.: Elkhart (MCZ); Virgo Co. Louisiana. Caddo Co.: Shreveport (UA); Evan- geline Co.: Lake Chicot State Park (TLE). Ohio. Ashtabula Co.: Conneaut (PU); Athens Co.: Athens (UMIN); Clermont Co.: Goshen (CNC, USNM) . Hamilton Co.: Cincinnati (UM); Licking Co.: He- born (MSU) ; Putnam Co. : Ottawa TWP, Sec. 85, Blanchard River (PU) ; Washington Co.: Maritta (MSU). Unrecorded locality in Ohio (OSU). Pennsylvania. Allegheny Co. (CU). Texas. Dallas Co.: Dallas (MSU). Unrecorded locality in Texas (MCZ) . Discussion. See above. 35. Loxandrus spinigrandis New Species Figs. 26; 83 A, B; 149 Holotype. S . Canal Zone, Barro Colorado Island, 1947. Zetek, Collector, Lot 47-4230 (USNM). Paratypes. 1 $ 5 $ 5 (USNM). Description. Color. Dorsal: head usually black sometimes red- dish brown to light yellow; pronotum black to reddish brown, lateral margins sometimes lighter in color than disk; elytra black to dark brown, a yellow spot near the humerus and towards the apex; apical spot may be faint or absent. Ventral: black to dark brown, apex of last abdominal segment yellow. Appendages: dark brown to yellow. Head. Clypeus: anterior margin straight. Epistomal suture: distinct, complete. Subgenal suture: usually distinct, sometimes obscure in females. Frontal sulci: shallow, broad, short. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: lobes nar- rowly rounded; tooth distinct, not as long as lobes, rounded at apex; furrows present shallow; fovae present or absent, when present small and shallow; microsculpture absent. Thorax. Pronotum: microsculp- ture absent; lateral margins evenly rounded, not sinuate posteriorly, punctate their entire length or at least posterior three quarters; poste- rior angles rounded; anterior marginal sulcus complete; posterior mar- ginal sulcus not reaching past basal impressions; median sulcus present, not reaching anterior marginal sulcus or posterior margin; basal im- pressions elongate, moderately impressed, punctate; area between impressions and between impressions and lateral margins sparsely punctate. Scutellum: microsculpture absent; apex pointed. N. B. En- tire ventral surface without microsculpture. Prosternum: impunctate. Proepisternum : punctate. Proepimeron: impunctate. Mesosternum: impunctate. Mesepisternum : punctate in anterior quarter. Metaster- Figs. 155-156. Geographical distribution maps. 155. L. erraticus Dej. 156. L. gibbus n. sp. 1972] Allen: Revision of Loxandrus LeConte 121 122 Entomologica Americana [Vol. 46, No. 1 num: punctate laterally. Metepisternum : punctate. Metepimeron: impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi grooved dorso- laterally. Elytra. Microsculpture on intervals absent; all striae dis- tinct, punctate, trough 1 isodiametric unit wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Ab- dominal sternites. Microsculpture absent; segment 1 punctate and rugose at base; segments 2-5 finely rugose laterally. Aedeagus. Me- dian lobe: not modified. Internal sac: everts medially, deflected to the right side and coiled towards the apex; two parallel rows of large spines extending the length of the sac; distinct scale groups absent; most of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring absent. Measurements. (7 specimens). TL — 7.6 (7.0-8.2); WH— 1.5 (1. 5-1.7); WP— 2.5 (2.2-2.7); WW- 3.3 (3. 1-3.6). Distribution. Central America. Localities, canal zone. Barro Colorado Island (USNM). Discussion. This species closely resembles L. discolor and L. whiteheadi in many external characters. It can readily be separated from these two species on the basis of the broadly explinate posterior lateral areas of the pronotum. This character is similar to the same condition that occurs in the distantly related L. saphyrinus. L. spini- grandis has been collected in the months of January, February, July, August, September and October. 36. Loxandrus discolor New Species Figs. 15; 29 A,B;82 A,B;149 Holotype. S. Canal Zone, Barro Colorado Island. May, 1941. James Zetek, Collector, collected at black light. Lot No. 41-20602 (USNM). Paratypes. 2 $ $ 2 2 2 (USNM). Description. Color. Dorsal: head, dark brown, labrum in female sometimes straw yellow; pronotum, disk dark brown, lateral margins light brown to straw yellow; elytra dark to light brown, a yellow spot near the humeri and towards the apex. Ventral: dark to reddish brown. Appendages: dark brown to yellow. Head. Clypeus: anterior margin straight. Epistomal and subgenal sutures: distinct, Figs. 157-158. Geographical distribution maps. 157. L. rectus (Say). 158. L. velocipes Casey, L. inferus n. sp. 1972] Allen: Revision of Loxandrus LeConte 123 <3 124 Entomologica Americana [Vol. 46, No. 1 complete. Frontal sulci: moderately deep, broad, short. Micro- sculpture: clypeus, frons and vertex isodiametric. Mentum: apex of lobes narrowly rounded; tooth distinct, rounded at apex; furrows shallow, broad; fovae present or absent; if present, small and puncture- like; microsculpture elongate. Thorax. Pronotum: Microsculpture absent; lateral margins almost evenly rounded their entire length, not sinuate posteriorly, coarsely and densely punctate in the posterior half to three quarters; posterior angles distinct, acute; anterior marginal sulcus complete; posterior marginal sulcus reaching well past basal impressions, almost complete in some specimens; median sulcus dis- tinct but not reaching anterior marginal sulcus or posterior margin; basal impressions deep, elongate, punctate; area between impressions punctate; area between impressions and lateral margins sparsely punc- tate. Scutellum: microsculpture isodiametric; apex pointed. N. B. En- tire ventral surface without microsculpture. Prosternum: impunctate. Proepisternum : punctate. Proepimeron: impunctate. Mesosternum: impunctate. Mesepisternum : punctate in anterior quarter to half. Metasternum: punctate laterally. Metepisternum : punctate. Metepi- meron: impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct, punctate, trough 3-4 isodiametric units wide; internal plica well developed; humeri rounded; micro- sculpture of epipleura absent. Abdominal sternites. Microsculpture absent; segment 1 rugose and punctate along the basal margin and toward the lateral margin; segments 2-3 weakly rugose laterally. Ae- deagus. Median lobe: not modified. Internal sac: everts medially, directed dorsally; no distinct scale groups present; most of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring absent. Measurements. (5 specimens). TL — 7.0 (6. 7-7. 4); WH — 1.4 (1.3-1. 5); WP— 2.2 (2. 1-2.4); WW— 3.2 (3. 1-3. 3). Distribution. Central America. Localities, canal zone. Barro Colorado Island (USNM). Discussion. L. discolor can be distinguished from L. whiteheadi by the more numerous and larger punctures along the lateral margins of the pronotum. The internal sac of the aedeagus is very similar to the internal sac of L. whiteheadi. However, the characters given in the key seem to be constant and serve to help distinguish these two species. Figs. 159-160. Geographical distribution maps. 159. L. pravitubus n. sp., L. circulus n. sp. 160. L. brevicollis (LeC.). 1972] Allen: Revision of Loxandrus LeConte 125 o 126 Entomologica Americana [Vol. 46, No. 1 L. discolor has been collected in April and May on Barro Colorado Island. 37. Loxandrus saphyrinus (Chaudoir) Figs. 19 A, B, C; 22; 36 A, B; 106; 164 Megalostylus saphyrinus Chaudoir 1843 : 766. Lectotype (here designated) . $ . The first specimen in a series of 3 specimens is a male and is designated the lectotype. The other two specimens are female. Louisiana is listed as the locality on the determination label. Description. Color. Entire body black, iridescent; appendages, dark to light brown, femora sometimes black. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: complete. Frontal sulci: abbreviated, broad, shallow. Microsculp- ture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes pointed; tooth large, shorter than lobes, apex rounded; furrows obscure; fovae distinct; microsculpture obscure, elongate. Thorax. Pronotum : microsculpture absent on disk, present along lateral mar- gins and in basal impressions; lateral margins narrowly reflexed in anterior two thirds, becoming broadly explinate in posterior third, punctate or not, evenly rounded, not sinuate posteriorly; posterior angles broadly rounded; anterior marginal sulcus incomplete; posterior marginal sulcus present, reaching to or only slightly past basal im- pressions; median sulcus present, not reaching anterior or posterior marginal sulci; basal impressions elongate, shallow but distinct, punc- tate or not; area between basal impressions usually punctate; area between basal impressions and lateral margin punctate or not. Scu- tellum: apex pointed; microsculpture mostly absent, sometimes vaguely present near apex, if so, elongate. Prosternum, Proepisternum, Proepimeron: impunctate; microsculpture absent. Mesosternum: impunctate; microsculpture elongate. Mesepisternum: punctate in the anterior three quarters; microsculpture vague, elongate. Meta- sternum : punctate toward lateral margins ; microsculpture present only in sulci and punctures. Metepistemum : punctate; microsculpture present only in sulci and punctures. Metepimeron: impunctate; micro- sculpture absent. Legs. Hind coxae: microsculpture usually absent, if present, isodiametric; tarsal segments 1-3 of middle and hind tarsi (sometimes 4 of hind tarsi) grooved dorso-laterally. Elytra. Micro- sculpture absent; striae 6 and 7 not as distinct as 1-5, trough 4-5 isodiametric units wide; internal plica well-developed; humeri rounded; Figs. 161-162. Geographical distribution maps. 161. L. nitidulus (LeC.). 162. L. crenatus LeC. 1972] Allen: Revision of Loxandrus LeConte 27 128 Entomologica Americana [Vol. 46, No. 1 microsculpture of epipleura absent. Abdominal sternites. Microsculp- ture, where present, isodiametric; segments 1-4 rugose and punctate laterally, less so posteriorly. Aedeagus. Median lobe: not modified. Internal sac: everts medially, turned dorsally; one scale group on the right lateral side of the sac, posterior to the midline (sometimes re- duced to a single scale); at least % of the sac’s surface covered with very small, weakly sclerotized scales, these sometimes becoming more distinct on the ventral surface at the base of the sac; sclerotized ring present. Measurements. (20 SS specimens). TL — 11.6(10.7- 12.3); WH— 2.5 (2.2— 2.6); WP— 3.8 (3.4-4.0); WW— 4.6 (4.0-4.9). Distribution. Southeastern United States, west to Oklahoma and South-central Texas. Localities. Alabama. Mobile Co.: Mobile (MCZ), Mt. Vernon (OSU); Randolph Co.: Wadley (USNM). *Chestong (OSU). Arkansas. Arkansas Co.: 5 mi. E. Dewitt (UA); Ashley Co.: 6 mi. W. Crossett (UA) ; Hempstead Co.: Hope (CAS) ; Washington Co.: 5 mi. S. Fayette- ville (MSU). *Southwest Arkansas, unrecorded locality (AM). Florida. Charlotte Co.: Punta Gorda (CAS); Hendry Co.: LaBelle (KU); Her- nando Co.: Brooksville (CAS, MCZ); Highlands Co.: Archbold Bio- logical Station, Lake Placid (CU, PSU); Hillsborough Co.: Plant City (FSPB); Tampa (CU, MCZ, MSU, PU, USNM); Leon Co.: Lake Jack- son (?), unrecorded locality (CU); Manatee Co.: Oneco (GB); Marion Co. (AM, CAS, CU); Orange Co.: Winter Park (MCZ); Osceola Co.: Kissimmee (USNM); Palm Co.: Canal Point (PU); Pinellas Co.: St. Petersburg (PU). Sarasota Co.: Sarasota (PU); Seminole Co.: Sanford (MCZ); Taylor Co.: Steinhatchee (USNM), Lake Okeechobee (PU). *Cedar Keys (USNM). *Oette (AM). *Ouecee (PU). Unrecorded locality in Florida (CAS, CNHM, CU, INHS, MCZ, OSU, USNM). Georgia. Thomas Co.: Thomasville (AM, USNM). Unrecorded locality (MCZ). Louisiana. East Baton Rouge Co. : Baton Rouge (MSU) ; Evan- geline Co. : Lake Chicot State Park (TLE) ; Vernon Co. : Rosepine (UA) . Unrecorded locality in Louisiana (AM, MCZ). Mississippi. Adams Co.: Natchez (CAS); Perry Co.: New Augusta (CU). *Lake View (CU). North Carolina. Johnston Co.: Clayton (NCSC) ; Hope Co.: Unrecorded locality (NCSC). South Carolina. Pickens Co.: Clemson (MSU). Ten- nessee. Unrecorded locality (UMIN). Texas. Brazos Co.: College Sta- tion (TAM); Calhoun Co.: Alligator Head (USNM); Colorado Co.: Columbus (MCZ, USNM), Skull Creek (UMIN). Discussion. L. saphyrinus is one of the larger Loxandrus species. It is very distinct and can readily be distinguished from other Lox- andrus species by the characters given in the key. I have collected it in very wet habitats under leaf debris along the margins of streams. I have also taken it under leaf debris in relatively dry habitats and at 1972] Allen: Revision of Loxandrus LeConte 129 black light traps. It has been collected in all months of the year except October and November. 38. Loxandrus lucens Chaudoir Figs. 37 A, B; 105; 163 Loxandrus lucens Chaudoir 1868:342. Lectotype, here selected. The second specimen in a series of 3. A male with a Texas locality label. The first specimen in the series is a male but has no data labels. The third specimen is a female from New Orleans and probably not part of the original type material. The type locality given by Chaudoir was Havre, Texas. Description. Color. Dorsal: black, shining, elytra iridescent. Ventral: black to dark brown. Appendages: dark to reddish brown. Head. Clypeus: anterior margin slightly concave. Epistomal and subgenal sutures: obscure, weak. Frontal sulci : almost absent, broad, shallow. Microsculpture: clypeus, weak, isodiametric; frons and ver- tex, distinct, isodiametric. Mentum: apex of lobes rounded; tooth distinct, rounded at apex; furrows absent or ill-defined; fovae absent; microsculpture slightly elongate. Thorax. Pronotum: microsculpture on disk absent, present in the reflexed portion of the pronotum; lateral margins evenly rounded their entire length, not sinuate posteriorly; posterior angles broadly rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past the basal fovae but incomplete; median sulcus reaching anterior sulcus but not posterior margin; basal impressions elongate, moderately deep, punctate; area between im- pressions sparsely punctate; area between impressions and lateral margins usually impunctate. Scutellum: microsculpture elongate; apex pointed. Prosternum: microsculpture distinct laterally, elongate; impunctate. Proepisternum : microsculpture absent; punctures weak, shallow. Proepimeron: microsculpture elongate; impunctate. Meso- sternum: microsculpture weak, elongate; impunctate. Mesepisternum : microsculpture usually absent; punctate in anterior half to three quar- ters. Metasternum: microsculpture distinct laterally, elongate; punc- tate laterally. Metepisternum: microsculpture absent; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface, if present, elongate. Tarsi: segments 1-3 of middle and hind tarsi and sometimes 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; striae 1-6 and 8 distinct, 7 less so, all striae punctate, trough 1-2 sometimes 3 isodiametric units wide; internal plica distinct; humeri rounded; micro- sculpture on epipleura absent. Abdominal sternites. Microsculpture isodiametric laterally, elongate medially; segment 1 punctate at base, 130 Entomologica Americana [Vol. 46, No. 1 laterally; segments 1-4 rugose at the base, laterally. Aedeagus. Me- dian lobe: not modified. Internal sac: everts to the left side; no distinct scale groups on the main body of the sac; most of the surface is covered with small weakly sclerotized scales; a secondary bulbous sac everts in the basal half of the main sac, apex with a distinct scale group; sclerotized ring present. Measurements. (7 specimens). TL — 9.1 (8.5-9. 7); WH— 1.9 (1. 8-2.1); WP— 3.0 (2.7— 3.2); WW— 3.6 (3. 3-3.9). Distribution. Southern United States. Localities. Alabama. Mobile Co.: Calvert (CAS). *Tumblin Gap (USNM), Arkansas. Arkansas Co.: E. Rydell, Bayou Meto (UA); Prairie Co.: (KU). Kentucky. Henderson Co.: Henderson (CU). Louisiana. Franklin Co.: Chase (UA); Orleans Co.: New Orleans (CAS) ($ only). Tennessee. Shelby Co.: Memphis (CAS). Unrecorded locality in Tennessee (UM). Texas. Calhoun Co.: Alligator Head (USNM) ($ only). Dallas Co.: Dallas (GB, MSU) (9$ only). Discussion. This species closely resembles L. uniformis in most external characters. The shape of the lateral margin of the pronotum serves to distinguish L. lucens from L. uniformis . This is a rather weak character and one that is hard to distinguish by one who is not familiar with Loxandrus. A more reliable identification can be made by examining the aedeagii of the two species. L. lucens has been collected in February, March, May, July and August. 39. Loxandrus uniformis New Species Figs. 38 A, B; 107; 165 Holotype. $. Alabama. Mobile, November 6, 1939. Van Dyke Col- lection (CAS). Paratypes. 5 $$ 1 9 (CU); 1 $ (MCZ); 1 $ (GB); 1 S (PU); 1 $ (INHS). Description. Color. Dorsal: black or dark brown, shining, elytra iridescent. Ventral: black to dark brown, shining. Append- ages: dark to light brown. Head. Clypeus: anterior margin slightly concave. Epistomal suture: weak but complete. Subgenal suture: present or absent. Frontal sulci: obscure, broad, shallow. Micro- sculpture: clypeus, frons and vertex weak, isodiametric. Mentum: apex of lobes rounded; tooth distinct, apex rounded; furrows absent or shallow; fovae absent; microsculpture slightly elongate. Thorax. Figs. 163-164. Geographical distribution maps. 163. L. lucens Chaud. 164. L. saphyrinus (Chaud.). 1972J Allen: Revision of Loxandrus LeConte 131 132 Entomologica Americana [Vol. 46, No. 1 Pronotum : microsculpture absent on disk, sometimes vaguely present along reflexed lateral margin; lateral margins more or less evenly rounded their entire length, not sinuate posteriorly; posterior angles narrowly rounded; anterior marginal sulcus usually incomplete; poste- rior marginal sulcus reaching to or slightly past basal impressions; median sulcus distinct but not reaching anterior or posterior marginal sulci; basal impressions moderately deep, elongate, punctate; area between basal impressions sparsely punctate; area between basal im- pressions and lateral margins usually impunctate. Scutellum: micro- sculpture isodiametric; apex pointed. Prosternum: microsculpture distinct toward lateral margin, elongate; impunctate. Proepisternum : microsculpture absent; punctate (sometimes weakly punctate). Pro- epimeron: microsculpture absent; impunctate. Mesosternum: im- punctate. Mesepisternum : microsculpture absent; anterior half punctate. Metasternum: microsculpture absent; punctate laterally. Metepisternum : microsculpture absent; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculp- ture on inner surface absent or very elongate. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct punctate, trough 2-3 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture distinct, elongate laterally, becoming absent medially; segments 1-4 and sometimes 5 punctate and rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position; four distinct scale groups present; most of the re- mainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present. Measurements. (8 specimens). TL — 9.5 (8.5-10.1); WH— 1.8 (1. 6-1.9); WP— 2.9 (2.6-3. 1); WW— 3.7 (3.4-3. 8). Distribution. Eastern United States. Localities. Alabama. Mobile Co.: Mobile (CAS, CU). Arkansas. Unrecorded locality (INHS). Ohio. Lucas Co. (PU). South Carolina. Sumpter Co.: Sumpter (GB). Virginia. Unrecorded locality (MCZ). Discussion. See discussion of L. lucens. L. uniformis has been collected in November and March. Figs. 165-166. Geographical distribution maps. 165. L. micans Chaud., L. uniformis n. sp. 166. L. duryi Wright. 1972] Allen: Revision of Loxandrus LeConte 133 c 134 Entomologica Americana [Vol. 46, No. 1 40. Loxandrus miniitus New Species Figs. 65 A, B; 128; 129 A, B, C; 175 Holotype. S . Florida. Sarasota Co., Myakka River State Park. March 24, 1954. George E. Ball (MCZ). Paratypes. 37 $ $ 33 $ $ (GB); 1 $ 1 $ (UA); 3 S $ (MCZ); 1 $ (USNM); 2 $ $ (CU); 4 $ $ (FSPB); 1 $ (UM); 1 $ (OSU) ; 4 $ S (CAS); 2 $ S (PU). Description. Color. Dorsal: black, shining. Ventral: usually wholly black, sometimes dark brown. Appendages: black to dark brown. Head. Clypeus: anterior margin straight or slightly concave. Epistomal suture: obscure but usually present. Subgenal suture: present or absent. Frontal sulci : almost absent, obscure, broad, shal- low. Microsculpture: clypeus, frons and vertex, isodiametric. Men- tum: apex of lobes narrowly rounded; tooth distinct, shorter than lobes; furrows obscure, shallow; fovae absent; microsculpture isodia- metric or slightly elongate. Thorax. Pronotum: microsculpture present, elongate on the disk, isodiametric anteriorly, along the reflexed lateral margins and in the basal impressions; lateral margins sparsely punctate posteriorly, evenly curved their entire length, not sinuate pos- teriorly; posterior angles rounded; anterior marginal sulcus complete; posterior marginal sulcus usually incomplete (vaguely complete in a few specimens) ; median sulcus reaching anterior marginal sulcus or not, never reaching posterior margin; basal impressions moderately deep, elongate, punctate; area between basal impressions and lateral margins usually impunctate. Scutellum: apex pointed; microsculpture isodia- metric. Prosternum: microsculpture elongate or absent medially, iso- diametric along the lateral margins; impunctate. Proepisternum and Proepimeron: microsculpture mostly isodiametric, sometimes elongate laterally; impunctate. Mesosternum: microsculpture slightly elongate; impunctate. Mesepisternum : microsculpture isodiametric; sparsely punctate anteriorly. Metasternum : microsculpture elongate or absent medially, isodiametric towards the lateral margins; impunctate. Met- episternum: microsculpture isodiametric; impunctate. Metepimeron: microsculpture elongate or absent; impunctate. Legs. Hind coxae: microsculpture on inner surface elongate. Tarsal segments: 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture absent on intervals; striae 7 not as distinct basally as 1-6; all striae punctate, trough 1-2 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of Figs. 167-168. Geographical distribution maps. 167. L. vitiosus n. sp., L. minor (Chaud.). 168. L. cincinnatiensis Casey. 1972] Allen: Revision of Loxandrus LeConte 135 136 Entomologica Americana [Vol. 46, No. 1 epipleura absent. Abdominal sternites. Microsculpture isodiametric laterally, elongate or absent medially; segment 1 sometimes punc- tate at the base; segments 1-3 rugose along the basal margin laterally; ambulatory setae sometimes present on second segment. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position; seven distinct scale groups present; two small sec- ondary sacs on the left side of the sac, everting near the sclerotized ring, both with a small group of scales at the apex. Most of the re- mainder of the sac’s surface is covered with very small, weakly sclero- tized scales; sclerotized ring present. Measurements. (20 specimens). TL— 6.4 (6.0-6.9); WH— 1.3 (1.2— 1.4); WP— 2.1 ( 1. 9-2.2) ;WW —2.7 (2. 6-2. 9). Distribution. Southeastern United States. Localities. Florida. Alachua Co.: Gainesville (FSPB); Hernando Co.: Brooksville (CAS); Highlands Co.: Highlands Hammock (FSPB); Hillsborough Co.: Tampa (MCZ, USNM); Manatee Co.: Oneco (GB); Marion Co.: Micanopy (CU); Pinellas Co.: Dunedin (PU, UM); Sara- sota Co.: Myakka River State Park (CU, FSPB, GB), Sarasota (PU), unrecorded locality (CAS, MCZ). Unrecorded locality in Florida (OSU). Georgia. Pierce Co.: Little Satilla River, at junction with U.S. - 301 (GB). Discussion. The smaller size and more rounded posterior angles of the pronotum serve to distinguish L. minutus from other species with which it might be confused. Positive identification, however, should be based on an examination of the male aedeagus. It has been col- lected in the months of January, February, March, April, June, October and November. 41. Loxandrus velocipes Casey Figs. 47; 99 A, B; 158 Loxandrus velocipes 1918:390. Holotype. $. District of Columbia. Type No. 47346 (USNM). Description. This species resembles L. inferus in all external characters. The reader is referred to the description of L. inferus. The two species differ only in the presence of one additional scale group on the right lateral surface of the internal sac in L. velocipes which is absent in L. inferus. Measurements. (20 specimens). TL — 9.8 (9.0-10.1); WH— 1.9 (1.7-2.0); WP— 3.0 (2.8-3.3); WW— 3.9 (3. 6-4.1). Distribution. Eastern United States. 1972] Allen: Revision of Loxandrus LeConte 137 Localities. Alabama. Mobile Co.: Calvert (CAS), Mobile (OSU), Mt. Vernon (CU); Tuscaloosa Co.: Tuscaloosa (GB). Arkansas. Ar- kansas Co.: White River National Wildlife Refuge, White River Chute (UA); Drew Co.: 5 mi. Northwest Dermott, 7 Devils Swamp (UA); Union Co.: Little Corne Bayou, Hy. 169 (UA). Southwest Arkansas, unrecorded locality (AM) . Illinois. Alexander Co. : Cashe River Bottom (RTB); Champaign Co.: Brownfield Woods (3 mi. N. Champaign) (UA); Cook Co.: Palos Park (UM). *Kahokia (MCZ). *Horseshoe Lake (INHS). Indiana. Posey Co. (PU) . Louisiana. Orleans Co.: New Orleans (MCZ). Unrecorded locality (UM). Minnesota. Houston Co. (UMIN). Missouri. Unrecorded locality (USNM). North Carolina. Robeson Co.: Lumberton (USNM). Tennessee. Shelby Co.: Memphis (UM). Virginia. Unrecorded locality (USNM) . Discussion. See above and discussion of L. inferus. 42. Loxandrus inferus New Species Fig. 158 Holotype. S. Virginia. Nelson Co., Rossyln, March 30, 1925. C. E. White, Collector (MCZ). Paratypes. 6 $$ (USNM); 2 $$ (MCZ); 1 $ (CU); 1 $ (OXU) ; 1 $ (INHS); 1 S (TLE) ; 1 $ (MSU); 1 $ (CAS). Description. Color. Dorsal: black, shining, elytra iridescent. Ventral: black shining. Appendages: yellow to dark brown. Head. Clypeus: anterior margin straight or slightly concave. Epistomal and subgenal sutures: weak. Frontal sulci: shallow, short, obscure. Microsculpture: clypeus, frons and vertex, distinct, isodiametric. Mentum: apex of lobes narrowly rounded; tooth large, apex truncate; furrows present; fovae absent; microsculpture distinct, isodiametric or slightly elongate. Thorax. Pronotum: microsculpture (usually) dis- tinct, elongate; lateral margins punctate along basal quarter, evenly rounded, not sinuate posteriorly; posterior angles subacute; anterior marginal sulcus complete; posterior marginal sulcus incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions elongate, narrow, deep, punctate; area be- tween basal impressions and between impressions and lateral margins punctate. Scutellum: microsculpture distinct, isodiametric; apex pointed. Prosternum: microsculpture isodiametric; impunctate. Proepisternum : microsculpture isodiametric; impunctate. Proepi- meron: microsculpture isodiametric; impunctate. Mesosternum: microsculpture elongate; impunctate. Mesepisternum: microsculpture elongate; anterior half punctate. Metasternum: microsculpture elongate; punctate laterally. Metepisternum: microsculpture elongate; punctate. Metepimeron: microsculpture weak, elongate; impunctate. 138 Entomologica Americana [Vol. 46, No. 1 Legs. Hind coxae: microsculpture on inner surface elongate. Tarsal segments 1-3 (and often 4) of middle and hind tarsi grooved dorso- laterally. Elytra. Microsculpture on intervals absent; all striae distinct, trough 3-4 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture isodiametric laterally, becoming elongate medially; segment 1 punctate along basal margin laterally; segments 1-4 usually rugose along basal margin laterally. Aedeagus. Median lobe: not modified. Internal sac: everts medially and is directed dorsally; 4 distinct scale groups are present; small weakly sclerotized scales cover most of remainder of the sac’s surface; sclerotized ring present. Measurements. (15 specimens). TL — 9.7 (9.1-10.1 ); WH —1.8 (1. 7-1.9); WP— 3.0 (2.7-3.2); WW— 3.8 (3. 6-4.0). Distribution. Eastern United States. Localities. Alabama. Mobile Co.: Calvert (CAS), Mt. Vernon (CU, OSU). District of Columbia. Washington (USNM, INHS). Georgia. Unrecorded locality (MSU). Louisiana. Evangeline Co.: Lake Chicot State Park (TLE); Orleans Co.: New Orleans (MCZ). Maryland. St. Marys Co.: Pinsy Pt. (USNM). Virginia. Nelson Co.: Rossyln (MCZ). Discussion. L. inferus and L. velocipes are two very closely related species. They appear to be alike in all external characters and can be separated only on the basis of characters associated with the male genitalia. L. velocipes has one scale group on the right lateral surface of the internal sac that is not found in L. inferus. The geo- graphical ranges of the two species overlap. L. inferus has been col- lected in the months of March, May, June and July. L. velocipes has been collected in the months of January, March, April, May, June, July and October. I have collected the latter species under very wet leaf debris in Arkansas and at a black light trap in Illinois. 43. Loxandrus remotus New Species Figs. 5; 70 A, B; 77 A, B; 148 Holotype. S. Canal Zone, Paraiso. May 24, 1911. August Busck (USNM). Paratypes. 1 <3 1 $ (USNM). Description. Color. Dorsal: usually black, sometimes dark brown, shining. Ventral: black to dark brown, shining. Appendages: black to dark brown to light yellow. Head. Clypeus : anterior margin slightly concave. Epistomal and subgenal sutures: distinct, complete. Figs. 169-170. Geographical distribution maps. 169. L. robustus n. sp., L. yeariani n. sp. 170. L. parallelus Casey, L. saccisecundaris n. sp. 1972] Allen: Revision of Loxandrus LeConte 139 170 L. PARALLELUS • L. SACCISECUNDARIS^ o 140 Entomologica Americana [Vol. 46, No. Frontal sulci: deep, broad, slightly diverging posteriorly. Microsculp- ture: clypeus, frons and vertex weak, isodiametric. Mentum: trans- verse; apex of lobes pointed; tooth distinct, apex obscurely indented, weak parallel carina on the tooth; furrows absent; fovae deep, distinct; microsculpture isodiametric and/or elongate. Thorax. Pronotum: microsculpture, elongate on disk, isodiametric along lateral margins and in basal impressions; entire surface impunctate; lateral margins feebly sinuate posteriorly; posterior angles acute; anterior marginal sulcus complete; posterior marginal sulcus absent; median sulcus almost reaching anterior marginal sulcus but not reaching posterior margin; basal impressions deep, elongate, continuous with a secondary poste- rior marginal sulcus, impunctate. Scutellum: microsculpture isodia- metric; apex pointed. Prosternum and Proepisternum : microsculpture elongate if present; impunctate. Proepimeron: microsculpture isodia- metric and/or elongate; impunctate. Mesosternum: microsculpture elongate; impunctate. Mesepisternum : microsculpture elongate if present; anterior half punctate. Metasternum: microsculpture absent; punctate laterally. Metepisternum : microsculpture absent; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct, punctate, trough 2-3 isodia- metric units wide; internal plica well developed; humeri rounded; microsculpture on epipleura absent. Abdominal sternites. Micro- sculpture elongate if present; segments 1-6 punctate laterally; segments 1-5 rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position; several distinct scale groups present; remainder of the sac’s surface covered with small weakly sclerotized scales; a small secondary sac everting on both the right and left lateral surface at the base of the primary sac; sclerotized ring absent. Mea- surements. (3 specimens). TL— 8.8 (8.5-9.2) ; WH— 1.7 (1.7-1. 8); WP— 2.7 (2.6-2.7); WW— 3.9 (3. 7-4.0). Distribution. Central America. Localities, canal zone. Corazal, La Chorrera, Paraiso (USNM). Discussion. This species is known from three specimens collected in the Canal Zone. The position of this species in the key does not indicate its true phylogenetic relationships. The reader is referred to the section on phylogeny for a further discussion. It has been collected in the months of April and May. Figs. 171-172. Geographical distribution maps. 171. L. florid anus LeC. 172. L. unilobus n. sp. 1972] Allen: Revision of Loxandrus LeConte 141 142 Entomologica Americana [Vol. 46, No. 1 44. Loxandrus brevicollis (LeConte) Figs. 14; 18 A, B; 48; 101 A, B, C, D; 160 Argutor brevicollis LeConte 1848:338. Holotype. 9. Type No. 5695. (MCZ) . There are three specimens in the type series. The first and second specimens are female and the third specimen is a male. None of the speci- mens have data labels and LeConte did not give a specific locality in the original description. I am here selecting Enterprise, Florida, as the type locality. See my discussion concerning types for my reasons for consider- ing this a valid species. Description. Color. Dorsal and ventral: black, shining. Ap- pendages: dark brown to reddish brown. Head. Clypeus: anterior margin slightly concave. Epistomal suture: distinct, complete. Sub- genal suture: obscure but complete. Frontal sulci: almost absent, shallow, broad. Microsculpture: clypeus, frons and vertex isodia- metric. Mentum: lobes narrowly rounded; tooth large, distinct; fur- rows shallow; fovae faintly present (in some specimens); microsculp- ture isodiametric or elongate. Thorax. Pronotum: microsculpture absent on disk, present along the lateral margins; lateral margins rounded in the anterior half becoming somewhat parallel posteriorly, appearing sinuate in some specimens; posterior angles acute; anterior marginal sulcus complete; posterior marginal sulcus reaching to basal impressions, incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions elongate, moderately deep, punctate; area between basal impressions punctate; area between basal impressions and lateral margins punctate or not. Scutellum: microsculpture isodiametric; apex pointed. Prosternum: microsculp- ture elongate medially, isodiametric laterally. Proepisternum : micro- sculpture elongate; punctate or not. Proepimeron: microsculpture elongate; impunctate. Mesosternum: microsculpture elongate; im- punctate. Mesepisternum: microsculpture absent; anterior three quarters punctate. Metasternum: microsculpture elongate if present; punctate laterally. Metepisternum: microsculpture absent; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface elongate if present. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct, punctate, -> Figs. 173-174. Geographical distribution maps. 173. L. agilis (Dej.), L. spinilunatus n. sp. 174. L. aduncus n. sp., L. rossi n. sp. 1972] Allen: Revision of Loxandrus LeConte 143 144 Entomologica Americana [Vol. 46, No. 1 trough 1-2 isodiametric units wide; internal plica distinct; humeri rounded; microsculpture of epipleura absent. Abdominal sternites. Microsculpture elongate medially on segments 2-6 and along the posterior margin of segments 3-6, isodiametric on other areas of the sternites; segment 1 sometimes punctate basally; segments 1-4 weakly rugose laterally. Aedeagus. Median lobe: not modified. Internal sac; everts medially and slightly directed dorsally; six distinct scale groups in the basal half of the sac; most of the sac’s surface in the basal half covered with very small weakly sclerotized scales, apical half more or less free of these small scales; a secondary sac everts on the left lateral surface; sclerotized ring present. Measurements. (20 specimens). XL— 10.6 (9.8-11.5); WH— 2.0 (1. 8-2.2); WP— 3.4 (3.2-3. 6); WW— 4.3 (4.0-4.6). Distribution. Eastern United States. Gulf of Mexico north to Illinois and Pennsylvania. Localities. Alabama. *Tumblin Gap (USNM). Arkansas. Arkan- sas Co.: White River National Wildlife Refuge (UA); Drew Co.: 3 mi. NW. Dermott, 7 Devils Swamp (UA); Greene Co.: Paragould (MSU); Howard Co.: 4 mi. W. Mineral Springs, Rte. 27 (UA); Lafayette Co.: Battle Lake, 7 mi. SW. Lewisville (UA); Union Co.: Little Corne Bayou, Hy. 169 (UA). Unrecorded locality in Arkansas (INHS). District of Columbia. Washington (INHS). Florida. Volusia Co.: Enterprise (USNM). Georgia. Unrecorded localities (MSU, USNM). Illinois. Jackson Co.: Carbondale (USNM); Johnson Co.: Goreville (MSU); Union Co.: Alto Pass (CNHM). Williamson Co.: Herrin (RTB). Indiana. Knox Co.: Unrecorded locality (PU). *Ent. Recon. Sta., Hovey Lake (PU). Louisiana. East Baton Rouge Co.: Baton Rouge (UA); Franklin Co.: Chase (UA). Massachusetts. Unrecorded locality (MCZ). Mississippi. George Co.: Lucedale (CU) ; Jones Co.: 4.2 mi. E. Laurel, Rte. 84. Bogue Home Creek (UA); Perry Co.: Richton (CU). Missouri. Butler Co.: Lake Wappapello, Poplar Bluff (CU); St. Louis Co.: St. Louis (UM). North Carolina. Robeson Co.: Lumberton (USNM). *Boardman (USNM). Oklahoma. Unrecorded locality (USNM). Pennsylvania. Daphin Co.: Harrisburg (MCZ); Delaware Co.: Essington (CAS). Unrecorded locality in Pennsylvania (MCZ, USNM). South Carolina. Anderson Co.: Clemson (MSU). Tennessee. Unrecorded locality (UMIN). State unknown (MCZ, RTB). Discussion. This is one of the larger and more distinct Loxandrus species. The males and females can usually be identified by using external characters. I have found it under wet leaf debris along the banks of streams and other bodies of water. It has also been collected at light traps. It has been collected in the months of February, March, April, May, June, July and August. 1972] Allen: Revision of Loxandrus LeConte 145 45. Loxandrus erraticus (Dejean) Figs. 45; 95 A, B; 155 Feronia erratica Dejean 1828:24. Holotype. $ . The first specimen in the second row of a series of 12 specimens. Bears the large green Dejean determination label, and a small green $ label (MNHP). Recorded from Boreal America. Other specimens in the series have locality labels from New Orleans (Louisiana?), and Cuba. Description. Color. Dorsal: black, shining, elytra iridescent. Ventral: black, sometimes dark brown, shining. Appendages: usually dark to light brown, sometimes femora black. Head. Clypeus: ante- rior margin slightly concave. Epistomal and subgenal suture: com- plete. Frontal sulci: obscure, broad, shallow. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes narrowly rounded; tooth distinct, apex broadly rounded; furrows present; fovae present or not; microsculpture isodiametric or slightly elongate. Thorax. Pronotum: microsculpture usually absent on disk, present along the reflexed lateral margins, isodiametric; lateral margins rounded in anterior half becoming somewhat parallel posteriorly, slightly sinuate near the basal angles, punctate posteriorly; posterior angles acute; anterior marginal sulcus complete or not; posterior marginal sulcus reaching to or slightly past basal impressions; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions moderately deep, elongate, punctate; area between basal furrows and between furrows and lateral margin punctate. Scutellum: microsculpture isodiametric; apex pointed. Prosternum, proepisternum, proepimeron: microsculpture elongate; impunctate. Mesosternum: microsculpture elongate; impunctate. Mesepisternum: microsculp- ture elongate if present; anterior three quarters punctate. Meta- sternum: microsculpture elongate where present; punctate laterally. Metepisternum : microsculpture elongate; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculp- ture on inner surface weak, elongate. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Mi- crosculpture on intervals absent; all striae distinct, punctate, trough 3-4 isodiametric units wide; internal plica well developed; humeri somewhat pointed; microsculpture on epipleura absent. Abdominal sternites. Microsculpture distinct, isodiametric laterally, becoming elongate medially; segment 1 punctate basally; segments 1-4 rugose along the basal margin laterally. Aedeagus. Median lobe: not modi- fied. Internal sac: everts in a median position; two distinct scale groups present, one elongate group in the apical half on the right lateral surface, and one large scale group almost encircling the entire sac; 146 Entomologica Americana [Vol. 46, No. 1 remainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present. Measurements. (20 specimens). TL —9.8 (9.2-10.2); WH— 1.9 ( 1. 7-2.0) ; WP— 3.0 (2.7-3.2) ; WW— 3.9 (3. 6-4. 2). Distribution. Southeastern United States, north to Illinois. Localities. Alabama. Mobile Co.: Calvert (CAS), Mobile (CU, MCZ, OSU), Mt. Vernon (CU); Tuscaloosa Co.: Tuscaloosa (GB). Arkansas. Arkansas Co.: 5 mi. E. Dewitt, Hy. 153 (UA); Greene Co.: Paragould (MSU); Hempstead Co.: Hope (MCZ); Howard Co.: 4 mi. W. Mineral Springs, Rte. 27 (UA). Jackson Co.: 5 mi. S. Newport (RTB); Prairie Co.: Rest Area, Hy. 40 at White River Bridge (UA); Pulaski Co.: Little Rock (MCZ). Unrecorded locality (INHS) . Florida. Manatee Co.: Oneco (GB); Putnam Co.: Crescent City (USNM). Georgia. Thomas Co.: Thomasville (OSU). Illinois. Jackson Co.: Carbondale (INHS); Richland and Lawrence Co.: Wabash Valley (MCZ); Vermillion Co.: Oakwood (INHS). *Goreham (GB, MCZ). Kentucky. *Saborn (MCZ). Louisiana. Orleans Co.: New Orleans (USNM, UM); Plaquemines Co.: Myrtle Grove (INHS). Unrecorded locality in Louisiana (MCZ, UM, USNM). Mississippi. George Co.: Lucedale (CU); Jones Co.: 2 mi. E. Laurel, Rte. 84, Bogue Home Creek (UA); Perry Co.: Richton (CU). Missouri. Butler Co.: Lake Wappa- pello, Poplar Bluff (CU). New Jersey. Atlantic Co.: Atlantic City (MCZ) ; Burlington Co.: Riverton (OSU). North Carolina. *Boardman (USNM). *Lumberton (USNM). * Washington (MCZ). Pennsylvania. Delaware Co.: Essington (CAS). *Delfor (USNM). South Carolina. Unrecorded localities (MCZ). State unknown (AM, RTB, MCZ, USNM, UMIN) . Discussion. This species is similar to L. gibbus in most external characters. Most of the specimens I have studied can be separated using the external characters presented in the key. The characters associated with the aedeagus are characteristic of the species. I have collected this species under leaf debris near the edge of streams and small pools of water. It has been collected in the months of March, May, June and December. 46. Loxantlrus gibbus New Species Figs. 6; 46; 96 A, B; 97; 156 Holotype. S. Alabama. Tuscaloosa, July 18, 1950. George E. Ball, Collector (MCZ). Paratypes. 5 SS (OSU); 3 $$ (CAS); 1 $ (UA). Description. Color. Dorsal: black, shining, elytra iridescent. Ventral: black to dark brown. Appendages: dark to reddish brown. Head. Clypeus: anterior margin slightly concave. Epistomal and 1972] Allen: Revision of Loxandrus LeConte 147 subgenal sutures: obscure, fine. Frontal sulci: almost absent, shal- low, broad, short. Microsculpture: clypeus, frons and vertex isodia- metric. Mentum: apex of lobes pointed; tooth distinct, rounded at apex; furrows and fovae absent; microsculpture elongate. Thorax. Pronotum: microsculpture on disk usually absent or very elongate if present, isodiametric along lateral margins; lateral margins rounded in anterior half, becoming oblique in posterior half, not sinuate, punctate in posterior quarter; posterior angles distinct, narrowly rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impressions but incomplete; median sulcus reaching anterior marginal sulcus but not posterior margin; basal impressions moderately deep, elongate, punctate; area between impressions and between impressions and lateral margins punctate. Scutellum: micro- sculpture isodiametric; apex pointed. Prosternum: microsculpture elongate, more distinct laterally; impunctate. Proepisternum : micro- sculpture elongate; weakly punctate. Proepimeron: microsculpture elongate; impunctate. Mesosternum: microsculpture elongate; im- punctate. Mesepisternum: microsculpture elongate; anterior three quarters punctate. Metasternum: microsculpture elongate where present; punctate laterally. Metepisternum: microsculpture very elongate; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi and segment 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct, punctate, trough 2-3 isodiametric units wide; internal plica distinct; humeri rounded or weakly dentate; microsculpture of epipleura absent. Abdominal sternites. Microsculpture distinct, iso- diametric laterally, elongate medially; segments 1-3 punctate along basal margin laterally; segments 1-4 weakly rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: sac, when everted, directed to the left side; 6 distinct scale groups present; a secondary bulbous sac near the midline on the left lateral surface, a scale group on one surface; most of the remainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present on the dorsal surface. Measurements. (10 specimens). TL — 8.6 (8.2— 9.2); WH— 1.5 (1.5— 1.6); WP— 2.6 (2.5-2.7); WW— 3.3 (3.2-3. 5). Distribution. Southern United States north to Missouri. Localities. Alabama. Mobile Co.: Calvert (CAS), Mt. Vernon (OSU), Mobile (OSU); Tuscaloosa Co.: Tuscaloosa (GB). *Salco (OSU). Arkansas. Ashley Co.: Hamburg (CNC); Hempstead Co.: Hope (CU); Union Co.: Little Corne Bayou, Hy. 169 (UA). Florida. Volusia Co.: Enterprise (USNM). Indiana. Virgo Co.: Unrecorded lo- 148 Entomologica Americana [Vol. 46, No. 1 cality(PU). Louisiana. Franklin Co. : Chase (UA). Unrecorded locality in Louisiana (USNM) . Missouri. St. Louis Co.: St. Louis (CAS) . North Carolina. *Boardman (USNM). Tennessee. Unrecorded locality (USNM). Discussion. Loxandrus gibbus is most similar to L. erraticus in external apearances. It is usually smaller and positive identification can be made by examining the male genitalia. It has been collected in the months of February, March, May, July, August and September. The L. nitidulus , L. robustus , L. cincinnatiensis , L. minor , and L. circulus Complex This group of medium size Loxandrus species closely resemble one another in external appearances and are closely related phylogeneti- cally. They have in common all external characters, especially a more or less quadrate pronotum with distinct, narrowly rounded hind angles. The aedeagi of the five species are quite similar also. Loxandrus nitidulus is presently known from more specimens and more localities than the other four species. The aedeagus of this species is most like L. circulus but has one less scale group on the internal sac. L. nitidulus has been collected in the months of March, April, May, June, July and October. Loxandrus circulus is known from only eight specimens. Six of the specimens are from the Washington, D. C. area (1 marked Md., the remainder D. C.), one from Wood Co., Ohio, and one specimen was taken in Bibb County, Alabama. The Alabama specimen lacks a small scale group on the ventral surface of the IS of the aedeagus that lies between two large scale groups (Fig. 102 D, SA 1). It also has a strong scale group on the left dorso-lateral margin not present in three of the specimens from Washington (Fig. 102 A, SA 2). Three of the four Washington specimens have a small group of scales on the ventral surface of the sac of the aedeagus that lies between the two larger groups (Fig. 102 D, SA 1). One specimen from Washington appears to be intermediate between the one from Alabama and the three remaining Washington specimens by having the small group of ventral scales represented as several weakly sclerotized scales. The specimen from Wood Co., Ohio, has the ventral scale group SA 1 (Fig. 102 D) absent as well as a larger scale group SA 3 (Fig. 102 B) Figs. 175-176. Geographical distribution maps. 175. L. taeniatus LeC., L. minutus n. sp. 176. L. cervicalis Casey, L. algidus n. sp., L. extendus n. sp. 1972] Allen: Revision of Loxandrus LeConte 149 150 Entomologica Americana [Vol. 46, No. 1 on the right lateral surface of the sac. However, the scale group SA 2 (Fig. 102 A) is distinct in the Wood Co. specimen. Because of the small number of specimens available and because of the seemingly intermediate condition of two specimens, I have chosen to call this group one species. Further collecting will no doubt resolve the prob- lem. Loxandrus minor and L. cincinnatiensis differ by only one scale group on the IS of the male aedeagus. L. minor is known from a num- ber of localities in the central part of the United States. It has been collected in April. L. cincinnatiensis is known from a number of localities and appears to be confined to areas west of the Appalachian Mountains. It has been collected in the months of March, April, May, June and November. Loxandrus robustus is known from several localities in Arkansas and from one locality in Indiana. The scale groups and their arrange- ment on the IS of the aedeagus of this species are very characteristic and cannot be confused with other species in this complex. This species has been collected in May, July and August. The Arkansas speci- mens were found under leaf debris. 47. Loxandrus nitidulus (LeConte) Figs. 12; 49; 103 A, B, C; 161 Argutor nitidulus LeConte 1848:239. Holotype. 2. Type number 5696. First specimen in a series of 4 specimens. Second specimen is a male. Specimens 3 and 4 are females (MCZ). Description. Color. Dorsal: black, shining; elytra iridescent. Ventral: black, shining. Appendages: black to dark or reddish brown. Head. Clypeus : anterior margin slightly concave. Epistomal and subgenal sutures: finely produced, complete. Frontal sulci: obscure, broad, shallow, short. Microsculpture: clypeus, frons and vertex: isodiametric. Mentum: apex of lobes narrowly rounded; Figs. 177-178. Geographical distribution maps. 177. All known local- ities for all Loxandrus species in the Continental United States. Ad . . . Adirondak Mountains; Ap . . . Appalachian Mountains; Ar . . . Arbuckle Mountains; CA . . . Cincinnati Arch; CP . . . Cumberland Plateau; DA . . . Driftless Area; IO . . . Illinois Ozarks; NU . . . Northeastern Uplands; Ou . . . Ouachita Mountains; OP . . . Ozark Plateau; PP . . . Piedmont Plateau; WI . . . Wichita Mountains. After Ross, 1965. 178. All known localities for all Loxandrus species occurring in Middle America and the West Indies. 1972] Allen: Revision of Loxandrus LeConte 151 FIG. 178 152 Ent omologica Americana [Vol. 46, No. 1 Fig. 179. Geographical distribution of the genus Loxandrus throughout the world. tooth distinct, apex rounded; furrows present; fovae present or absent; microsculpture elongate. Thorax. Pronotum: microsculpture on disk absent, present along reflexed lateral margin, isodiametric; lateral margins evenly rounded, not sinuate posteriorly, sparsely punctate posteriorly; posterior angles subacute; anterior marginal sulcus com- plete; posterior marginal sulcus reaching slightly past basal furrows, incomplete; median sulcus reaching anterior sulcus but not posterior margin; basal impressions moderately deep, elongate, punctate; area between impressions punctate; area between impressions and lateral margins sparsely punctate. Scutellum: microsculpture isodiametric; apex pointed. Prosternum: microsculpture elongate; impunctate. Proepisternum: microsculpture elongate; punctate or not. Proepi- meron: microsculpture elongate if present; impunctate. Meso- sternum: microsculpture elongate; impunctate. Mesepisternum: microsculpture elongate; anterior three quarters punctate. Metaster- num: microsculpture absent or obscure; punctate laterally. Mete- pimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct, punctate, trough 2-3 isodiametric units wide; internal plica well developed; 1972] Allen: Revision of Loxandrus LeConte 153 humeri rounded; microsculpture on epipleura absent. Abdominal sternites. Microsculpture isodiametric laterally, becoming more elon- gate medially and on posterior segments and absent medially on segments 2-5; segments 1-3 sparsely punctate laterally; segments 1-5 weakly rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts in a median position, directed dorsally; 6 distinct scale groups; a band of scales in the basal half completely encircling the sac; most of the remainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present. Measurements. (20 specimens). TL— 9.1 (7.6-10.4); WH— 1.7 ( 1. 4-2.0) ; WP— 2.9 (2. 3-3. 3); WW— 3.6 (3.0-4.1). Distribution. Eastern United States, south to the Gulf Coast, north to Illinois and Indiana, west to Iowa and Arkansas. Localities. Alabama. Mobile Co.: Mobile (MCZ, OSU), Mt. Ver- non (CU, OSU) ; Tuscaloosa Co.: Tuscaloosa (GB). Arkansas. Calhoun Co.: Bayou Moro, Hy. 160 (UA). Unrecorded locality in Arkansas (MCZ, INHS). Florida. Unrecorded locality (MCZ). Illinois. Calhoun Co.: Kampsville (USNM, MCZ). ^Bureau (CAS). *Kahokia (MCZ). Unrecorded locality in Illinois (INHS, MCZ, MSU). Indiana. Elkhart Co.: Elkhart (CAS); Knox Co.: Unrecorded locality (CAS, PU); Viggo Co.: Unrecorded locality (CAS). Iowa. Johnson Co.: Iowa City (USNM). Unrecorded locality in Iowa (USNM). Louisiana. Union Co.: Farmerville (UA). Unrecorded locality in Louisiana (UM). Mississippi. George Co.: Lucedale (CU). Missouri. St. Louis Co.: St. Louis (INHS). Tennessee. Shelby Co.: Memphis (USNM). Discussion. See above. 48. Loxandrus robustus New Species Figs. 53; 118 A, B; 169 Holotype. S. Arkansas. Lonoke Co., Carlisle; Liebeck Collection (MCZ). Paratypes. 2 $ $ (MCZ); 1 $ (UM); 1 <5 (CU); 6 S $ (UA); 1 $ (PU). Description. Color. Dorsal: black, shining; elytra iridescent. Ventral: black or dark brown. Appendages: dark to reddish brown. Head. Clypeus : anterior margin slightly concave. Epistomal and sub- genal sutures: distinct, complete. Frontal sulci: obscure, shallow, short. Microsculpture: clypeus, frons and vertex distinct, isodiametric. Mentum: apex of lobes pointed; tooth distinct, apex rounded; furrows shallow; fovae absent; microsculpture isodiametric and elongate. Thorax. Pronotum: microsculpture on disk absent or very weak, distinct and isodiametric along the reflexed lateral margin; lateral margin evenly rounded, not sinuate posteriorly, punctate posteriorly; 154 Entomologica Americana [Vol. 46, No. 1 posterior angles narrowly rounded; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impressions but incom- plete; median sulcus reaching anterior sulcus but not posterior margin; basal impressions moderately deep, elongate, punctate; area between impressions sparsely punctate; area between impressions and lateral margins sparsely punctate or not. Scutellum: microsculpture isodia- metric; apex pointed. Prosternum: microsculpture isodiametric laterally, elongate medially. Proepisternum : microsculpture isodia- metric; impunctate. Proepimeron: microsculpture elongate; impunc- tate. Mesosternum: microsculpture slightly elongate; impunctate. Mesepisternum : microsculpture elongate; anterior three quarters punctate. Metasternum: microsculpture elongate where present; punctate laterally. Metepisternum: microsculpture elongate; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface weak, elongate. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture absent on intervals; all striae distinct, punctate, trough 2-3 isodiametric units wide; internal plica distinct; humeri rounded; microsculpture of epipleura isodiametric. Abdominal ster- nites. Microsculpture isodiametric laterally, elongate medially; segment 1 sparsely punctate laterally; segments 1-5 rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts medially and is directed dorsally; 4 distinct scale groups present; most of remainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present. Measurements. (5 specimens). TL — 9.6 (9.4— 9.8); WH— 1.7 (1.7-1. 8); WP— 2.9 (2.8-3.0); WW— 3.9 (3. 8-4.0). Distribution. Southern United States. Localities. Arkansas. Aransas Co.: White River National Wildlife Refuge, White River Chute Area (UA); Ashley Co.: 6 mi. W. Crossett (UA); Howard Co.: 4 mi. W. Mineral Springs, Rte. 27 (UA); Prairie Co.: Interstate 40 at White River Bridge (UA); Lonoke Co.: Carlisle (MCZ). Unknown localities in Arkansas (CU, MCZ). Indiana. Dubois Co. : Locality unknown (PU) . Discussion. See above. 49. Loxandrus cincinnatiensis Casey Figs. 52; 116; 168 Loxandrus cincinnatiensis Casey 1924:80. Holotype. $. Type No. 47355. (USNM). 1 female paratype. Description. Color. Dorsal: black, shining: elytra iridescent. Ventral: black or dark brown. Appendages: legs black or dark 1972] Allen: Revision of Loxandrus LeConte 155 brown; palpi and antennae dark to light brown. Head. Clypeus: anterior margin straight. Epistomal suture: finely produced, complete. Subgenal suture: obscure or absent. Frontal sulci: obscure, broad, shallow. Microsculpture: clypeus, frons and vertex isodiametric, weak. Mentum: apex of lobes rounded: tooth distinct, apex rounded; furrows present, shallow; fovae distinct; microsculpture elongate. Thorax. Pronotum: microsculpture on the disk absent, isodiametric along the reflexed lateral margins; lateral margins evenly rounded, not sinuate posteriorly, sparsely punctate posteriorly; posterior angles narrowly rounded; anterior marginal sulcus complete; posterior mar- ginal sulcus reaching past basal impressions but incomplete; median sulcus reaching anterior marginal sulcus or not, not reaching posterior margin; basal impression moderately deep, elongate, sparsely punctate; area between impressions sparsely punctate; area between impressions and lateral margins sparsely punctate or not. Scutellum : microsculp- ture isodiametric; apex pointed. Prosternum: microsculpture more distinct laterally, elongate; impunctate. Proepisternum : microsculp- ture elongate; obscurely punctate or not. Proepimeron: microsculp- ture elongate; impunctate. Mesosternum: microsculpture obscure, elongate; impunctate. Mesepisternum : microsculpture elongate; ante- rior three quarters punctate. Metasternum: microsculpture distinct laterally, elongate; punctate laterally. Metepisternum : microsculpture elongate; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner surface absent. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct, punctate, trough 3-4 isodiametric units wide; internal plica well developed; humeri rounded; microsculpture on epipleura absent. Abdominal sternites. Microsculpture isodiametric laterally, elongate or absent medially; segments 1 and 2 sparsely punctate laterally; seg- ments 1-3 weakly rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts towards the left, slightly directed ven- trally; 5 distinct scale groups present; the right lateral surface is al- most entirely covered by an ill-defined group of elongate scales; most of the remainder of the sac’s surface covered by small, weakly sclero- tized scales; sclerotized ring present. Measurements. (20 specimens). TL— 9.2 (8.1-10.1); WH— 1.7 (1.4— 1.9); WP— 2.9 (2.4-3. 3); WW —3.7 (3.2-4.1 ). Distribution. Eastern United States from the Gulf of Mexico north to Illinois, Ohio west to Missouri, Arkansas and Texas. Localities. Alabama. Mobile Co.: Mobile (OSU), Mobile or Mt. Vernon (MCZ) ; Tuscaloosa Co. : Tuscaloosa (GB). Arkansas. Jackson Co.: 5 mi. S. Newport (RTB); Pulaski Co.: Little Rock (MCZ). Illinois. 156 Entomologica Americana [Vol. 46, No. Alexander Co.: Cairo (CU); Champaign Co.: Urbana, Brownfield Woods (UA), Trelease Woods (RTB); Vermillion Co.: Kickapo State Park (RTB). Indiana. Knox Co.: Unrecorded locality (PU) . Kentucky. Mason Co.: Maysville (CU). Louisiana. Orleans Co.: New Orleans (MCZ); Plaquemines Co.: Narin (USNM). Unknown localities in Lou- isiana (UM, USNM). Mississippi. George Co.: Lucedale (CU). Missouri. St. Louis Co.: St. Louis (MCZ). Ohio. Hamilton Co.: Cin- cinnati (MCZ, UM, OSU). South Carolina. Unrecorded locality (USNM). Tennessee. Lake Co.: Reelfoot Lake (RTB). Shelby Co.: Memphis (USNM). Unknown locality in Tennessee (UM) . Texas. Dal- las Co.: Dallas (MSU). No state and no locality (MCZ). Discussion. See above. 50. Loxandrus minor (Chaudoir) Figs. 51; 115; 117; 167 Megalostylus minor Chaudoir 1843 : 766. Holotype. $ . The first specimen in a series of two specimens is a male and bears a red type label. Louisiana is given as the type locality. The second specimen in the series is a female from Texas. Description. Color. Dorsal: black, shining; elytra iridescent. Ventral: black to dark brown. Appendages: dark to light brown. Head. Clypeus : anterior margin slightly concave. Epistomal and sub- genal sutures: distinct, complete. Frontal sulci: obscure, broad, shallow, short. Microsculpture: clypeus, frons and vertex, isodia- metric; weak on the clypeus. Mentum: apex of lobes narrowly rounded; furrows absent; fovae deep, distinct; microsculpture elongate. Thorax. Pronotum: microsculpture on disk absent, distinct, isodia- metric along reflexed lateral margins; lateral margin evenly rounded in anterior half, oblique in posterior half, not sinuate; posterior angles subacute; anterior marginal sulcus complete; posterior marginal sulcus reaching past basal impressions but incomplete; median sulcus reaching anterior sulcus but not posterior margin; basal impressions moderately deep, elongate, punctate; area between basal impressions and between impressions and lateral margins sparsely punctate. Scutellum: micro- sculpture isodiametric; apex pointed. Prosternum: microsculpture elongate, more distinct laterally; impunctate. Proepistemum : micro- sculpture isodiametric; punctures weakly present or not. Proepimeron: microsculpture elongate and isodiametric; impunctate. Mesosternum: microsculpture elongate or absent; anterior three quarters punctate. Metasternum: microsculpture elongate if present; punctate laterally. Metepisternum : microsculpture elongate; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculp- 1972] Allen: Revision of Loxandrus LeConte 57 ture on inner surface absent distally, obscurely present (elongate) proximally. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on inter- vals absent; all striae distinct, punctate, trough 2-4 isodiametric units wide; internal plica distinct; humeri rounded; microsculpture of epi- pleura isodiametric. Abdominal sternites. Microsculpture distinct, isodiametric laterally, elongate medially; segments 1-3 obscurely punctate laterally; segments 1-4 rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts medially and is slightly turned to the left side; 6 distinct scale groups present; most of the remainder of the sac’s surface covered with small weakly sclerotized scales; sclerotized ring present. Measurements. (2 specimens). TL — 9.0 (8.4— 9.6); WH— 1.7 (1. 6-1.7); WP— 2.9 (2.8-3.0); WW- 3.7 (3.6— 3.7). Distribution. Southern United States. Localities. Arkansas. Jackson Co.: 5 mi. S. Newport (RTB); Hempstead Co.: Hope (MCZ). Illinois. Alexander Co.: Cairo (CU). Kansas. Douglas Co.: Lawrence (CAS). Louisiana. Unknown locality (UM). Discussion. See above. 51. Loxandrus circulus New Species Figs. 10; 50; 102 A, B, C, D, E; 159 Holotype. $ . District of Columbia. Roland Hayward, Collector (MCZ). Paratypes. 1 $ (GB); 1 S (USNM);2 $$ (INHS). Description. Color. Dorsal: black, shining; elytra iridescent. Ventral: black, shining. Appendages: black to dark or reddish brown. Head. Clypeus: anterior slightly concave. Epistomal sulcus: distinct, complete. Subgenal suture: obscure, finely produced. Frontal sulci: obscure, broad, shallow, short. Microsculpture: clypeus, frons and vertex isodiametric. Mentum: apex of lobes rounded; furrows and fovae absent; microsculpture isodiametric. Thorax. Pronotum: mi- crosculpture on disk usually absent, elongate if present; lateral margins evenly rounded, not sinuate posteriorly, sparsely punctate posteriorly; posterior angles subacute; anterior marginal sulcus complete; posterior marginal sulcus reaching to just past basal impressions; median sulcus reaching anterior marginal sulcus but not posterior margin; basal im- pressions moderately deep, elongate, punctate; area between impres- sions sparsely punctate; area between impressions and lateral margins impunctate. Scutellum: microsculpture isodiametric; apex pointed. Prostemum: microsculpture elongate medially, isodiametric laterally; 158 Entomologica Americana [Vol. 46, No. 1 impunctate. Proepisternum : microsculpture elongate; punctate. Pro- epimeron: microsculpture elongate; impunctate. Mesosternum: microsculpture elongate; impunctate. Mesepisternum: microsculpture absent; anterior three quarters punctate. Metasternum: microsculpture usually absent, elongate if present; punctate laterally. Metepisternum : microsculpture absent; punctate. Metepimeron: microsculpture absent; impunctate. Legs. Hind coxae: microsculpture on inner sur- face absent, elongate and/or isodiametric. Tarsi: segments 1-3 of middle and hind tarsi and 4 of hind tarsi grooved dorso-laterally. Elytra. Microsculpture on intervals absent; all striae distinct, punc- tate, trough 2-3 isodiametric units wide; internal plica well developed; humeri weakly dentate; microsculpture on epipleura absent. Abdom- inal sternites. Microsculpture distinct, isodiametric laterally, elongate medially; segments 1-3 punctate laterally; segments 1-6 rugose laterally. Aedeagus. Median lobe: not modified. Internal sac: everts medially, directed dorsally; 8 or 9 distinct scale groups present; most of the remainder of the sac is covered with small weakly sclerotized scales; sclerotized ring present; see discussion for comments on varia- tion. Measurements. (5 specimens). TL — 9.1 (8.8-9.9);WH — 1.7 (1.7— 1.9); WP— 2.9 (2.8-3. 3); WW— 3.6 (3.4-4.0). Distribution. Eastern United States, Washington, D. C., south to Alabama. Localities. Alabama. Bibb Co. : Unrecorded locality (GB). District of Columbia. Washington (CNC, INHS, MCZ, USNM). Maryland. Un- recorded locality (UM). Ohio. Wood Co.: Unrecorded locality (PU) . Discussion. See above. 52. Loxandrus velox (Dejean) Feronia velox Dejean 1828:245. Lectotype, here designated. The first specimen, a male, in a series of four specimens in the Dejean collection in the Paris Museum. The first specimen bears a small green label. Le- Conte is given as the collector. Three additional specimens, two females and one male, are in the series. Description. Color. Dorsal and ventral: black. Legs: dark brown. Antennae: dark brown. Palpi: dark to light brown. Head. Clypeus: weakly concave. Epistomal and subgenal sutures: not visible. Frontal sulci: obscure, short, shallow. Mentum: apex of lobes rounded; tooth not visible; furrows and fovae absent. Thorax. Pro- notum: microsculpture if present weak and elongate; lateral margins not visible; posterior angles rounded; anterior marginal sulcus com- plete; posterior marginal sulcus reaching past basal impressions but 1972] Allen: Revision of Loxandrus LeConte 159 not complete; median sulcus weak, reaching anterior marginal sulcus but not posterior margin; basal impressions moderately deep, elongate; entire pronotum impunctate. Ventral surface: impunctate; micro- sculpture mostly absent but where visible, elongate. Scutellum: apex acutely pointed. Legs. Hind coxae: microsculpture on inner surface appears to be absent. Elytra. Microsculpture on intervals absent; all striae, except base of 7, distinct, punctate; trough appears to be 1 iso- diametric unit wide; internal plica distinct; humeri rounded. Abdom- inal sternites. Microsculpture not visible; segments 1-2 weakly rugose basally; impunctate. Aedeagus. Median lobe: apex indented. In- ternal sac: everts in a median position; 7 scale groups present; a secondary sac everts apically; sclerotized ring present on the dorsal surface. Measurements. None. Distribution. “Boreal America” (MNHP). Discussion. See above. POSTULATED ANCESTRAL AND DERIVED STATES OF CHARACTERS IN THE GENUS LOXANDRUS There are several basic tenets that must be followed in working out the phylogenetic relationships of a group of organisms. The fol- lowing list is a brief summary of the more important tenets that should be followed. These tenets and the basic philosophy relating to them are discussed in greater detail by Ross (1937, 1956, 1959, 1965, 1967) andHennig (1966). (1) The derived and ancestral states of at least some characters are postulated after a study of taxa believed to be closely related to the taxon under consideration. (2) Utilizing these characters a hypothetical ancestor is postulated for the group with which one is working. (3 ) It must be assumed that derived states of characters which are found in the group of organisms under study came about through a logical and orderly progression from ancestral to derived. (4) The various species of the group are compared with one another and those species sharing derived characters in common are tentatively assumed to have risen from a common ancestor. (5) On a strictly logical basis, species are grouped into distinctive phylogenetic lines arising from a common ancestor only on the basis of shared characters. (6) Derived characters must originate from ancestral characters and not the opposite. (7) One must strive to account for all the derived conditions of the characters the least number of times. 60 Entomologica Americana [Vol. 46, No. As mentioned previously, in order to ascertain the ancestral and derived states of various characters in Loxandrus it was necessary to determine the state of the characters in closely related groups. The closely related groups utilized in this study are listed below. A detailed diagnosis of each group can be found in Allen, 1969. Tribe AMARINI Genus Amara Bonelli 1809 A. cupreolata Putz. 1866 A. impuncticollis Say 1823 Tribe AGONINI Genus Calathus Bonelli 1809 C. ( Calathus ) opaculus LeC. 1854 Genus Agonum Bonelli 1809 A. ( Circinalia ) pallipes Casey 1920 A. ( s . str .) decorus (Say) 1823 A. (s. str.) extensicollis (Say) 1823 A. (s. str.) punctijormis (Say) 1823 Tribe PTEROSTICHIN1 Genus Stomis Schellenberg 1806 S. pumicatus Panzer 1796 Genus Trigonognatha Motschoulsky 1857 T. cor acinus (Say) 1825 Genus Abaris Dejean 1831 A. aequenoctialis Chaudoir 1852 Genus Oxycrepis Reiche 1843 O. intercepta Chaudoir 1873 Genus Piesmus LeConte 1852 P. monedulus German 1824 Genus Dyschromus Chaudoir 1835 D. nitidipennis Putzeys 1846 Genus Pterostichus Bonelli 1809 P. ( Cylandrocharis ) rostrata (Newman) 1838 P. ( Cylandrocharis ) grandiceps (LeConte) 1848 P. ( Holciophorus ) lama Mentries 1844 P. ( Pherypes ) tarsalis LeConte 1873 P. ( Hypherpes ) californicus (Dejean) 1828 P. ( Hypherpes ) congestus Menetres 1844 P. ( Hypherpes ) validus Dejean 1831 P. ( Hypherpes ) brunneus Dejean 1831 P. ( Monoferonia ) diligendus (Chaudoir) 1866 P. ( Gastrellaris ) honestus (Say) 1825 1972] Allen: Revision of Loxandrus LeConte 161 P. ( Poecilus ) chalcites Say 1823 P. ( Poecilus ) lucublandus Say P. ( Poecilus ) occidentalis Dejean P. ( Poecilus ) texanus LeConte 1863 P. ( Bothiopterus ) pennsylvanicus (LeConte) 1863 P. ( Bothiopterus ) adst rictus (Eschscholtz) 1823 P. ( Bothiopterus ) sp. P. ( Lophoglossus ) sp. P. ( Euferonia ) coracina Newman 1838 P. ( Euferonia ) sp. P. ( Refonia ) moesta (Say) 1823 P. ( Metamelanius ) ebeninus Dejean 1828 P. ( Melanius ) caudicalis Say 1823 P. ( Melanius ) corvinus Dejean 1828 P. ( Melanius ) luctuosus Dejean 1828 P. ( Dysidius ) purpuratus LeConte 1852 P. ( Dysidius ) mutus Say 1823 P. ( Argutor ) sp. P. ( Abacidus ) hamiltoni (Horn) 1880 P. ( Abacidus ) permundus (Say) 1834 P. ( Abacidus ) sculptilis (LeConte) 1852 P. ( Gastrosticta ) subacuta Casey 1918 P. (^. Air.) herculaneus Mannerheim 1843 P. (a. Air.) mandibularis Kirby 1850 P. (a. Air.) adoxus Say 1825 P. (a. Air.) stygicus Say 1823 P. (a. Air.) relictus Newman 1838 Genus Evarthrus LeConte 1852 E. sordalis LeConte 1848 E. seximpressus LeConte 1848 E. nonnitens LeConte 1873 E. incisus LeConte 1873 Armed with the information concerning the state of characters in groups closely related to the genus Loxandrus and using the philo- sophical principles and basic tenets previously set forth it is now possible to postulate the ancestral and derived state of some characters found in Loxandrus. The external surface of all areas and sclerites usually has a charac- teristic microsculpture in groups related to Loxandrus. This micro- sculpturing can be an isodiametric pattern (Figs. 11 A, B; 16 A, B, C; 20 A, B, C; 21 ) or an elongate pattern (Figs. 13; 14; 15; 17; 18 A, B; 19 A, B, C; 22), the latter consisting of short fine parallel lines. In either case it is distinct. The prosternum, proepisternum, proepimeron, mesepisternum, metasternum, metepisternum, and metepimeron usu- ally have microsculpturing of the elongate type. This elongate type 162 Entomologica Americana [Vol. 46, No. that is distinct is considered the ancestral state for these sclerites in Loxandrus. If the microsculpture is isodiametric, reduced or absent in species of Loxandrus it is considered derived. The head, mesosternum, pronotum, scutellum, and elytra usually have microsculpturing of the isodiametric type. This is considered the ancestral state for these morphological parts and an elongate micro- sculpture or the absence of microsculpturing is considered derived for these areas in Loxandrus. The sclerites on the ventral surface of the thorax can be either punctate or impunctate. In the majority of the groups related to Loxandrus the following condition exists: (1) the prosternum, pro- episternum, proepimeron, mesosternum, mesepimeron, and metepi- meron are impunctate (Figs. 20 A, B, C); (2) the mesepisternum (anterior % to %), and the lateral area of the metasternum are punctate (Figs. 19 A, B, C). Therefore, in the first group of six sclerites, the impunctate state is considered ancestral and a punctate state is derived, and in the second group of two sclerites the punctate state is considered ancestral and an impunctate state derived in Loxandrus. In most of the genera and subgenera examined the intercoxal proc- ess of the prosternum is not margined. The unmargined state of the intercoxal process is considered ancestral and the margined state is considered derived in Loxandrus. The color of most of the species examined was entirely black or dark brown. When another color other than black or dark brown is encountered among the species of Loxandrus it is considered derived. In most genera and subgenera closely related to Loxandrus the internal plica on the lateral, apical margin of each elytron is present and well developed. This state is considered ancestral. When the internal plica is reduced or absent among species of Loxandrus it is considered a derived state of the character. The internal sac of the aedeagus of the males in the genera and subgenera closely related to Loxandrus has various structures present or absent, i.e. scales, scale groups, and/or sclerites. These various arrangements can be divided into the following five generalized divi- sions: (1) the surface of the membraneous sac can be completely devoid of scales, scale groups and sclerites; (2) the surface of the membraneous sac can have scales covering varying amounts of the surface but no distinct scale groups or sclerites (Fig. 82 A); (3) the surface of the membraneous sac can be devoid of scale groups but have certain areas sclerotized (Fig. 94 A); (4) the surface of the membraneous sac can be devoid of sclerotized areas but possess scale groups (Fig. 10); (5) the surface of the membraneous sac can have a combination of sclerotized areas and scale groups (Fig. 10). 1972] Allen: Revision of Loxandrus LeConte 163 The information derived from the study of genera and subgenera closely related to Loxandrus indicates that the ancestral state of the internal sac of the aedeagus of male Loxandrus was a sac with scales present and sclerites absent. Whether or not the scales were in well defined groups is not clear. In some species of Loxandrus there occurs a sclerotized ring, usually near the apex of the internal sac. I believe that this ring is associated with the gonopore because in some species I have been able to trace the ejaculatory duct to this ring and I believe it is connected to the ring (Fig. 10 - SR). I have not been able to find the gonopore in those species of Loxandrus without a sclerotized ring. Therefore I have some reservations as to what the true relationship of the sclero- tized ring is to the ejaculatory duct, i.e. sclerotized gonopore or not. Among those groups I have studied the sclerotized ring is a unique, derived adaptation in Loxandrus. In most of the genera and subgenera examined the tooth of the mentum is short and not as long as the lobes of the mentum. In some species of Loxandrus the lobes of the mentum are short and the tooth is as long as the lobes or almost as long as the lobes. This gives the mentum a transverse appearance (Figs. 138; 139; 140). This trans- verse state is considered the derived condition in Loxandrus. In all of the groups studied ambulatory setae were absent on the second abdominal sternite. This is considered the ancestral condition. When ambulatory setae do occur on the second abdominal sternite it is considered a derived condition. It should be noted here that Loxandrus species from other parts of the world have been included where material was available for study. These species have not been incorporated into the key nor has a description been provided. These species and others will be treated in subsequent papers. The following is a list of species represented in the phylogeny but not treated in the taxonomic section of this paper. Their general area of distribution is indicated. Loxandrus longifonnis Sloane 1898 Australia Loxandrus amplicolis Sloane 1903 Australia Loxandrus subiridescens M’Leay 1869 Australia Loxandrus australiensis Sloane 1895 Australia Loxandrus rufilabris Castleneau 1867 Australia Loxandrus atronitens M’Leay 1869 Australia Loxandrus major Darlington 1962 New Guinea Loxandrus medius Darlington 1962 New Guinea Loxandrus latus Darlington 1962 New Guinea Loxandrus subrectus Darlington 1962 New Guinea Loxandrus negrei Straneo 1964 South America Loxandrus tucumanus Dejean 1831 South America Loxandrus dubius Curtis 1839 South America 164 Entomologica Americana [Vol. 46, No. 1 THE PHYLOGENETIC RELATIONSHIPS AMONG THE SPECIES OF LOX ANDRUS The following discussion annotates Figures 180, 181 and 182. Based on the information concerning the ancestral and derived states of various characters in the genus Loxandrus it is possible to postulate certain characteristics that the progenitor of this group probably pos- sessed as follows. The microsculpture of prosternum, proepisternum, proepimeron, mesepisternum, metasternum, and metepimeron was distinct, fine, elongate parallel lines (Figs. 19 A, B, C). Microsculp- ture of head, mesosternum, pronotum, scutellum, and elytra was dis- tinct, isodiametric (Figs. 16 A, B, C; 20 A, B, C). The prosternum, proepimeron, mesosternum, mesepimeron, and metepimeron were impunctate (Figs. 20 A, B, C). The mesepisternum (anterior % to %) and the lateral areas of the metasternum were punctate (Figs. 19 A, B, C). The trough of the striae on the elytra were 1-2 isodiametric units wide (Figs. 11 A, B; 13). The intercoxal process of the prosternum was not margined. The mentum was not transverse and did not possess fovae. The second abdominal sternite lacked ambulatory setae. The color was a uniform black or dark brown. The internal sac of the aedeagus was devoid of sclerites and of a sclerotized ring toward the apex (Fig. 9). The sac probably possessed scales on the surface but there was only one defined scale group, if any. The internal plica near the apex of the elytra was present and well developed. The progenitor of Loxandrus is represented by ancestor 1 . Three lineages seem to have arisen from ancestor 1. Loxandrus infimus , one lineage arising from ancestor 1, appears to have changed little from the ancestor. The second lineage arising from ancestor 1 gave rise to ancestor 2 in which the microsculpture on the elytra became reduced to fine elongate lines or became altogether absent (Figs. 13, 14), a character which has apparently evolved twice in Loxandrus as we shall presently see. The third lineage arising from ancestor 1 gave rise to ancestor 1 1 in which a ring process near the apex of the internal sac of the aedeagus began to develop (Figs. 91 A; 93 A). Ancestor 2 gave rise to three lineages. Loxandrus elongatus ap- pears to have changed little from the ancestor. Loxandrus dubius is the end result of a second lineage arising from ancestor 2 in which two sclerites developed on the internal sac of the aedeagus (Figs. 73 B - Sl, S2). The third lineage arising from ancestor 2 resulted in ancestor 3 in which the microsculpture on the elytra was entirely absent. -> Figs. 180-182. Hypothetical phylogeny of Loxandrus species. Three parts. 1972] Allen: Revision of Loxandrus LeConte 165 i to ancestor II 166 Entomologica Americana [Vol. 46, No. i LONGIFORMiS 1972] Allen: Revision of Loxandrus LeConte 167 168 Entomologica Americana [Vol. 46, No. 1 Loxandrus nocticolor and Loxandrus rectangulus are two existent species which appear to have changed little from ancestor 3. Ancestor 3 also gave rise to a lineage in which fovae on the mentum were devel- oped (Figs. 138, 139, 140). Ancestor 4 is represented in our present day fauna by Loxandrus celeris and Loxandrus cubanus. These two species appear to have changed little from ancestor 4. Ancestor 4 also gave rise to a lineage in which the microsculpture on the surface of all ventral sclerites became greatly reduced if not absent and which re- sulted in ancestor 5. Ancestor 5 is represented in our present day fauna by Loxandrus subiridescens and Loxandrus remotus. These two species appear to have changed little from ancestor 5. Ancestor 5 also gave rise to a lineage in which the microsculpture on the pronotum was lost and which resulted in ancestor 6. Ancestor 6 is represented in our present day fauna by Loxandrus subrectus and Loxandrus latus. These two species appear to have changed little from ancestor 6. Ancestor 6 also gave rise to a lineage which resulted in ancestor 7 in which a unique character among Lox- andrus appeared. The character was the development of orange or dark yellow spots on each elytron, one near the humeri and one towards the apex (Fig. 2 A). Ancestor 7 gave rise to at least four and possibly five phylogenetic lineages. Loxandrus spinigrandis , Loxandrus tropicus , and Loxandrus lepidus are the end results of three of these lineages and appear to have changed little from the ancestor. Loxandrus discolor , a fourth lineage that may have arisen from ancestor 7, also possesses most of the ex- ternal characters attributed to ancestor 7, but the internal sac of the male aedeagus (Fig. 82 A) is very similar to Loxandrus whiteheadi (Figs. 86 A, B). Occasional specimens of Loxandrus discolor also have ambulatory setae present on sternite 2 of the abdomen, a char- acter which apparently arose one other time in a far-distantly related Loxandrus group. The phylogenetic position of Loxandrus discolor is thus questionable at this time. A fifth lineage arising from ancestor 7 in which the punctures on the mesepisternum were lost resulted in ancestor 8 (Fig. 20 B). Ancestor 8 seems to have given rise to two evolutionary lines. Loxandrus whiteheadi is the end result of one of these lines and ap- pears to have changed little from its ancestor. In the second line arising from ancestor 8 the punctures on the metepisternum (Fig. 20 A) were lost, resulting in ancestor 9. Ancestor 9 appears to have given rise to 3 phylogenetic lineages. Loxandrus balli, one line arising from ancestor 9, is a species that seems to have changed little from the ancestor. Loxandrus elnae arose 1972] Allen: Revision of Loxandrus LeConte 169 from ancestor 9 and has one notable external character that is different from its hypothetical ancestor. In Loxandrus elnae the spots on each elytron have become elongate, thus forming lines near the lateral margins of the elytra. The internal sac of the aedeagus of Loxandrus elnae must also be mentioned (Figs. 90 A, B). It is quite similar to Loxandrus floridanus (Figs. 122 A, B), a distantly related species. This similarity is very striking but I feel that at the present time the evidence indicates that the common ancestry of Loxandrus elnae and Loxandrus floridanus is not very close. In the third line arising from ancestor 9 there developed a small group of scales near the apex of the internal sac that resulted in ancestor 10 (Figs. 87 A; 89). Ancestor 10 gave rise to two lineages. One lineage resulted in Loxandrus fraus and the other lineage resulted in Loxaudrus unispinus. These two species appear to have changed little from their ancestor. Thus far, none of the species discussed have possessed a sclerotized ring near the apex of the internal sac. The third phylogenetic line arising from ancestor 1 resulted in a short series of species that illus- trate what I believe to be the gradual development of this sclerotized ring. Hypothetical ancestor 1 1 and 3 species of our present day stock represent the first step in the development of the sclerotized ring. Ancestor 11 gave rise to several phylogenetic lineages. In one lineage arising from ancestor 11 the ring on the internal sac of the aedeagus became larger and was weakly sclerotized but was not com- pletely enclosed inside the sac of the aedeagus (Fig. 94 A - Rl) and the microsculpture on the elytra became reduced or obscure, fine, parallel lines (Figs. 13, 14, 15). This lineage resulted in ancestor 12. The remaining lines that arose from ancestor 1 1 resulted in Loxandrus pactinullus , Loxandrus negrei , and Loxandrus sculptilis. The ring on the internal sac of the aedeagus of males of the above three species is turned inward and covered by the membraneous sac (Figs. 91 A; 93 A - Rl). I have been unsuccessful in trying to force the ring to the surface without destroying the sac. The ring is weakly sclerotized if at all. It does have enough rigidity that when compressed with a pair of forceps it will return to its original shape. The median lobe and internal sac of the aedeagii of the males of the three species under discussion show no other distinctive morphological characters. The mentum in these three species is transverse. That is, the tooth is as long as or almost as long as the lobes of the mentum. This is ap- parently a derived character state. The remainder of the external morphology is rather nondescript. Thus, it is presently impossible to discuss any further the relationship of the species Loxandrus pacti- nullus, Loxandrus negrei , and Loxandrus sculptilis. 170 Entomologica Americana [Vol. 46, No. 1 Ancestor 12 gave rise to two lineages. Loxcmdrus tucumanus ap- pears to have changed little from its ancestor. In a second line arising from ancestor 12 that resulted in ancestor 13 the ring on the internal sac of the aedeagus completed its development into a form that is common today in many of our Loxcmdrus species. The ring was fully sclerotized and not enclosed by the internal sac (Fig. 95 A - R1 ). Ancestor 13 gave rise to three lineages. Loxandrus longiformis , a species representing one lineage arising from ancestor 13, appears to have changed little from its hypothetical ancestor. The aedeagus of this species is unknown to me and its phylogenetic position may change when a male specimen is studied. In a second lineage arising from ancestor 13 the posterior angles of the pronotum became rounded (Figs. 51, 52). This character appears to have evolved several times in Loxandrus. This lineage resulted in ancestor 22. In the third lineage arising from ancestor 13 the microsculpture on the proepisternum became reduced or absent resulting in ancestor 14. Ancestor 14 apparently gave rise to 3 distinct lineages and two species that have changed little from the ancestor. Loxandrus gibbus and Loxandrus erraticus are the two species that are similar to their ancestor. In another lineage arising from ancestor 14 the microsculp- ture on the proepisternum was entirely lost and resulted in ancestor 17. In a second distinct lineage arising from ancestor 14 the hind angles of the pronotum became rounded (Figs. 43 A, B; 44 A, B). This evolutionary pathway resulted in Loxandrus rectus and Loxandrus pravitubus. Both species have a characteristic scale pattern on the internal sac but I am unable to ascertain which of these two patterns represents the ancestral condition. In the third lineage arising from ancestor 14 the scale groups on the surface of the internal sac of the aedeagus became arranged in a characteristic pattern and gave rise to ancestor 15. Hypothetical ancestor 15 is characterized by the development of two elongate groups of scales present on the ventral, basal portion of the internal sac of aedeagus (Fig. 102 D - SA2). This ancestor, an- cestor 15, gave rise to two phylogenetic lineages. In one lineage the resulting species changed little from their ancestor. These species are Loxandrus velocipes and Loxandrus inferus. In the second lineage arising from ancestor 15 there developed a group of scales on the ventral surface in the center of the apical half of the sac (Fig. 101 C - SA1 ) which resulted in ancestor 16. Ancestor 16 gave rise to one species that differs very little from it, Loxandrus brevicollis , and one distinct phylogenetic lineage. In the distinct phylogenetic lineage arising from ancestor 1 6 there developed a wide ring of small scales completely encircling the basal half of the 1972] Allen: Revision of Loxandrus LeConte 171 internal sac and a small group of elongate scales near the sclerotized ring of the internal sac (Figs. 102 A, B). These developments re- sulted in Loxandrus nitidulus and Loxandrus circulus. Ancestor 17, which we have previously noted as arising in a lineage emanating from ancestor 1 4 through the loss of the microsculpture on the prosternum, gave rise to several phylogenetic lineages. Four spe- cies arose from ancestor 17 and appear to have changed very little from it. The four species are Loxandrus amplicolis, Loxandrus cre- natus, Loxandrus saphyrinus and Loxandrus lucens. I have not seen a male of Loxandrus amplicolis and I place the species here only tenta- tively. It should also be noted that the sclerotized ring of Loxandrus crenatus is very different from the other Loxandrus species in this group (Fig. 104 - Rl). In another line arising from ancestor 17 the in- tercoxal process of the prosternum became margined resulting in an- cestor 21. In the last lineage arising from ancestor 17 there developed on each side of the ventro-lateral surface in the basal quarter of the in- ternal sac a round or elongate group of scales that resulted in ancestor 18 (Figs. 108 C; 110 B; 109 C, D, E, F - SA1). We have previously noted a similar group of scales arising in a lineage arising from ancestor 14 and resulting in ancestor 15. In this lineage, however, the group was always elongate. Ancestor 1 8 gave rise to two lineages. The first lineage resulted in Loxandrus uniformis , a species that apparently changed little from its ancestor. In the second lineage arising from ancestor 18 the basal, ventral scale group on the left side of the sac lost its elongate form by becoming larger and more oval in form (Fig. 108 C - SA1) resulting in ancestor 19. Ancestor 19 also gave rise to two lineages. One lineage, repre- sented in our present day fauna by Loxandrus micans , seems to have changed little from its ancestor. In the second lineage arising from ancestor 19 there developed an additional group of scales on the ventral surface in the center of the basal half of the internal sac (Figs. 109 E; 110C-SA2). This lineage resulted in ancestor 20 which in turn gave rise to Loxandrus duryi and Loxandrus vitiosus. These spe- cies have characteristic scale patterns on the internal sac but I am unable to discern the ancestral state. Ancestor 21, which arose from a phylogenetic line emanating from ancestor 17 previously discussed, gave rise to several phylogenetic lineages. Four of these lineages resulted in Loxandrus rufilabris, Lox- andrus atronitens, Loxandrus major , and Loxandrus medius. In a fifth lineage arising from ancestor 21 the punctures on the pronotum were lost. This resulted in Loxandrus australiensis. The internal sacs of the aedeagii of Loxandrus major (Fig. Ill B), 172 Entomologica Americana [Vol. 46, No. 1 Loxandrus rufilabris (Fig. 112) and Loxandrus medius (Figs. 114 B, C), originating from ancestor 21 (a male is unknown for Loxandrus amplicolis and therefore questionable) are characteristically devoid of distinct organized scale groups. This apparently represents a lost character after the main evolutionary stream of Loxandrus had de- veloped characteristic scale groups on the internal sacs. Ancestor 22 arose from ancestor 12 through the development of rounded posterior angles of the pronotum as mentioned previously. It should be noted here that this is the second time this adaptation has occurred in Loxandrus. Previously it occurred in a lineage arising from ancestor 14 and resulting in Loxandrus rectus and Loxandrus pravitubus. Ancestor 22 gave rise to at least two and possibly three lineages. Loxandrus yeariani is a species that has apparently changed little from the hypothetical ancestor 22 from which it arose. Its placement here in the phylogeny is tentative because it possesses some characters that would suggest other relationships. A second line arising from ancestor 22 resulted in ancestor 23. In this line there developed two small parallel groups of scales on the dorsal surface of the sac at the base (Figs. 116; 117; 118 A, B-SA1). The third line arising from an- cestor 22 resulted in ancestor 24. In this lineage there developed a tendency for ambulatory setae to be present on the second abdominal sternite. This character is expressed in many specimens of the resulting species but not in all. Ancestor 23 apparently gave rise to three species, Loxandrus minor , Loxandrus cincinnatiensis and Loxandrus robustus , that have changed little from the ancestor. Ancestor 24 gave rise to two lineages. In one lineage punctures on the metepisternum, and mesepisternum were lost (Figs. 20 A, B). This resulted in ancestor 26. In the second lineage arising from an- cestor 24 there developed a secondary sac on the left lateral surface of the primary sac of the males (Figs. 120 A, B; 121 A, C - SS). This resulted in ancestor 25 which in turn gave rise to two species repre- sented in our present day fauna, Loxandrus saccisecundaris and Lox- andrus parallelus. These two species have evidently changed little from hypothetical ancestor 25. Ancestor 26, which arose from ancestor 22 through the loss of punctures on the mesepisternum and metepisternum, gave rise to several phylogenetic lines. In one line arising from ancestor 26 there devel- oped on the left lateral surface of the internal sac a characteristic group of scales. In some species the group is elongate (Fig. 122 B - SA1) while in others it is more rounded (Fig. 127 A - SA 1). The group does, however, appear to have the same general position in all the 1972] Allen: Revision of Loxandrus LeConte 173 species evolving past this point in the phylogeny. This evolutionary pathway led to ancestor 29. In a second lineage arising from ancestor 26 the median lobe of the aedeagus became modified and resulted in ancestor 27. In addition to these two lineages there arose from an- cestor 26 two species that have apparently changed little from their ancestor. These two species are Loxandrus floridanus and Loxandrus unilobus. Ancestors 27 and 28 and the three species resulting from them, Loxandrus velox, Loxandrus aduncus and Loxandrus agilis, represent a phylogenetic sequence in which the median lobe of the aedeagus be- came highly modified. Based on the general appearance of the mor- phology I have arranged these three species in what appears to be the evolutionary sequence beginning with the simplest, Loxandrus velox (Figs. 134, A, B), an intermediate form, Loxandrus aduncus (Figs. 124 A, B), and the most complex, Loxandrus agilis (Fig. 125). The reader is cautioned that this is the only place in this phylogeny in which this kind of criterion is used. Note also that the ambulatory setae on the second abdominal sternite in Loxandrus velox have not been verified. Ancestor 29 which arose from ancestor 26, as previously discussed, gave rise to two distinct phylogenetic lineages and two additional species. The two species Loxandrus rossi and Loxandrus taeniatus would appear to have changed little from their ancestor. In another lineage arising from ancestor 29 there developed on the internal sac a crescent-shaped group of scales on the lateral surface of the left side resulting in ancestor 32 (Figs. 132 A; 133 B - SA1). Ancestor 29 also give rise to yet another lineage in which developed one elongate group of scales in the apical % of the internal sac near the sclerotized ring, more or less on the ventral surface (Figs. 129 C; 130 C; 131 A - SA1). This resulted in ancestor 30. Ancestor 30 gave rise to two lineages. One lineage is represented by Loxandrus minutus, a species that appears to have changed little from its ancestor. In the second lineage arising from ancestor 30, additional groups of scales developed near the sclerotized ring on the internal sac of the aedeagus and this resulted in ancestor 31 (Figs. 130 A, B, D; 131 A,B-SA2). Ancestor 3 1 gave rise to two species Loxandrus cervicalis and Lox- andrus extendus. The former species, Loxandrus cervicalis , appears to have changed very little from its ancestor while in the latter species, Loxandrus extendus , the lateral margins of the metasternum have become punctate (Fig. 19 C). Ancestor 32 which arose from ancestor 29 as previously dis- cussed gave rise to two phylogenetic lineages. One lineage resulted in 174 Entomologica Americana [Vol. 46, No. 1 Loxandrus algidus, a species that seems to have changed little from its ancestor. In the second lineage arising from ancestor 32 there was an addition of a large group of scales that is partially enclosed by the crescent- shaped scale group (Fig. 132 A-SA2). This evolutionary line resulted in Loxandrus spinilunatus. Based on the information I now possess, I believe that the above discussion is the one most plausible concerning the evolutionary history of the species comprising the genus Loxandrus. ZOOGEOGRAPHY At the present time there appear to be two general schools of thought concerning methods to be employed in the study of the zoo- geography of animals. One group of zoogeographers tends to utilize the “working principles” set forth by Darlington (1957). A second group of zoogeographers places more emphasis on first studying the phylogenetic relationships of taxa. Once these phylogenetic relation- ships are postulated the present day geographical ranges and data from historical geography are utilized to form hypotheses concerning the past dispersals of the organisms. W. H. Henig and H. H. Ross are two lead- ing proponents of the second method. In the following discussion essentially the latter approach has been followed. Two factors greatly hinder a discussion of the historical zoogeog- raphy of Loxandrus. These are ( 1 ) an inadequate knowledge of the distribution of most species, and (2) the sympatric distribution of many species, especially species pairs. Loxandrus in North America. Current data concerning phylogenetic relationships and distribution patterns, indicate that there are two basic faunal groups of Loxandrus in North America. One fauna is found primarily in the Southeastern United States while the second fauna is found primarily in the Middle American- West Indies region of North America. A few species are common to both areas. A study of phylogeny and distribution patterns as presented in Fig. 183 indicates the following sequence of events in the past dispersal of the genus Loxandrus. Those species that are more closely related to the hypothetical ancestor of the genus Loxandrus (ancestor 1) are now found in Middle America and in South America — L. infimus , L. dubius , L. eiongatus , L. sculptilis , L. negrei , L. pactinullus and L. tucumanus. The fauna now endemic to the Southeastern United States apparently arose from hypothetical ancestor 12 and resulted in ancestor 13. Ancestor 13 gave rise to three lineages that resulted in ancestor 14, ancestor 22, and one existent species (L. longiformis known only from Australia). 1972] Allen: Revision of Loxandrus LeConte 175 a ■ a a ■ ■ a a ■ >a a 1^ |, IId?l§sll5lll»ilt3l 5Ii?3ilili513imi!II3ISS§5 illlSilUSilissIsS M UJ Fig. 183. Hypothetical phylogeny of Loxandrus species and their general geographical ranges. Aust . . . Australia; MA . . . Middle America; USA . . . United States of America; WI . . . West Indies. Ancestor 14 gave rise to several lineages that eventually resulted in 16 species endemic to the Southeastern United States. Ancestor 14 also gave rise to a lineage, via ancestor 17, that apparently colonized Australia. This lineage evolved into 6 distinct species endemic to the Australian continent. Ancestor 22, which as previously noted arose from ancestor 14, gave rise to at least two phylogenetic lineages and possibly three. The two evident evolutionary lines arising from ancestor 22 prolifer- ated and eventually resulted in 16 species. Fifteen of these species are found only in the Southeastern United States. One of the 16 species, L. floridanus, is also found in the Bahama chain of West Indies; L. yeariani also may have arisen from ancestor 22. This latter species is presently known only from Mexico. The phylogenetic line that resulted in the large majority of our present day Middle America- West Indies fauna also contains taxa endemic to South America and Australia. This lineage, originally emanating from ancestor 1, continues through a moderately long series of hypothetical ancestors, ancestors 2-10, and has resulted in the evolution of 19 species. There are 11 species endemic to Middle America, 3 to Australia, 1 to South America, 1 to the West Indies (Cuba) while 2 occur in both Middle America and the Southeastern United States and 1 is found in Middle America and the West Indies. It should be noted that the majority of the Middle American species, 9 species, evolved from a phylogenetic lineage originally emanating from hypothetical ancestor 6. 176 Entomologica Americana [Vol. 46, No. The factors that led to the geographical isolation of populations and subsequent evolution of the species that now comprise the genus Loxandrus in North America are not at all evident. Freitag and Ball (in Freitag 1969) have suggested that “The analysis of the distribution of pairs of closely related species (sister species) seems to be the most fruitful approach to historical zoogeography at the local level. Their distributions are likely to provide the clearest evidence of location of former barriers and patterns of phylogeny. Of course this is true only if the two sister species are largely allopatric.” Examination of the phylogeny presented in Fig. 183 and the distribution maps, Figs. 144-176, indicates that there are two species pairs in the Southeastern United States and three species pairs in Mid- dle America that are allopatric. They are listed as follows: South- eastern United States — saccisecundaris-parallelus’, spinilunatus-algidus. Central America — fraus- unispinus; nocticolor-rectangulus; dubius- elongatus. The two species pairs found in the Southeastern United States indi- cate a barrier or barriers that divided a once single large population into an east and west population that have now evolved into distinct species. In both cases, saccisecundaris-parallelus , spinilunatus-algidus , the barrier may well have been the Mississippi Embayment on the Mississippi River Basin as it served as an outlet for glacial flood waters. In the case of the spinilunatus-algidus pair the barrier could have been the Allegheny Mountain Range although I doubt this since L. spini- lunatus has not been found east of the Mississippi River. Evidence for barriers that divide populations into north and south segments is almost absent among Loxandrus species. None of the species pair groups indicate this type of division. Only in the case of L. duryi is there an indication of distinct north and south populations (Fig. 166). In this species the northern population is found in Indi- ana, Ohio, and Pennsylvania, while the southern population is found in Western Louisiana and Eastern Texas. Ross (1967) pointed out “that the inter- American and circum- Caribbean area has been one of the world’s most prolific hotbeds of evolution and speciation.” This author goes on to say “It is now be- coming more and more probable that Central America and the West Indies have been connected at times, and broken up into several iso- lated land masses at other times, at least since mid-Cretaceous, and possibly with the fracturing reaching its maximum degree during mid- Tertiary.” It is also known that the inter- American land masses have been alternately connected and broken with the South American continent during the geological history of this area. This type of history could easily account for the division and isolation of a single 1972] Allen: Revision of Loxandrus LeConte 177 population into two segments that would have led to the evolution of the three species pairs found in Middle America and South America — fraus-unispinus , nocticolor-rectangulus, dubius-elongatus. The discussion of the past dispersal patterns of the genus Lox- andrus and the analysis of the sister species pairs are only as strong as the phylogeny. The zoogeographic patterns and the phylogenetic relationships as presented above seem to be compatible in most respects but there are two areas that cause concern at the present time. Both problems have to do with the Australian fauna. Firstly, the evidence presented in this paper indicates that the present day Australian fauna has been derived from three separate phylogenetic lineages. This is in direct conflict with the rule of parsimony. It would be much simpler if it were possible to account for the Australian Lox- andrus fauna being derived from a single phylogenetic line. Secondly, those species that are more closely related to the hypothetical ancestor of the genus Loxandrus are found in Middle America and in South America. This would indicate that the genus Loxandrus may have its origin in the New World. This is directly opposite from the views of Darlington (1962) who felt that the genus had originated in Australia and spread to the New World early in geological history. Elucidation of both the South American and Australian faunas should help to solve these problems. Loxandrus and the insect fauna of the Eastern United States. Van Dyke (1939) postulated that the coleopterous fauna of the Eastern United States had been derived from faunal regions outside this area. In other words he indicated that our present day beetle species migrated into the Eastern United States. Ross (1953) has already pointed out the fallacy of this argument. Numerous other workers have also contributed data showing that not only migration but also the evolution of “new” species from resident populations in the Eastern United States have been a common occurrence for millions of years. In fact much of the entomological literature concerning insect zoogeography in North America has dealt with speciation and the physical factors contributing to speciation, especially the effect of Pleistocene glaciation on our Eastern United States fauna. It is to this phenomenon of Pleistocene glaciation and its effect on the insect fauna of the Eastern United States that I would now like to direct a few comments. There seems to be no question that Pleistocene events had a pro- found effect on the evolution of the insect fauna of the Eastern United States. The question is rather “how did the insect fauna react to these Pleistocene events?” 178 Entomologica Americana [Vol. 46, No. 1 Two opposing theories have been postulated concerning the effect of Pleistocene glaciation and climates on the biota of the Eastern United States. Proponents of the first theory believe that the climate of the Pleistocene had a profound effect, shifting large segments of the fauna and flora of the Eastern United States several hundred miles southward. Opposing this theory is the idea that the biota in front of the glaciers was little disturbed for any great distance. Howden (1969) has summarized much of the entomological literature dealing with the Pleistocene climate and fauna in the Eastern United States. From How- den’s review article, from other literature (mostly consulting directly the papers he cites) and from my work with two warm-adapted beetle groups ( Loxandrus and the carabid tribe Morionini) there seems to be a measure of truth in each of the two above-mentioned theories. The crucial factor in whether or not North American Pleistocene climates had a displacement effect on a particular group of insects revolves around the question of whether the group is primarily adapted to cooler climates and therefore has a more northern range or whether it is adapted to warmer climates and has a more southern range. The following brief statements summarize the ways in which an insect population may have reacted to the momentous events of the Pleistocene Age. (1) Certain populations were displaced to a more southern range as the glacier itself advanced. As the ice retreated, relict southern populations were left behind as other portions of the population moved northward. These two populations subsequently became separated by ecologically inclement barriers and evolved independently of one another. Ross (1965) has given several examples of this occurring among the caddisflies. (2) Some groups with a more southern range were not displaced but their populations were frag- mented by related Pleistocene events. The evolution of these groups was essentially “an in situ development” (Howden 1969). (3) Pop- ulations could have been entirely destroyed. (4) Populations could have been displaced southward and then retreated northward without leaving relict populations. Fossil evidence would be the only means of detecting type 3 and 4 reactions. (5) Populations could have been displaced southward then retreated northward leaving a relict popu- lation, the two populations then becoming geographically isolated but neither evolving into a different species ( Allocapnia pygmaea , Ross 1965). (6) The population was seemingly unaffected by Pleistocene events ( Moirion monilicornis , Allen 1968). Figs. Ill and 178 illustrate the known locality records for all species in the genus Loxandrus. Figs. 144 to 176 illustrate the distri- bution of the individual species of Loxandrus. When the distribution patterns of the individual species are examined it is found that of the 34 1972] Allen: Revision of Loxandrus LeConte 179 TABLE IV. General distribution of Loxandrus species occurring in the Eastern United States Species Endemic to an Species Occurring in the Species Endemic to the Area North of the Coastal Flood Plain & in Coastal Flood Plain Coastal Flood Plain Northern Areas L. parallelus L. rossi L. circulus L. agilis L. lucens L. aduncus L. pravitubus L. taeniatus L. rectus L. crenatus L. nitidulus L. cervicalis L. duryi L. floridanus1 L. erraticus L. minutus L. velocipes L. micans L. saphyrinus L. unilobus L. cincinnatiensis L. algidus L. minor L. rectangulus1 L. vitiosus L. gibbus L. robustus L. brevicollis L. celeris1 L. uniformis L. extendus L. velox2 L. spinilunatus2 L. saccisecundaris2 1 Species occurring in the West Indies and/or Mexico. 2 Species whose range is not definitely known. species of Loxandrus found in the Eastern United States 10 are en- demic to the coastal flood plains. Two species are found in the coastal flood plains but also have ranges extending into the West Indies or Mexico. One species is endemic to an area outside the coastal flood plain, 1 8 species occur both in and outside the coastal flood plain and the exact range of 3 species is unknown. These data as presented here and in Table IV seem to indicate that Loxandrus is a taxon in which many of the species are more or less adapted to warmer climates. Effects of Pleistocene events on this genus are elusive. There is no doubt that the paucity of known locality records contributes to this problem. A second factor may be due to Loxandrus being a relatively old taxon, the members of which have not experienced a great amount 180 Entomologica Americana [Vol. 46, No. 1 of speciation for some time. More information concerning this ques- tion as well as the many others that have been raised will hopefully be forthcoming as work continues on the insect fauna in North America. ACKNOWLEDGMENTS I would like to thank Dr. Herbert H. Ross for his encouragement and guidance during this study. I am grateful to the Bach Fund of the National Academy of Science and to the Society of the Sigma Xi for grants which aided my work. I would like also to thank Dr. George Ball, Dr. Phillip J. Darlington, Jr., and Dr. Milton W. Sanderson for their encouragement and advice. Dr. W. C. Yearian was especially helpful in making the photographs. The following individuals kindly lent me specimens: Dr. Richard D. Alexander, The University of Michigan; Dr. Ross H. Arnett, Pur- due University; Dr. Ross T. Bell, University of Vermont; Dr. W. Wayne Boyle, The Pennsylvania State University; Mr. W. J. Brown, Canadian National Collection of Insects; Dr. Horace R. Burke, Texas A & M University; Dr. George W. Byers, The University of Kansas; Dr. O. L. Cartwright, Smithsonian Institution; Dr. Wilbur R. Enns, University of Missouri; Mr. Terry L. Erwin, University of Alberta; Dr. S. W. Frost, The Pennsylvania State University; Dr. Oswaldo Giannotti, Institute Biologico; Mr. T. F. Hlavac, Museum of Compar- ative Zoology; Dr. Karl V. Krombein, Smithsonian Institution; Mr. Hugh B. Leech, California Academy of Sciences; Mr. Jaque Negre, Paris Museum of Natural History; Mr. Gerald R. Noonan, California State College; Dr. Charles Oehler, Cincinnati Museum of Natural History; Dr. L. L. Pechuman, Cornell University; Mr. R. D. Pope, British Museum (Natural History); Dr. Jerome G. Rozen, Jr., The American Museum of Natural History; Dr. Frederick W. Stehr, Uni- versity of Minnesota; Dr. Charles A. Triplehorn, The Ohio State University; Patricia Vaurie, The American Museum of Natural His- tory; Dr. Rupert L. Wenzel, Chicago Natural History Museum; Mr. D. W. Whitehead, University of Alberta; Dr. Robert Woodruff, Florida Department of Agriculture; Dr. David A. Young, Jr., North Carolina State University. REFERENCES Allen, R. T. 1968. A synopsis of the tribe Morionini in the Western Hemisphere with descriptions of two new species (Coleoptera: Carab- idae). Carib. J. Sci., 8(3-4) : 141-163. 1972] Allen: Revision of Loxandrus LeConte 181 1969. A preliminary study of the phylogenetic relationships among species of the genus Loxandrus. Unpublished doctoral dis- sertation. University of Illinois Library. Urbana. Ball, G. E. 1962. Carabidae (Latreille, 1810). {In. Arnett, Ross H., Jr. The Beetles of the United States. The Catholic University of America Press; Washington, D. C. 55-181 ). 1966. A revision of the North American species of the subgenus Cryobius Chaudoir {Pterostichus, Carabidae, Coleoptera). Opusc. Ent. Supp., 28: 1-166. Bates, H. W. 1871. Notes on Carabidae, and descriptions of new species. Ent. Monthly Mag., 7 : 194. 1881. Biologia Centrali-Americana, Insecta, Coleoptera, Carabi- dae, Cicindelidae. 1(1): 1-152. 1891. Additions to the Carabideous fauna of Mexico, with remarks on some of the species previously recorded. Trans. Ent. Soc. Lond., 39: 223-278. Blackburn, T. 1890. Notes on Australian Coleoptera, with descriptions of new species. Part XI. Proc. Linn. Soc. N. S. Wales, (2) 5: 707-746. 1892. Notes on Australian Coleoptera, with descriptions of new species. Part XI. Proc. Linn. Soc. N. S. Wales, (2) 7: 65-152. 1907. Further notes on Australian Coleoptera with descriptions of new genera and species, XXXII. Trans. Roy. Soc. S. Austral., 27 : 91-182. Blatchley, W. S. 1918. Some new or scarce Coleoptera from Western and Southern Florida. Canad. Entom., 50: 416-423. Casey, T. L. 1918. Studies among some of the American Amarinae and Ptero- stichinae. Mem. Coleoptera, 8: 224-393. 1924. Additions to the known Coleoptera of North America. Mem. Coleoptera, 9: 1-347. Chaudoir, Maximilien 1843. Carabiques nouveaux. Bull. Soc. Imp. Nat. Moscow, 16: 671-795. 1868. Observations synonymiques sur les carabiques de l’Amerique septentrionale et descriptions d’especes nouvelles de ce pays. Rev. Mag. Zool. , Ser. 2, 20: 161-345. 182 Entomologica Americana [Vol. 46, No. 1 Darlington, Jr., P. J. 1934. New West Indian Carabidae, with a list of the Cuban species. Psyche , 41: 66-131. 1935. West Indian Carabidae II. Itinerary of 1934; Forests of Haiti; new species; and a key to Colpodes. Psyche , 42: 167-215. 1938. Loxandrus infimus Bates in Texas. Psyche, 45: 1. 1953. West Indian Carabidae (Coleoptera). The Bahama species. Amer. Mus. Novitates, no. 1650: 1-16. 1957. Zoogeography: The geographical distribution of animals. John Wiley and Sons, Inc., New York. 657 p. 1962. The Caribid beetles of New Guinea. Part 1. Cicindelinae, Carabinae, Harpalinae through Pterostichini. Bull. Mus. Comp. Zool., 126: 319-564. Dejean, P. F. M. A. 1825. Species general des Coleopteres. 1. Paris, pp. I-XXX, 1-563. 1828. Species general des Coleopteres. 3. Paris, pp. I-VII, 1- 556. 1831. Species general des Coleopteres. 5. Paris. PP. I-VIII, 1- 883. Freitag, Richard 1969. A revision of the species of the genus Evarthrus LeConte. Quaest. Ent., 5: 89-212. Hennig, W. 1966. Phylogenetic Svstematics. Univ. of Illinois Press, Urbana, 111. 263 pp. Howden, H. F. 1963. Speculations on some beetles, barriers and climates during the Pleistocene and pre-Pleistocene periods in non-glaciated portions of North America. Sept. Zool, 12: 178-201. Illies, J. 1965. Phylogeny and zoogeography of the Plectoptera. Ann. Rev. Entomol., 10: 117-140. LeConte, J. L. 1848. A descriptive catalogue of the Geodephagous Coleoptrea. Ann. Lyc. Nat. Hist. (New York), 4: 173-474. 1852. Synopsis of the species of Pterostichus and allied genera, inhabiting temperate North America. Proc. Acad. Nat. Sci. Phila., 5: 225-256. 1878. Additional descriptions of new species. In: E. A. Schwartz, The Coleoptera of Florida. Proc. Amer. Phil. Soc. 17. Phila., 5: 353-472. 1972] Allen: Revision of Loxandrus LeConte 183 Lindroth, C. H. 1966. The ground-beetles of Canada and Alaska. Part 4. Opusc. Ent., 29:409-648. Lindroth, C. H. and Freitag, R. 1969. North American ground-beetles (Coleoptera, Carabidae, excluding Cicindelidae) described by Thomas Say: Desig- nation of Lectotypes and Neotypes. Psyche, 6(3): 326-361. Macleay, W. 1888. The Insects of Kings Sound and its vicinity I. Proc. Linn. Soc. N. S. Wales (2nd series), 3(2): 443-480. Moore, B. P. 1965. Studies on Australian Carabidae (Coleoptera) 4. - The Pterostichinae. Trans. R. Ent. Soc. Lond., 117(1): 1-32. Motschulsky, V. 1865. Enumeration des nouvelles especes de Coleoptera rapportes de ses Voyages 4 e article, suite. Bull. Soc. Nat. de Moscow, 38(2): 227-313. Ross, H. H. 1937. A generic classification of the Nearctic sawflies. Illinois Biol. Monographs, 15(2): 1-173. 1953. On the origin and composition of the Nearctic insect fauna. Evolution, 7: 145-158. 1956. Evolution and classification of the mountain caddis flies. Univ. Illinois Press, Urbana, 111., 213 pp. 1959. A survey of the Empoasca fabae comples. Ann. Entomol. Soc. Amer., 52(3): 304-316. 1965. Pleistocene events and insects, pp. 583-596. In H. E. Wright, Jr., and D. G. Frey (ed.). The Quaternary of the United States. Princeton University Press, Princeton. 1967. The evolution and past dispersal of the trichoptera. Ann. Rev. Ent., 12: 169-206. Ross, H. H., Decker, G. C., and Cunningham, H. B. 1965. Adaptation and differentiation of temperate phylogenetic lines from tropical ancestors in Empoasca. Evolution, 18 (4): 639-651. Say, T. 1823. Description of insects of the families of Carabeii and Hydro- canthari of Latreille, inhabiting North America. Trans. Amer. Phil. Soc. 2: 1-109. Sloane, T. G. 1898. On Carabidae from West Australia sent by Mr. A. M. Lea (with descriptions of new genera and species, synoptic tables, etc.). Proc. Linn. Soc. N. S. Wales, 23: 444-520. 184 Entomologica Americana [Vol. 46, No. 1 1903. Studies in Australian entomology, No. XII, New Carabidae (Panageini, Bembidiini, Pogonini, Platysmatini, Platynini, Lebini with re visional lists of genera and species, some notes on synonymy, etc.). Proc. Lin. Soc. N. S. Wales , 27: 566- 642. Straneo, S. L. 1949. Nuovi Pterostichini (Coleoptera, Carabidae, Bull. Inst. Sci. Not. Belg. ) 25(41), 13 pp. 1951. On some Central and South American Pterostichini (Cole- optera, Carabidae) in the Museum of Comparative Zoology. Psyche, 58: 1-18. 1953. Nuovi Pterostichini VII (Coleoptera, Carabidae). Coriana, Genoa, 1(36), pp. 1-12, 5 figs. Tschitscherine, T. 1898. Materiaux pour servir a l’etude de Feroniens. IV. Horae Soc. Ent. Ross., 32: 1-224. 1900. Note sur le Platysmatini du Museum d’Historie Naturelle de Paris (4). Ibid., 34: 220-248. 1901. Note sur quelques Platysmatini nouveaux ou peu connus. Ibid., 35: 49-72. 1903. Nouvelles especes de la tribu de Platysmatini. Ibid., 36: 38-64. Van Dyke, E. C. 1939. The origin and distribution of the coleopterous insect fauna of North America. Proc. 6th Pacific Sci. Congr. Pacific Sci. Assoc., 4: 255-268. Wright, J. F. 1939. A new Loxandrus (Coleoptera, Carabidae) from Cincinnati, Ohio. Bull. Brook. Ent. Soc., 34: 257-258. INSTRUCTIONS TO AUTHORS 1. Manuscripts must be typed, double-spaced throughout, on standard bond paper, with margins of IV2 inches on all sides and pages numbered consecutively. 2. A title page should be supplied with the title of the paper centered at the top of the page and the author’s name (or authors’ names) centered below it. 3. Page one of the manuscript should include the title, centered at the top of the page, the author’s name centered below the title and followed by a superscript footnote number, a brief table of contents, a brief abstract, and a footnote at the bottom of the page giving the author’s present professional titles and affiliations. 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Emerson, O. Park, T. Park, and K. P. Schmidt 1949. Principles of animal ecology. Philadelphia and London, W. B. Saunders Co., 837 pp. Kistner, D. H. 1969. The biology of termitophiles. In Biology of termites, vol. 1 (ed. K. Krishna and F. M. Weesner). New York and London, Academic Press, Inc., chap. 17, pp. 525-557. 6. Keys should be typed on a separate sheet of paper, not run into the text, and provided with a formal title. Each couplet should be numbered, the numbers aligned flush left. The first words of all couplet enries should be aligned, and run-over lines indented and aligned. The number or name which ends each couplet entry should be aligned flush right and the space leading up to it filled in with leaders (periods separated by spaces). Examples : 4. Transverse fascia on corium always present and straight 5 Transverse fascia on corium irregular, not straight; or fascia absent 8 5. 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The dimensions of the printed page (4" X 61/4") should be kept in mind in the preparation of illustrations, which should be large enough to that details will be clear after reduction. All figures should be numbered consecutively, in the order of their mention in the text, with no distinction between text figures, plates, maps and photographs. All figures should be identified on the reverse side with the author’s name and the figure number. The approximate places where figures are to be inserted in the text should be indicated by the author in the margin of the manuscript. Legends for figures should be typed, double spaced, on a separate sheet of paper. 9. Tables should be typed on separate sheets of paper, double spaced, and the places where they are to be inserted in the text indicated by the author in the margin of the manuscript. A brief title should be typed above each table, and tables should be numbered consecu- tively (with arabic numerals) in the order of their mention in the text. 10. The metric system should be used for all measurements. The names of authors of species and geographical names (except in journal titles) should not be abbreviated. 11. Authors will be expected to pay a page charge of $15 per printed page toward publication costs. This fee will be waived for authors who do not have funds for publication. Reprints are not given gratis. They must be ordered from the printer at the time that proofs are returned. Extensive changes in proofs, other than correction of printer’s errors, will be charged to authors. Entomologica Americana REVIEW OF THE GENUS STERPHUS PHILIPPI (DIPTERA: SYRPHIDAE). PART I. F. Christian Thompson PUBLISHED BY THE NEW YORK ENTOMOLOGICAL SOCIETY INCORPORATING THE BROOKLYN ENTOMOLOGICAL SOCIETY Vol. 46, No. 2 August 8, 1973 The New York Entomological Society (incorporating the Brooklyn Entomological Society) publishes two journals: Entomologica Ameri- cana (irregularly, according to the availability of acceptable papers) and the Journal of the New York Entomological Society (quarterly). Monographs and longer papers (minimum length approximately 70 manuscript pages) are published in Entomologica Americana. Shorter papers may be submitted to the Journal. Manuscripts should be sent to the Editor, Entomologica Americana, Department of Entomology, The American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. Entomologica Americana was first published by the Brooklyn Entomolog- ical Society in April, 1885. Publication was suspended after six volumes, resumed in 1926, again suspended in 1964 with Volume 44, and resumed in 1969 with Volume 45. Since Entomologica Americana is published irregularly, subscription is by volume, not by year and costs $9.00 per volume. Correspondence regarding subscriptions and other non-editorial matters should be addressed to Publication Business Manager, New York Entomological Society, Department of Entomology, The American Museum of Natural History. Back issues may be purchased through Hafner Press, a Division of Macmillan Publishing Company, Incorporated, 866 Third Avenue, New York, New York, 10022. EDITOR Lee H. Herman ASSISTANT EDITOR David H. Miller Mailed August 8, 1973 Entomologica Americana is published for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Vol. 46, No. 1 was published on August 1, 1972 Entomologica Americana Vol. 46, No. 2, 1973, pp. 185-240 REVIEW OF THE GENUS STERPHUS PHILIPPI (DIPTERA: SYRPHIDAE). PART I. F. Christian Thompson1 Table of Contents Introduction 185 The Genus Sterphus 186 Key to the species of Sterphus 191 The Subgenus Telus 193 The Subgenus Sterphus 199 The Subgenus Ceriogaster 207 Abbreviations 236 Acknowledgments 237 References 238 Abstract: The genus Sterphus Philippi is reviewed. Twenty- eight species are recognized in this genus and are grouped into three subgenera and eleven species groups. Part I includes de- scriptions of all subgenera, key to and description of all species except those of the fascithorax group. Figures of some of the significant features of these taxa are also provided. One new subgenus and seven new species are described. INTRODUCTION During the course of a study of the Neotropical milesine genera (Thompson 1972) a number of new species of Sterphus , Crepidomyia Shannon and related genera were found. Some of these species display intermediate states of the characteristics traditionally used to separate Sterphus, Crepidomyia, Tatuomyia Shannon, Senoceria Hull and Mutillimyia Hull; thus, all these genera were synonymized. However, the descriptions of these new species were delayed until a complete review of all the known species could be prepared. No revision or key to the species of Sterphus or its subdivisions has ever been pub- lished and the species descriptions are widely scattered in the literature. 1 Research Fellow, Department of Entomology, The American Museum of Natural History, New York, New York 10024. [185] 186 Entomologica Americana [Vol. 46, No. 2 Genus Sterphus Philippi Sterphus Philippi, 1865, p. 737. Type species, Sterphus autumnalis Philippi (original indication) =coeruleus Rondani. Subsequent refer- ences: Kertesz, 1910, (cat. citation). — Shannon, 1926, p. 45 (cat. citation). — Shannon & Aubertin, 1933, p. 155 (descrip, note; distr. data). — Hull, 1949, p. 369 (descript.). — Thompson, 1972, p. 153 (descript.). Stherphus : Fluke, 1957, p. 106 (misspelling). Head : higher than long; face bare, usually either completely pollinose or pollinose only on sides, mostly shiny only in subgenus Telus , usually carinate, with medial and lateral carinae, frequently with carinae very strong and straight, with medial carina slightly concave beneath antennal bases and swollen laterally near the oral margin in subgenus Sterphus , rarely tuberculate; cheeks broad, as broad as or broader than long; facial grooves long, extending about two-thirds the way to the antennal bases; facial stripes indistinct; frontal prominence low, above the middle of the head; front of male short, slightly longer than vertical triangle; front of female broad, as broad as or broader at antennal bases than long; vertical triangle of male long, about twice as long as broad; ocellar triangle always dis- tinctly before posterior margin of eyes. Eyes bare, usually narrowly holoptic in male, narrowly dichoptic in males of fascithorax group, dichoptic only in males of telus and stimulans. Antennae usually elongate, slightly shorter than face, sometimes only half as long as face; third segment elongate to orbicular; arista long, longer than antenna. Thorax : longer than broad, with short sparse pile, without long thick woolly pile, may have pair of medially interrupted transverse golden pollinose bands on anterior part of notum; mesokatepistema with separate dorsal and ventral pile patches; mesoanepimera with posterior portion bare; meropleurae bare; metasterna usually bare, pilose only in cybele and always developed; post-metacoxal bridge usually incomplete, complete only in fascithorax group, metathoracic pleurae usually bare, with a few hairs in plagiatus ; metathoracic spiracle usually enlarged, in some species larger than third antennal segment; plumulae absent to elongate; scutellum usually with ventral pile fringe, without ventral pile fringe only in fascithorax group, with or without distinct emarginate apical rim. Legs : hind trochanters frequently with spurs; hind femora usually slightly swollen, straight ventrally, with two rows of very strong ventral spines; hind tibiae frequently with apical spurs. Wings: marginal cell open; apical cell closed and petiolate; anterior cross vein 1973] Thompson: Review of the Genus Sterphus Philippi 187 Figs. 1-7. Heads of Sterphus , lateral view; 1. coeruleus (Rondani); 2. incertus, n. sp. (holotype); 3. fulvus, n. sp. (holotype); 4. shannoni, n. sp.; 5. intermedius, n. sp. (holotype); 6. telus, n. sp. (holotype); 7. stimulans , n. sp. (holotype). 188 Entomologica Americana [Vol. 46, No. 2 at or beyond middle of discal cell, always greatly oblique; anterior margin of wings may be dark. Abdomen : usually elongate, rarely petiolate; batesi group with long petiole; coarc tatus and fasci thorax groups with short petiole; constriction always on second segment. Male genitalia :2 cerci long pilose, rectangular, usually rounded on dorsal margin, medium sized, without special modification; surstyli usually of typical Xylota form, with elongate curved and tapering dorsal lobe which is densely covered with short appressed pile, with sparsely pilose or bare broad and apically expanded ventral lobe, in chloropygus group with both dorsal and ventral lobes greatly expanded, in stimulans with ventral lobe expanded, in tinctus with ventral lobe greatly reduced; 9th sternum frequently with small membranous opening on dorsolateral surface near base of superior lobe; lingula short or long, absent in stimulans ; superior lobes broad, broadly and completely fused to sternum, pilose, with large ventral teeth or processes; ejaculatory apodeme umbrella shaped except rod shaped in stimulans and woodorum ; aedeagus consisting of large curved dorsal lobe, paired elongate and tapering lateral lobes and flattened ventral lobe, with posterior margin of dorsal lobe flared into two lateral flaps, with tips of lateral lobes usually between base of these flaps, and with ventral lobe beneath and/or between the lateral lobes, with lateral lobes di- vided in chloropygus group, batesi and genuinus complex, with lateral lobes absent in stimulans. Discussion: A discussion of the relationships and distinctive characteristics of Sterphus has been included in my review of the Neotropical milesine genera (Thompson, 1972, p. 156) and thus is not repeated here. With nothing known of the immature stages of Sterphus and with undoubtedly less than half the probable number of species described, it is premature to discuss the phylogeny of the group in detail. How- ever, to justify my classification (Table I) it is necessary to provide at least a general phylogeny of Sterphus. Such a general outline (Diagram 1) can be developed from the study of two morphoclines, cline A — facial color and shape and cline B — abdominal shape and number of hind tibial spurs. In cline A, the facial color darkens, from primitive reddish or orange (Ap & A0) thru partially black (Ai) to completely black (A2) and the facial carinae develop, from strongly tuberculate and concave above (Ap), thru slightly tuberculate but still concave above (A0) and slightly tuberculate and concave above 2 The male genitalia are described as they appear in lateral view, the same perspective as used for the figures. 1973] Thompson: Review of the Genus Sterphus Philippi 189 TABLE I Classification of the Genus Sterphus Philippi Subgenus Telus Thompson telus Thompson Subgenus Sterphus Philippi 1. stimulans group stimulans Thompson 2. coeruleus group coeruleus (Rondani) aurijrons Shannon Subgenus Ceriogaster Williston 3. incertus group incertus Thompson 4. auricaudatus group auricaudatus (Williston) 5. chloropygus group chloropygus (Schiner) intermedius Thompson woodorum Thompson 6. cybele group cybele (Hull) 7. plagiatus group plagiatus (Wiedemann) shannoni Thompson genuinus (Williston) trier epis (Shannon) 8. fascithorax group fascithorax (Williston) transversus (Walker) transversus (Hine) spinosa (Shannon) scutellatus (Curran) panamensis (Curran) arethusa (Hull) aureopila (Hull) funebris (Hull) rudis (Hull) 9. tinctus group tinctus (Fluke) 10. coarctatus group coarctatus (Wiedemann) 1 1 . batesi group batesi (Shannon) fulvus Thompson (Ax-2), to straight and carinate (A3). In cline B the abdomen becomes constricted and petiolate, from parallel-sided (Bp), thru slightly constricted (B0) and with a short petiole (B^, to with a long petiole (B2). Also in cline B the hind tibial spurs de- velop, from no spurs (B0) thru one apical spur (Bi) to two apical spurs (B2). On the basis of this phylogeny, I have recognized three subgenera; Telus for the species telus ; Sterphus , s. s., for the coeruleus and stimulans groups; and Ceriogaster , for all the other species groups. Telus is considered the plesiomorphic sister-group to Sterphus + Ceriogaster. Character state A0, slightly tuberculate and carinate face, demonstrates the monophyly of the combined group, Sterphus + Ceriogaster. The monophyletic nature of Ceriogaster is defined by character state A1? partially black and slightly tuberculate and concave face. The recognization of Mutillimyia (= auricaudatus group), Senoceria (= coarctatus group), Crepidomyia (= plagiatus group) or Tatuomyia (= batesi group) is not justified because such an action would leave Ceriogaster a paraphyletic and symplesiomorphic group. 190 Entomologica Americana [Vol. 46, No. 2 Telus Sterphus Ceriogaster Diagram 1. Phylogeny of the subgenera and species groups of Sterphus Philippi. Subgenera listed across the top, species groups listed verti- cally and apomorphic character states on the bottom. Explanation of characters is in the text. The only weakness in this arrangement is that no apomorphic char- acter state has been demonstrated for the nominate subgenus, Sterphus. Thus there is the strong possibility that Sterphus , s.s., is a symplesio- morphic group. This possibility would require that a subgenus be recognized for stimulans , if Ceriogaster is to be continued to be recognized as a subgenus. In this case, the enlarged meta thoracic spiracle would be used as the defining apomorphic character of the coeruleus group. However, I consider the completely pollinose faces of both the stimulans and coeruleus groups to be a synapomorphous condition and define my concept of the nominate subgenus, Sterphus, by it (see discussion under the subgenus Sterphus). My recent key to Neotropical milesine genera (Thompson, 1972) should be modified to include the new subgenus Telus. MODIFICATION OF NEOTROPICAL MILESINI GENERA KEY (THOMPSON, 1972, P. 148) 1-14, the same, no change. 15. Face straight, with distinct keels (fig. 4) Sterphus ( Ceriogaster ) Williston Face either concave or tuberculate, never straight 16 1973] Thompson: Review of the Genus Sterphus Philippi 191 16. Hind femora spindle-shaped, greatly enlarged on apical half, slender on basal half and apex (Fig. 11) Sterphus {Telus) Thompson Hind femora slender, if enlarged, then uniformly enlarged 17 17. Apical cell petiolate, with petiole as long as or longer than humeral crossvein 18 Apical cell not petiolate, closed at wing margin 19 18. Face bright yellow or orange in ground color Sterphus ( Sterphus ) Philippi Face dark in ground color 20 19. Scutellum with apical emarginate rim; abdomen oval-elongate Philippimyia Shannon Scutellum without apical emarginate rim; abdomen petiolate _ Odyneromyia Shannon and Aubertin 20. Face straight or outwardly sloping above, with epistoma strongly produced below; hind femora greatly enlarged Crioprora Osten-Sacken Face concave or tuberculate, not as above; hind femora slender ____ 21 21. Face usually with tubercle; if without facial tubercle, then either with strongly constricted abdomen or wings bicolored, anterior edge dark and posterior part light Valdivia Shannon Face concave; abdomen not strongly constricted and wings never bicolored 22 22. Anterior crossvein distinctly before middle of discal cell; males with dichoptic eyes Hemixylota Shannon & Aubertin Anterior crossvein at or beyond middle of discal cell; males with holoptic eyes Xylota Meigen KEY TO THE SPECIES OF STERPHUS PHILIPPI A. Postmetacoxal bridge complete; occiput and anterior edge of mesono- tum with transverse row of short black spines fascithorax (Williston) group Postmetacoxal bridge incomplete; occiput (except in plagiatus ) and anterior edge of mesonotum never with a row of such spines„__ 1 1. Abdomen petiolate, minimal width usually less than % maximal width (figs. 19-21) 2 Abdomen not petiolate, if slightly constricted, then minimal width never less than % maximal (fig. 18) 6 2. Hind tibiae with apical spurs 4 Hind tibiae without apical spurs 3 3. Hind legs orange except brownish coxae and apical two tarsal seg- ments, with black femoral spines; abdominal segments without bands of golden tomentose-like pile; face straight, epistoma not produced tinctus (Fluke) Hind legs bicolored, coxae, trochanters, femora and apical tarsal seg- ments black, tibiae and basal tarsal segment orange, with black femoral spines; abdominal segments without apical bands of 192 Entomologica Americana [Vol. 46, No. 2 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. tomentose-like pile; face with tuberculate swelling below (fig. 2) incertus, new species Hind legs dark, with yellow femoral spines; abdominal terga with apical bands of thick tomentose-like golden pile; face with epistoma distinctly produced forward .... auricaudatus (Williston) Hind tibiae with two apical spurs; abdominal petiole long, about three-fourths as long as second segment (fig. 21) 5 Hind tibiae with single apical spur; abdominal petiole short, about quarter as long as second segment (fig. 20) coarctatus (Wiedemann) Anal cell bare except apical quarter microtrichose (fig. 17); middle legs orange fulvus, new species Anal cell almost completely microtrichose (fig. 16); middle legs black batesi (Shannon) Abdomen red; wings completely dark except for apical light band telus, new species Abdomen dark; wings hyaline or almost completely hyaline 7 Face shiny medially, pollinose laterally, mostly black, never com- pletely yellow or orange 1 1 Face completely pollinose, orange, yellow or black in ground color 8 Cheeks and most of frons orange, all of frontal triangle in males and lower third in females orange 10 Cheeks and frons black 9 Legs black; mestasterna bare 16 Anterior four legs orange; metasterna pilose cybele (Hull) Thoracic pile reddish brown; bases of tibiae light reddish brown .. aurifrons Shannon Thoracic pile white and black; tibiae completely black coeruleus (Rondani) Mesonotum with longitudinal stripes of golden appressed pile on posterior half ___ 12 Mesonotum without longitudinal stripes of golden pile 13 Abdomen with bands of golden tomentose-like pile on third thru fourth (males) or fifth (females) terga; wings diffusely yellowish brown shannoni, new species Abdominal terga without apical golden pile bands; wings with dis- tinct diagonal brown stripe (fig. 13) plagiatus (Wiedemann) Hind trochanter and tibiae without spurs or processes 3 Hind trochanter and/or tibiae with distinct spurs or processes (figs. 10, 12) ... 14 Transverse suture golden pollinose; hind tibiae without apical spur .... genuinus complex Transverse suture without golden pollinosity; hind tibiae with long apical spur (fig. 10) : 15 1973] Thompson: Review of the Genus Sterphus Philippi 193 15. Mesonotum with front and sides silvery-gold pilose; frontal triangle of male shiny black; hind trochanter of male with simple spur woodorum, new species Mesonotum black and yellow pilose, completely black pilose above wings and on postalar calli; frontal triangle of male golden pol- linose; hind trochanter of male with a bilobed spur intermedius, new species 16. Face orange, concave, slightly tuberculate (fig. 7); hind tibiae with- out spurs stimulans, new species Face black, straight (as in fig. 5); hind tibiae with apical spur chloropygus (Schiner) Subgenus Telus, new subgenus Type species: Sterphus {Telus) telus, new species Head : face tuberculate; mostly shiny, pollinose only under antennae; antennae short, third segment orbicular, arista about as long as face; eyes of male dichoptic, separated by about length of third antennal segment. Thorax : metathoracic spiracle slightly larger than third antennal segment; scutellum non-emarginate. Wings: dark except for apical orange spot; apical crossvein just beyond middle of discal cell. Legs : (fig. 11), hind trochanter without spur; hind femora greatly enlarged on apical half, spindle-shaped, with patch of thick pile on apical half, with small dorsoapical tubercle on inner side; hind tibiae with slight medial swelling, with two apical spurs, with outer spur longer than inner, with medial patch of thick pile. These characteristics of the hind leg may be restricted to the male sex. Abdomen : elongate, slightly broader apically. Discussion: The combination of a xylotine body form with a tuberculate face is an unique feature in the Milesini and will distinguish the subgenus Telus from other milesine groups. The structure of the hind leg of Telus is unique among syrphids. The hind leg of Cacoceria Hull is similar but it lacks a dorsoposterior apical tubercle and has only one spur on the tibia. Cacoceria with its elongate antennae, bifurcate third antennal segment and pilose metasterna can hardly be confused with Telus. Telus with its tuberculate face is considered the plesiomorphic sister-group to Sterphus {Sterphus + Ceriogaster) . The name Telus is derived from the Greek, Telos, meaning end and should be treated as masculine. Since Telus is the plesiomorphic sister-group to Sterphus + Cerio- gaster and is amply distinct, it would be feasible to recognize Telus as a genus rather than a subgenus of Sterphus. While this action is clearly justified and would perhaps better illustrate the phylogenetic 194 Entomologica Americana [Vol. 46, No. 2 Figs. 8-17. fig. 8. Head of Xylota ventralis Walker (holotype), lateral view; figs. 9-11. Hind legs, lateral view; 9. Xylota ventralis Walker (holotype); 10. Sterphus intermedius, n. sp. (paratype); 1973] Thompson: Review of the Genus Sterphus Philippi 195 relationships involved, I have decided against recognition of Telus as a genus on utilitarian grounds, i.e., a broader genus is of greater utility and intelligibility to more people than would be two smaller genera. Crowson (1970, pp. 47-56, 298), Darlington (1971, pp. 147, 148) and others have presented a strong case against excessive splitting of genera and for the use of broader, more utilitarian genera, with the use of subgeneric categories by specialists, if necessary. Sterphus (Telus) telus, new species Type locality: Ecuador, Azuay Province, Tarqui, 2800 m. Holotype $ CNC. Head: (fig. 6), face slightly concave below antennae with a strong tubercle above epistoma, dark reddish brown medially, black laterally and beneath antennae, shiny except silvery pollinose band under antennae; cheeks black, shiny, black pilose; front black except dark yellowish red on lower quarter, shiny medially, silvery pollinose laterally and posteriorly, black pilose; frontal lunule yellowish red; vertical triangle black, lightly silvery pollinose, black pilose; occiput black, lightly silvery pollinose, black pilose with a few light brownish gold hairs on upper half. Eyes broadly dichoptic, separated by dis- tance equal that between anterior ocellus and posterior ocelli. Anten- nae dark brown except lighter at base of third segment; third segment roughly oval, as long as broad, slightly longer below than above; arista dark reddish brown except black at base, about as long as face. Thorax : dark bluish black except dark reddish on humeri and postalar calli, black pilose except white pilose on proepisterna and on mesokatepisterna between front and middle coxae and light golden brown on anterior part of humeri, dull grayish pollinose except for shiny spots on upper posterior corner of mesoanepisterna and medially to notopleura and dark black and brown pollinose pattern on meso- notum; mesonotal pollinose pattern as figured (fig. 34); scutellum black, grayish pollinose, black pilose except golden subscutellar fringe; <- 11. telus, n. sp. (holotype); fig. 12. Enlarged view of base of hind leg of Sterphus intermedius, n. sp. (holotype), biased lateral view; figs. 13-17. Wings of Sterphus, brown or blackish coloration indi- cated by large dots, microtrichia indicated by small dots as well as by the large dots; 13. plagiatus (Wiedemann), only extent of brownish coloration indicated; 14. coarctatus (Wiedemann) (Typic popu- lation); 15. coarctatus (Wiedemann) (Panama population); 16. batesi (Shannon); 17. fulvus, n. sp. (holotype). en = trochanteral spur, enlarged view. 196 Entomologica Americana [Vol. 46, No. 2 Figs. 18-22. fig. 18-21. Abdomens of Sterphus (Ceriogaster) , dorsal and lateral views; 18. genuinus (Williston); 19. tinctus (Fluke); 20. coarctatus (Wiedemann); 21. batesi (Shannon); fig. 22. Male genitalia of Sterphus batesi (Shannon), lateral view, t = 9th tergum and associated structures, s = 9th sternum, a = aedeagus and apodeme. 1973] Thompson: Review of the Genus Sterphus Philippi 197 Fig. 23. Male genitalia of Sterphus coeruleus (Rondani), lateral view, a = aedeagus and apodeme, e = ejaculatory apodeme, s = 9th ster- num, t = 9th tergum and associated structures. plumulae black; squamae white with black fringe; halters brown with yellow stalk. Legs : black except reddish femoral-tibial joints, black pilose except with a few white hairs intermixed on coxae, trochanters and bases of femora. Wings’, black except for yellowish-orange sub- apical spot, which extends from wing margin at end of subcosta to just beyond third vein; completely microtrichose. Abdomen', elongate, slightly broader at apex than base, red except first segment black, black pilose except white pilose on first and basal two-thirds of second segment. Male genitalia : (fig. 25), cerci rectangular, rounded dorsally, with slight indentation on dorsal margin. Surstyli: dorsal lobe narrow, elongate, gently arched on dorsal mar- gin, strongly sinuous on ventral margin and slightly turned in medially at apex; ventral lobe elongate, broaden on apical three-fourths and with blunt apical margin. Ninth sternum without dorsolateral opening; lingula short, broad, and tri-lobed medially; superior lobe with dorso- apical margin evenly curved and densely covered with very short pile, with ventroapical portion hook-like and ending in short outer tooth and longer inner tooth, with ventral margin with a number of small teeth and larger single basal tooth, with lateral flap covering all but large basal tooth. Aedeagus with two dorsolateral processes at base; dorsal lobe large, produced posteriorly; lateral lobes elongate, narrow, 198 Entomologica Americana [Vol. 46, No. 2 Figs. 24-25. Male genitalia of Sterphus, lateral view; 24. stimulans, n. sp. (holotype); 25. telus , n. sp. (holotype). a = aedeagus and apodeme, s = 9th sternum, so = tip of 9th sternum, ventrolateral view, t = 9th tergum. broadly tapering to point; ventral lobe flat, long, reaching almost to end of dorsal lobe. Ejaculatory apodeme umbrella shaped. Material Examined; Ecuador, Azuay Prov., Tarqui, 2800 m., 11 Mar. 1965, L. Pena, 1 S (CNC, holotype). 1973] Thompson: Review of the Genus Sterphus Philippi 199 Discussion: Sterphus telus is easily distinguished from all other species of Sterphus by its bright red abdomen. Subgenus Sterphus Philippi Head : face bright orange, completely pollinose, slightly to strongly concave above, with either large or small tuberculate swelling below, facial carinae weak; third antennal segment roughly oval, as long as broad; males with holoptic or dichoptic eyes. Thorax and abdomen dark bluish or greenish black, either iridescent or dull; humeri with white pollinose spot on medial margin; scutellum with weakly emarginate rim; meta thoracic spiracle either large or small; wings without dark anterior margin, slightly fumose; legs dark, hind legs simple except hind trochanter usually spurred in males, hind femora not enlarged; abdomen elongate oval. Discussion: Sterphus is readily separated from both Telus and Ceriogaster by its dark coloration and bright orange pollinose face. The facial shape of Sterphus with its strong concavity beneath the antennae, is also characteristic. As suggested above, Sterphus may be a symplesiomorphic group. The enlarged metathoracic spiracle of aurifrons and coeruleus, a synapomorphic character, clearly suggests that these two species form a monophyletic group but stimulans does not have an enlarged spiracle. Most of the other character states mentioned in the descrip- tion must be considered plesiomorphic when compared to those of Ceriogaster. However, if the extensively shiny face of Telus is ac- cepted as the primitive condition of the face, then the completely pollinose face of Sterphus can be used to demonstrate the mono- phyletic nature of the subgenus. This evaluation of the character states is not unreasonable since most species of Ceriogaster have laterally pollinose faces (intermediate condition) and only batesi and chloropygus have an almost or completely pollinose face. The facial pollinosity of Ceriogaster species would support the idea of a trend from the primitive shiny condition to the derived pollinose condition. coeruleus group A synapomorphic character state of the coeruleus group is the enlarged metathoracic spiracle. The structure of aedeagus is also unique to the group. Sterphus (Sterphus) coeruleus (Rondani) Xylota coerulea (Rondani), 1863, p. 8 (also, 1864, p. 8 (1863)). Type locality: Chile. Type 8 ? 200 Entomologica Americana [Vol. 46, No. 2 Sterphus coeruleus: Kertesz, 1910, p. 308 (cat. citation). — Shannon, 1926, p. 46 (descript, note; distr. recs.). — Porter, 1932, p. 190 (distr. rec.). — Shannon & Aubertin, 1933, p. 156 (distr. recs., fig. head of male, lateral). — Porter, 1937, p. 42 (distr. rec.). — Stuardo, 1946, p. 127 (cat. citation). — Fluke, 1957, p. 107 (cat. citation). — Etcheverry & Shenefelt, 1962, p. 208, figs. 5(3 genitalia), 10 (2 genitalia). — Pino, 1962, p. 51, figs. 12, 13 (heads), 14 (antenna), 15 (mouthparts), 16 (wings), 17 (legs). — Etcheverry, 1963, p. 80 (descript., synonymy, distr. data). — Thompson, 1971, p. 526 (fig., head of male, lateral). Sterphus autumnalis Philippi, 1865, p. 738, 782. Type locality: Chile, Prov. Valdivia. Types ? MNHN Santiago. Sterphus antennalis Philippi, 1865, p. 737. Incorrect original spelling of autumnalis Philippi (First reviser, Lynch Arribalzaga, 1892, p. 191). Xylota aurifacies Schiner, 1868, p. 360. Type locality: Chile. Types VMNH. Subsequent references: Kertesz, 1910, p. 297 (cat. citation). — Shannon, 1926, p. 50 (questioned generic placement of). — Shan- non & Aubertin, 1933, p. 123 (note on status). — Stuardo, 1946, p. 130 (cat. citation). — Fluke, 1957, p. 157 (cat. citation). — Etcheverry, 1963, p. 110 (citation). New Synonym. Head : (fig. 1), face orange, orange pollinose, weakly carinate, slightly concave beneath antennae, slightly swollen below; cheeks orange, shiny, with a few black and white hairs intermixed posteriorly; frontal triangle of male orange, orange pollinose and pilose; front of female shiny orange on lower third, bluish-black and white pollinose on upper two-thirds, black pilose; frontal lunule orange; vertex bluish black, lightly brownish pollinose, black pilose; occiput bluish black, white pollinose, white pilose below becoming intermixed with black pile above. Antennae black, black pilose; third segment roughly oval, as long as broad; arista black, about as long as face. Thorax : metallic bluish black; pleurae white pollinose, white pilose except black pile intermixed on mesoanepisterna and upper half of mesoanepimera and all black pile on upper posterior corner of mesoanepisterna; mesonotum brown pollinose except for shiny lateral and sublateral stripes and shiny patch in front of scutellum, light brownish yellow and black pilose on anterior half, black pilose posteriorly; scutellum shiny, black pilose except white subscutellar fringe. Legs : simple except spur on hind trochanter of male, black, black pilose except white pilose as follows: front and middle coxae, lateral apical patch on hind coxae, about basal half on front and middle femora, basal three-fourths of hind femora, slightly more extensive on posterior than anterior surface of femora, and tips of tibiae. Plumulae brownish, squamae white with black margin and 1973] Thompson: Review of the Genus Sterphus Philippi 201 fringe. Metathoracic spiracle much larger than third antennal seg- ment. Halters black with orange stem. Wings : diffusely blackish, completely microtrichose. Abdomen : metallic bluish, shiny except dull pollinose as fol- lows: basal margin of first tergum, narrowly along basal margin of second tergum medially, apical half of second tergum with narrow medial connecting stripe to basal band, and apical third of third tergum; pile appressed black except long white as follows: on all sterna and first tergum, in lateral triangular patches on basal three- fourth of second tergum with these patches touching medially, and on basal third of lateral margins of third and fourth terga. Male genitalia : (fig. 23), cerci rectangular, with shallow indentation on dorsal margin. Surstyli: dorsal lobe narrow, elongate, slightly curved medially at apex, with dorsal margin arched and ventral margin recurved except basal portion straight, with a few long marginal hairs on dorsal edge, elsewhere short pilose; ventral lobe broad, elongate, with rounded apical margin. Ninth sternum with small dorsolateral membranous opening; lingula very short, reduced to small convex lip. Superior lobes with dorsoapical margin angulate, with ventroapical portion hook-like and ending in a long narrow outer tooth and small inner tooth, with two small teeth near base of apical hook-like process, with ventral margin with one small tooth, with large lateral flap extending posteroventrally in form of large tooth. Aedeagus: dorsal lobe large, produced dorsally, with well developed lateral flaps and with medial production between flaps; lateral lobes narrow, tapering into long thin points; ventral lobe with large lateral flaps which extend posteriorly into sharp point. Ejaculatory apodeme umbrella shaped. Material Examined: Argentina: Neuquen, San Martin de los Andes, Oct. 1952-Jan. 1953, F. H. Walz, 1 $ (FCT). chile: 2 $ (AMNH); Valparaiso, A. Faz, 1 S (USNM); Valparaiso, E. P. Reed, 1 2 (AMNH); Valparaiso, Limache, A. Faz, 1 $ (AMNH); Valparaiso, Pichilemis, Jan. 1925, Reed, 1 2 (FCT); Arauco, Peral, 1-2 Jan. 1966, Flint & Cekalovic, 1 S (USNM); Malleco, Angol, 4 Feb. 1951, J. Suarlez, 1 $ (USNM), 27 Mar. 1930, 1 2 (AMNH), 2 Apr. 1951, J. Suarlez, 1 3 (USNM), 6 Apr. 1926, 1 2 (AMNH), 5 Oct. 1937, Gekwall, 1 2 (USNM), 20 Oct. 1946, F. Trina, 1 2 (AMNH); Llanquihue, Casa Panque, Dec. 1926, R. & E. Shannon, 1 $ (USNM); Chiloe, Castro, 20 Dec. 1926, R. & E. Shannon, 13,22 (USNM); Chiloe, Isla de Chiloe, Ancud, Apr. 1941, P. A. Berry, 1 3, 2 2 (USNM); Magallanes, Wellington Island, Puerto Eden, 7-15 Dec. 1962, P. J. Darlington, 1 2 (MCZ); Magallanes, Magellan Strait, 25 Mar. 1925, 2 2 (AMNH). In addition, I briefly 202 Entomologica Americana [Vol. 46, No. 2 examined 36 S 9 from Chile, ranging from Santiago to Aysen, in the Canadian National Collection. Discussion: S. coeruleus is readily separated from most syrphids by its shiny metallic blue body color with a bright orange face and front. Both Philippimyia cyanocephala and Stilbosoma rubiceps are very similar in appearance but can be readily separated by their dis- tinctive generic characters and also, in S. rubiceps by the orange vertex, and in P. cyanocephala by the metallic blue face. A discussion of the distinguishing characters between S. coeruleus and the other species of the subgenus have been included under those species, aurifrons and stimulans. The name “ Xylota aurifacies Bigot” was first published by Schiner in the discussion of the relationships of his new species, Mallota xylotaeformis (“. . . ich kenne einige exotische Xyloten, wie z. B. Xylota aurifacies Big. aus Chile, bei welechen die Cubitalader schon ziemlich stark eingebogen ist, . . This name was never used by Bigot. Since Schiner’s citation, the name aurifacies has been usually carried in the literature as available, either attributed to Bigot or Schiner but of uncertain applicability (i.e. nomen dubium). Vockeroth and I have looked for material of “ Xylota aurifacies ” in the Bigot Collection at both Oxford and the British Museum (Natural History) without finding any. However, among the S. coeruleus material in the American Museum of Natural History there is an old specimen labelled uXylota aurifacies Schiner” by Curran.3 This determined specimen lead me to suspect that there was similarly determined material in the Naturhistorisches Museum Wien where Schiner did his work. Ruth Lichtenberg of that museum informs me that there are two such specimens among the S. coeruleus material in their col- lections, one labelled as “X. aurifacies n. sp. Bigot, Chili, Schiner 1869” and the other as “aurifacies B. Alte Sammlung, Philippi Chili 1870.” Schiner’s statement was probably based on the first of these two specimens and thus that specimen should be considered as the type of aurifacies. Until a careful examination of all the S. coeruleus material in the Naturhistorisches Museum Wien is made to ascertain which specimens could have been available to Schiner at that time, I hesitate labelling this specimen as either the type or lectotype. There is no question that the name aurifacies must now be synonymized under coeruleus. 3 During the 1920’s Curran acquired (? through exchange with Hans Zerny?) an extensive collection of syrphid material from the Naturhistorisches Museum Wien. Thus the American Museum has cotypes and other authenti- cated specimens of many of Wiedemann’s and Schiner’s species. 1973] Thompson: Review of the Genus Sterphus Philippi 203 Figs. 26-27. Male genitalia of Sterphus, lateral view except so; 26. woodorum, n. sp. (holotype); 27. intermedius, n. sp. (holotype). a = aedeagus and base of apodeme, s = 9th sternum, so = tip of 9th sternum from ventral view with slight lateral bias, ss = surstyle, t = 9th tergum and associated structures. 204 Entomologica Americana [Vol. 46, No. 2 Sterphus (Sterphus) aurifrons Shannon Sterphus aurifrons Shannon, 1926, p. 46. Type locality: Chile, Juan Fernandez Islands (see below). Type 8 BM(NH), Subsequent References: Shannon & Aubertin, 1933, p. 156 (descript, note, distr. data). — Stuardo, 1946, p. 127 (cat. citation). — Fluke, 1955, p. 42 (descript. $, distr. recs.). — Fluke, 1957, p. 107 (cat. citation). — Etcheverry, 1963, p. 79 (descript., synonymy, distr. data). Material Examined: chile: Juan Fernandez Islands, Masa- tierra, Plazoleta del Yunque, 12 February 1951, P. J. Kuschel, 1 8 (AMNH). Discussion: S. aurifrons is very similar to coeruleus in all respects except as follows: 1) the thoracic pile is completely reddish- brown instead of a mixture of black and white; 2) the mesonotum is completely reddish-brown pollinose with pollinosity completely obscuring the ground color instead of lightly brownish pollinose with a pair of submedial and sublateral shiny vittae and pollinosity not obscuring the bluish-black ground color; 3) the abdominal pile is brownish yellow and black instead of white and black; 4) the abdomen has a more greenish metallic lustre instead of steel blue lustre; 5) the dorsal lobe of surstyle (fig. 30ss) is more evenly curved on the ventral margin, not strongly angulate near the base; 6) the ventral lobe of the surstyle (fig. 30ss) has a slight ventroapical notch, not evenly rounded; and 7) the dentation of the superior lobe (fig. 30s) is quite different, the apical hook-like process is much broader with the inner tooth greatly reduced, the two small teeth at the base of the hook-like process are absent and the lateral flap is not as strongly produced. The type locality of S. aurifrons was given by Shannon as “Chile (J. Fernz),” and later in Diptera of Patagonia and South Chile (Shan- non and Aubertin, 1933) it was stated that “precise locality unre- corded.” However, the only known material of the species besides the holotype was collected on Juan Fernandez Islands (Fluke 1955) and this strongly suggests that Shannon confused a contraction of / uan Fernandez for a collector’s name. Since there is no indication on the locality label of the type that “J. Fernz” is either a location or collector it should be assumed to be a locality and that the type locality of Sterphus aurifrons be restricted to the Juan Fernandez Islands, where the species is known to exist. stimulans group Some of the apomorphic character states of stimulans, the only species in the stimulans group, that may be of use in defining the 1973] Thompson: Review of the Genus Sterphus Philippi 205 Fig. 28-30. Male genitalia of Sterphus, lateral view; 28. shcinnoni Thompson, n. sp. (paratype), 29. tinctus (Fluke), 30. aurifrons Shannon, a = aedeagus and apodeme, s = 9th sternum, ss = surstyle, t = 9th tergum and associated structures. 206 Entomologica Americana [Vol. 46, No. 2 group, are: 1 ) the loss of the ventral lobe of the surstyle; 2) the lack of lateral lobes on the aedeagus; and 3) the elongate and bifid ventral lobe of the aedeagus. Sterphus (Sterphus) stimulans, new species Type locality: Bolivia, Chipiriri. Holotype $ CNC. Head : (fig. 7), face orange, golden pollinose, deeply concave beneath antennae, with distinct but low tubercle below; cheeks black, shiny on anterior two-thirds, silvery pollinose on posterior third, white pilose; frontal lunule black; frontal and vertical triangle black, brown- ish black pollinose, black pilose; occiput black, silvery pollinose and white pilose on lower half, brownish black pollinose and black pilose on upper half. Eyes dichoptic in male, separated by distance equal to that between anterior ocellus and posterior ocelli; antennae black, black pilose; third segment roughly oval, as long as broad; arista black, much longer than face. Thorax : bluish black, completely dull pollinose, pleurae with pile same as in coeruleus; mesonotum black pilose except for a few light hairs on anterior margin and notopleurae; scutellum dull pol- linose, black pilose except white subscutellar fringe; halters black; squamae white with black margin and fringe; plumulae light brown. Legs : black, simple, without spur on trochanters or tibiae, with pile same as in coeruleus except all white pile on hind coxae. Wings: diffusely light black, completely microtrichose. Abdomen : bluish black, dull pollinose except shiny narrowly on lateral margins of terga; mostly yellow pilose, longer on sterna and lateral margins of terga, black pilose on third and fourth terga except broad basal triangular corner patches of yellow pile. Male genitalia : (fig. 24), cerci rectangular, with dorsal margin rounded; surstyli broad, about as broad as long, with narrow curved dorsal ridge (= dorsal lobe), with large ventral extension (= ventral lobe) beneath this ridge, with margins of ventral extension straight except for basoventral and posteroventral lobes; ninth sternum without dorso- lateral opening; lingula absent; superior lobe with dorsoapical margin angulate, very short pilose narrowly along apical margin, with two lateral flaps, with outer flap with evenly curved margin and inner flap with serrate ventral edge; aedeagus with dorsal lobe produced posteriorly and with long anterior process, with lateral lobes absent, with ventral lobe greatly elongate and bifid; ejaculatory apodeme rod-shaped. Material Examined: Bolivia: Chipiriri, Sept. 1962, T. Stein- bach, 1 $ (holotype, CNC). 1973] Thompson: Review of the Genus Sterphus Philippi 207 Discussion : S. stimulans, with its orange face, simple legs, and bluish-black coloration, is quite distinct from all other species of Sterphus except coeruleus and aurifrons. S. stimulans can be con- trasted with coeruleus and aurifrons as follows : 1 ) the cheeks and lower front (all of frontal triangle in male) is black, not orange; 2) the face is strongly concave above, not slightly concave (fig. 7); 3) the frontal pile in the male is black, not orange; 4) the frontal lunule is black, not orange; 5) the metathoracic spiracle is much smaller than the third antennal segment, not much larger; 6) the metathoracic spiracle has a black fringe, not a white fringe; 7) the lateral margins of the abdomen has a long golden pile, not an alternatively mainly black and white pile; and 8) the hind coxae are completely white pilose, not mainly black pilose ( coeruleus ) or golden (aurifrons) . S. stimulans also differs from coeruleus in that the scutellum is dull pollinose, not shiny and the mesonotum is completely dull pollinose, not with shiny longitudinal stripes. It differs from aurifrons in that its mesonotum is black pilose not tawny brown pilose. Subgenus Ceriogaster Willis ton Ceriogaster Williston, 1888, p. 285. Type species, fascithorax Williston (monotypy). Subsequent references: Kertesz, 1910, p. 308 (cat. cita- tion).— Shannon, 1926, p. 50 (note on relationship to Tatuomyia) . — Curran, 1934, p. 405 (notes, synonymy of Zonemyia) . — Hull, 1949, p. 367 (descript.) . — - Fluke, 1957, p. 104 (cat. citation). New status. Zonemyia Shannon, 1925, p. 109. Type species, spinosa Shannon (orig- inal designation). Subsequent references: Shannon, 1927, p. 4 (key ref.). — Shannon, 1928, p. 573 (relationship of Lepidostola to genus). — Shannon & Aubertin, 1933, p. 139 (key ref.). Crepidomyia Shannon, 1926, p. 47. Type species, tricrepis Shannon (original designation). Subsequent references: Curran, 1934, p. 404 (note). — Hull, 1949, p. 368 (descript.). — -Fluke, 1957, p. 105 (cat. citation). — Thompson, 1971, p. 524 (synonymy). Tatuomyia Shannon, 1926, p. 48. Type species, batesi Shannon (original designation). Subsequent references: Hull, 1949, p. 367 (descript.). — Fluke, 1957, p. 105 (cat. citation, synonymy of Senoceria) . Senoceria Hull, 1930, p. 144. Type species, spinifemorata Hull (original designation) =: coarctatus Wiedemann. Subsequent reference: Hull, 1949, p. 367 (descript., as subgenus of Tatuomyia) . Mutillimyia Hull, 1943, p. 139. Type species, Ceriogaster auricaudata Williston (original designation). Subsequent references: Hull, 1949, p. 372 (descript.). — Fluke, 1957, p. 109 (cat. citation). Willistonomyia Hull, 1943, p. 140. Incorrect original spelling of Mutil- limyia. 208 Entomologica Americana [Vol. 46, No. 2 Head : face usually black, orange in cybele and partially orange in incertus , usually with strongly developed facial carinae, carinae weak in incertus and chloropygus group, usually straight and without tubercle, weakly tuberculate only in incertus and with epistoma pro- duced only in auricaudatus , usually pollinose laterally and shiny medially, completely pollinose only in cybele and chloropygus ; third antennal segment usually elongate, usually 1 Vi times or more as long as broad, chloropygus group (and probably incertus) with oval third segments; eyes of male holoptic or narrowly dichoptic in fascithorax group. Thorax : black or brownish black, usually dull pollinose, fre- quently with yellow pollinose markings, only shiny in batesi and incertus ; metathoracic spiracle either large or small; scutellum with or without apical emarginate rim; wings frequently with dark anterior margins, otherwise hyaline except plagiatus with diagonal brown stripe. Legs : frequently with hind femora enlarged, with or without trochanteral or tibial spurs. Abdomen : ranging from parallel sided to strongly petiolate. Discussion: The subgenus Ceriogaster can be divided into two sets of species groups: those groups with weakly carinate faces and oval third antennal segments, and those with strongly carinate faces and elongate third antennal segments. The first division, a symplesio- morphous grouping, includes only the incertus , auricaudatus and chloropygus groups; whereas all the other groups fall into the second division. The groups of the first division can be easily separated by their facial shape, and the groups of the second division by abdominal shape (figs. 18-21). Other differences between these species groups are listed in the following Table. TABLE II Character table for the species groups of Sterphus ( Ceriogaster ) Character states Species groups I 2 3 4a 4b 4c 5 6 7 8 9 10 incertus - ? - - - - ? - 0 ? - - auricaudatus - - + - - + - - 0 + - - chloropygus - - - - - - + + 0 + - - cybele - - - - - - ? - 2 ? - + plagiatus - + - - - - + + 0-2 - + + fascithorax + + - - - + - + 0 - + + tinctus + + + - - + - - 0 - + + coarctatus + + + - + + + + 1 - + + batesi + + + + + + - r 2 + - + 1973] Thompson: Review of the Genus Sterphus Philippi 209 TABLE II (continued) List of species group characters 1 ) Cheeks linear, about four times as long as broad ( + ) ; about twice as long as broad (-). 2) Third antennal segment elongate, usually two times as long as broad ( + ) ; oval, about as long as broad (-) . 3) Wing with dark anterior margin ( + ); without dark anterior margin (-). 4) a) Abdomen strongly petiolate, with long petiole ( + ); not so (-). b) Abdomen strongly petiolate, with long or short petiole ( + ); not so (-). c) Abdomen slightly (or strongly petiolate) ( + ); not so (-). 5) Males with spurs on hind trochanter ( + ); unspurred (-). 6) Hind femora enlarged ( + ); simple (-). 7) Hind tibiae with spurs, one (1), two (2), or none (0). 8) Male frontal triangle shiny ( + ); or golden pollinose (-). 9) Mesonotum with golden pollinose markings ( + ); without (-). 10) Face with strong facial carinae ( + ); not so (-). incertus group The incertus group can be easily separated from all other species groups by its partially orange and slightly tuberculate face. Other diagnostic characters for the group have been listed in Table II. Sterphus (Ceriogaster) incertus, new species Type locality: Venezuela, La Negra, Paramo. Holotype 2 MZUSP Head : (fig. 2), face black medially, orange laterally, light yel- lowish pollinose on orange areas, shiny on black areas, slightly con- cave beneath antennae, with slight tuberculate swelling above epistoma; cheeks black, shiny, light pollinose posteriorly, yellow pilose posteriorly; frontal lunule yellow; front black, shiny and bare on lower half, brownish pollinose on upper half, golden brown pilose on upper half with a few black hairs intermixed; vertex black, lightly brownish pollinose, black pilose; occiput black, silvery pollinose, yellow pilose below becoming darker above. Antennae with first two segments reddish orange, with black pile; third segment missing. Thorax : black except reddish brown postalar calli; pleurae silvery pollinose, yellow pilose except black pilose on upper posterior corner of anepisterna; mesonotum shiny except brownish pollinose narrowly on sides and across front edge between humeri and with medial posteriorly directed spur of pollinosity, reddish brown pilose. Scutellum shiny black, with reddish brown pilose with a few longer black hairs intermixed, with subscutellar fringe yellow. Squamae 210 Entomologica Americana [Vol. 46, No. 2 dirty white, with dark brown fringe; plumulae and halters brownish orange. Legs : black except orange tibiae and basal two tarsal seg- ments, yellow pilose except black pilose on apical third of femora and medial third of tibiae and apical four tarsal segments. Wings: pale yellowish brown, completely microtrichose. Abdomen : black, greenish bronze pollinose except shiny lateral margins and fifth segment. Pile long and erect yellow on sterna, lateral margins of all terga, all first tergum, second and third terga except for posterio-medial triangular patch of appressed black pile, and narrowly on anterior margins of fourth and fifth terga; short appressed black pilose on second and third terga in form of posterio- medial triangular patch on posterior half and three-fourths respectively of second and third terga, and all fourth and fifth terga except yellow pilose on lateral and anterior margins. Material Examined: Venezuela: La Negra, Paramo, August 1948, Lichy, 1 $ (holotype, MZUSP). Discussion: S. incertus differs from the typical species of the subgenus Ceriogaster in that its: 1) face is orange laterally; 2) face is slightly concave beneath the antennae and has a distinct tuberculate swelling above the epistoma; 3) transverse sutures are shiny; and 4) tibiae and apical two tarsal segments are orange. S. incertus is repre- sented by a single female which has the third antennal segments miss- ing and for these reasons I have named this species, incertus. auricaudatus group The auricaudatus group is readily distinguished from all other species groups by its anteriorly produced epistoma. Other diagnostic characters for the group have been listed in Table II. Sterplms (Ceriogaster) auricaudatus (Williston) Ceriogaster auricaudata Williston, 1892, p. 73, pi. 2, figs. 10 (habitus), 10a (head). Type locality: Mexico, Guerrero, Omiltemi, 8000 ft. Types <3 $ BM (NH). Subsequent references: Kertesz, 1910, p. 308 (cat. citation). — Shannon, 1926, p. 50 (note on placement in Temnostomini) . — Curran, 1934, p. 405 (note, key ref.). MutiUimyia auricaudata : Hull, 1943, pp. 137, 140 (described as mutillid mimic). — Fluke, 1957, p. 109 (cat. citation). Head : face black except for orange spot on either side of epistoma, shiny medially, yellowish-white pollinose laterally; cheeks shiny black, with some yellow pile posteriorly; front black except for small narrow yellow triangle above antennal bases, shiny about antennal bases, elsewhere yellowish-white pollinose, yellow pilose; 1973] Thompson: Review of the Genus Sterphus Philippi 211 vertex black, yellow pilose; occiput black, white pollinose, yellow pilose. Antennae reddish brown, about three-fourths as long as face; third segment quadrate, slightly longer than wide, about as long as first two segments; arista yellow, slightly longer than antenna. Thorax : black, light pollinose, silvery pollinose on pleurae, brownish pollinose on mesonotum except velvety black pollinose on transverse suture, short yellow pilose except longer golden pilose in front of pos talar calli; scutellum black, lightly brownish pollinose, white pilose. Legs : white pilose, coxae and femora blackish except yellowish apically on all femora and also basally on hind femora; tibiae and tarsi dark reddish brown; hind femora with double row of ventroapical short yellow spines. Halters brownish yellow with dark head. Squamae dirty white with brown fringe. Plumulae brownish. Wings: brownish anteriorly and hyaline posteriorly, micro- trichose except bare anterior quarter of anal cell. Brown extends back to anal and discal cell basad to anterior crossvein, beyond anterior crossvein brown area bordering on spurious vein and extending diagonally from tip of spurious vein to apical corner of first posterior cell (also see Williston’s figure for exact extent of brownish anterior margin of wings). Abdomen : petiolate, minimal width about one-third maximal width; first tergum opaque velvety black except shiny on sides; second tergum opaque velvety black except two large lateral shiny metallic triangles on petiole, with triangles connected medially; remainder of terga black, lightly whitish-yellow pollinose; pile of dorsum short yel- low except thick, opaque golden on apical margin of first, second, and third terga, and all fourth and fifth terga. Venter black, with short appressed white pile except black medially on third and fourth and all of fifth sterna. Material Examined: Mexico: Guerrero, Omiltemi, 8000 ft., 12 1# (cotypes, BM(NH)). Discussion: S. auricaudatus is readily separated from all other species of Sterphus by its anteriorly produced epistoma. In appear- ance it is similar to S. tinctus (Fluke) but it is easily distinguished by the characters given in the key. The female cotype was figured in color in the original publication. However, a few discrepancies between the female cotype and figure should be noted: 1) the epistoma is produced more in the type; 2) the antennae are longer than shown, reaching almost to the epistoma; 3) too much yellow is shown on front, there is only a small narrow triangle of yellow extending back from the frontal lunule; and 5) the color on the tips of the femora is not reddish but yellowish. 212 Entomologica Americana [Vol. 46, No. 2 chloropygus group Sterphus chloropygus, woodorum and intermedius are very closely related, being separated from all other species of Sterphus by the structure of their hind tibiae, scutellum and male genitalia. The hind tibia (fig. 10) of the males, with its subbasal medial keel and apical keel-like spur, is unique among syrphids. In the females these struc- tures are less pronounced, the apical spur does not have the keel-like basal portion and the subbasal medial keel is much lower. All the members of chloropygus group have a distinct apical emarginate rim on the scutellum, this rim being absent or very weak and indistinct in all other species of Sterphus. The surstylus with its large ventral extension to the dorsal lobe is also distinctive among Sterphus species. Thus, the chloropygus group is clearly monophyletic. Sterphus (Ceriogaster) chloropygus (Schiner) Xylota chloropyga Schiner, 1868, p. 366. Type locality: “Columbian” (m Venezuela) . Lectotype $ VMNH. Kertesz, 1910, p. 298 (cat. citation). — Shannon, 1926, p. 50 (questioned generic placement). — Fluke, 1957, p. 157 (cat. citation). Male. — Head : black; face straight, slightly convex about epistoma, completely silvery gold pollinose; cheeks shiny, with a few yellow hairs posteriorly; frontal triangle silvery gold pollinose, with some black hairs at junction of eyes; frontal lunule reddish brown; vertical triangle dull black, black pilose; occiput light silvery pollinose, thickly yellow pilose below and sparsely black pilose on upper quarter. Antennae black, short, about three-fourths as long as face; third segment trapezoidal, ventral margin slightly longer than dorsal; arista black, about IV2 times as long as face. Thorax : black, dull brownish pollinose (there appears to be a trace of a pair of faint, medial, light brownish, pollinose vittae as in intermedius) , black pilose except yellow pilose on propleurae; meta- thoracic spiracle distinctly larger than third antennal segment, with white fringe; scutellum with emarginate rim, shiny bluish black, black pilose including subscutellar fringe; squamae brownish with brown margin and fringe; plumulae yellow. Wings: brownish, completely microtrichose. Legs: black except reddish brown basal two tarsal segments of anterior legs and only first tarsal segment of hind leg, black pilose; hind trochanter with long flatten spur; hind femora only slightly enlarged; hind tibiae with ventral keel on basal third and long apical keel-like spur. Abdomen: orange, except brownish black first tergum and medially on second, third and basal half of fourth terga, long yellow 1973] Thompson: Review of the Genus Sterphus Philippi 213 pilose except shorter black pilose on dark areas of first thru third terga, shiny except dull on first thru third terga. Female. — Quite similar to male except for normal sexual dimorphism and front shiny except for silvery pollinose spot on eye margin half way between antennae and vertex, black pilose on upper half; mesonotum with a few scattered golden hairs intermixed with black pile; tarsi much darker and hind tarsi all black; abdomen much darker, sterna dark reddish brown, terga mostly dull black, shiny bronze on lateral margin of second, third and all of fourth terga, with yellow pile on sterna and lateral margins of second, third, and all of fourth terga, rest black pilose. Material Examined: Venezuela: 1864, Lindig, HI? (cotypes of chloropygus , VMNH). Discussion: S. chloropygus is readily distinguished from both woodorum and intermedius by its completely pollinose face, black pilose legs and mesothorax. I have examined the types of chloropygus Schiner, both of which are labelled “Lindig, 1864, Venezuela,” “ chloropygus , Alte Sammlung.” The male is hereby designated lectoype and has been so labelled. The type locality is thereby re- stricted to Venezuela. Schiner in his part of “Reise der oster- reichischen Fregatte Novara ” described a number of species as “ aus ColumbierT and types of all the syrphids so described have been found to be Lindig material from Venezuela as are the chloropygus cotypes. Thus, it is apparent that Schiner’s “ Columbien ” included Venezuela. Sterphus (Ceriogaster) woodorum, new species Type locality: Venezuela, Distrito Federal, Parque Nacional Avila, 2000 m. Holotype 8 CNC. Head : black; face straight, shiny medially, silvery- white pollinose laterally, bare; cheeks shiny, white pilose; frontal triangle of male shiny except very narrowly silvery white pollinose along eye margins, with a few black hairs at junction of eyes; frontal lunule brownish black; front of female shiny on lower half, silvery- white pollinose and black pilose on upper half; vertex sparsely silvery pollinose anteriorly and bronze pollinose posteriorly, black pilose; occiput white pollinose and pilose below becoming bronze pollinose and black pilose on upper third. Antennae short, about two-thirds as long as face, black pilose; third segment trapezoidal, dorsal edge about half as long as ventral; arista yellow, about twice as long as antenna. Thorax : black, mainly bronze pollinose, predominately yellowish silver pilose with some black pile intermixed on all sclerites except on 214 Entomologica Americana [Vol. 46, No. 2 mesokatepisterna, with disc of mesonotum completely short black pilose in female; mesonotum with distinct pair of light brownish pol- linose, medial vittae and a pair of indistinct, grayish pollinose, sub- lateral vittae behind suture; pile on margins of mesonotum and scutellum thicker, opaque and brighter in color; scutellum with dis- tinct apical emarginate rim, with thick white ventral fringe, without apical spines, with intermixed black pile much longer than light colored pile; mesonotum without black spine-like pile above wing bases; meta thoracic spiracle distinctly larger than third antennal seg- ment; halters orange; plumulae orange; squamae light orange with upper margin dark brown and lower margin orange, with fringe orange. Wings: light brownish, completely microtrichose. Legs: black and dull pollinose except hind femora shiny bronze to bluish black, reddish brown femoral-tibial joints and basal segment of middle and hind tarsi; white pilose except black pilose in male as follows: apical posterior third and ventral margin of front femora, apical posterior tip and ventral margin of middle femora, apical half of hind femora, intermixed on middle and hind coxae, medial posterior third of front tibiae, ventral and posterior surface of middle tibiae, all of hind tibiae, apical four tarsal segments of all legs; legs more exten- sively white pilose in female; hind trochanter with short sharp papilla in male, papilla reduced in female; hind femora greatly swollen, with large tuberculate swelling on basal third of inner side of male; hind tibiae with ventral keel on basal third and long apical keel-like spur in males, these structures reduced in female but still quite distinct. Abdomen: shiny bronze except dull pollinose medially on second and third terga in males and first thru third in females, long bronze- yellow pile on shiny areas and appressed black on dull areas. Male genitalia: (Fig. 26), cerci elongate, produced posterodorsally. Surstyli: dorsal lobe very broad, broader than long, with curved dorsal and ventral ridges, with slight mediodorsal projection, with a few longer hairs on anterodorsal edge, with ventroapical edge angulate and extending slightly beneath ventral lobe; cleft between dorsal and ventral lobes short and broad; ventral lobe short, broad, broader than long, with fairly extensive marginal pile, with broadly rounded apical margin. Ninth sternum with dorsolateral membranous opening; lingula short, broadly triangular, with apex blunt; superior lobe sparsely pilose, with dorsoapical margin angulate, with large lateral excavation extending dorsally from ventral margin, with single large tooth on ventral margin. Aedeagus: dorsal lobe produced posteriorly, with large lateral flaps; large, long cleft between dorsal and lateral lobes; lateral lobes divided into basal and medial processes, with basal 1973] Thompson: Review of the Genus Sterphus Philippi 215 process directed ventrally and with blunt apex, with medial process curved posteriorly and with pointed apex. Ejaculatory apodeme rod- shaped, with anterior end slightly expanded. Material Examined: Venezuela: D. F., Parque Nac. Avila, 2000 m, 28 Feb. 1971, G & M Wood, 1 $ (holotype), 1 $ (allo- type) (CNC). Discussion: The species is named after its collectors, Dr. Monty Wood, of the Canada Department of Agriculture, and his wife. S. woodorum differs from the other species of the chloropygus group in that: 1) the arista is yellowish orange, not black; 2) the frontal triangle of the male is shiny, not golden pollinose; and 3) the lateral margins of the mesonotum are silvery golden pilose. Sterphus (Ceriogaster) intermedius, new species Type locality: Bolivia, Cochabamba, Chapare, Limbo, 2200 m. Holotype $ MZUSP Head’, (fig. 5), black; face straight beneath antennae, slightly convex above epistoma, shiny except broadly silvery pollinose on sides; cheeks shiny, with a few light yellowish hairs posteriorly; frontal triangle light golden pollinose with two black hairs on each side near eye contiguity; frontal lunule orange; vertical triangle light pollinose, black pilose; occiput silvery pollinose, light yellow pilose except with black pile intermixed on upper part. Antennae black, short, slightly more than half as long as face; third segment trapezoidal, ventral margin about twice as long as dorsal; arista black, about as long as face. Thorax : black, brownish pollinose except for faint pair of medial light brownish pollinose vitta on mesonotum; pile yellow and black intermixed, except all black pilose on dorsal posterior corner of mesoanepisterna, above wings and on postalar calli; metathoracic spiracle slightly larger than third antennal segment, with white fringe; scutellum with emarginate rim, black with metallic luster, black and yellow pilose, with yellow ventral fringe; halters orange; squamae light brownish orange, with dark brown margin and fringe; plumulae orange. Legs’, black, yellow pilose except black pilose as follows: apical posterior tip of front femora, apical third of middle femora, apical half of hind femora, both middle and hind tibiae, apical four of front and middle tarsal segments and all hind tarsal segments; hind femora only slightly enlarged; hind trochanter (fig. 12) with short curved bilobed spur; hind tibiae (fig. 10) with large apical spur and ventral keel on basal third. Wings: light brownish, completely microtrichose. Abdomen : elongate, parallel-sided, metallic bluish black, shiny 216 Entomologica Americana [Vol. 46, No. 2 except for dark brown pollinose bands on apical half of second and third terga, with pollinose bands not reaching lateral margins and bands of second terga with narrow medial projection reaching to anterior margin; pile long, erect yellow except appressed black on pollinose areas. Male genitalia : (fig. 27), quite similar to woodorum except as follows : 1 ) dorsal lobe of surstyli without mediodorsal projection; 2) cleft between the dorsal and ventral lobes much longer; 3) posteroventral edge of dorsal lobe of surstyli rounded, not angulate; 4) ventral lobe of surstyli larger and produced more into a postero- dorsal lobe; 5) dorsolateral membranous opening on ninth sternum triangularly shaped, not round; 6) lingula longer and with apex rounded; 7) superior lobe without lateral excavation; 8) superior lobe with two large ventral teeth which are hidden in lateral view; 9) dorsoapical margin of superior lobes more rounded, not angulate; 10) superior lobe with pile sparse except for dense patch near basoventral corner; 11) basolateral process of aedeagus ending in sharp point, not blunt point; 12) cleft between lateral and dorsal lobes of aedeagus very short; and 13) ejaculatory apodeme umbrella- shaped. Material Examined: Bolivia: Cochabamba, Chapare, Limbo, 2200 m, November 1953, Martinez, 1 $ (Holotype, MZUSP); Cochabamba, October 1965, Steinbach, 1 S (paratype, CNC). Discussion: The name intermedius, is an adjective from latin, alluding to the position of the species and its group in the phylogeny of Sterphus. S. intermedius is distinguished from the other species of the chloropygus group in that: 1) it lacks the abundant semiopaque golden pile on the tip of the abdomen; 2) the tarsi are all black; and 3) the male hind trochanter has a bilobed process. Also, the second tergum has two large shiny spots and the third tergum has a broad basal shiny band whereas both woodorum and chloropygus have only the lateral margins of these terga shiny. cybele group The cybele group can be easily separated from all other species groups by its pilose metasterna. Other diagnostic characters for the group have been listed in Table II. Sterphus (Ceriogaster) cybele (Hull) Crepidomyia cybele Hull, 1951, p. 184. Type locality: Peru, Chan- chamayo. Type $ Hull. Subsequent reference: Fluke, 1957, p. 106 (cat. citation). 1973] Thompson: Review of the Genus Sterphus Philippi 217 Head : face bright yellowish orange, yellow pilose, golden pol- linose, without medial shiny stripe; cheeks black, shiny, yellow pilose; front black, shiny except brownish golden pollinose across middle, yellow pilose; vertex shiny black, black pilose; occiput black, yellowish pollinose, brassy yellow pilose throughout. Antennae brownish black, third segment roughly oval, about as long as broad, about as large as metathoracic spiracle. Thorax : black, mesonotum with broad, medial, posteriorly furcate, faint, reddish pollinose vitta and still more faint similar ones sublate rally; lateral margins of mesonotum and medial vitta on anterior half with appressed, opaque, brassy yellow pilose; pleural and scutellar pile brassy yellow; remainder of mesonotum with short black pile, without short black bristle-like hair above wings; scutellum with distinct apical emarginate rim, subscutellar fringe thick, brassy yellow; metasterna with long distinct pile; squamae and halters pale orange. Legs : Anterior legs orange except brownish orange coxae and trochanters, with similarly colored pile; hind coxae black; hind trochanter reddish brown; hind femora yellowish brown basally, becoming reddish through middle, still darker at apices, golden pilose except black pilose on apical quarter, with nine black ventral spines; hind tibiae nearly blackish, but actually dark reddish sepia, with black pile except for short streak of golden hair basolaterally and medio- apically, with two stout, short spinous apical processes; hind tarsi yellowish brown. Wings: almost uniformly pale yellowish brown with costal cells and first basal cell light yellow with pterostigma deep yellow except apically; microtrichose except very narrowly bare behind Ax. Abdomen : dorsum black except slightly reddish on medial sides of second and third terga, dull medially on second thru fourth terga, shiny with brassy yellow to purplish iridescence on all first tergum and laterally on second thru fourth terga, with shiny laterally areas in form of medially pointed triangles; with appressed short black pile on dull areas and long erect golden pile on shiny areas; venter white pilose, with first sternum reddish brown, with second and basal three-fourths of third sterna orange, with apical quarter of third and all of fourth and fifth sterna black. Material Examined: peru: Chanchamayo, 20 Jan. 1949, J. Schunke, 1 2 (holotype, Hull). Colombia: Cundinamarca, Monter- redondo, 19 Jan. 1957, 1 2 (Cornell University). Discussion: S. cybele appears to be a typical member of the subgenus Ceriogaster except that the metasterna are pilose and the face is yellowish orange in ground color. The first character state is 218 Entomologica Americana [Vol. 46, No. 2 unique in Sterphus (usually considered as a primitive state in other syrphids) but the second character state is found in the two primitive subgenera, Telus and Sterphus. The question of whether cybele represents a case of the retention of some primitive traits in a otherwise highly derived fly or convergence in those derived character states, cannot be decided without the study of additional characters, such as those of the male genitalia. Until this question is answered, I have taken the most parsimonious alternative and have tentatively included cybele in Sterphus (Ceriogaster) . plagiatus and jascithorax groups The transverse pollinose markings on the mesonotum of these groups are unique among the taxa of the Xylota group of genera as well as of the tribe Milesini (see Thompson, 1972). The similar pollinose markings in Temnostoma ( Temnostoma group) are quite different and many other characters strongly support my groupings and thereby the contention that this is a convergent character state in Temnostoma. Thus the possession of transverse pollinose mesonotal markings in both the jascithorax and plagiatus groups is synapomorphy and demonstrates the monophyly of this combined grouping. The character states previously used to separate the jascithorax group as a genus are only specialized states easily derived from those of a plagiatus- type ancestor: 1 ) the medial facial keel becomes convex below; 2) the frontal and vertical triangles are elongate and the eyes separate slightly; 3 ) the occiput and mesonotum acquire a row of short spines (a few spines are already present on occiput of plagiatus ); 4) the meta thoracic epimera extends to complete the post- metacoxal bridge; 5) the scutellum loses its ventral pile fringe; and 6) the abdomen becomes slightly constricted. Since the unique character states of the jascithorax group could have been derived from a plagiatus- type ancestor and the group does share synapomorphic character states (facial keels and color, antennal shape, pollinose markings, etc.) with other Sterphus species, the jascithorax group is included in Sterphus and placed as the sister group to the plagiatus group. To maintain the jascithorax group as a separate genus in its traditional sense, would require one of two courses of action: 1) one would have to accept the hypothesis that the jascithorax group arose independently of Sterphus , but that the two are sister groups and that the apparent similarities in various character states are con- vergences due to homoiology (Griffiths, 1972:24) ; or 2) if one accepts my phylogenetic analysis of the group, then one would be required 1973] Thompson: Review of the Genus Sterphus Philippi 219 to elevate almost all of the other species groups here recognized to generic rank to avoid either paraph yletic or polyphyletic genera. I have rejected both these alternatives, the first on the basis of the rule of parsimony and the second on utilitarian grounds (see discus- sion under Telus ) — my arrangement does not involve any con- vergences in character states and utilizes only one medium size genus instead of ten very small genera. plagiatus group The plagiatus group roughly corresponds to the original limits of Shannon’s genus Crepidomyia, but three species groups, which would have been undoubtedly included by Shannon in his genus, have been left out of the plagiatus group. The chloropygus , tinctus, and incertus groups all lack the pollinose bands on the mesonotum and further differ by: the short antennae in the chloropygus and probably incertus groups; the slightly petiolate abdomen in the tinctus group; and the slightly concave and tuberculate face in the incertus group. Sterphus (Ceriogaster) plagiatus (Wiedemann) Xylota plagiata Wiedemann, 1830, p. 98. Type locality: Brazil. Type $ VMNH. Subsequent reference: Kertesz, 1910, p. 304 (cat. citation). Crepidomyia plagiata : Shannon, 1926, p. 48 (descript, note, distr. rec. — Brazil). — Curran, 1934, p. 404 (note on type, distr. rec. — Guyana). — Doesburg, 1963, p. 19 (distr. rec. — Surinam); 1966, p. 98 (distr. rec. — Surinam). — Fluke, 1957, p. 106 (cat. citation). Zelima plagiata : Sack, 1941, p. 116 (also, 1951, p. 112) (distr. rec. — Peru). Head : black; face shiny medially, silvery pollinose laterally; cheeks shiny anteriorly, white pilose, silvery pollinose posteriorly; front bare and sparsely white pollinose on lower half, yellow pilose and thickly white pollinose below becoming browner near vertex on upper half; frontal lunule dark reddish brown; vertex brownish pol- linose, yellow pilose; occiput white pollinose, white pilose below becoming yellower above, with row of short thick spine-like hairs above; antennae elongate, slightly longer than face, brownish black, yellow pilose with a few longer black bristle-like hairs on ventral and dorsal margins of basal two segments; third segment about twice as long as broad, narrowed apically into blunt tip; arista yellowish on basal half, brown apically; distinctly longer than antennae. Thorax : black; pleurae sparsely silvery pollinose, short white pilose, with patch of long hairs behind metathoracic spiracle; mesono- 220 Entomologica Americana [Vol. 46, No. 2 turn mainly blackish pollinose and short, appressed, black pilose except as follows: long white pilose on notopleurae, shorter white pilose on humeri, two lateral golden pilose vittae stretching from basal corners of scutellum and posterior corners of postalar calli to level of wing bases, with patch of short thick spine-like black pile above wing bases, silvery pollinose on mesial sides of humeri and on trans- verse sutures, dark brown pollinose on medial ends of transverse suture, with faint medial and sublateral grayish pollinose vittae; scutellum with appressed black pile except thickly golden pilose on margin. Halters white; squamae white with brown margin and fringe. Legs : front legs dark brownish black, white pilose except black pile on anterior tip of femora, ventral half of tibiae, and all tarsi; middle legs similar to front legs except tawny basal three tarsal segments with white pile; hind legs brownish black except tawny femora, femora short black pilose except white pilose laterally on basal half and dorsal fourth, tibiae sparsely white pilose; tarsi black pilose; hind tibiae with strong apical spur. Wings: (fig. 13), hyaline except for diagonal brown stripe along posterior edge of second basal cell, across base of third posterior and discal cell, through basal two-thirds of apical cell to wing margin; microtrichose except bare behind spurious vein in first basal cell and basad to branching of Rs. Abdomen : black; venter sparsely silvery pollinose; first three sterna white pilose and fourth and fifth sterna black pilose; first tergum silvery pollinose, long white pilose; second tergum brownish black pollinose except shiny reddish laterally on basal seven-eights, shiny portion with long white pile, rest with appressed black pile; third tergum shiny grayish silvery pollinose basally and dull brownish- black pollinose apically shiny area reaching from basal third medially to apical quarter laterally, dull area with short appressed black pile and shiny areas with short white pile; fourth and fifth terga shiny grayish silvery pollinose, short appressed black pilose. Material Examined: Mexico: 1 2 (USNM). panama: Canal Zone, Barro Colorado, 1 Aug 1924, N. Banks, 1 2 (MCZ). British guiana: Kamakusa, 1922, H. Lang, 2 2 (AMNH & HULL). Ecuador: Napo, Napo River, Coca, 250m, May 1965, L. Pena, 3 2 (CNC) ; Napo, Pano River, 12 Feb 1923, F. X. Williams, 1 2 (USNM); Pastaza, Napo River, Pompeya, 14-22 May 1965, L. Pena, 1 2 (CNC). peru: Madre de Dios, Avispas, 400 m, 1-15 Oct 1962, L. Pena. 1 2 (CNC); Middle Rio Ucayali, 18 Nov 1923, H. Bassler, 1 2 (AMNH). brazil: Amazonas, Rio Caiary-Uaupes, 1906, H. Schmidt, 1 2 (AMNH); Amazonas, Rio Cauburi, Boca do Tucano, Nov 1965, E. Dente, 1 2 (MZUSP); Amapa, Rio Felias, 1973] Thompson: Review of the Genus Sterphus Philippi 221 28 July 1959, J. Lane, 1 2 (MZUSP); Amapa, Rio Amapari, 9-10 July 1957, J. Lane, 3 2 (FCT & MZUSP). Discussion: S. plagiatus differs from all other Sterphus species by the presences of a few long hairs behind the metathoracic spiracle and by a diagonal dark stripe on an otherwise hyaline wing (fig. 13). S. plagiatus is very closely related to shannoni but it can be con- trasted as follows: 1) the scutellum has a marginal band of thick, opaque golden pile and lacks spines, not a fine white subscutellar fringe and strong short black spines; 2) the hind femora are tawny, with an appressed black pile on the apical half, not orange with an orange pile; 3) the abdomen has a short appressed black pile, not bands of thick, golden, tomentose pile on the tip of the abdomen; 4) the abdomen has shiny metallic bands on the third tergum, rather than being completely dull black; 5) the first basal cell is bare behind the spurious vein, not microtrichose. Character states 1 and 4 can also be used to distinguish plagiatus from the genuinus complex. Also, S. plagiatus is unique in the plagiatus group because of its short row of black spines on the upper fourth of the occiput. Sterphus (Ceriogaster) shannoni, new species Type locality: Brazil, Santa Catarina, Nova Teutonia. Holotype $ FCT. Head : (fig. 4), black; face shiny medially, golden pollinose laterally; cheeks shiny anteriorly, silvery white pollinose posteriorly, white pilose; frontal lunule reddish brown; frontal triangle of male golden pollinose; front of female mainly shiny on lower half, with medial vitta of golden pollinosity and narrowly whitish pollinose laterally on lower half, light brownish pollinose and yellow pilose on upper half; vertex brown pollinose with some light yellowish pol- linosity around anterior part of ocellar triangle, yellow pilose; occiput silvery white pollinose, yellow pilose. Antennae elongate, slightly longer than face, yellow pilose, light to dark brown on basal two segments, lighter orange brown on apical segment; third segment elongate, about twice as long as broad, with blunt tip but with dorsal margin slightly but distinctly concave before tip; arista yellow, only about as long as antenna. Thorax : black; pleurae sparse silvery pollinose, short yellow pilose except for a few long golden hairs on apical posterior edge of mesoanepisterna; mesonotum mainly dark brownish pollinose and short appressed black pilose, except as follows: long yellow pilose on notopleurae, shorter yellow pilose on humeri and between them, two lateral golden pilose vittae stretching from posterior corner of 222 Entomologica Americana [Vol. 46, No. 2 postalar calli to level of wing bases, with patch of short thick spine- like black pile above wings, golden pollinose on mesial sides of humeri and transverse suture, dark brown velvety pollinose behind golden pollinose, with faint medial and two sublateral grayish pollinose vittae. Scutellum short appressed black pilose, with sparse white ventral fringe, with indistinct apical emarginate rim, with a few very short apical spines. Squamae white with brown margin and fringe; plumulae brown; halters orange. Wings \ light yellowish orange on anterior basal half, elsewhere light brownish; completely microtrichose. Legs: yellow pilose; front legs dark brownish black; middle legs slightly more reddish brown; hind coxae and trochanter dark reddish brown; hind femora and tibiae tawny, light reddish brown; hind tarsi darker; hind trochanter of male with long spur; hind tibiae of both sexes with apical lateral spur and with subapical medial spur in male. Abdomen : black; dorsum dull brownish pollinose except shiny on sides of first and second terga, long white pilose laterally on first and second terga, short yellow appressed pilose elsewhere and with thicker opaque golden pile in form of transverse bands on apex of third tergum and basal margin and apical half of fourth tergum. Venter shiny on first and second sterna, light pollinose on rest of sterna, yellow pilose. Male genitalia : (fig. 28), cerci broad, rounded dorsally, with small medial tuberculate swelling. Surstyli: dorsal lobe elongate, narrow, with dorsal margin angulate and ventral margin evenly curved, with apex blunt, with many long hairs medially on dorsal edge; ventral lobe broad, broader than long, with rounded apical margin. Ninth sternum without dorsolateral opening, with large lateral membranous area above base of lingula; lingula very long and narrow, ending in small upturned point. Superior lobe with dorsoapical margin rounded, without apical hook-like process, ex- tensively pilose, with two subbasolateral and one basolateral processes, with all these blunt-tipped and extending toward lingula. Aedeagus with dorsal lobe produced posteriorly, with lateral lobes broad and short, with lateral lobes not extending between lateral flaps of dorsal lobe, with ventral lobe produced on basal section and with apical section flatten. Ejaculatory apodeme umbrella-shaped. Material Examined: brazil: Parana, Caviuna, Sept. 1947, A. Mailer 1 $ (AMNH) Sao Paulo, Campinas, Mar. 1924, F. X. Wil- liams, 1 $ (USNM) ; Sao Paulo, Villa Americana, Feb. 1924, F. X. Williams, 1 2 (USNM); Sao Paulo, Rio Claro, Adutora, Sept. 1940, 1 $ (USNM) Sao Paulo, S. Sebastiao, A. A. Barbiellini, 1 $ (USNM); Sao Paulo, Sto. Amaro, 8 May 1944, Oct. 1950, Feb. 1973] Thompson: Review of the Genus Sterphus Philippi 223 1950, J. Lane, 2 $ 1 9 (MZUSP); Santa Catarina, Nova Teutonia, Sept.-May, 191 4 $ (AMNH, CNC, FCT, MCZ, MZUSP, Hull, BM(NH); holotype male, allotype female, both Nov. 1971, in FCT; other paratypes). Argentina: Cordoba, Dept. San Javier, La Paz, 15-31 Dec. 1928, C. Bruch, 1 5 (USNM). Discussion: S. shannoni is quite easily separated from all other species of Sterphus by its golden tomentose pile bands on the tip of the abdomen. For discussion of the other differences between shan- noni, plagiatus and the genuinus complex, see the discussion under those species. S. shannoni is named for Raymond Corbet Shannon, who needs no introduction to syrphids workers. genuinus complex S. genuinus (Willis ton) and trier epis (Shannon) are very closely related and apparently only distinguishable by a few characteristics of the male genitalia. These genitalic differences are as follows: 1) the apical hook of the superior lobe is rounded dorsally and is sharp and simple apically in genuinus (fig. 31s) but in trier epis it is angulate dorsally and is blunter and notched apically (fig. 32s); 2) the distribution of setae on the superior lobe is more uniform in genuinus (fig. 31s) but in tricrepis it is reduced (fig. 32s); 3) the ventral portion of medial lobe of aedeagus is broad and perpendicular in genuinus (fig. 31a) but in tricrepis it is thin and recurved (fig. 32a) ; 4) the lower lobe of the surstyle is short and somewhat angulate posteriorly in genuinus (fig. 3 It) but in tricrepis it is longer and more curved posteriorly (fig. 32ss); and 5) the left surstyle is only produced slightly on the lateral side (fig. 31ssd), but in tricrepis it is greatly produced laterally (fig. 32ssd). The following description and discussion applies to both species. Head : black; face shiny medially, white to silvery yellow pollinose laterally in female, golden pollinose laterally in male; cheeks shiny anteriorly, silvery white pollinose posteriorly, white pilose; frontal lunule orange; front of female shiny on lower half, white pollinose and yellow pilose on upper half; frontal triangle of male golden pol- linose; occiput silvery white pollinose, white pilose below becoming yellower above with some black hairs intermixed on upper third. Antennae shorter than face, brownish black except yellow arista, black pilose; third segment obtusely pointed, about IV2 times as long as broad; arista about twice as long as antenna. Thorax : black; pleurae sparse silvery pollinose, short white pilose except for a few long black hairs on apical posterior tip of mesoan- 224 Entomologica Americana [Vol. 46, No. 2 Figs. 31-32. Male genitalia of Sterphus, lateral view except ssd dorsal view; 31. genuinus (Williston), 32. tricrepis (Shannon), a = aedeagus and base of apodeme (32a is only ventral prolongation of lateral lobe), s = 9th sternum, ss = surstyle, ssd — surstyle dorsal view, t = 9th tergum. 1973] Thompson: Review of the Genus Sterphus Philippi 225 episterna; mesonotum mainly brownish pollinose with faint medial and two sublateral grayish pollinose vittae, with transverse sutures and mesial sides on humeri golden pollinose, appressed black pilose except for long white pilose on notopleurae, with black spine-like hairs above wings; scutellum brownish pollinose, short appressed black pilose, with very indistinct apical emarginate rim, with long white ventral fringe, with six to eight short strong marginal spines. Squamae white with brown margin and fringe; plumulae brown; halters yellow. Wings : very light brownish black, microtrichose except bare in front of Rs and behind spurious vein on basal half of first basal cell. Legs : black; mainly black pilose, white pilose as follows: on coxae, on basal two-thirds of anterior and all posterior side of front and middle femora, on basal half of both anterior and posterior sides of hind femora, dorsal half of front and middle tibiae; hind trochanter with short apical spur in female and longer one in male. Abdomen : (fig. 18), black, mainly dull grayish black pollinose, shiny on sides of first thru third terga and medially on first and third sterna and on all of second sterum; second tergum frequently with pair of small lateral orange spots in male; long white pilose on shiny areas including all of first and third sterna, short yellow subappressed pilose on apical half of fourth tergum, appressed black pilose else- where on dorsum and long black pilose on fourth sterna; male genitalia with yellow pile. Male genitalia : (figs. 31-32), cerci quadrate with basal portion flattened, with dorsal margin evenly rounded. Surstyli: dorsal lobe elongate, broad, straight, not curved ventrally or medially, with blunt apex, on left side slightly to greatly produced basolaterally, with some long marginal pile on dorsal edge; ventral lobe broad, short, either with angulate or rounded apex. Ninth sternum with small dorsolateral membranous opening; lingula short, broadly triangular, with apex rounded; superior lobe with dorsoapical margin either angulate or rounded, densely or sparsely pilose, with apical portion hook-like, with either single sharp point or bifid point. Aedeagus: dorsal lobe large, arched produced dorsally; lateral lobes divided into short posteriorly directed process and large ventrally directed process, with posteriorly directed process with blunt tip and ventrally directed process extending into sharp point; ventral lobe produced ventrally on basal portion. Ejaculatory apodome umbrella- shaped. Discussion: S. genuinus and tricrepis are very similar to shan- noni and plagiatus but can be separated from both species by their 1) lack of longitudinal stripes of appressed golden pile on the posterior mesonotum; 2) shorter and blunt tipped third antennal 226 Entomologica Americana [Vol. 46, No. 2 segment; 3) black pile on the upper posterior corner of the meso- pleurae; 4) spurs on the hind trochanter in the female, and 5) lack of well developed spurs on the hind tibiae. The S. genuinus complex further differs from shannoni in its 1) lack of golden tomentose pile on the abdomen, 2) black hind femora, and 3) hyaline wings. Other differences between genuinus and plagiatus are listed in the discussion of the latter species. Sterplius (Ceriogaster) tricrepis (Shannon) Crepidomyia tricrepis Shannon, 1926, p. 47. Type locality: Peru, Rio Charape. Type $ USNM. Subsequent references: Fluke, 1950, p. 453 (synonymy under genuinus Williston). Crepidomyia darlingtoni Hull, 1944, p. 40. Type locality: Cuba, Eastern Oriente, Mounts north of Imias, 3-4000 ft. Type $ MCZ. Subse- quent reference: Fluke, 1957, p. 106 (cat. citation). New synonymy. ? Crepidomyia dion Hull, 1951, p. 183. Type locality: Peru, Chancha- mayo. Type $ Hull. Subsequent reference: Fluke, 1957, p. 106 (cat. citation). Discussion: When Hull described darlingtoni he stated that it was “related to tricrepis Shannon but very distinct in the smoky wing apex.” I have examined the holotype of darlingtoni including the male genitalia and can find no other differences between it and tricrepis. Thus, I feel the slight difference in the intensity of the color of the wing apex, tricrepis having light brownish black wings, is only of trivial importance. I have also examined the type of dion Hull and can find no differences between it and the females of the genuinus complex. Since the females of the genuinus are indistinguishable, I have tentatively synonymized dion under tri- crepis on the basis of geographical distribution, that is, all the males of the genuinus complex examined from Peru, the type locality of dion , have been tricrepis. Material Examined: cuba: Pico Turquino, Cumbre, 22 July 1922, S. C. Bruner & C. H. Ballou, 1 9 (AMNH); Eastern Oriente, Mounts north of Imias, 3-4000 ft., 25-28 July 1936, P. J. Darling- ton, 1 <3 (holotype of darlingtoni , MCZ). dominica: Pont Casse, April 1965, 12 Oct.-30 Nov. 1964, P. J. Spangler & R. Davis, 8 9 (USNM), 27-30 Nov. 1964, P. J. Spangler, 1 <3 (USNM). peru: Rio Charape, 14 Sept. C. H. T. Townsend, 1 <3 (holotype of tricrepis, USNM); Piches & Perene Valleys, 2000-3000 ft., 1^29 (paratypes of tricrepis , USNM). Bolivia: Cochabamba, Oct. 1965, Steinbach, 1 <3 (CNC); El Limbo, 65°36'W 17°07'S, 2200 m, Nov. 1963, F. Steinbach, 1 <3 (CNC). 1973] Thompson: Review of the Genus Sterphus Philippi 227 Sterphus (Ceriogaster) genuinus (Williston) Xylota genuina Williston, 1888, p. 284. Type locality: Brazil, Rio de Janeiro. Type 9 AMNH. Subsequent references: Kertesz, 1910, p. 301 (cat. citation). — Shannon, 1926, p. 50 (suggested placement in Crepidomyia) . Zelima genuina : Sack, 1941, p. 116 (also, 1951, p. 112) (distr. rec. — Peru [= tricrepisl ] ) . Crepidomyia genuina : Fluke, 1950, p. 453, fig. 19 (male genitalia) (synonymy of tricrepis ; distr. recs. — Brazil, Argentina); 1957, p. 106 (cat. citation). Material Examined: costa rica: Cartago, Turrialba, La Suiza, 1 Oct., 13 Nov., P. Schild, 2 $ (USNM). Venezuela: Mt. Duida, 28 Dec. 1928, 16 Jan. 1929, Tate, 2 $ (AMNH). brazil: Minas Gerais, Passa Quatro, 30 March 1916, 1 $ (USNM); Rio de Janeiro, Itatiaya, 1200 m, Feb. 1941, R. C. Shannon, 7 8 5 $ (USNM); Rio de Janeiro, Itatiaya, Maromba, 23 Dec. 1935, 11 April 1940, J. F. Zikan, 2 8 (MZUSP & FCT); Rio de Janeiro, Dist. Federal, Jan. 1938, 1 8 (USNM); Sao Paulo, C. Jordao, Jan. 1954, J. Lane, 1 8 (MZUSP); Santa Catarina, Nova Teutonia, March 1966, Sept. 1966, 1 <5 1 $ (CNC). Argentina: Tucuman, Villa Nougues, 13 Jan. 1929, Jan. 1929, R. A. Jaynes, 3 (USNM). fascithorax group Face with medial carina slightly concave below antennal pits and slightly convex below that concavity; cheeks narrow; frontal triangle of male about two-thirds as long as vertical triangle; vertical triangle of male long, more than twice as long as broad at occiput; occiput with transverse row of short strong spines on upper half; eyes with an oblique transverse impressed groove at level of antennal pits, narrowly dichoptic in male; third antennal segment always elongate, two or more times as long as broad; anterior edge of mesonotum with transverse row of short strong spines; postmeta- coxal bridge complete; metathoracic spiracle small; scutellum with- out ventral pile fringe; abdomen slightly petiolate. Discussion: The species of this group will be reviewed in a separate paper. The group as a whole is quite distinctive, with its 1) complete postmetacoxal bridge, 2) lack of a scutellar fringe, 3) row of spines on the occiput and the mesonotum, and 4) modified medial facial keel. It is hardly likely that the species of the fascithorax group will be confused with any other Sterphus species. 228 Entomologica Americana [Vol. 46, No. 2 tinctus group The tinctus group is readily distinguished by its abdominal shape (fig. 19) and the lack of hind tibial spurs. Other diagnostic characters for the group have been listed in Table II. Sterphus (Ceriogaster) tinctus (Fluke) Crepidomyia tincta Fluke, 1950, p. 453, fig. 20 (head), 21 (wing). Type locality: Brazil, Santa Catarina, Nova Teutonia. Type $ AMNH. Subsequent reference: Fluke, 1957, p. 106 (cat. citation). Head : black; face shiny medially and broadly golden pollinose laterally in male, almost completely shiny in female, with only very narrow stripe of pollinosity on border of cheek in female; cheek shiny, linear, white pilose; frontal lunule orange; frontal triangle of male golden pollinose; front of female with lower half shiny, with upper half dark brownish pollinose, with pollinosity lighter at junction of shiny area, yellow pilose; vertex brownish pollinose, yellow pilose; occiput sparsely yellowish brown pollinose, white pilose below becoming yellower above. Antennae brownish orange, black pilose, longer than face in female, about as long as face in male; third segment about IV2 times as long as broad in female, slightly longer than broad in male; arista orange, more than twice as long as antenna. Thorax : brownish black; mesonotum mostly dull brownish pol- linose, with transverse sutures black velvety pollinose, with faint grayish medial pollinose vitta, grayish pollinose on mesial sides of humeri, short appressed brownish yellow pilose except black pilose medially on posterior half; without black spine-like pile above wings; pleurae brownish yellow pollinose, short yellow pilose; scutel- lum appressed yellow pilose, with indistinct apical emarginate rim, with yellow ventral fringe, without spines; squamae and halters orange; plumulae reddish brown. Wings: light yellowish brown, dark brown on anterior half; microtrichose except bare anterior third of anal cell. Legs : mainly orange, slightly darker on anterior legs of male, yellowish to orange pilose, front coxae brownish black on basal four-fifths; middle and hind coxae brownish basally, front tarsi with apical four segments brownish black, middle and hind tarsi with apical two segments dark; tarsi with dark pile on dark segments, hind trochanter with very short stubbly papilla. Abdomen : elongate, slightly constricted on second segment, minimal width about two-thirds of maximal width (two-thirds in female; three-fifths in male); dark brownish black, dull brownish black pollinose, mainly short yellow to golden pilose, slightly longer 1973] Thompson: Review of the Genus Sterphus Philippi 229 pile basolaterally. Male Genitalia', (fig. 29) cerci quadrate, with dorsal margin slightly rounded; dorsal lobe of surstyli elongate, broad, angulate, with broad apex, apparently without long marginal pile; ventral lobe of surstyli greatly reduced, completely bare; ninth sternum with small dorsolateral membranous opening; lingula short, very broad, with apex broadly truncate; superior lobe with dorso- apical margin rounded, sparsely pilose, without apical hook-like process, with large lateral excavation extending up from ventral margin; aedeagus with dorsal lobe produced posterodorsally, and arched, with long and wide cleft between dorsal and lateral lobes, with lateral lobes broad and tapering to long sharp apex, with ventral lobe flattened; ejaculatory apodeme umbrella-shaped. Material Examined: brazil: Santa Catarina, Nova Teutonia, August-January, F. Plaumann, 21 $ 2, (FCT, BM(NH), & AMNH ( including holotype ) ) . Discussion: S. tinctus is quite similar to S. auricaudatus in overall appearance, both species have slightly petiolate abdomens, general brownish black color, brown anterior edge of the wing, and simple legs. However, S. tinctus can be easily separated by its straight black face and lack of golden tomentose-like pile on the abdomen. Also, the possession of the apomorphic black and straight face character states clearly demonstrates that the similarities between S. auricaudatus and tinctus are due to convergence. coarctatus group The coarctatus group can be easily separated from all other species groups by its abdominal shape (fig. 20) and single hind tibial spur. Other diagnostic characters for the group have been listed in Table II. Sterplius (Ceriogaster) coarctatus (Wiedemann) Xylota coarctata Wiedemann, 1830, p. 100. Type locality: Brazil. Type 2 SMF. Subsequent reference: Kertesz, 1910, p. 298 (cat. citation). Tatuomyia coarctata : Shannon, 1926, p. 49 (differential diagnosis from batesi ). — Fluke, 1957, p. 105 (cat. citation). Zelima coarctata: Sack, 1941, p. 116 (also, 1951, p. 112) (distr. rec. — Peru). Senoceria spinifemoratas Hull, 1930, p. 145. Type locality: Brazil. Type $ ANSP. Head : face black, shiny medially, golden pollinose laterally; cheeks black, shiny, white pilose; frontal lunule orange; frontal triangle of male golden pollinose, with a few marginal yellow hairs; 230 Entomologica Americana [Vol. 46, No. 2 Figs. 33-34. Fig. 33. Male genitalia of Sterphus coarctatus (Wiedemann), lateral view except sd dorsal view; 34. Mesonotal pollinose pattern of Sterphus telus Thompson, a — aedeagus, s = 9th sternum, t = 9th tergum and associated structures (with outline of tip of lower lobe of left surstyle). 1973] Thompson: Review of the Genus Sterphus Philippi 231 front of female shiny on lower half, silvery pollinose and white pilose on upper half; vertex black, silvery pollinose in front of ocellar triangle, brown pollinose on ocellar triangle, yellow pilose; occiput black, white pollinose, white pilose below becoming yellower above; antennae tawny, except dark third segment, with white pile on medial surface and black pile on lateral surface, slightly longer than face; third segment elongate, with blunt tip, twice as long as broad; arista orange, about three-fourths longer than face. Thorax: black; pleurae lightly silvery pollinose, short yellow pilose; mesonotum dark grayish pollinose except as follows: golden pollinose on transverse sutures and mesial sides of humeri, dark brown pollinose behind the areas of golden pollinosity, sparsely light golden pollinose in form of medial longitudinal stripe with splits into two stripes behind the transverse suture and two broad sub- medial longitudinal stripes starting behind transverse sutures; meso- notal pile short appressed golden in front of transverse sutures, short appressed golden in form of anteriorly pointed triangular patch on postalar and adjacent mesonotum; scutellum black, dark grayish pollinose, appressed yellow pilose, subscutellar fringe golden yellow; squamae white, with brown margin and fringe; halters dark reddish brown with yellow stalk; plumulae light brownish yellow. Legs’, dark reddish brown, white pilose except black pilose on dorsal edge on apical quarter of hind femora; hind tibiae with single short spur; hind trochanter of male with short spur. Wings: (fig. 14), with anterior portion metallic purplish black, hyaline behind; with dark color extending back to spurious vein, second basal cell light yellowish brown; microtrichose except bare narrowly along anterior edge of anal cell, apical three-fourths of alula, behind and in small spot in front of Ax. Abdomen: (fig. 20), black, except red petiole, dark grayish pollinose except shiny first segment and petiole; pile long white on first segment and lateral margin of second tergum, long black on third and fourth sterna, short appressed black with a few scattered yellow hairs on second and third terga, short appressed yellow with a few scattered black hairs on fourth tergum; fifth tergum of female with short appressed black pile. Male genitalia: (fig. 33), cerci slightly expanded dorsally, with dorsal margin rounded. Surstyli: dorsal lobe elongate, straight except dorsal and ventral margins slightly sinuous, with some long marginal pile on dorsal edge, with apex broadly rounded; ventral lobe broad, almost square, with posteroapical corner of left ventral lobe slightly produced; ninth sternum without dorsolateral opening, with large lateral lobe, with 232 Entomologica Americana [Vol. 46, No. 2 medial side of this lobe short pilose; lingula short, broadly triangular, with apex rounded; superior lobe with dorsoapical margin rounded, with apical hook-shaped process, with four large teeth on ventral margin; aedeagus with dorsal lobe produced posterodorsally, with lateral lobes small and broadly rounded apically, with ventral lobe long; ejaculatory apodeme umbrella-shaped. Material Examined: panama: Chorrera, 6 June 1948, 1 $ 1 2 (MCZ). Guyana: Essequibo River, Moraballi Creek, 8 Sept. 1929, Oxford University Expedition, 1 2 (BM(NH)). brazil: Amazonas, Rio Tucano, Serra da Neblina, 230 m, 4 Dec. 1965, E. Dente, 2 $ (MZUSP); 24 July 1936, Lange de Morretes, 1 2 (MZUSP); “Freireiss” [=G. W. Freyreiss], 1 2 (holotype of coarctatus , SMF); H. B. Merill, 1 $ (holotype of spinifemorata, ANSP). Discussion: S. coarctatus is easily separated from all other described Sterphus by its abdominal shape and other species group characters as discussed above. I have examined a pair of coarctatus from Panama which differs from all the other material studied in that 1 ) the wings are bordered with dark brown, not metallic purplish black and 2) the anal cell is extensively bare (fig. 15). Since I can find no other differences (including genitalic differences) between these Panamanian specimens and the others I am attributing the above mentioned differences to geographic variation. batesi group The synapomorphic character states of the batesi group are the long abdominal petiole and the presence of two apical spurs on the hind tibiae. Sterphus (Ceriogaster) batesi (Shannon) Tatuomyia batesi Shannon, 1926, p. 48. Type locality: Brazil, Ega. Type $ BM(NH). Subsequent reference: Fluke, 1957, p. 105 (cat. citation). Head : black; face silvery pollinose except for small medial triangular spot on facial carina under antennal base; cheeks shiny, white pilose; frontal triangle shiny, bare; frontal lunule dark brown- ish orange; vertical triangle silvery pollinose below ocellar triangle, shiny above ocellar triangle, with a few short yellowish hairs on ocellar triangle; occiput silvery pollinose, white pilose below be- coming yellower above, with a few black hairs intermixed on upper third. Antennae dark brown, as long as face, black pilose; third 1973] Thompson: Review of the Genus Sterphus Philippi 233 segment about three times as long as broad, with tip blunt; arista yellowish, almost twice as long as antennae. Thorax : black; dorsum shiny except silvery pollinose on mesial side of humeri and dark brownish pollinose between humeri, black pilose except white pilose on notopleurae, with pile greatly reduced and appressed; pleurae silvery pollinose, white pilose; metathoracic spiracle about half as large as third antennal segment is long; halters white with brown head; squamae white with brown margin and fringe; scutellum shiny, black pilose except yellowish subscutellar fringe. Legs : mainly black, except yellowish brown on apical third of hind femora and basal three middle tarsal segments. Wings: (fig. 16), bicolored, brownish black anteriorly, hyaline posteriorly; black area extending to anal cell, to discal cell before anterior cross- vein and to diagonal line from anterior crossvein to apex of apical cell; almost completely microtrichose, except bare narrowly on both sides of basal half of anal vein, on apical corner of anal cell, and on spot in front of and behind vein and on alula. Abdomen : (fig. 21), black except reddish brown petiole; shiny except brownish black pollinose on apical third of second segment, black pilose except white pilose on first segment and petiole of second segment, with dorsal pile greatly reduced and appressed. Male genitalia : (fig. 22), cerci broadly expanded and flattened dorsally, with narrow basal neck; dorsal lobe of surstyli elongate, broad, with dorsal margin angulate, apical margin truncate; ventral lobe of surstyli broad, broader than long, with rounded apical margin; ninth sternum with large dorsolateral membranous opening; lingula short, blunt apically; superior lobe with dorsoapical margin rounded, with anteroventrally directed process on posteroventral margin, with small basoventral lateral tooth; aedeagus with dorsal lobe postero- dorsally produced and arched, with lateral lobes divided into long thin and ventrally produced basolateral process and broader curved anteriorly directed process, with ventral lobe long and produced on basal half; ejaculatory apodeme umbrella-shaped. Material Examined: brazil: Amazons, Saunders Collection, 1 $ (OXF). Discussion: The specimen on which the above description was based agrees with Shannon’s original description of batesi in ail particulars except that the basal three segments of the middle tarsi are dark orangish brown, not black. However, Dr. Hippa has compared the above specimen with the holotype of batesi and states that the two are conspecific. For a discussion of the differences between batesi and fulvus check under the latter. 234 Entomologica Americana [Vol. 46, No. 2 Sterphus (Ceriogaster) fulvus, new species Type locality: Brazil, Sao Paulo, Salesopolis, Boracea. Holotype $ MZUSP. Head : (fig. 3), face black medially, reddish brown laterally, shiny medially, white pollinose laterally; cheeks dark reddish brown, with a few white hairs posteriorly; frontal lunule orange; front dark reddish brown, shiny and bare on lower half, brownish black pol- linose and yellow pilose on upper half except white pollinose along eye margins; vertex dark reddish brown, dark brownish black pol- linose, yellow pilose; occiput black, white pollinose, white pilose below becoming yellow above. Antennae elongate, almost as long as face, dark orange, orange pilose except for a few black bristle- like hairs on dorsal and ventral margins of first two segments; third segment rectangular, twice as long as broad; arista orange on basal third, dark apically slightly longer than antennae. Thorax : dark brown, except slightly reddish on humeri and postalar calli, uniformly brownish pollinose except white pollinose on mesial side of humeri and indistinct light brownish pollinose medial and two submedial vittae, short yellow pilose, without short black spine-like pile above wings. Legs : coxae dark brown ex- cept orange on ventral portion of hind coxae, yellow pilose; tro- chanters dark reddish brown, yellow pilose; femora tawny, yellow pilose; anterior tibiae and tarsi dark brownish black, black pilose except with yellow pile intermixed on dorsal surface; middle tibiae tawny except with brown, broad, apical band, yellow pilose; middle tarsi tawny except dark brownish apical two segments, yellow pilose; hind tibiae and tarsi tawny except dark brownish apical two segments, yellow pilose. Halters tawny, head not dark; squamae brownish. Wings: (fig. 17), brownish anteriorly, hyaline posteriorly, with ex- tent of brown area same as in batesi, microtrichose except mostly of anal cell and behind anal cell bare and alula bare. A bdomen : dark brown except tawny petiole, yellow pilose except black pilose on fifth segment and intermixed black pile on sterna. Material Examined: brazil: Sao Paulo, Salesopolis, Boracea, 24-30 January 1953, Collectors — Carrera, Vanzolini, Oiticica, Pear- son; 1 9 (holotype, MZUSP). Discussion: S. fulvus is very similar to batesi Shannon but as its name implies it is a dull brownish fly whereas batesi is a shiny black fly. The principal differences between fulvus and batesi are as follows: 1) the antennae are dark orange, not black; 2) the mesonotum is pollinose and brownish, not shiny black; 3) the postalar calli are golden pilose, not black; 4) the pile above wings 1973] Thompson: Review of the Genus Sterphus Philippi 235 is orange and long, not black and bristle-like; 5) the scutellum is golden pilose, not mainly black; 6) the head of the halters are orange, not brownish black; and 7) abdominal terga 3 and 4 are pollinose, brownish and with golden pile, not shiny black with black pile. S. julvus can also be separated from batesi by its more extensively bare anal cell (fig. 17) and less swollen hind femora. “Xylota” ventralis Walker Xylota ventralis Walker, 1858, p. 96. Type locality: Aru Islands. Type $ BM(NH). Subsequent references: Kertesz, 1910, p. 307 (cat. citation). — Shannon, 1926, p. 50 (“apparently belongs to . . . Crepi- domyia”). Head : (fig. 8), black; face bare, carinate, with slight con- cavity, black with metallic bluish lustre, white pollinose except shiny V-shaped medial area; cheeks shiny; front shiny except with broad pollinose band across medial half, light yellow pilose; vertex shiny, light yellow pilose; ocellar triangle slightly anterior to posterior margin of eyes; occiput completely white pollinose, yellow pilose. Antennae black; third segment elongate, twice as long as second. Thorax : black, with metallic bluish luster except orange postalar calli and scutellum; pleurae lightly white pollinose and short yellow pilose, anterior mesoanepisterna bare; posterior mesoanepimera bare; barrette and hypopleurae bare; metathoracic pleurae bare; meta- sterna pilose and developed; postmetacoxal bridge incomplete; scutel- lum with distinct emarginate rim, with ventral pile fringe (single row) ; mesonotum without pollinose markings except white pollinose along anterior margin, short appressed black pilose except yellow pilose as follows: on notopleurae, humeri, along transverse suture, on and in front of postalar calli and scutellum, and in medial area in front of and between transverse sutures; squamae white with black border and fringe; plumula brownish. Wings: with slight brownish tinge; stigma brown; marginal cell open; apical cell petiolate, with petiole longer than humeral crossvein; anterior crossvein slightly before middle of discal cell, slanted; anal cell petiole straight, slightly longer than petiole of apical cell; marginal crossveins disjunctive, not contin- uous, with external spurs at their bases; microtrichose except bare as follows: basal quarter of costal cell, area between Rs and Rl, anterior half and basal quarters of second basal cell, narrowly along anterior edge of anal cell, along posterior edge of anal cell and in front Ax. Legs: anterior four legs light orange except darker reddish orange on trochanters and apical tarsal segments, yellowish pilose, with coxae thickly white pollinose; hind coxae dark reddish brown with 236 Entomologica Americana [Vol. 46, No. 2 metallic bluish luster, sparsely white pollinose, white pilose; hind trochanters dark reddish brown; hind femora (fig. 9) evenly swollen, orange on basal and apical quarter, darker reddish orange medially, yellow pilose; hind tibiae orange on basal half darker reddish orange apically, yellow pilose; hind tarsi brownish orange, black pilose above, orange below. Abdomen : metallic bluish black, appressed white pilose laterally on first three terga (color of pile on medial area cannot be seen because wings are folded on the area ) , fourth tergum appressed white pilose on basal half and black pilose apically; fifth tergum all black pilose. Material Examined: Aru Island, A. R. Wallace, 1 9 (holo- type, BM(NH) ). Discussion: When Shannon described his genus Crepidomyia he referred “ Xyloto ” ventralis Walker to it. I have examined the type of ventralis and find that it definitely does not belong to Sterphus. “ Xylota ” ventralis probably deserves separate generic status. How- ever, without a study of the characters of the male genitalia of ventralis , it is difficult to properly place ventralis within the present classification of the Xylota group. “ Xylota ” ventralis appears similar to Hardimyia Ferguson. It differs from elongata (Hardy), the unique species of Hardimyia , as follows: 1) the second antennal segment is not elongate, but about half as long as the third segment, not longer than the third segment; 2) the face has a shiny V- shaped area, not completely pollinose; 3) the scutellum is orange, not black; and 4) the hind femora are completely reddish, not black apically. This species will key to Neplas in my key (Thompson, 1971) to syrphid genera with carinate faces. Xylota ventralis differs from Neplas as follows: 1) the face is nearly straight, not concave; 2) the pleurotergum is not carinate; 3) the abdominal sterna are normal, not greatly narrowed; and 4) the hind femora are not as strongly swollen as in Neplas nor are the hind tibiae strongly arcuate. ABBREVIATIONS Museums and Private Collections AMNH — American Museum of Natural History, New York ANSP — Academy of Natural Sciences, Philadelphia BM(NH) — British Museum (Natural History), London CNC — Canadian National Collection, Ottawa 1973] Thompson: Review of the Genus Sterphus Philippi 237 FCT Hull MCZ MNHN MZUSP SMF USNM VMNH — F. Christian Thompson Collection — Frank M. Hull Collection, Oxford, Mississippi — Museum of Comparative Zoology, Cambridge — Museo Nacional de Historia Natural, Santiago — Museu de Zoologia, Universidade de Sao Paulo — Senckenbergisches Museum, Frankfurt am Main — United States National Museum, Washington — Naturhistorisches Museum, Vienna Used in synonymy cat. citation descrip, note descript, distr. data distr. rec. (s) key ref. — catalog citation — descriptive note — description (full) — distribution data — distribution record (s) — placement in a key ACKNOWLEDGEMENTS I would like to thank the following: Dr. H. Hippa for comparing my S. coarctatus material with the type; Dr. F. M. Hull for the loan of the Sterphus types in his personal collection; Dr. L. V. Knutson, USD A, ARS, Systematic Entomology Laboratory, Washing- ton, for the loan of material and permission to study and dissect the type of S. trier epis Shannon; Dr. R. Lichtenberg, Naturhistorisches Museum, Vienna, for the loan of the types of S. chloropygus Schiner and information on types of S. aurifacies (Schiner); Dr. J. F. Lawrence, Museum of Comparative Zoology, Cambridge, for the loan of material and permission to study and dissect the type of S. darlingtoni (Hull); Dr. W. W. Morris, Academy of Natural Sciences, Philadelphia, for the loan of the type of S. spinijemoratus (Hull); Dr. N. Papavero, Museu de Zoologia, Universidade de Sao Paulo, for the loan of material; Mr. K. G. V. Smith, British Museum (Natural History), London, for his notes on the types in his custody; Dr. W. Tobias, Senckenbergisches Museum, Frank- furt am Main, for the loan of the type of S. coarctatus (Wiedemann); Dr. J. R. Vockeroth, Canada Department of Agriculture, Entomology Research Institute, for the loan of material; and Dr. P. Wygodzinsky for his critical comments on this paper and other things too numerous to mention. 238 Entomologica Americana [Vol. 46, No. 2 REFERENCES Crowson, R. A. 1970. Classification and Biology. New York, Atherton Press, ix+ 350 pp. Curran, C. H. 1934. Diptera of Kartabo, Bartica District, British Guiana. Bull. Amer. Mus. nat. Hist. 66: 287-532. Darlington, P. J., Jr. 1971. The carabid beetles of New Guinea. Part IV. General Con- siderations; Analysis and History of Fauna; Taxonomic Sup- plement. Bull. Mus. comp. Zool. 142(2): 129-337. Doesburg, P. H. van. 1963. Preliminary list of Syrphidae known from Suriname and British and French Guiana. Stud. Fauna Suriname [=Natu- urwet. Stud. Suriname] 5(28): 1-33. 1966. Syrphidae from Suriname. Additional records and descriptions. Stud. Fauna Suriname [=Natuurwet. Stud. Suriname] 9(35): 61-107. Etcheverry, M. 1963. Descripciones originales, sinonimia y distribution geografica de las especies de al familia Syrphidae (Diptera) en Chile. Publ. Centro Estud. Ent. 5: 1—141. Etcheverry, M. & R. D. Shenefelt. 1962. A preliminary study of the genitalia of Chilean Syrphidae. Verhandl. XI Internatn. Kongr. Ent. 1960 1: 207-214. Fluke, C. L. 1950. Some new tropical syrphid flies with notes on others. Acta Zool. Lilloana 9: 439-454. 1955. Los Insectos de las Islas Juan Fernandez. 18. Syrphidae (Diptera). Revta Chilena Ent. 4: 39-43. 1956-57. Catalogue of the family Syrphidae in the Neotropical region (Diptera). Revta Brasil. Ent. 6: 193-268, 7: 1-181. Hull, F. M. 1930. Some new species of Syrphidae from North and South America. Trans. Amer. ent. Soc. 56: 139-148. 1943. The genus Ceriogaster Williston (Syrphidae). Revta Soc. Ent. Argentina 12: 137-140. 1949. The morphology and inter-relationship of the genera of syrphid flies, recent and fossil. Trans, zool. Soc. London 26: 257-408, 25 figs. 1951. Some New World Xylotinae (Diptera: Syrphidae). Pan- Pacific Ent. 27: 183-186. Kertesz, K. 1910. Catalogus dipterorum hucusque descriptorum. Vol. 7, Lipsiae, Budapestini [^Leipzig, Budapest], 470 pp. 1973] Thompson: Review of the Genus Sterphus Philippi 239 Lynch Arribalzaga, F. 1892. Dipterologia Argentina (Syrphidae). An. Soc. Cien. Argentina 33: 189-199. Philippi, R. A. 1865. Aufzahlung der chilenischen Dipteren. Verhandl. K. -k. Zool. -Bot. Gesell. Wien 15 ( Abhandl. ): 595-782. Pino, G. 1962. Estudios de algunos caracteres morfologicos de nueve especies de Syrphidae Chilenos de diferentes generos (Diptera). Publ. Centro Estud. Ent. 4: 45-58. Porter, C. E. 1932. Acerca de algunos Insectos Chilenos. Revta Chil. Hist. nat. 36: 190-193. 1937. Acerca de la distribucion geografica de algunos Insectos Chilenos. Revta Chil. Hist. nat. 41 : 41-42. Rondani, C. 1863. Diptera exotica revisa et annotata. Modena, 99 pp., 1 pi., (also published as “Dipterorum species et genera aliqua exotica” 1864, Arch. Zool. anat. Fis., Modena (1863) 3(1): 1-99, pi. 5.). Sack, P. 1941. Syrphidae, pp. 97-120. In Titschack, E., Beitrage zur Fauna Perus. Band I, 344 pp. [Most copies of this publication were destroyed during World War II. It was reissued as Sack, 1951. see Zool. Rec. 1951, 88(13): 132 (#2537)]. 1951. Syrphidae, pp. 93-116. In Titschack, E., Beitrage zur Fauna Perus. Band II, 344 pp. Schiner, I. R. 1868. Diptera. In Reise der osterreichischen Fregatte Novara. Zool., Vol. 2, Abt. 1, B., Wien, 388 pp. Shannon, R. C. 1925. Some American Syrphidae (Diptera). Proc. ent. Soc. Wash. 27(5): 107-112. 1926. Review of the American xylotine syrphid-flies. Proc. U. S. natn. Mus. 69(9): 1-52. 1927. A review of South American two-winged flies of the family Syrphidae. Proc. U. S. natn. Mus. 70(9): 1-34. 1928. Two new and remarkable Syrphidae in the Museo Nacional de Historia de Buenos Aires. An. Mus. nac. Hist, nat., Buenos Aires, 34: 573-576. Shannon, R. C. and D. Aubertin. 1933. Syrphidae. pp. 120-170. IN British Museum (Natural History), Diptera of Patagonia and South Chile. Pt. 7, fasc. 3, pp. 117-175. Stuardo, C. 1946. Catalogo de los Dipteros de Chile. Santiago, Chile, Min. Agric., 251 pp. 240 Entomologica Americana [Vol. 46, No. 2 Thompson, F. C. 1971. The genus Nepenthosyrphus with a key to world genera of Tropidiini. J. Kansas ent. Soc. 44: 523—534. 1972. A contribution to a generic revision of the Neotropical Milesinae (Diptera: Syrphidae). Arq. Zool. 23(2): 73-215. Walker, F. 1858. Catalogue of the Dipterous insects collected in the Aru Islands by Mr. A. R. Wallace, with descriptions of new species. J, Linn. Soc. London (Zool.) 3: 77-110. WlLLISTON, S. W. 1891-2. Fam. Syrphidae. In Godman, F. D. and Salvin, O., eds., Biologia Centrali- Americana. Zoologia-Insecta-Diptera, Vol. 3, 127 (pp. 1-56, 1891; 57-79, 1892). INSTRUCTIONS TO AUTHORS 1. 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The entry should appear, indented, beginning on the line below the auhor’s name with the information in the following order: date, title of article (with only the first word and proper nouns capital- ized), name of journal (underlined, for italics, with conventional title capitalization, and abbreviated according to the list published by the current Biological Abstracts or Chemical Abstracts or spelled out in full if not listed), volume number (in arabic numerals, followed by a colon), and page numbers; OR date, title of book (with only the first word and proper nouns capitalized), city of publication, publisher, volume number if any (preceded by “vol.”), and the number of pages in the book (followed by the abbreviation “pp.”). Authors are wholly responsible for accuracy and completeness of references. Examples follow: Beal, R. S., Jr. 1954. A revision of the species included in the genus Novelsis (Coleoptera: Dermesti- dae). Trans. Amer. Entomol. Soc., 80: 73-90. Allee, W. D., A. E. Emerson, O. Park, T. Park, and K. P. Schmidt 1949. Principles of animal ecology. Philadelphia and London, W. B. Saunders Co., 837 pp. Kistner, D. H. 1969. The biology of termitophiles. In Biology of termites, vol. 1 (ed. K. Krishna and F. M. Weesner). New York and London, Academic Press, Inc., chap. 17, pp. 525-557. 6. Keys should be typed on a separate sheet of paper, not run into the text, and provided with a formal title. Each couplet should be numbered, the numbers aligned flush left. The first words of all couplet enries should be aligned, and run-over lines indented and aligned. The number or name which ends each couplet entry should be aligned flush right and the space leading up to it filled in with leaders (periods separated by spaces). Examples: 4. Transverse fascia on corium always present and straight 5 Transverse fascia on corium irregular, not straight; or fascia absent 8 5. 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Entomologica Americana THE ORTHOTYLINAE AND PHYLINAE (HEMIPTERA: MIRIDAE) OF SOUTH AFRICA WITH A PHYLOGENETIC ANALYSIS OF THE ANT-MIMETIC TRIBES OF THE TWO SUBFAMILIES FOR THE WORLD Randall Tobias Schuh PUBLISHED BY THE NEW YORK ENTOMOLOGICAL SOCIETY INCORPORATING THE BROOKLYN ENTOMOLOGICAL SOCIETY Vol. 47 February 27, 1974 The New York Entomological Society (incorporating the Brooklyn Entomological Society) publishes two journals: Entomologica Ameri- cana (irregularly, according to the availability of acceptable papers) and the Journal of the New York Entomological Society (quarterly). Monographs and longer papers (minimum length approximately 70 manuscript pages) are published in Entomologica Americana. Shorter papers may be submitted to the Journal. Manuscripts should be sent to the Editor, Entomologica Americana, Department of Entomology, The American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024. Entomologica Americana was first published by the Brooklyn Entomolog- ical Society in April, 1885. Publication was suspended after six volumes, resumed in 1926, again suspended in 1964 with Volume 44, and resumed in 1969 with Volume 45. Since Entomologica Americana is published irregularly, subscription is by volume, not by year and costs $9.00 per volume. Correspondence regarding subscriptions and other non-editorial matters should be addressed to Publication Business Manager, New York Entomological Society, Department of Entomology, The American Museum of Natural History. Back issues may be purchased through Hafner Press, a Division of Macmillan Publishing Company, Incorporated, 866 Third Avenue, New York, New York, 10022. EDITOR Lee H. Herman ASSISTANT EDITOR David C. Miller Mailed February 27, 1974 Entomologica Americana is published for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Vol. 46. No. 2 was published on August 8, 1973 Entomologica Americana Vol. 47, 1973, pp. 1-332 THE ORTHOTYLINAE AND PHYLINAE (HEMIPTERA: MIRIDAE) OF SOUTH AFRICA WITH A PHYLOGENETIC ANALYSIS OF THE ANT-MIMETIC TRIBES OF THE TWO SUBFAMILIES FOR THE WORLD1 Randall T. Schuh2 Table of Contents Introduction 4 Methods and Materials 5 Terminology 8 Nomenclature 8 Part 1. The Orthotylinae and Phylinae of South Africa Historical review 9 Checklist of genera and species 10 Keys to subfamilies of Miridae 14 Keys to genera of Orthotylinae and Phylinae south of the Sahara .... 15 Subfamily Orthotylinae 26 Tribe Halticini 26 N amaquacapsus, new genus 26 Nanniella Reuter 28 Halticus Hahn 29 Nichomachini, new tribe 29 Nichomachus Distant 29 Pseudonichomachus, new genus 35 Tribe Orthotylini 38 Cyrtorhinus Fieber 38 Felisacodes Bergroth 40 “The Orthotylus complex” 41 Pseudambonea, new genus 42 Pseudoloxops Kirkaldy 45 Pseudopilophorus, new genus 47 Zanchiella, new genus 49 Zanchius Distant 58 Subfamily Phylinae 66 Tribe Hallodapini 66 Acrorrhinium Noualhier 66 Azizus Distant 80 Carinogulus, new genus ; 81 Formicopsella Poppius 89 1 Modified from a doctoral dissertation completed in the Biological Sci- ences Group, University of Connecticut, with support from grants from the National Science Foundation and the University of Connecticut Research Foundation. 2 Current address: P. O. Box 518, Ferndale, California 95536. HI 2 Entomologica Americana [Vol. 47 Hallodapus Fieber 91 Laemocoris Reuter 103 Myombea China and Carvalho 104 Pangania Poppius 104 Skukuza , new genus 108 Systellonotopsis Poppius 111 Systellonotus Fieber 112 Trichophorella Reuter 114 Trichophthalmocapsus Poppius 117 Aeolocoris Reuter, extralimital 121 Boopidella Reuter, extralimital 121 Diocoris Kirkaldy, extralimital 122 Leucophoropterini, new tribe 123 Karoocapsus, new genus 123 Tytthus Fieber 135 Tribe Phylini 136 Austropsallus, new genus 136 Brachycranella Reuter 145 Capecapsus, new genus 146 Coatonocapsus, new genus 148 Denticulophallus, new genus 155 Ellenia Reuter 157 Eminoculus, new genus 160 Lamprosthenarus Poppius 164 Lasiolabopella, new genus 165 Lepidocapsus Poppius 167 Leptoxanthus Reuter 169 Macrotylus Fieber 170 Natalophylus, new genus 173 Odhiamboella, new genus 175 Parapseudosthenarus, new genus 177 Parasciodema Poppius 179 Plagiognathidea Poppius 182 Psallus Fieber 182 Pseudosthenarus Poppius 183 Sthenarus-Campylomma 189 Stoebea, new genus 190 Widdringtoniola, new genus 194 Tribe Pilophorini 196 Aloea Linnavuori 196 Ambonea Odhiambo 201 Neoambonea, new genus 204 Parambonea, new genus 207 Paramixia Reuter 210 Pilophorus Hahn 232 Zoogeography 232 Part 2. A Phylogenetic Analysis of the Ant-mimetic Tribes OF THE ORTHOTYLINAE AND PHYLINAE FOR THE WORLD History of the classification of the Miridae 254 Analysis of characters and a revised classification 256 1974] Schuh: South African Orthotylinae and Phylinae 3 The relationships of the Orthotylinae and Phylinae 268 Subfamily Orthotylinae 269 Tribe Halticini 273 Nichomachini, new tribe 275 Tribe Orthotylini 277 Subfamily Phylinae 286 Tribe Hallodapini 292 Leucophoropterini, new tribe 303 Tribe Phylini 307 Tribe Pilophorini 311 Misplaced genera 315 Genera incertae sedis 318 Acknowledgments 319 References 320 Abstract: This study is divided into two parts: 1) a faunal study of the Orthotylinae and Phylinae of South Africa; and 2) a phylogenetic analysis of the tribal classification of the two subfamilies for the world. Much new information on the rela- tionships of these taxa is derived from the South African fauna. In South Africa the Orthotylinae and Phylinae are repre- sented by 54 genera, 20 of which are described as new, and 103 species, 81 of which are described as new. Of the genera occurring in South Africa, 15 percent of the Orthotylinae and 34 percent of the Phylinae are ant mimetic. Generic keys are presented for the Orthotylinae and Phylinae of Africa south of the Sahara; keys to the species are given for those genera oc- curring in South Africa. A zoogeographic analysis reveals that the fauna of South Africa is composed of endemic, tropical African, Paleotropical, pantropical, and cosmopolitan elements. The endemic element is the largest and is concentrated in the Southwest Cape and along the Drakensberg Escarpment. Three tribes are recognized in the Orthotylinae — the Halti- cini, the Nichomachini, new tribe, and the Orthotylini. The Pilophorini, which have been placed in the Orthotylinae by all previous authors, are moved to the Phylinae, in which four tribes are recognized — the Hallodapini, the Leucophoropterini, new tribe, the Phylini, and the Pilophorini. The classification of the Miridae is discussed briefly. A morphological descrip- tion and discussion of the phylogenetic and zoogeographic re- lationships are presented for each tribe. A discussion of each included genus is given for the primarily ant mimetic tribes Nichomachini, Hallodapini, Leucophoropterini, and Pilophor- ini. In the remaining tribes only those genera which must be moved relative to their placement in the Carvalho System of Classification are discussed in detail. 4 Entomologica Americana [Vol. 47 INTRODUCTION This study had its origin in 1967-1968 when I had the oppor- tunity to spend nearly a year in the field in South Africa collecting and observing the Miridae of the area. It soon became evident that the South African fauna was not only large and varied, but very poorly known, and that it contained a number of elements, knowl- edge of which was fundamental to a more mature understanding of the phylogeny and zoogeography of the Miridae as a whole. My decision to approach the work in the format presented here has been based on the following premises: 1) that the current bases for placing genera in the Phylinae and Orthotylinae were un- satisfactory; and 2) that the many ant-mimetic genera in the two subfamilies appeared to have been placed in the taxonomic hier- archy on the basis of shared mimetic characters of a superficial nature rather than on features of a more fundamental phylogenetic significance. Therefore, this paper is divided into two sections: a faunal and biological study of the Orthotylinae and Phylinae of South Africa with an analysis of the taxonomic relationships and distributions of the genera and species of the region; and an anal- ysis of the phylogeny, taxonomy, and zoogeography of the ant- mimetic tribes and genera of the two subfamilies, not only for South Africa, but for the entire world. The Orthotylinae and Phylinae are especially intriguing phylo- genetically because 1) a relationship between the two subfamilies has been observed by most students of the Miridae (Reuter, 1905a; Slater, 1950; Wagner, 1955; Kelton, 1959b; Leston, 1961), but has never been carefully documented; 2) the subfamily Orthotylinae has for some time been known to contain a number of possibly un- related elements (Slater, 1950; Kelton, 1959b), but these have never been comprehensively reviewed; and 3) large numbers of ant-mimetic genera occur in both subfamilies but the phylogenetic relationships among these genera and their relationships to non- mimetic genera are very poorly understood. In South Africa two genera (15 percent) in the Orthotylinae and 14 genera (34 percent) in the Phylinae are ant mimetic. On a world basis approximately 20 (17 percent) of the orthotyline genera and 53 (24 percent) of the phyline genera are ant mimetic. This high proportion of mimetic genera in South Africa, particu- larly in the Phylinae, identifies the area as one of active ant-mimic evolution and one suitable for studies involving the phenomenon. In addition, the extremely diverse nature of the South African flora, 1974] Schuh: South African Orthotylinae and Phylinae 5 in light of the strong host plant specificity of the Miridae, enhances the area’s attractiveness as one in which to develop a greater sys- tematic knowledge of the Orthotylinae and Phylinae and to begin phylogenetic studies. METHODS AND MATERIALS Throughout this study I have employed standard taxonomic procedures. All measurements were taken with an ocular microm- eter and are in millimeters. The line drawings were made with the aid of a squared grid at 150x using a Leitz binocular dissecting mi- croscope and all are reproduced to the same scale. The phallus is illustrated in a lateral view with the phallobase to the left, unless otherwise indicated. Consistency of orientation was attained by view- ing the phallobase laterally and superimposing the two sides of the bilaterally symmetrical structure. In almost all cases this presents the most useful view of the vesica for comparative purposes. In a few cases other views of the vesica are given where the conventional orientation does not show the most useful structural details. The left clasper and phallotheca in the Phylinae are all illustrated in a lateral perspective. The right clasper is always drawn in such a view as to expose the greatest surface area. Consistency of orientation is more difficult with the claspers of the Orthotylinae but the view is usually obvious or explained in the figure captions. The posterior wall and sclerotized rings of the females are drawn showing their orientation relative to the ovipositor valves. Where genitalic struc- tures are similar for all species of a given genus the structures under consideration are illustrated for only one or a limited number of species. All photographic illustrations were made with a Wild M-4 photomicrographic apparatus at 12.5 X on Kodak Panatomic-X 35 mm. film and reproduced to the same scale. I have relied heavily on the Carvalho Catalogue of the Miridae of the World (Carvalho 1952a; 1958a, b; 1960). This invaluable work brings together taxonomic references on the Miridae up to 1955. A list of taxa treated in the South African faunal study is given below. These names are often cited in abbreviated form elsewhere in the text to conserve space. For each of these taxa I have included a complete citation for the original description. For genera treated in detail in the tribal analysis, a complete citation is included only for those not included in the Carvalho Catalogue. No references 6 Entomologica Americana [Vol. 47 are given between the date of original description (if before 1955) and 1955 unless they pertain to new synonymy or were omitted from the Carvalho Catalogue. All South African locality data are listed in a somewhat abbreviated form, except in the case of new species where the holotype data are transcribed from the specimen labels as accurately as possible, with the exception of abbreviated words. Collectors are listed by surname in parentheses at the end of the locality data. If no collectors are listed and the dates are 1967- 1968 the specimens were collected by J. A. Slater, M. H. Sweet, S. T. Slater, R. T. Schuh, and in some cases J. Munting. Abbre- viations of collections where specimens are deposited are listed in parentheses at the end of the locality data. All keys are strictly utilitarian and I have made no attempt to show phylogenetic relationships in them. Where possible, if a coup- let breaks out a higher category, this is indicated. In many cases genera are keyed out more than once. I have tried to avoid the use of sexual characters or those that require dissections, but in some cases this has not been possible. The graphs used to depict distributional patterns follow Wy- godzinsky (1966). As in Wygodzinsky’s work, the hatched boxes represent endemic genera and the numbered boxes indicate more widespread genera and show the number of faunal regions in which a genus occurs. Papua is included in the Oriental Region, because the very limited number of taxa known from that area appear to have their strongest affinities with the mainland Oriental fauna. The Ethiopian Region includes Africa south of the Sahara and Madagascar. At the end of each tribal analysis is included a section entitled ‘'Discussion of individual genera”. For the Orthotylini, Halticini, and Phylini I have included only ant-mimetic genera or genera that must be moved from their position in the Carvalho classification; the latter are marked with an asterisk. A complete generic listing is given for the primarily ant-mimetic tribes Nichomachini, Hallo- dapini, Leucophoropterini, and Pilophorini. A terminal listing of genera incertae sedis and “misplaced genera” is also included for the Orthotylinae and Phylinae as defined in the Carvalho Catalogue. The geographic area included in the South African faunal study encompasses the Republic of South Africa, Swaziland, Lesotho, and South West Africa. Some taxa from outside this area are discussed where they are either closely related to genera or species from South Africa or where it is probable that they may eventually be found there. They are placed at the end of the appropriate genus or tribe. 1974] Schuh: South African Orthotylinae and Phylinae 7 Data for this research were gathered from several sources. A large number of specimens were collected by J. A. Slater, M. H. Sweet, and myself from October 1967 to May 1968. During this time we worked intensively in most areas of South Africa except the arid western regions and South West Africa. The major ad- ditional sources of information for South Africa were the South African National Collection of Insects, Pretoria, the South African Museum, Cape Town, the Transvaal Museum, Pretoria, the Lund University Zoological Institute, Lund, Sweden, and the British Mu- seum (Natural History), London. Many Reuter and Poppius types important to this study are in the Helsinki Museum and I have relied heavily on this extremely valuable collection. Poppius (1914a; 1921) indicated that some of his type specimens were deposited in collections other than Hel- sinki. Most important were the Paris Museum and the Berlin- Humboldt Museum. I have determined, with the generous assis- tance of all curators concerned, that in fact most of the specimens which Poppius studied were never placed in the museums indicated. This was apparently a result of the outbreak of World War I. The specimens in question are all deposited in Helsinki and for the most part are in excellent condition. Where appropriate I have corrected Poppius’ original statements and given the present location of the types. In many cases where original series of two, three, or four specimens were indicated, now only a single specimen can be found. The whereabouts of the remaining specimens are unknown, for I could not locate them after much searching. In all cases where I have been able to find only a single specimen of what was originally a co-type series of two or more specimens, I have designated the specimen as a lectotype. Institutional and private collections from which I have borrowed material are listed below with the abbreviations used in the locality records in the text. AMNH BM (NH) CAS HM JAS American Museum of Natural History, New York British Museum (Natural History), London California Academy of Sciences, San Fran- cisco, California Helsinki Zoological Museum, Helsinki, Finland J. A. Slater Collection, University of Con- necticut, Storrs, Connecticut 8 Entomologica Americana [Vol. 47 Lund University Zoological Institute, Lund University, Lund, Sweden Museum National D’Histoire Naturelle, Paris, France R. T. Schuh Collection South African Museum, Cape Town, South Africa South African National Collection of In- sects, Pretoria, South Africa Transvaal Museum, Pretoria, South Africa United States National Museum, Washing- ton, D. C. TERMINOLOGY Cobben (1968) discussed pretarsal nomenclature in the He- miptera and concluded on morphological grounds that the struc- tures arising between the tarsal claws in the Miridae are not true arolia and proposed the use of the term parempodia. For pseud- arolia Cobben used the term pulvilli. Goel and Schaefer (1970) came to the same conclusions as Cobben, apparently independently, for they did not cite him. Although there is justification for the argument that use of the terms parempodia and pulvilli in the Miridae is unnecessary sub- stitution for the long-standing terms arolia and pseudarolia (Knight, 1918), the arguments for the change seem stronger. Cobben (1968) noted that true arolia do exist in the Hemiptera, notably in the Amphibicorisae. If the term arolia is used in non-Amphibicorisan families, a new term will have to be coined for the true arolia. The term pseudarolia is used only in the Miridae to apply to apparently homologous structures called pulvilli in most groups. Little con- fusion should arise in the Miridae because the change involves only a simple substitution of terms and not a reevaluation of the mor- phology of the structures (see also discussion under Paramixia , below). I have adopted the terminology of Kelton (1959b) for the male genitalia and that of Slater (1950) and Davis (1955) for the fe- male genitalia. Other terminology is more or less standard in the modern hemipterological literature. NOMENCLATURE Although the Carvalho Catalogue of the Genera of Miridae of the World (Carvalho, 1952a) was issued long before the most re- cent International Code of Zoological Nomenclature (1964), the LU PM RTS SAM SANC TM USNM 1974] Schuh: South African Orthotylinae and Phylinae 9 subfamily and tribal names applied by Carvalho in the Orthotylinae and Phylinae are correct. Many of these names were changed be- fore 1961 to agree with the oldest included genus. All have received general acceptance subsequently and therefore are valid under Ar- ticle 40A of the Code. I am considering publication of names in the Carvalho Cata- logue (1958a, b) to represent introduction into the primary zoo- logical literature. If this was not done many of the names pub- lished by Poppius (1914a) would have to be considered as forgotten under Article 23b of the Code or they would have to be referred to the International Commission. Most previously described taxa in this paper illustrate the general inapplicability of Article 23b, particularly in entomology, and I have therefore not strictly fol- lowed the Code on this point. PART 1. THE ORTHOTYLINAE AND PHYLINAE OF SOUTH AFRICA HISTORICAL REVIEW The only monographic treatment of the Miridae of the Ethiopian Region is Poppius’ “Die Miriden der Athiopischen Region” (1912; 1914a). In this work Poppius detailed the knowledge of the Miridae of Africa, Madagascar, and the adjacent islands of the Atlantic and Indian Oceans. Previous to this only a handful of small papers had treated the African Miridae. Poppius (1914a) recorded 41 genera, 19 of which were de- scribed as new, for the Orthotylinae and Phylinae south of the Sa- hara. From 1914 to 1955 only five new genera and 13 new species of Orthotylinae and Phylinae were described from Africa, and very little additional information was added to the literature regarding previously recorded species. With the inception of the work of T. R. Odhiambo in 1958, the status of the African Miridae began to change rapidly. From 1955 to 1971, nine new genera and 50 new species of Orthotylinae and Phylinae were described from Africa south of the Sahara. The total number of genera in these two sub- families now recorded from Ethiopian Africa, including those de- scribed below, stands at 68. The publication of the South African Animal Life Series be- ginning in 1955 has greatly increased our knowledge of many groups of animals in South Africa, and it is certainly the most significant contribution to South Africa entomology in recent years. Unfortu- nately the mirid chapter (Carvalho, et al., 1960) was based solely on the original Brinck and Rudebeck collections and is therefore 10 Entomologica Americana [Vol. 47 incomplete. The neglected nature of the South African fauna is revealed in the fact that previous to 1960 and the publication of the Miridae section of the South African Animal Life Series, only nine genera and nine species of Orthotylinae and Phylinae were recorded from the region. Carvalho, et al. (1960) recorded 11 genera, none of which were described as new, and 12 speices, one of which was described as new. In the following treatment, of the 54 genera I record from South Africa, 20 (37%) are described as new and only 17 were previously recorded from the area. Of the 103 species, 81 (79%) are described as new and 14 were previously recorded from the region (species of Orthotylus , Psallus, and Campylomma-Sthenarus are not included in these figures). In addition, there are several apparently undescribed genera available in collections, but in all cases the material is either inadequate or its relationships too ob- scure to allow for description at the present time. CHECKLIST OF GENERA AND SPECIES Subfamily Orthotylinae Tribe Halticini Namaquacapsus, new genus melanostethoides, new species Nanniella Reuter Halticus Hahn Nichomachini, new tribe Nichomachus Distant minutus , new species rufescens, new species sloggetti Distant sweeti, new species Pseudonichomachus , new genus capeneri, new species mimeticus, new species Tribe Orthotylini Cyrtorhinus Fieber melanops Reuter Felisacodes Bergroth bry ocarina (Poppius) Orthotylus- complex Pseudambonea, new genus capeneri , new species Pseudoloxops Kirkaldy transvaalensis , new species 1974] Schuh: South African Orthotylinae and Phylinae 11 Pseudopilophorus, new genus capeneri, new species Zanchiella, new genus bowkeriae, new species capensis, new species ericae, new species natalensis, new species sweeti, new species Zanchius Distant alba, new species buddleiae, new species leucosideae, new species nigrolineatus, new species Subfamily Phylinae Tribe Hallodapini Acrorrhinium Noualhier brincki Carvalho and Becker capensis, new species drakensbergensis, new species jormicarium (Poppius) incrassata, new species monticola, new species muntingi, new species oudtshoornensis, new species Azizus Distant oculatus (Poppius) Carinogulus, new genus hobohmi, new species kochi, new species transvaalensis, new species varii , new species Formicopsella Poppius regneri Poppius Hallodapus Fieber albofasciatus ( Motschulsky ) pseudosimilis, new species quadrimaculatus, new species similis (Poppius) transvaalensis, new species dispar (Odhiambo), new combination, extralimital poseidon (Kirkaldy), extralimital vittatus (Odhiambo), new combination, extralimital Laemocoris Reuter Myombea China and Carvalho bathycephala China and Carvalho 12 Entomologica Americana [Vol. 47 Pangania Poppius fasciatipennis Poppius chnous (Odhiambo), new combination, extralimital Skukuza, new genus slateri, new species zeugma (Odhiambo), new combination, extralimital Systellonotopsis Poppius bifasciatus Poppius Systellonotus Fieber brincki, new species Trichophorella Reuter australis, new species Trichophthalmocapsus Poppius australis, new species hessei, new species pilosus Poppius, extralimital pumilis (Odhiambo), new combination, extralimital Aeolocoris Reuter, extralimital alboconspersus Reuter Boopidella Reuter, extralimital fasciata Reuter Diocoris Kirkaldy, extralimital agalestus Kirkaldy Leucophoropterini, new tribe Karoocapsus, new genus bifasciatus, new species brunneus, new species flavomaculatus, new species middelburgensis, new species obscurus, new species occidentalis, new species pulchrus, new species trifasciatus, new species Tytthus Fieber parviceps (Reuter) Tribe Phylini Austropsallus, new genus albonotum, new species drakensbergensis, new species helichrysi, new species middelburgensis, new species saniensis, new species senecionus, new species Brachycranella Reuter viridipunctata (Stal) 1974] Schuh: South African Orthotylinae and Phylinae 13 Capecapsus , new genus tradouwensis, new species Coatonocapsus, new genus johannsmeieri, new species pallidus, new species sweeti, new species transvaalensis, new species Denticulo phallus, new genus adenandrae, new species Ellenia Reuter obscuricornis (Poppius) Eminoculus, new genus drosanthemi, new species hirsutus, new species Lamprosthenarus Poppius near sjostedti Poppius Lasiolabopella, new genus capeneri, new species Lepidocapsus Poppius rubrum, new species Leptoxanthus Reuter flavomaculatus Reuter Macrotylus Fieber hemizygiae, new species niger, new species Natalophylus, new genus heteromorphae, new species Odhiamboella, new genus solani (Odhiambo), new combination Parapseudosthenarus, new genus buchenroederae, new species Parasciodema Poppius albocoxa, new species nigrofemur , new species nitens, Poppius Plagiognathidea Poppius Ps alius Fieber Pseudosthenarus Poppius ater Poppius grossus, new species namaquaensis, new species rozeni, new species Sthenarus-Campylomma nigricornis (Poppius) ( Sthenarus ), extralimital Stoebea, new genus barbertonensis, new species 14 Entomologica Americana [Vol. 47 elginensis , new species plettenbergensis, new species Widdringtoniola, new genus kirstenboschiana, new species Tribe Pilophorini Aloea Linnavuori, (in press) australis, new species samueli, new species Ambonea Odhiambo mutiroi, new species rustenbergensis, new species Neoambonea, new genus cynanchi, new species slateri, new species Parambonea, new genus transvaalensis, new species Paramixia Reuter australis, new species suturalis Reuter Pilophorus Hahn pilosus Odhiambo KEY TO THE SUBFAMILIES OF MIRIDAE1 1. Ocelli present Isometopinae Ocelli absent 2 2. Parempodia fleshy, convergent or divergent apically 3 Parempodia hair like, parallel 4 3. Parempodia distinctly divergent apically; pronotal collar usually present, generally rounded and separated from the remainder of the pronotum by a furrow (see however Stenodemini) Mirinae Parempodia convergent apically, usually recurved (lyre shaped); pronotal collar seldom present Orthotylinae and Phylinae (in part) 4. Pulvilli present, sometimes large, but often small and difficult to see, either free except at base or less commonly adnate to claw over entire length 5 Pulvilli absent; claws long and slender or strongly toothed basally; if claws strongly toothed, pronotal collar always present and rounded; if claws long and slender, pronotal collar either present or absent 7 5. Pulvilli arising from ventral surface of claws, usually small; tarsi linear; membrane with two cells Phylinae (in part) Pulvilli arising from inner surface of claws, usually enlarged, flat- 1 Modified from Carvalho (1955a). The Palauocorinae are not included. For figures of parempodia types see Carvalho (1955a), Knight (1923; 1941; 1968) and Wagner (1961). 1974] Schuh: South African Orthotylinae and Phylinae 15 tened, adnate to claw only at base; tarsi and membrane vari- able 6 6. Membrane with one cell; tarsi thickened distally Bryocorinae Membrane with two cells; tarsi linear - Dicyphinae 7. Claws strongly toothed basally; dorsum usually heavily punctate Deraeocorinae Claws not toothed basally, usually long and slender; dorsum punc- tate or impunctate Cylapinae KEYS TO THE GENERA OF ORTHOTYLINAE AND PHYLINAE OF AFRICA SOUTH OF THE SAHARA The following keys are divided into three basic sections. The first deals with genera that have fleshy, apically convergent, recurved parempodia, and also genera with weakly fleshy parempodia that are slightly convergent apically; this includes the Orthotylinae, Pilo- phorini, and certain genera in the Phylini. The second section deals with those genera that have hair-like parallel parempodia and also genera with weakly fleshy convergent parempodia that are also in- cluded in the first section of the key; this includes only the Hallo- dapini, Leucophoropterini, and Phylini. The third section deals with brachypterous forms in the Orthotylinae and Phylinae. Those genera of the Orthotylinae and Phylinae known to occur in Africa south of the Sahara that are not included in the keys are Atractotomus Fieber, Bibundiella Poppius, Brachycranella Reuter, Chaetocapsus Poppius, Dimorphocoris Reuter, and Leptoxanthus Reuter; Mar- morodapus Schmitz keys to Trichophorella Reuter. Genera that are followed by the name of the author are not generally treated in detail elsewhere in the paper. Genera with convergent parempodia 1. Ant mimetic, brown or black, with light maculae or transverse fasciae on hemelytra, at least at base of cuneus; lateral corial margins always sinuate 2 Non-mimetic (see however Pilophorus ), color variable, without distinct light maculae or fasciae; lateral corial margins only rarely sinuate 3 2. Pronotum strongly constricted just anterior to middle, posterior lobe tumid, anterior lobe narrow, neck-like (Fig. 7); scutellum without long erect hairs Pseudonichomachus Pronotum not strongly constricted near middle, posterior lobe tumid, anterior lobe short, not distinctly neck-like (Fig. 5); scutellum with a few long erect hairs Nichomachus 3. Pronotum very heavily punctured; dorsum entirely black 4 Pronotum not heavily punctured; coloration of dorsum variable 5 16 Entomologica Americana [Vol. 47 4. Antennal segment 2 enlarged, spindle shaped, greater in diameter than segment one, densely covered with semierect stout hairs Millerimiris Carvalho Antennal segment 2 cylindrical, not enlarged, diameter slightly less than that of segment one, with fine reclining pubescence Nanniella 5. Entire body and all appendages with long erect black hairs; pro- notum swollen posteriorly; cuneal incisure deep; red and black N amaquacapsus Pubescence not as above; pronotum and color variable 6 6. Head short, wide, flattened anteroposteriorly, concave behind; pos- terior margin of vertex finely carinate, head tending to obscure anterior margin of pronotum 7 Head not particularly short, flattened, or concave behind, posterior margin of vertex not finely carinate, head not obscuring anterior margin of pronotum 14 7. Lateral corial margins distinctly sinuate; hemelytra with a few transverse patches of sericeous appressed, scale-like hairs Pilophorus Lateral corial margins never sinuate, either straight or convex; hemelytra never with transverse patches of scale-like hairs, although often with decumbent, sericeous, wooly hairs 8 8. Dorsum with single type of pubescence, without flattened or wooly hairs 9 Dorsum with reclining setiform hairs and decumbent, sericeous, wooly hairs 10 9. Entirely black, including all appendages; antennal segment 2 about four-fifths width of head across eyes Parambonea Not entirely black; antennal segment 2 subequal to width head across eyes Pseudambonea 10. Antennal segment 2 distinctly laminate, broadest medially Druthmarus Distant Antennal segment 2 cylindrical, not flattened 11 11. Relatively small species about 3.5 mm. long; basically cream col- ored; pronotum, apex of corium, and sometimes cuneus, red Aloea Size variable; coloration never light with distinct red markings 12 12. Entirely black, excluding appendages Neoambonea Body not entirely black 13 13. Abdominal venter with decumbent, wooly, sericeous hairs; vesica in male U-shaped, gonopore subapical (Fig. 318) Ambonea Abdominal venter with only reclining setiform hairs, without wooly sericeous hairs; vesica forming nearly complete coil, gonopore apical (Figs. 332, 335) Paramixia 1974] Schuh: South African Orthotylinae and Phylinae 17 14. Head convexly rounded behind eyes, eyes removed from anterior margin of pronotum by distance about equal to or greater than diameter of antennal segment 2 (Fig. 14); body elongate, flattened; hemelytra often hyaline or subhyaline 15 Head not convexly rounded behind eyes, posterior margin of eyes contiguous with anterior margin of pronotum or nearly so; body not particularly elongate or flattened; hemelytra seldom hyaline 19 15. Clavus usually with distinct row of punctures more or less parallel to claval suture (see however Zanchiella sweeti and Z. ericae ) 16 Clavus without row of punctures as above 17 16. Antennal segment 1 about as long as width of head across eyes; all appendages very long Felisacodes Antennal segment 1 about as long as interocular space ..... Zanchiella 17. Antennal segment 1 longer than width of head across eyes, con- siderably enlarged, about three times diameter of segment 2, densely covered with stout hairs Uleana Carvalho Antennal segment 1 shorter than width of head across eyes, not greatly enlarged or more than one and a half times diameter of segment 2, without conspicuous stout hairs 18 18. Eyes set forward on head, removed from anterior pronotal mar- gin by about half (or nearly so) diameter of eye (Fig. 19) Zanchius Eyes removed from anterior margin of pronotum by about diam- eter of antennal segment 2 (Fig. 9) Cyrtorhinus 19. Antennal segment 1 about as long as width of interocular space plus one eye, distinctly enlarged, rather densely covered with erect or semierect often dark hairs about as long as diameter of segment; coloration usually red and white or almost entirely red Pseudoloxops Antennal segment 1 usually about as long as width of interocular space, only moderately enlarged, pubescence short, inconspic- uous, reclining or decumbent; coloration variable 20 20. Clypeus, juga, and lora brown or black, highly polished, shining, strongly contrasting in texture and usually in coloration with remainder of head 21 Clypeus, juga, and lora not strongly contrasting with remainder of head in texture and coloration, although sometimes black and shining 22 21. Males elongate, lateral corial margins nearly straight; females ovate, brachypterous, hemelytra just covering abdomen; male genital capsule without ventral keel Capecapsus Ovate; lateral corial margins distinctly convex; both sexes macrop- terous; male genital capsule with keel ventrally Ellenia 18 Entomologica Americana [Vol. 47 22. Small, length 3.0 mm. or less; body black; metafemora conspicu- ously enlarged; height of head below eyes greater than height of eye Halticus Size and color variable, body never totally black; metafemora not conspicuously enlarged; height of head below eyes about two- thirds height of eye or less 23 23. Parempodia recurved (Orthotylinae) ; usually elongate, green, al- though sometimes red or brown; antennal segment 3 about two- thirds length of segment 2; usually over 3.5 mm. long Orthotylus and Pseudorthotylus Poppius Parempodia not recurved (Phylinae); usaully ovate or under 3.5 mm. long; antennal segment 3 less than two-thirds length of segment 2 24 24. Dorsum with only reclining, black, setiform hairs; frons strongly convex Widdringtoniola Dorsum with reclining setiform hairs and clumps of wooly seri- ceous hairs Stibaromma Odhiambo Genera with hair-like parallel parempodia 1. Pronotum often strongly narrowed anteriorly, often with a distinct flattened collar at least as wide as diameter of antennal segment 2 (see Karoocapsus) ; hemelytra usually dark with one or more light maculae or fasciae contrasting with basic coloration; often strongly ant mimetic; pulvilli always minute (Hallodapini, Leuco- phoropterini in part) 2 Pronotum sometimes narrowed anteriorly and with flattened collar; if pronotal collar present, eyes substylate or stylate; hemelytra never with contrasting maculae or fasciae; seldom appearing ant mimetic; pulvilli usually minute, sometimes long and free apically or long and fused with claw (Phylini, Leucophoropter- ini) 24 2. Anterior pronotal margin finely carinate, upturned, not in the form of flattened collar; head, pronotum, and scutellum with reclining, dark, setiform hairs and appressed, flattened, seri- ceous, scale-like hairs (the latter are very easily rubbed off); usually with two or three yellow maculae strongly contrasting with dark hemelytra (Figs. 42-49) Karoocapsus Anterior pronotal margin always in the form of flattened collar, at least as wide as diameter of antennal segment 2; vestiture of dorsum variable, never with appressed, scale-like, sericeous hairs; coloration variable, maculae or fasciae usually white if present 3 3. Vertex produced into spine above clypeus (Figs. 23, 24) Acrorrhinium 4 Vertex not produced into spine 1974] Schuh: South African Orthotylinae and Phylinae 19 4. Eyes set far forward on head, removed from anterior margin of pronotum by distance equal to at least diameter of eye (Figs. 36, 37, and 38) 5 Eyes contiguous with anterior margin of pronotum or nearly so, not removed by more than distance equal to diameter of an- tennal segment one (Figs. 26, 30, and 39) 7 5. Scutellum produced into sharp spine about as high as pronotum Myombea Scutellum flat or rounded, not produced into spine 6 6. Antennae inserted at about midpoint of anterior margin of eyes, fossae nearly contiguous with eyes F ormicopsella Antennae inserted below ventral margin of eyes, fossae removed from eyes by distance equal to diameter of antennal segment 1 Skukuza 7. Hemelytra with at least one light macula or fascia contrasting with dark background; basic coloration never solid black 8 Elemelytra without contrasiing maculae or fasciae or if with fascia then basic coloration solid black 20 8. Head concave behind; posterior margin of vertex carinate, pro- duced posteriorly over pronotal collar; scutellum strongly pro- tuberant (Fig. 25) 9 Head either weakly convex behind forming short neck, or neck ob- solete and posterior margins of eyes contiguous with anterior margin of pronotum; posterior margin of vertex not carinate (although sometimes with low rounded transverse ridge be- tween eyes); scutellum sometimes protuberant 10 9. Gula with distinct longitudinal carina; gula at least as long as diameter of antennal segment one; southern Africa Carinogulus (in part) Gula without carina; length of gula less than diameter of antennal segment one, buccal cavity sometimes contiguous with pro- sternum; northern Africa Glaphyrocoris Reuter and Hypomimus Findberg 10. Gula with distinct longitudinal carina; gula at least one and a half times length of diameter of antennal segment one; scutellum strongly protuberant; eyes removed from anterior margin of pronotum by distance about equal to diameter of antennal seg- ment one; dorsum with some long, erect hairs Carinogulus (in part) Gula without distinct carina, length variable; scutellum occasionally protuberant or spiniform; eyes either contiguous with or slightly removed from anterior margin of pronotum; dorsal vestiture variable 1 1 11. Head greatly elongated dorsoventrally; gula almost vertical, nearly as long as height of eye; hemelytral fascia broad laterally, nar- rowed mesially, forming transverse hourglass-shaped marking 20 Entomologica Americana [Vol. 47 on corium; metatibiae slightly to strongly flattened Diocoris and Gampsodema Odhiambo Head only moderately elongated dorsoventrally, length of gula less than one-half height of eye, or head not at all elongated and gula very short 12 12. Length of gula about one-half height of eye, gula nearly vertical Systellonotus Gula never longer than diameter of antennal segment one, often obsolete and buccal cavity contiguous with prosternum 13 13. Dorsum with long, erect hairs nearly as long or longer than diam- eter of antennal segment one, and with or without shorter de- cumbent hairs 14 Dorsum never with long, erect hairs although occasionally with short, erect hairs, and always with short, decumbent hairs ____ 18 14. Species always with wing-edge stridulatory mechanism; lateral cor- ial margin always lacking projecting hairs and possessing fine serrations (sometimes not visible even under high magnifica- tion); inner surface of metafemora pebbled or otherwise mod- ified into stridulatory plectrum, always glabrous; at least meta- tibiae with very long spines 15 Species without wing-edge stridulatory mechanism; lateral corial margins usually with projecting hairs, always lacking serrations; inner surface of metafemora may be finely granulose, never distinctly pebbled or otherwise modified into stridulatory plec- trum, usually with distinct pubescence; metatibiae without ex- tremely long spines 17 15. Eyes of males very large (Figs. 33, 34), much larger than those of females, occupying nearly entire sides of head and reach- ing almost to bucculae; lateral corial margin distinctly sinuate; corium with single transverse fascia medially; metatibiae usually broadened medially, spindle-shaped Trichophthalmocapsus Eyes in males not extremely large, only slightly larger than those of females, genal area exposed; lateral corial margin and hem- elytral markings variable; metatibia usually cylindrical 16 16. Scutellum in males usually spiniform; females brachypterous, pro- notum strongly swollen, scutellum swollen, not spiniform as in males Laemocoris Scutellum in males more or less flat; females usually brachypter- ous, pronotum and scutellum not highly modified, similar in structure to males Hallodapus (in part) 17. Small species, always less than 4.5 mm. long; posterior margins of eyes contiguous with anterior margin of pronotum Hallodapus (in part) Larger species, about 4.5 mm. long; posterior margins of eyes slightly removed from anterior margin of pronotum Systellonotopsis I1 1974] Schuh: South African Orthotylinae and Phylinae 21 18. Eyes in males very large, occupying nearly entire sides of head and reaching almost to bucculae Boopidella Eyes in males not extremely large, genal area exposed (eyes of fe- males only slightly smaller that those of males) 19 19. Small species, always less than 4.5 mm. long ____ Hallodapus (in part) Larger species, at least 5.0 mm. long Pangania 20. Basic coloration velvety black, sometimes with broad white hem- elytral fascia Syngonus Bergroth Basic coloration not black, usually light brown, often mottled .... 21 21. Slender bodied, ratio of total length to greatest width at least 3:1 Trichophorella More heavy bodied than above, ratio of total length to width 2.75: 1 or less 22 22. Antennal segment 3 longer than segment 2 Kapoetius Schmitz Antennal segment 2 longer than segment 3 23 23. Metalegs, antennae, and most of dorsum marmorate Aeolocoris Legs, antennae, and dorsum more or less solid color, hemelytra sometimes with a few light markings Azizus 24. Eyes stylate, width of head across eyes nearly equal to or greater than width of posterior margin of pronotum; pronotum ante- riorly with wide flat collar; black Eminoculus Eyes not distinctly stylate, sometimes substylate; head rarely as wide as posterior margin of pronotum; pronotum never with flattened collar; color variable, but if eyes substylate never solid black 25 25. Dorsum heavily punctate; black 26 Dorsum never heavily punctate; if black seldom highly polished; head rarely very short 27 26. Antennal segment 2 enlarged, greater in diameter than segment 1 Millerimiris Carvalho Antennal segment 2 linear, of smaller diameter than segment 1 Lamprosthenarus 27. Antennal segment 2 conspicuously enlarged distally, diameter about equal to or greater than diameter of segment 1 and of much greater diameter than that of segments 3 and 4 28 Antennal segment 2 not conspicuously enlarged, seldom more than two-thirds diameter of segment 1, not more than 2 times diam- eter of segments 3 and 4 29 28. Antennal segment 2 thickened distally to about two times proximal diameter; black Rakula Odhiambo Antennal segment 2 nearly uniformly thickened over entire length; reddish Lepidocapsus 29. Eyes substylate, head transverse, width across eyes nearly as great as maximum width of pronotum; body densely covered with flattened, appressed, scale-like hairs 30 22 Entomologica Americana [Vol. 47 Eyes not substylate; dorsum, if at all, only partially covered with appressed, scale-like hairs, although sometimes rather densely covered with decumbent wooly pubescence 31 30. Antennal segment one short, cylindrical; antennae lacking scale- like hairs; tibiae without dark spines; coloration not entirely black, hemelytra with large light areas Lasiolabopella Antennal segment one long, increasing in diameter distally; anten- nae with scale-like hairs; tibiae with dark spines; black Lasiolabops Poppius 31. Pulvilli enlarged, flattened, free from claw apically, reaching to about apex of claw; clypeus prominent 32 Pulvilli usually very small, occasionally enlarged and fused with nearly entire ventral surface of claw 33 32. Dorsum rather densely covered with semierect or reclining, dark, heavy, setiform hairs Denticulophallus Dorsum with only fine reclining hairs or with reclining hairs and decumbent wooly pubescence Macrotylus 33. Head including eyes concave behind, vertex finely carinate; anterior margin of pronotum obscured by posterior margin of head; small species, under 3.5 mm. long, body ovoid Sthenarus-Campylomma Head and eyes either convex or only weakly concave behind; pos- terior margin of vertex sometimes finely carinate and obscuring anterior margin of pronotum; if species under 3.5 mm. long, head never concave behind and not obscuring anterior margin of pronotum; species usually over 4.0 mm. long, moderately elongate 34 34. Dorsum with two distinct types of pubescence, usually with re- clining setiform hairs and decumbent, somewhat flattened, wooly, sericeous pubescence, or very seldom with some clumps of scale-like, appressed sericeous hairs 40 Dorsum with single type of pubescence, never with wooly sericeous hairs 35 35. Pulvilli large, fused with almost entire ventral surface of claw Parasciodema Pulvilli minute 36 36. Antennal segment 2 about 1.4 times as long as width of posterior margin of pronotum; elongate black species; labium just sur- passing procoxae Natalophylus Antennal segment 2 equal to or less than width of posterior margin of pronotum; shape, color, and labial length variable 37 37. Small, light colored species; dorsum with reclining black hairs or inconspicuous light hairs 38 At least pronotum dark; vestiture usually conspicuous, never only of black hairs 39 1974] Schuh: South African Orthotylinae and Phylinae 23 38. Dorsum with only reclining, black, setiform hairs; frons convex, clypeus not visible from above; parempodia weakly fleshy, convergent apically Widdringtoniola Dorsum with decumbent, fine, light hairs; frons produced anteriorly, clypeus prominent as viewed from above; parempodia hair-like and parallel Plagiognathidea 39. Ratio of length of head to width of head about 1:1.25; all femora light at least proximally Tytthus Ratio of length of head to width of head 1:4; all femora dark at least proximally Odhiamboella 40. Head, pronotum, and scutellum (also mesepisterna and metepi- sterna) with flattened, appressed, sericeous, scale-like hairs; hem- elytra mostly with reclining setiform hairs except for a few scale-like hairs along claval suture; antennal segment 2 at least as long as width of posterior margin of pronotum and up to 1.6 times pronotal width; elongate species, usually with yellow maculae contrasting with dark background, sometimes solid brown or with inconspicuous maculae (Figs. 42-49) Karoocapsus Dorsum including hemelytra usually rather densely covered with decumbent, sericeous, wooly hairs and semierect or reclining setiform hairs, never with scale-like hairs as above; length of antennal segment 2 and coloration variable 41 41. Small, 3.5 mm. long or less; light colored with variable markings (Fig. 82); labium very long, reaching to middle of abdomen; antennal segment 2 at least as long as width of posterior mar- gin of pronotum Stoebea Usually over 3.5 mm. long; coloration either light or dark, if light usually with many dark spots on dorsum or unicolorous and labium reaching to or surpassing mesocoxae; if dark usually brown or black, unicolorous and labium short, reaching only slightly past procoxae at most; length of antennal segment 2 variable 42 42. Labium short, not or only slightly exceeding posterior margin of procoxae; brown or black 43 Labium long, surpassing mesocoxae; usually light with heavily spotted dorsum; if labium just surpassing procoxae, clypeus, juga, and lora black, highly polished 45 43. Clypeus, juga, and lora at and below level of antennal bases black or dark brown, highly polished, shining, remainder of head dull or only weakly shining; parempodia weakly fleshy, con- vergent apically Capecapsus Clypeus, juga, and lora not highly polished or differing in texture from remainder of head; parempodia hair-like, parallel 44 44. All tibiae and antennae black; male genitalia as in Figs. 289-292 Parapseudosthenarus 24 Entomologica Americana [Vol. 47 All tibiae light with black spines with black bases; male genitalia as in Figs. 293-312 Pseudosthenarus 45. Clypeus, juga, and lora at and below level of antennal bases, black, highly polished, contrasting with remainder of head 46 Clypeus, juga and lora unicolorous with surrounding areas of head and of same texture 47 46. Parempodia hair-like, parallel; vesica of male usually forming a loop (Figs. 238, 241-243); dorsum usually with heavy black seti- form hairs; females brachypterous; male genital capsule without ventral keel Coatonocapsus Parempodia weakly fleshy, convergent apically; vesica in male not forming a loop (Fig. 248); dorsum with fine, dark, setiform hairs; females macropterous; male genital capsule with ventral keel Ellenia 47. Membrane marmorate; antennal segment 2 enlarged distally, clavate A napsallus Odhiambo Membrane not marmorate, unicolorous or cells differing in col- oration from remainder of membrane; antennal segment 2 lin- ear or nearly so, not clavate 48 48. Large species, macropterous forms over 4.0 mm. long; dorsum with long, semierect setiform hairs; females occasionally brachypterous; parempodia hair-like, parallel Austropsallus Smaller species, less than 4.5 mm. long; dorsum with reclining, relatively short, setiform hairs; females macropterous; parem- podia often slightly fleshy and weakly convergent apically ____ 49 49. Wooly, sericeous hairs on dorsum in distinct patches; vesica long, forming complete loop Stibaromma Odhiambo Wooly sericeous hairs on dorsum (sometimes absent) more or less uniformly distributed; vesica short (e.g., Fig. 248) Psallus Genera with submacropterous and brachypterous forms 1. Hemelytra greatly reduced, about the same length as pronotum or less, covering only base of abdomen (Figs. 6 and 37); ant mimetic _ 2 Hemelytra reduced, either just covering abdomen or covering about half of abdomen, but never less than one and a half times length of pronotum (Figs. 32, 57, and 61); ant mimetic or not __ 4 2. Eyes removed from anterior margin of pronotum by distance greater than diameter of eye; anterior lobe of pronotum small Skukuza Eyes contiguous with anterior margin of pronotum; anterior lobe of pronotum variable 3 3. Scutellum swollen, protuberant; parempodia fleshy, convergent apically, recurved Nichomachus Scutellum flat, not swollen; parempodia hair-like, parallel Karoocapsus 1974] Schuh: South African Orthotylinae and Phylinae 25 4. Hemelytra covering about half of abdomen, often truncate poste- riorly (Figs. 24 and 65); sometimes ant mimetic 5 Hemelytra just covering abdomen, often appearing fully winged in absence of comparison with macropterous forms (Fig. 57) 9 5. Eyes stylate, head conspicuously produced laterally beyond antero- lateral angles of pronotum; shining black; coleopteroid Eminoculus Eyes not stylate although may be protuberant; not entirely black; not coleopteroid 6 6. Frons with distinct spine projecting over clypeus (Fig. 24) Acrorrhinium Frons without spine as above 7 7. Eyes removed from anterior margin of pronotum by distance equal to at least diameter of eye; ant mimetic Formicopsella Eyes contiguous with anterior margin of pronotum 8 8. Pronotum with posterior lobe strongly swollen, elevated high above hemelytra; scutellum swollen Laemocoris Pronotum with posterior lobe not strongly swollen; scutellum nearly flat (Fig. 32) Hallodapus 9. Labium short, not or only slightly exceeding procoxae 10 Labium long, reaching at least to mesocoxae, occasionally to mid- dle of abdomen 12 10. Clypeus, juga, and lora nearly black, highly polished, contrasting with dull surface of remainder of head; parempodia weakly fleshy, convergent apically Capecapsus Clypeus, juga, and lora not strongly contrasting with remainder of head in color and surface texture; parempodia hair-like and parallel 1 1 11. Tibiae light with black spines with black bases Pseudosthenarus Tibiae black with black spines Pcirapseudosthencirus 12. Clypeus, juga, and lora black, highly polished, shining, strongly contrasting with remainder of head Coatonocapsus Clypeus, juga, and lora neither black nor strongly shining, not con- trasting with remainder of head in color and surface texture 13 13. Antennae with numerous erect, black hairs about three times as long as diameter of segment on which they occur A ustropsallus Antennae without long, erect, black hairs 14 14. Hemelytra cream, pronotum, scutellum, and macula at cuneal fracture red; parempodia fleshy, convergent apically, recurved Aloe a Dorsum generally light with reddish, greenish, or brownish mark- ings; parempodia hair-like and parallel Stoebea 26 Entomologica Americana [Vol. 47 SUBFAMILY ORTHOTYLINAE TRIBE HALTICINI Namaquacapsus, new genus Macropterous Male: Robust; entire dorsum, thoracic pleura, abdominal venter, femora, and tibiae densely covered with erect black hairs about two and a half times as long as tibial diameter; antennal segment one with a few, erect, fine spines, segments 2, 3, and 4 with short reclining pubescence and a few long, erect, black hairs; labium with short erect hairs. Head short, deflexed; eyes weakly granular, about half height of head; frons convex between antennal bases; antennal segment one moderately enlarged, swollen medially, segment 2 with proxi- mal half narrow, distal half enlarged to about one and a half times diameter of proximal half, approaching diameter of segment one, segments 3 and 4 subequal in diameter (4 missing in holotype), slightly less than proximal diameter of segment 2; clypeus large, flattened dorsally, rounded ventrally; bucculae narrow; buccal cav- ity large, broad; gula about half length of distal diameter of anten- nal segment 2; pronotum with flattened collar about as wide as prox- imal diameter of antennal segment 2; lateral pronotal margins nearly straight posteriorly, broadly rounded anteriorly; calli obsolete; pro- notum with anterior lobe short, depressed behind collar, posterior lobe elevated, inflated, mesoscutum obscured by pronotum; scutel- lum strongly convex, clavi steeply declining laterally from scutellum and commissure; corium rounded transversely, lateral margins strongly convex; cuneal incisure very deep, fracture perpendicular to longitudinal axis of body; cuneus strongly convex laterally; mem- brane with two cells; tibiae without rows of tiny, closely spaced spines; tarsal claws relatively short, weakly curved; parempodia fleshy, convergent apically, recurved; pulvilli minute. Male Genitalia: Figures 100, 101. Vesica membranous. Female unknown. Type Species: Namaquacapsus melanostethoides , new species. This genus is named for Namaqualand, the region of the type locality of Namaquacapsus melanostethoides. Namaquacapsus is placed in the Halticini because of the flat- tened pronotal collar, dorsoventrally elongated head, dark color- ation, and the structure of the male genitalia. The genus appears to be most closely related to Orthocephalus Fieber, from Europe and the Mediterranean, by virtue of the heavy vestiture and body 1974] Schuh: South African Orthotyi.inae and Phylinae 27 shape. Namaquacapsus shows specialized features within the Hal- ticini, particularly in the type of vestiture and the reddish coloration of the hemelytra; the former condition may be an adaptation to an extremely arid environment. Namaquacapsus melanostethoides, new species Figures 1, 100, 101 Macropterous Male: Basic coloration deep castaneous; an- terior two-thirds and apex of corium and anterior two-thirds of cuneus red. Head, pronotum, scutellum, antennae, labium, legs, and ab- dominal venter polished, shining; remainder of body, including an- terior lobe of pronotum dull; eyes glabrous. Posterior margin of vertex nearly straight, with broad rounded carina; antennae removed from margins of eyes; labium just sur- passing mesocoxae; pronotum with anterior margin sinuate, pos- terior margin straight across mesoscutum, broadly rounded later- ally; abdomen reaching to apex of corium; metatarsal segments 1 and 3 subequal in length, segment 2 about two-thirds length of seg- ment 3. Measurements: Total length 5.36, maximum width 2.16, length head .60, width head .76, interocular space .60, length pro- notum 1.00, width pronotum 1.92, length scutellum .64, width scu- tellum .96, length corium 2.36, length clavus 2.00, length cuneus 1.04, width cuneus .92, length claval commissure 1.04, distance apex commissure-apex membrane 2.36, length metatibia 1.92; length antennal segments 1 — .32, 2 — 1.30, 3 — .88, 4 — .38 (from para- type); length labial segments 1 — .42, 2 — .42, 3 — .20, 4 — .40. Male Genitalia: Figures 100, 101. Holotype: Macropterous male, south Africa: Cape Prov- ince, Kamieskroon, Namaqualand, Museum Staff, Sept. 1930 (SAM). Paratype: 1 macropterous male, same data as holotype (RTS). This species is named for its similarity to Melanostethus Stal (Lygaeidae) in general coloration. Namaquacapsus melanostethoides can be recognized by its dark red and almost black coloration and extremely long, dense, black pubescence. No host or ecological data are available. 28 Entomologica Americana [Vol. 47 Nanniella Reuter Nanniella Reuter, 1904, p. 6. Nanniella was described by Reuter (1904) for a single species, N. chalybea Reuter, from Kinshasa; Poppius (1914a) added N. reuteri Poppius from “Nyassa.” Both authors placed the genus in the Halticini. Nanniella was synonymized with Falconia Distant by Carvalho (1952a), but no explanation was given for the action. Comparison of type specimens of F. poetica Distant, the type spe- cies of the genus, from South America, and Nanniella from Africa, indicates that there is indeed a very close superficial resemblance between the two genera. However, a careful examination reveals that in fact they are much less closely related than general facies would indicate. The parempodia in both Falconia and Nanniella are apically convergent and recurved. The female genitalia, how- ever, are diagnostic for the two genera: in Falconia the posterior wall possesses well developed K-structures characteristic of the Orthotylini; in Nanniella the posterior wall is a simple sclerotized plate and lacks K-structures, a feature which in combination with the convergent parempodia suggests that the genus belongs to the Halticini. Additional characters supporting this tribal placement for Nanniella are: the solid black coloration; the flattened, rather broad, pronotal collar; the club-shaped right clasper; the simple membranous vesica; and the dorsoventrally elongated head. Nan- niella does not have noticeably enlarged hind femora, which are characteristic of most members of the Halticini. Nanniella can be recognized by its heavily punctate, shining, black dorsum, vertical head with protuberant eyes removed from the anterior margin of the pronotum, and flattened pronotal collar. It is most closely related to Acratheus Distant from India, which is also heavily punctate on the dorsum. Acratheus has a light cuneus and membrane whereas in Nanniella the entire dorsum is black. The available material suggests that several closely related spe- cies of Nanniella are present in Africa. Three specimens are known from South Africa which agree generically with the type series of N. chalybea deposited in the Musee Royal de FAfrique Central, and a specimen identified by Poppius as chalybea in the Helsinki Museum which is from Kinshasa. A male from Sarnia, Natal (Fig. 2), deposited in the British Museum (Natural History), and a female from Umkomaas, Natal, deposited in the South African National Collection of Insects, have completely brown antennae and a band of dense wooly hairs immediately posterior to the pro- 1974] Schuh: South African Orthotylinae and Phyunae 29 notal collar. A female from Albert Falls, Umgeni River, Natal, deposited in the Lund University Collection (see Carvalho, et al., 1960), has dark antennae, but lacks the band of wooly hairs on the pronotum and has distinctly brown tibiae, whereas in other specimens from South Africa the tibiae are light. Specimens of N. chalybea from Kinshasa, Congo, have the first antennal segment light and lack the band of wooly hairs on the pronotum. At the present time it seems undesirable to assign names or describe new species until a careful study of Nanniella and Acratheus is undertaken. Halticus Hahn Halticus Hahn, 1832, p. 113. Only two species of the cosmopolitan genus Halticus are cur- rently recorded from Africa (Carvalho, 1958b). A male specimen from Satara Camp, Kruger National Park, Transvaal, deposited in the J. A. Slater Collection, probably represents a new species. It has the second antennal segment about three times as long as the first, the femora generally black, and the posterior margin of the vertex narrowly white. NICHOM ACHINI , new tribe Nichomachus Distant Nichomachus Distant, 1904a, p. 104. Nichomachus can be characterized by the following redescrip- tion of the genus. Macropterous Male: Elongate, ant mimetic; head, pronotum, and scutellum granulose or rugulose, dull or weakly shining; corium and clavus mostly pruinose; basal fifth of corium and clavus, cuneus, cell of membrane, legs, and venter polished, shining; pronotum, scutellum, cuneus, femora, and antennal segment 1 with short, de- cumbent, sericeous hairs; scutellum, clavus, and corium with a few long, erect, shining hairs; antennal segments 2, 3, and 4 with dense short vestiture; tibiae with a few semierect spines about as long as tibial diameter. Head nearly vertical, concave behind; vertex depressed between eyes, posterior margin carinate; frons transversely rugose, weakly convex, with faint longitudinal sulcus; eyes large, occupying entire sides of head; antennae inserted just above ventral margin of eyes, fossae contiguous with eyes; antennal segment 1 slightly enlarged, 30 Entomologica Americana [Vol. 47 Figs. 1-6. Halticini, Nichomachini. Fig. 1. Namaquacapsus melano- stethoides, male, holotype. Fig. 2. Nanniella near chalybea, male (Sarnia, Natal). Fig. 3. Nichomachus minutus, male, holotype. Fig. 4. Nichomachus rufescens , male, holotype. Fig. 5. Nichomachus sweeti, male, holotype. Fig. 6. Nichomachus sweeti, female (Schoe- mannspoort, Cape Province). 1974] Schuh: South African Orthotylinae and Phylinae 31 segments 2 and 3 subequal in diameter, about three-fourths diam- eter of segments 1 and 4; labium just surpassing posterior margin of metasternum; pronotum with anterior lobe about one-fifth length of entire pronotum, forming poorly defined neck, anterior margin carinate, upturned, posterior lobe tumid; mesoscutum inclined ante- riorly; scutellum strongly inflated, bulbous; lateral corial margins weakly sinuate, reflexed ventrally on anterior half, cuneal incisure obsolete; membrane with two cells; abdomen long, slender, just sur- passing apex of cuneus; parempodia fleshy, convergent apically, re- curved; pul villi minute. Male Genitalia: Figures 96, 97. Left clasper long, slender, with a barb apically, basal lobe with spine formed of long, stiff, erect hairs; left clasper situated as in Figure 93A; right clasper extremely small, lanceolate; vesica membranous, with minute spines. Brachypterous Female: See Nichomachus sweeti. Female Genitalia: Figures 98, 99. Ring glands very small, contorted; posterior wall a simple sclerotized plate, oriented cepahlo- caudad. Nichomachus is most closely related to Pseudonichomachus, but can be separated from it by the form of the anterior lobe of the pronotum, which is short in the former and long and neck-like in the latter. Distant (1904a) related Nichomachus to Systellon- otus. Carvalho (1952a) placed the genus in the Pilophorini. The structure of the male and female genitalia and the type of sexual dimorphism in Nichomachus show no close relationship to the Pi- lophorini or Systellonotus , but suggest that the genus belongs to a distinct evolutionary line within the Orthotylinae. I have there- fore placed it in a new tribe with several other Ethiopian genera (see tribal classification below). A brachypterous female from Swartberg Pass, 25 miles north of Oudtshoorn, Cape Province and deposited in the J. A. Slater Collection, may be congeneric with Nichomachus. This specimen has an hourglass-shaped pronotum which is, however, very much different from the pronotum of N. sweeti. An additional female specimen from Mkuze Game Reserve, Natal, which is deposited in the J. A. Slater Collection, also appears to be related to Nichoma- chus. It is submacropterous, with only the connexival region of the dorsally flattened abdomen being exposed. The head is very sim- ilar in structure to that of N. sweeti , but the pronotum is hourglass- shaped, and similar in structure to that of the abovementioned fe- male from the Swartberg Pass. The four described species of Nichomachus are all from South- 32 Entomologica Americana [Vol. 47 ern Africa. The genus does not seem to be associated with the tropical African vegetative element but with the Southwest Cape related flora. Key to macropterous specimens of Nichomachus 1. Dorsum mostly red or reddish brown, except for white hemelytral maculae 2 Dorsum nearly uniform dark brown or black except for white hem- elytral maculae 3 2. Scutellum dark brown, contrasting with reddish pronotum and cor- ium; clavi with white transverse macula at about midpoint of claval commissure sloggetti Scutellum unicolorous with pronotum and corium; clavi without white macula rufescens (Fig. 4) 3. Large black species, 4.80 mm. long; distal fifth of antennal segment 3 usually white sweeti (Fig. 5) Smaller, dark brown species, 4.24 mm. long; antennal segment 3 unicolorous brown minutus (Fig. 3) Nichomachus minutus, new species Figure 3 Macropterous Male: Basic coloration dark brown or cas- taneous; most of corium and clavus lighter brown than remainder of body; membrane smoky brown; hemelytra with white macula on basal third of corium, on calvi at about midpoint of claval com- missure, and on basal third of cuneus; posterior margin of met- episternum white; scutellum and cuneus strongly shining. Measurements: Total length 4.24, maximum width 1.14, length head .36, width head .80, interocular space .34, length pro- notum .70, width pronotum .98, length scutellum .46, width scutel- lum .46, length corium 1.74, length clavus 1.16, length cuneus .60, width cuneus .44, length claval commissure .76, distance apex com- missure-apex membrane 1.78, length metatibia 1.80, length an- tennal segments 1 — .30, 2 — 1.02, 3 — .80, 4 — .62; length labial segments 1 — .36, 2 — .24, 3 — .52, 4 — .42 Male Genitalia: Basic structure as in N. sweeti. Female unknown. Holotype: Macropterous S, south Africa: Cape Province, 4 mi. W. Gydo Pass summit, N. of Ceres, 26 Jan. 1968, J. A. & S. Slater, T. Schuh, M. H. Sweet (SANC). This species is named for its relatively small size. Nichomachus minutus most closely resembles N. sweeti but is much smaller and is generally brown rather than black. The type locality was a dry sandy wash with macchia vegetation. 1974] Schuh: South African Orthotylinae and Phylinae 33 Nichomachus rufescens, new species Figure 4 Macropterous Male: Basic coloration bright brownish orange; corium at level of apex of scutellum with broad, transverse, trans- parent band; corium with complete, brown, transverse fascia contigu- ous with posterior margin of transverse band above; basal two-fifths of cuneus white, apical three-fifths castaneous; membrane light smoky brown; abdomen yellowish basally, castaneous apically; an- tennal segments 1 and 2 and metafemur only appendages present on holotype. Measurements: Total length 4.80, maximum width 1.22, length head .38, width head .92, interocular space .26, length pro- notum .74, width pronotum 1.22, length scutellum .64, width scutel- lum .72, length corium 2.20, length clavus 1.68, length cuneus .80, width cuneus .48, length claval commissure .68, distance apex com- missure-apex membrane 2.26; length antennal segments 1 — .36, 2 — 1.42, 3 — ?, 4 — ?; length labial segments 1 — .52, 2 — .56, 3 — .42, 4— .56. Male Genitalia: Basic structure as in N. sweeti. Female unknown. Holotype: Macropterous 8, south Africa: Cape Province, Bulshoek, Clw., S.A.M., 12-56 (SAM). This species is named for its reddish coloration. Nichomachus rufescens is most similar to N. sloggetti and also resembles Pseudonichomachus capeneri. The absence of the white transverse macula on the posterior portion of the clavus in rufescens will separate it from sloggetti. The generic differences in the shape of the pronotum will separate rufescens from P. capeneri. Nichomachus sloggetti Distant Nichomachus sloggetti Distant, 1904a, p. 104. Distant’s original description from a single macropterous male and the preceding key will distinguish N. sloggetti from congeneric species. The coloration of sloggetti is very similar to that of N . rufescens and also resembles closely that of Pseudonichomachus capeneri (see rufescens discussion). Male Genitalia: Basic structure as in N. sweeti. Only two male specimens of N. sloggetti are known: the holo- type from Deelfontein, Cape Province, deposited in the British Mu- seum (Natural History), and an individual from Zomerkomst, Po- litzi, Transvaal, 23.X.64, deposited in the South African National 34 Entomologica Americana [Vol. 47 Collection of Insects. A large number of species from the South African National Collection of Insects (see e.g. Karoocapsus ) are known from Middelburg, Cape Province, and also from Zomer- komst, Politzi, Transvaal. These localities are very different flo- ristically (Acocks, 1951), and there is some question about the accuracy of labeling. The occurrence of N. sloggetti in Deelfontein and Middelburg, would be more logical than in Deelfontein and Politzi because of the greater similarity of available habitats at Mid- delburg and Deelfontein. This situation merits further investigation to be sure that distributions within South Africa are not being in- terpreted incorrectly. Nichomachus sweeti, new species Figures 5, 6, 96-99 Macropterous Male: Body generally black or nearly so; hemelytra with white maculae on basal third of corium, on clavi at about midpoint of claval commissure, and on basal third of cu- neus; posterior margin of metepisternum white; distal sixth of an- tennal segment 3 yellow. Measurements: Length 4.80, maximum width 1.36, length head .34, width head .92, interocular space .34, length pronotum .82, width pronotum 1.16, length scutellum .52, width scutellum .56, length corium 2.14, length clavus 1.40, length cuneus .66, width cuneus .54, length claval commissure .90, distance apex commissure- apex membrane 2.24, length metatibia 2.10, length antennal seg- ments 1 — .26, 2 — 1.10, 3 — .80, 4 — .60; length labial segments 1 — .40, 2— .42, 3— .40, 4— .44. Male Genitalia: Figures 96, 97. See generic description. Brachypterous Female: Strongly ant-mimetic, brachypter- ous; black; transverse fascia at level of apex of scutellum, posterior margin of hemelytra laterally, posterior margin of metepisternum, metacoxae distally, and antennal segment 3 white. Entire body granulose or rugulose, weakly shining, with scat- tered, short, decumbent, shining hairs; abdomen also with scat- tered, long, erect, shining hairs; scutellum with several long, erect, shining hairs; antennae, tibiae, and tarsi with decumbent shining hairs. Head broad, concave behind, frons convex; eyes small; poste- rior margin of vertex slightly concave and broadly carinate; anten- nal segment 1 only very slightly enlarged, segment 2 tapering dis- tally to slightly less than diameter of segment 1, segments 3 and 4 subequal in diameter, slightly greater than diameter of segment 1; 1974] Schuh: South African Orthotylinae and Phylinae 35 antennae inserted just below eye and mesad of eye by distance nearly equal to length of antennal segment 1; labium just surpass- ing metacoxae; pronotum with narrow depressed collar, anterior lobe greatly swollen, posterior lobe about one-quarter length of anterior lobe, collar-like, posterior margin straight; mesoscutum strongly inclined anteriorly; scutellum inflated, nearly conical; hem- elytra undifferentiated, lateral margins nearly straight, posterior margin sinuate, posterolateral angles forming acute projections; abdomen narrow basally, greatly expanded medially, pointed api- cally; all femora swollen distally; metatibiae bowed. Measurements: Total length 4.48, maximum width 1.60, length head .32, width head 1.08, interocular space .66, length pronotum .94, width pronotum .95, length scutellum .32, width scutellum .44, length hemelytra .90, length metatibia 2.10, length antennal segments 1 — .28, 2 — 1.00, 3 — .68, 4 — .62; length labial segments 1 — .50, 2 — .50, 3 — .40, 4 — .56. Female Genitalia: Figures 98, 99. See generic description. Holotype: Macropterous 8, south Africa: Cape Province, Schoemannspoort, 10 mi. N. of Oudtshoorn, elevation 1200 ft., 18 Nov. 1967, M. H. Sweet (SANC). Paratypes: Cape Province — 3 macropterous 8 8, 2 brachyp- terous 2 2, same data as holotype; 1 macropterous 8 , Cape Penin- sula, Noordhoek Beach, 23 Jan. 1968 (JAS, RTS). This species is named for Dr. Merrill H. Sweet, of Texas A. & M. University, who collected most of the known specimens. Nichomachus sweeti most closely resembles N. minutus but can be recognized by its larger size and much darker coloration. The resemblance of the female of N. sweeti to species of Cre- matogaster ants from both Schoemannspoort and Noordhoek Beach is remarkable. The Noordhoek Beach specimen of sweeti was taken under Helichrysum crispum (L.) D. Don. (Compositae). Pseudonichomachus, new genus Macropterous Male: Ant mimetic; head, pronotum, and scutellum smooth, weakly shining; anterior two-thirds of clavus, ad- jacent corium, cuneus, and cell of membrane strongly shining; re- mainder of corium pruinose; membrane dull; dorsum with scattered, decumbent, short hairs; antennae with dense short vestiture; tibiae with scattered semierect spines about as long as tibial diameter. Head nearly vertical, concave behind; frons weakly convex, transversely rugose, longitudinally sulcate; posterior margin of ver- tex with low, broad carina; eyes occupying entire sides of head; 36 Entomologica Americana [Vol. 47 antennae inserted just above ventral margin and very close to eyes; antennal segment 1 slightly swollen, segments 2, 3, and 4 subequal in diameter, slightly less than diameter of segment 1; labium reach- ing or just surpassing mesocoxae; pronotum strongly constricted just anterior to middle, with narrow depressed collar, anterior lobe about two-thirds width of head, neck-like, posterior lobe inflated, nearly hemispherical; mesoscutum inclined anteriorly; scutellum bulbous; lateral corial margin strongly sinuate, anterior half strongly reflexed ventrally; clavus raised along commissure to nearly height of scutellum; cuneal incisure obsolete; membrane with single cell; abdomen narrow, reaching almost to apex of membrane; parempodia fleshy, convergent apically, recurved; pulvilli minute. Male Genitalia: Figures 93A-95. Basic structure as in Nichomachus. Females unknown. Type Species: Pseudonichomachus mimeticus, new species. This genus is named for its close relationship to Nichomachus. Pseudonichomachus differs from Nichomachus by the former having a much longer, more neck-like, anterior pronotal lobe than the latter. Both known species of this genus are from South Africa and are ground living. Pseudonichomachus capeneri, new species Figure 7 Macropterous Male: Basic coloration reddish brown; apex of clavus, anterior quarter of corium, legs, antennae, and labium weakly castaneous; cuneus deep castaneous; abdomen light ba- sally, castaneous apically; membrane smoky gray brown; hemelytra with white transverse maculae on anterior third and posterior third of corium, on basal quarter of cuneus, and on clavus at about mid- point of claval commissure; posterior margin of all epistrena above coxae and distal margin of all trochanters white. Head and pronotum rather strongly shining. Vertex slightly depressed between eyes, labium reaching to mid- dle of mesocoxae; all appendages except antennal segments 1 and 2 and mesofemora and metafemora and tibiae missing in holotype. Measurements: Total length 4.80, maximum width 1.24, length head .38, width head 1.00, interocular space .24, length pro- notum .88 (anterior lobe .28, posterior lobe .60), width pronotum 1.24, length scutellum .78, width scutellum .78, length corium 2.00, length clavus 1.60, length cuneus .82, width cuneus .50, length 1974] Schuh: South African Orthotylinae and Phylinae 37 claval commissure .66, distance apex commissure-apex membrane 2.12, length metatibia 2.18, length antennal segments 1 — .32, 2 — .96, 3 — ?, 4 — ?; length labial segments 1 — .34, 2 — .34, 3 — .40, 4— .44. Male Genitalia: Basic structure as in P. mimeticus. Holotype: Macropterous S, south Africa: Transvaal , Elandshoek 11.1956, A. L. Capener (SANC). Paratypes: Transvaal — 2 macropterous Zoutpansberg District, Khalarha District, 3700 ft., 23. iv. 1954, at light, very open bush and grass veld (Balfour-Browne) (BM[NH]). This species is named for Mr. A. L. Capener of Pretoria, who has been one of the most active collectors of Miridae in South Africa over the last two decades and has therefore made available for this study a tremendous amount of invaluable material. Pseudonichomachus capeneri can be separated from P. mimet- icus, the only other described species in the genus, by its bright orangish to reddish brown coloration; mimeticus is dark brown to nearly black. Pseudonichomachus mimeticus, new species Figures 8, 93A-95 Macropterous Male: General coloration blackish brown; head and anterior lobe of pronotum, corium at level of claval com- missure, and femora, castaneous; distal third of antennal segment 3, broad band on distal half of mesotibiae and metatibiae, transverse fascia on corium at level of apex of scutellum (just reaching onto clavus), narrow transverse marking on clavus at midpoint of claval commissure, and narrow, anteriorly inclined, transverse bands two- thirds of width of corium (reaching lateral corial margin) at apex of clavus, dull white; anterior quarter of cuneus, posterior margin of all mesepisterna above coxae, and distal margin of trochanters white; all tarsi and abdominal sternite 2 light brown. Head, pronotum and scutellum weakly shining. Labium just surpassing mesocoxae. Measurements: Total length 4.08, maximum width 1.10, length head .20, width head .82, interocular space .26, length pro- notum 1.06 (anterior lobe .40, posterior lobe .66), width pronotum 1.10, length scutellum .62, width scutellum .58, length corium 1.66, length clavus 1.26, length cuneus .76, width cuneus .40, length claval commissure .60, distance apex commissure-apex membrane 1.70, length metatibia 1.86; length antennal segments 1 — .26, 2 — 1.04, 38 Entomologica Americana [Vol. 47 3 — .80, 4 — .35; length labial segments 1 — .36, 2 — .32, 3 — .32, 4— .38. Male Genitalia: Figures 93A-95. Basic structure as in Nichomachus. Female unknown. Holotype: Macropterous 3, south Africa: Transvaal , top Magoebaskloof, 12 Dec. 1967, J. A. & S. Slater, T. Schuh, J. Munt- ing (SANG). Paratypes: Cape Province — 1 macropterous 3, Cape Town (Bridwell). Transvaal — 1 macropterous 3, same data as holotype; 1 macropterous 3, 22 mi. S. Barberton, 4900 ft. elevation, 24 Mar. 1968 (USNM, J AS, RTS). This species is named for its ant-like appearance. See discussion under P. capeneri for separation of species. TRIBE ORTHOTYLINI Cyrtorhinus Fieber Cyrtorhinus Fieber, 1858, p. 313. Cyrtorhinus was monographed by Carvalho and Southwood (1955). The genus presently contains six species distributed pri- marily in the Old World tropics with one species common to Europe and North America. Cyrtorhinus melanops Reuter Figures 9, 102, 103 Cyrtorhinus melanops Reuter, 1905a, p. 6. — Carvalho, Dutra, and Becker, 1960, pp. 459-460, 475 (in part). Cyrtorhinus melanops can be most easily recognized by the apically convergent, recurved parempodia, orthoty line- type male genitalia (Figs. 102, 103), the dark head, pronotum, and scutellum, the light hemelytra, and the shape of the head and body (Fig. 9). No biological information is available for this species, but it is possible that it is primarily predatory, as is Cyrtorhinus caricis (Fallen) (Southwood and Leston, 1958). Cyrtorhinus melanops is known only from Ethiopia and South Africa (Carvalho, 1958b). Carvalho et al. (1960), incorrectly recorded specimens of Tytthus parviceps from 10 miles north of Matatiele, Cape Province, as C. melanops (see also T. parviceps). Specimens Examined: south Africa: Cape Province — 1 macropterous 3, Kokstad, 6. III. 51 (Brinck and Rudebeck). Trans- 1974] Schuh: South African Orthotylinae and Phylinae 39 Figs. 7-12. Nichomachini, Orthotylini. Fig. 7. Pseudonichomachus capeneri, male, holotype. Fig. 8. Pseudonichomachus mimeticus, male, holotype. Fig. 9. Cyrtorhinus melanops , male (Rustenburg, Transvaal). Fig. 10. Felisacodes bryocorina, male (Grootvaters- bosch Forest Reserve, Cape Province). Fig. 11. Pseudoambonea capeneri, female, holotype. Fig. 12. Pseudoloxops transvaalensis, male (Messina, Transvaal). vaal — 1 macropterous S, Hartebeespoort Dam, 20 mi. W. Pretoria, 30 October 1967; 3 macropterous $ $, Little Sabie River, Sabie, 29 Nov. 1967; 1 macropterous $, Rustenburg, 1-4-11-1957 (Ca- pener); 2 macropterous S$, Rustenburg, 1-5-1957 (Capener); 1 40 Entomologica Americana [Vol. 47 macropterous $, Rietfontein, 4.12.06 (SANC, TM, LU, JAS, RTS). Felisacodes Bergroth Rhodesiella Poppius, 1914a, pp. 64-65 (preocc.). Felisacodes Bergroth, 1926, p. 64 (new name). — Odhiambo, 1967, p. 1681. Madagascariella Carvalho, 1953a, pp. 44-45. New Synonymy. Odhiambo (1967) noted that until the male genitalia of Fe- lisacodes were examined the correct tribal placement of the genus could not be determined. I have dissected both the male and female genitalia of F. bryocorina (Poppius) from South Africa. Those of the male are typical of the Orthotylini, with a membranous vesica lacking spiculi. The female has well developed K-structures which unquestionably places Felisacodes in the Orthotylini. Zanchiella is the most closely related genus. Odhiambo (1967) examined the holotype of Madagascariella longipides Carvalho and noted that it and Felisacodes were ex- tremely closely related, if not congeneric. I have also compared specimens of F. bryocorina with the holotype of M. longipides in the Paris Museum. The structure and coloration of the two are very similar, the most obvious difference being a more strongly rugose pronotum in Madagascariella. I do not believe this feature to be generically significant and therefore synonymize Madagascariella Carvalho with Felisacodes Bergroth. List of described species of Felisacodes bryocorina Poppius (Rhodesiella) , 1914a, p. 65. Rhodesia; South Africa. dibuora Odhiambo (Felisacodes) , 1967, pp. 1681-1683. Cameroon. longipides Carvalho (Madagascariella), 1953, pp. 44-45. Madagascar. Felisacodes bryocorina (Poppius) Figure 10 Rhodesiella bryocorina Poppius, 1914a, p. 65. Felisacodes bryocorina Carvalho, Dutra, and Becker, 1960, pp. 462- 463. Felisacodes bryocorina can be distinguished from all other South African mirids by the elongate body (total length 4.12 mm., maxi- mum width 1.04 mm.), the hyaline hemelytra, the row of punctures 1974] Schuh: South African Orthotylinae and Phylinae 41 on the clavus parallel to the claval commissure, and the extremely long appendages, with antennal segment 1 being longer than the width of the head. F. bryocorina can be separated from F. dibuora, the only other species of F elisacodes in Africa, in that dibuora has a light colored scutellum and bryocorina has a dark scutellum, which is unicolorous with the posterior lobe of the pronotum. Three males and one female of F. bryocorina are in the British Museum (Natural History). I have selected a male as the lectotype. It bears the labels: “S. Rhodesia, Chirinda, 12.VI.1911, Swynner- ton” and “LECTOTYPE Rhodesiella bryocorina Poppius, det. R. T. Schuh.” The only host plant record for this species is Plectranthus fruti- cosus L’Hes. (Labiatae). The plants were growing in a heavily shaded forest. Specimens Examined: south Africa: Cape Province — 40 macropterous 8 8, 23 macropterous 2$ (1 nymph in alcohol), Grootvatersbosch For. Res., 14 mi. N. Heidelburg, 5 Feb. 1968 (Adults and nymphs on Plectranthus fruticosus L’Hes.); 2 macrop- terous 2$, Port St. Johns, Pondoland, Sept. 1923 (Turner); 1 macropterous 8, Storms River Mouth, 13 Feb. 1968; 1 macropter- ous 8, Tsitsikama Forest, Stormsrivierpiek, 13.1.51 (Brinck and Rudebeck). Natal — 1 macropterous 8, Kloof, 1500 ft., Aug. 1926 (Turner) (SANC, BM[NH], TM, SAM, HM, LU, USNM, JAS, RTS). “The Orthotylus complex” Several groups of species that can be placed in Orthotylus Fieber or closely related genera are present in South Africa. The only com- prehensive work on Orthotylus is that of South wood (1953) which is restricted to the British species. This work is unfortunately of limited use outside of Europe for it does not define the genus on a world basis and the subgenera of Southwood are based only on European species. Lindberg (1951; 1953) has dealt extensively with the species of Orthotylus from the Canary Islands and segre- gated Canariocoris Lindberg from Orthotylus. Knight (1968) de- scribed several new species of Melanotrichus Reuter, which he con- sidered as a distinct genus, from the western United States. The extreme variation found in Orthotylus is described in part by Southwood (1953) and can be judged also by the number of generic synonyms associated with the genus (see Carvalho, 1958b). Orthotylus is probably cosmopolitan, although Carvalho does not record it from South America. Poppius (1914a) listed only four 42 Entomologica Americana [Vol. 47 species from Africa (including Chlorosomella geniculata Reuter). Very little has been done to advance our knowledge of Orthotylus in Africa since Poppius’ work. In the collections I have examined from South Africa there are approximately 20 species that can be assigned to Orthotylus and related genera. Several distinct groups of species exist, and the type of character variation in them is difficult to understand. At least two species from South Africa appear to be related to the European genus Pachylops Fieber. The claspers of the males are modified and bizarre in one species, which has a slender, elon- gate labium, while the other species has much more conventional male genitalia, and a short apically thickened labium, very similar to Pachylops species from Europe. The coloration of these species is essentially brownish or reddish and the dorsum is polished and shining. Two specimens very close to Chlorosomella (= Orthotylus) geniculata Reuter are known from Politzi, Transvaal. A long series of light green males from light traps, primarily at Grootfontein, Middelburg, Cape Province, probably represents a single rather variable species. These specimens have the black, scale-like hairs of the subgenus Orthotylus ( Melanotrichus ); they also have claspers that are of a type that occurs in several species that lack the black scale-like hairs. Two small groups of light green species, with the clasper type found in the “ Melanotrichus ” species mentioned above, can be dis- tinguished on labial length. Eight additional species which vary in characters of the eyes, beak length, male genitalia, and general body shape have also been examined. One of the most common “Orthotylus” species is velvety green and lives on Acacia. Two specimens appearing to be closely related to this species are known from Djab, South West Africa and are de- posited in the Transvaal Museum; they have the hemelytra velvety red instead of green. A very small species with a dark head, pronotum, and scutellum and light hemelytra is known from Malips Drif, Transvaal. Pseudambonea, new genus Macropterous Female: Head nearly vertical, body thick- set; pronotum distinctly transversely rugose, head, scutellum, and hemelytra smooth, head with scattered, semierect, light hairs about as long as tibial diameter; remainder of dorsum with decumbent light hairs about as long as tibial diameter; antennae with short, light 1974] Schuh: South African Orthotylinae and Phylinae 43 hairs about as long as diameter of segment 3; antennal segment 1 with a few, erect, fine, light spines on inner surface; labium with some short, erect, light hairs; thoracic pleura glabrous; femora with some decumbent light hairs and a few very long, fine, erect hairs on ven- tral surfaces; tibiae and tarsi with reclining, short, light hairs and a few semierect light spines about the length of tibial diameter; ab- domen with reclining light hairs about as long as tibial diameter. Head strongly declivous, slightly wider than anterior margin of pronotum; eyes occupying about half height of head, not noticeably granular; antennae inserted at level of ventral margin of eyes; an- tennal segment 1 moderately enlarged, segment 2 distinctly tapering proximally, distally about three-fourths diameter of segment 1, seg- ments 3 and 4 subequal in diameter, equal to proximal diameter of segment 2 (about three-fourths distal diameter of segment 2); clyp- eus large, rounded transversely, strongly curved posteroventrally; bucculae small; buccal cavity short; gula short, length about equal to diameter of antennal segment 1; pronotum only very slightly nar- rowed anteriorly, anterior margin finely carinate, upturned, lateral margins nearly straight; pronotum flattened, calli indistinct; entire posterior margin of pronotum slightly upturned; mesoscutum con- cealed beneath pronotum; scutellum very slightly elevated; hem- elytra strongly convex transversely; lateral corial margins strongly and somewhat irregularly convex, hemelytra widest at level just posterior to midpoint of cl aval commissure; cuneal incisure deep, fracture at right angles to longitudinal axis of body; cuneus and membrane strongly declivous; membrane with two cells; femora narrow; only metatibiae with longitudinal rows of tiny closely- spaced spines; claws stout, strongly curved, thickened basally; par- empodia fleshy, convergent apically, recurved; pulvilli minute. Female Genitalia: Figure 106. Posterior wall with well develoed K-structures. Macropterous Male: Structurally similar to female but more elongate. Male Genitalia: Figures 104, 105. Vesica membranous with sclerotized spiculi. Type Species: Pseudambonea capeneri, new species. This genus is named for its very close resemblance to Ambonea Odhiambo. Pseudambonea appears superficially to be closely related to Ambonea in the Pilophorini. The structure of the posterior wall of the female, in particular, indicates that there is actually no close relationship between the two genera. This contention is supported 44 Entomologica Americana [Vol. 47 also by the structure of the male genitalia. The genus can be most easily recognized by the compact body form, the posteriorly concave, strongly declivous, broad head, the single type of pubescence, the orthotyline pretarsal structures and the type of male and female gen- italia. Ambonea can be easily separated from Pseudambonea in that it has wooly pubescence as well as setiform hairs on the dorsum. Pseudambonea capeneri, new species Figures 11, 104-106 Macropterous Female: Dorsum generally light brown or tan; broad median longitudinal band on head including clypeus (excluding posterior magin of vertex), pronotum very broadly on either side of midline, and clavi (except anteriorly along suture) brown; posterior half of hemelytra and cuneus mesally more or less strongly suffused with brown or reddish brown; membrane smoky brown; antennal segment 1, proximal half of antennal segment 2, labium, and legs including procoxae cream; distal half of antennal segment 2, antennal segments 3 and 4, and all tarsal segments 3 dark brown; metafemora with broad red band distally; mesocoxae and metacoxae, thoracic pleura, and most of abdominal venter red- dish brown; anterior half of abdominal segment 9 light. Entire body highly polished and shining. Posterior margin of vertex with distinct raised carina; antennal fossae removed from anterior margins of eyes by distance equal to distal diameter of antennal segment 2; labium reaching apex of meso- coxae; anterior pronotal margin weakly sinuate, posterior margin straight across scutellum, very broadly rounded laterally; posterior margin of membrane cells broadly rounded; abdomen just surpass- ing apex of cuneus; metatarsal segment 2 slightly longer than seg- ment 1, segment 3 about 2 times length of segment 1. Measurements: Total length 3.36, maximum width 1.92, length head .16, width head 1.04, interocular space .60, length pro- notum .64, width pronotum 1.40, length scutellum .52, width scu- tellum .72, length corium 1.64, length clavus 1.36, length cuneus .64, width cuneus .56, length claval commissure .72, distance apex commissure-apex membrane 1.52, length metatibia 1.48; length antennal segments 1 — .32, 2 — 1.08, 3 — .52, 4 — .40; length labial segments 1 — .34, 2 — .32, 3 — .20, 4 — .24. Female Genitalia: Figure 106. Macropterous Male: Appearing uniformly gray brown; col- oration may be result of teneral condition of only known male specimens. 1974] Schuh: South African Orthotylinae and Phylinae 45 Male Genitalia: Figures 104, 105. Holotype: Macropterous 2, south Africa: Cape Province, 16 mi. north of Steytlerville, 24.X.64, A. L. Capener (SANC). Paratypes: 3 macropterous 2$, 2 macropterous SS, same data as holotype (SANC, RTS). This species is named for the collector, Mr. A. L. Capener. As the only known species in the genus, P. capeneri can be recognized by the characters noted in the generic discussion. Pseudoloxops Kirkaldy Loxops Fieber, 1858, p. 314 (preocc.). Pseudoloxops Kirkaldy, 1905, p. 268 (new name). Pseudoloxops Kirkaldy is widely distributed in the southern Palearctic and Old World tropics including the Southwest Pacific. It currently includes 16 described species. The genus can be rec- ognized by the following combination of characters: body ovoid; coloration usually carmine-red and yellow- white; frons bluntly pro- duced; antennal segment 1 and dorsum with long, shaggy pubes- cence; parempodia convergent apically, recurved; and male and female genitalia of the “Orthotylus- type”. The shaggy pubescence is often badly rubbed. Several specimens probably representing three new species of Pseudoloxops, in addition to the species described below, are known from South Africa. They are: a male from Satara Camp, Kruger National Park, deposited in the J. A. Slater Collection; a female from Malelane, Transvaal, deposited in the Transvaal Museum, probably the same species as the Satara specimen; a female from Port St. Johns, Cape Province, deposited in the British Museum (Natural History), probably closely related to the above specimens, and a female from Keiskama Hoek, King Williams Town District, Cape Province, deposited in the South African National Collection of Insects, distinct from all of the above specimens. Pseudoloxops transvaalensis, new species Figures 12, 107, 108 Macropterous Male: Red as follows — pronotum, mesoscu- tum, clavus, corium, apical half of cuneus, veins of membrane, mid- line of frons, vertex very narrowly, juga, antennal segment one, thoracic pleura, distal two-thirds of metafemora, and venter of ab- domen laterally; vertex, frons, scutellum (heavily suffused with red medially), antennal segment 2 (segments 3 and 4 missing from 46 Entomologica Americana [Vol. 47 holotype), thoracic sternum, all coxae, profemora and mesofemora entirely, metafemora proximally, all tibiae, all tarsi, and abdominal venter, irregularly, cream; basal half of cuneus orange. Entire body surface smooth, dull; dorsum with moderately dense, semierect, wooly hairs about as long as IV2 times diameter of an- tennal segment 2; pronotum, scutellum, clavus, and corium with some decumbent, flattened, sericeous hairs (in patches on clavus and corium); antennal segment 1 with many long, shaggy, dark hairs as on dorsum, segment 2 with very short, decumbent, light pubes- cence; labium with some erect, short, light hairs; femora with re- clining hairs and a few long, fine, erect hairs on ventral surfaces; tibiae with only very sparse, fine, reclining light hairs and a few semierect light spines slightly longer than tibial diameter; thoracic pleuron glabrous; abdominal venter with long, reclining, light hairs. Eyes very large, strongly protuberant, occupying nearly entire sides of head as viewed from above, vertex flat, nearly horizontal, posterior margin poorly defined, ecarinate; frons transversely rugose, produced into broad, blunt, rounded projection occupying entire space between antennal bases as viewed from above; anterior mar- gins of eyes strongly sinuate; antennae inserted at middle of anterior margins of eyes, fossae contiguous with eyes; antennal segment 1 distinctly enlarged, somewhat swollen medially, segment 2 cylindri- cal, slightly more than half diameter of segment 1; bucculae only slightly expanded; buccal cavity reaching prosternum; labium reach- ing to trochanteral joint of mesocoxae; pronotum with anterior mar- gin finely carinate, upturned, sinuate, lateral margins nearly straight, distinctly convergent anteriorly, posterior margin excavated across mesoscutum; calli obscure, with shallow longitudinal depression be- tween them; mesoscutum broadly exposed, about half length of flat scutellum; lateral corial margins straight, parallel; cuneal incisure obsolete, fracture at right angles to corial margin; metatibiae only with longitudinal rows of tiny closely spaced spines; metatarsal seg- ments 1 and 2 subequal in length, segment 3 slightly longer than segment 2; claws strongly curved, broad basally; parempodia fleshy, convergent apically, recurved; pulvilli minute. Measurements: Total length 3.68, maximum width 1.44, length head .36, width head .84, interocular space .32, length pro- notum .44, width pronotum 1.24, length scutellum .64, width scu- tellum .92, length corium 1.84, length clavus 1.44, length cuneus .60, width cuneus .30, length claval commissure .84, distance apex commissure-apex membrane 1.44, length metatibia 1.96, length antennal segments 1 — .48, 2 — .80, 3 — .28, 4 — .32. 1974] Schuh: South African Orthotylinae and Phylinae 47 Male Genitalia: Figures 107, 108. Vesica membranous with sclerotized spiculi. Macropterous Female: Similar to male but with the eyes smaller and vertex relatively wider; pronotum more flattened than in male, posterior margin not excavated; lateral corial margins slightly convexly rounded. Female Genitalia: Posterior wall with well developed Re- structures. Holotype: Macropterous S, south Africa: Transvaal , Clau- diushoop, 11 mi. N. Dendron, 15.12.65, M. Johannsmeier (SANC). Paratypes: Transvaal — 1 macropterous S, Messina, XII-30-I- 2-1957 (Capener); 1 submacropterous 2, Punda Milia, KNP., 16.1.65 (Hoffman) (SANC, RTS). Additional Specimens: South West Africa — 1 macropterous 2, Abachaus, IX. 1946 (Hobohm) (TM). This species is named for its occurrence in the Transvaal. P. transvaalensis is characterized by long wooly pubescence, a bluntly produced frons, deep red coloration with the basal half of the cuneus orange, and the structure of the male genitalia. The specimen from Abachaus, South West Africa, is in very poor con- dition and therefore has not been included in the paratype series. Nothing is known of the biology of this species. Pseudopilophorus, new genus Macropterous Female: Elongate, ant mimetic; entire body smooth, dull; pronotum indistinctly transversely rugose; body with moderately dense, sericeous, decumbent pubescence; antennae with very fine, short, semiappressed vestiture; femora and tibiae with de- cumbent hairs. Head vertical, elongate dorsoventrally, frons and gula nearly parallel; eyes large, somewhat reniform in lateral view, contiguous with anterior margin of pronotum; posterior margin of vertex cari- nate, slightly arched above dorsal margin of eyes; vertex depressed just anterior to posterior margin; frons nearly flat; antennae in- serted just above ventral margin of eyes; antennal segment 1 only slightly enlarged, segment 2 tapering proximally, distal diameter about equal to diameter of segment 1, segment 3 about equal to proximal diameter of segment 2, segment 4 of slightly smaller diam- eter than segment 3; clypeus large; labrum compressed laterally; gula about length of antennal segment 1; buccal cavity large, elon- gate oval; labium very short; pronotum campanulate, flat, somewhat 48 Entomologica Americana [Vol. 47 inclined posteriorly; mesoscutum exposed, flattened; scutellum con- vex; hemelytra with lateral margins strongly sinuate, narrowest at about middle of corium; clavi elevated along commissure; cuneal fracture angled anteromedially; membrane with 2 cells; profemora and tibiae of conventional structure; mesofemora slightly larger proximally than distally; mesotibiae flattened laterally, tapered, greatest width about one-third distance from femoral joint; meta- femora and tibiae similar in structure to mesolegs but much longer and conspicuously bowed; mesotibiae and metatibiae with longitu- dinal rows of tiny closely spaced spines; parempodia fleshy, apically convergent, recurved; pulvilli minute; abdomen strongly constricted basally. Female Genitalia: Figures 109, 112. Posterior wall with well developed K-structures; sclerotized rings with lateral margins strongly infolded. Macropterous Male: Very similar to female. Male Genitalia: Figures 110, 111. Vesica membranous with long sclerotized spiculum. Type Species: Pseudopilophorus capeneri , new species. This genus is named for its resemblance to Pilophorus. Pseudopilophorus superficially resembles Pilophorus\ however, the structure of the male and female genitalia place it in the Ortho- tylini, rather than in the Pilophorini. This is the only ant-mimetic genus in the Orthotylini known to occur in Africa and it is not ob- viously related to any other described genus. Pseudopilophorus can be recognized by its ant-mimic appearance, coloration pattern, con- vergent recurved parempodia, and male and female genital structures. Pseudopilophorus capeneri, new species Figures 13, 109-112 Macropterous Female: Posterior two-thirds of pronotum, scutellum (except as noted below), thoracic pleura and venter, and abdomen slate gray; head, anterior half of pronotum, protibiae on dorsal and ventral surfaces, antennal segment 3, distal half of an- tennal segment 4, labial segments 1 and 2, labrum, and dorsal stripe on profemora orangish to mahogany; proximal half of antennal seg- ment 4, labial segments 3 and 4, procoxae, profemora, lateral sur- faces of protibiae, mesotibiae distally, all tarsi, metacoxae, oval mac- ulae covering most of posterior half of scutellum and most of cunei cream to light yellow; remainder of legs castaneous to black; corium and clavus generally gray brown; membrane and posteromesial por- 1974] Schuh: South African Orthotylinae and Phylinae 49 tion of cuneus smoky gray; much of body surface with dull whitish bloom. Mesial margins of eyes straight, diverging only slightly ventrally in anterior view; antennal fossae nearly contiguous with eyes; labium just attaining middle of mesosternum; anterior margin of pronotum straight, posterior margin sinuate, concave across mesoscutum; ab- domen not quite attaining apex of membrane; posterior margin of large membrane cell broadly rounded; metatarsal segments subequal in length. Measurements: Total length 4.96, maximum width 1.44, length head .20, width head 1.04, interocular space .48, length pro- notum .84, width pronotum 1.32, length scutellum .96, length cor- ium 2.44, length clavus 1.92, length cuneus .92, width cuneus .52, length claval commissure 1.12, distance apex commissure-apex mem- brane 1.96, length metatibia 4.20; length antennal segments 1 — .32, 2 — 1.76, 3 — 1.28, 4 — .72; length labial segments 1 — .30, 2 — .32, 3 — .28, 4— .32. Female Genitalia: Figures 109, 112. Male Genitalia: Figures 110, 111. Holotype: Macropterous $, south Africa: Transvaal, Tza- neen, 11-16 Dec. 1963, A. L. Capener (SANC). Paratypes: Macropterous $, 10 macropterous $9, same data as holotype (1 specimen — host plant Terminalia sericea ). Swazi- land—Eranchi, XII-15-31-1954 (Capener) (SANC, JAS, RTS). This species is named for Mr. A. L. Capener. As the only species in the genus, F. capeneri can be recognized by the characters noted in the generic discussion. This species has been taken on Terminalia sericea Burch. (Combretaceae), but no other biological information is available. Zancliiella, new genus Macropterous Male: Small, elongate, elliptical, or nearly parallel sided; head, pronotum, and scutellum smooth; pronotum weakly transversely rugulose; hemelytra hyaline or subhyaline; dor- sum with shining or dull, moderately long, semierect hairs; head broad, narrowed behind eyes; eyes large, granular, with or without short hairs; vertex weakly convex; antennae inserted slightly below middle of anterior margin of eyes which are more or less emarginate; antennal segment 1 slightly enlarged distally, segment 2 of slightly smaller diameter than segment 1, segments 3 and 4 subequal in diameter, about two-thirds diameter of segment 2; bucculae weakly 50 Entomologica Americana [Vol. 47 developed; gula short; labium long; pronotum with very narrow collar; calli low, indistinct; lateral margins of pronotum slightly con- cave; mesoscutum narrowly exposed, separated from scutellum by distinct, deep, transverse impression; scutellum usually broadly con- vexly elevated; lateral margins of hemelytra usually broadly con- vexly rounded; cuneal incisure very shallow, fracture slightly angled anteromedially; clavus with row of very fine punctures adjacent to scutellum and commissure and also row of punctures parallel to claval suture; two cells in membrane, inner vein nearly parallel to inner margin of clavus; abdomen reaching approximately to base of membrane; metatibiae with several longitudinal rows of tiny, closely- spaced spines; all tibiae with a few scattered, light, reclining spines about length of tibial diameter; tarsal claws strongly curved; par- empodia fleshy, apically convergent, recurved; pulvilli minute. Male Genitalia: Vesica membranous, without spiculi. Macropterous Female: Structurally similar to macropterous male; eyes slightly smaller and vertex correspondingly wider in fe- male than in male. Female Genitalia: Posterior wall with well developed Re- structures. Type Species: Zanchiella bowkeriae , new species. This genus is named for its close resemblance to Zanchius Distant. Zanchiella can be recognized by the generally hyaline hemelytra with a row of punctures along the claval suture (see however Z. ericae ) and the relatively short appendages. It is most closely re- lated to Felisacodes and Zanchius in Africa.. The row of punctures on the clavus, the shape of the head, and the hyaline hemelytra, ally Zanchiella to Felisacodes ; the general body form, particularly the structure of the hemelytra, and the type of vestiture, relate Zanchi- ella to Zanchius. The convergent recurved parempodia, the mem- branous vesica in the male, and the posterior wall with well devel- oped K-structures in the female support the placement of Zanchi- ella in the Orthotylini. Key to species of Zanchiella 1. Basic coloration greenish (some specimens yellowish), cuneus red, contrasting with remainder of hemelytra; hemelytra not hyaline ericae (Fig. 16) Basic coloration sometimes greenish, but cuneus and remainder of hemelytra never contrastingly colored; hemelytra hyaline or subhyaline - 2 1974] Schuh: South African Orthotylinae and Phylinae 51 2. Clavus dark brown except for narrow light area along lateral mar- gin; frons, clypeus, and pronotum partially red; veins of mem- brane cells dull red capensis (Fig. 15) Clavus entirely light or with dark marking only on posterior third and along mesial margin; frons, clypeus, pronotum, and veins of membrane not red 3 3. Anterior third of pronotum tan, posterior two-thirds very dark brown to black; posterior third of clavus suffused with dark brown natalensis (Fig. 17) Pronotum and clavus nearly unicolorous light yellow or green .... 4 4. Basic coloration light yellow; hemelytra with distinct brown fascia between apex of claval commissure and base of membrane .... bowkeriae (Fig. 14) Basic coloration of hemelytra light greenish; head, pronotum, and scutellum light brownish; transverse fascia on hemelytra obsolete or very faint sweeti (Fig. 18) Zanchiella bowkeriae, new species Figure 14 Macropterous Male: Head, pronotum, scutellum, hemelytra, thoracic venter, and legs very light yellowish with darker markings as noted below; pronotum lateroventrally and posterolaterally and scutellum anterolaterally suffused with brown; corium with irregular brown transverse macula between apex of clavus and base of mem- brane; inner margin of costal vein and lateral margin of cuneus ob- scurely suffused with green; eyes black; antennal segment 1 and distal third of segment 2 dark brown, proximal two-thirds of seg- ment 2 light brown or yellow, distal half of segment 2 with broad reddish band; antennal segments 3 and 4 light brown; membrane smoky brown, veins slightly darker; abdomen very light green to nearly white. Hemelytra hyaline, weakly shining. Frons slightly produced between eyes; anterior margin of eyes conspicuously emarginate at antennal bases; labium just attaining distal end of metacoxae; punctures on clavus small but distinct. Measurements: Total length 4.52, maximum width 1.34, length head .30, width head .66, interocular space .24, length pro- notum .52, width pronotum 1.02, length scutellum .46, width scu- tellum .62, length corium 1.98, length clavus 1.30, length cuneus .81, width cuneus .40, length claval commissure .72, distance apex commissure-apex membrane 2.00, length metatibia 2.20; length antennal segments 1 — .42, 2 — 1.50, 3 — .82, 4 — approx. .74; length labial segments 1 — .28, 2 — .30, 3 — .40, 4 — .38. 52 Entomologica Americana [Vol. 47 Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous $, south Africa: Transvaal , 22 mi. S. Barberton, 4900 ft. elevation, 24 Mar. 1968, T. Schuh, J. A. & S. Slater, M. Sweet (Adults on Bowkeria cymosum McOwan) (SANG). Paratypes: 4 macropterous $$, 3 macropterous 2 2, same data as holotype (SANC, JAS, RTS). This species is named for the host plant genus Bowkeria. Zanchiella bowkeriae is most closely related to Z. capensis and Z. natalensis. It can be most easily recognized by the light colored pronotum in combination with the conspicuous dark fascia on the posterior half of the corium. This species is known only from the type locality on Bowkeria cymosum McOwan (Scrophulariaceae) . The host genus is endemic to South Africa (Phillips, 1951). Zanchiella capensis, new species Figure 15 Macropterous Male: Basic coloration very light yellowish; frons, clypeus, basal third of antennal segment 1, pronotum dorsally (except for a rounded, light, central marking and the posterior quarter), mesoscutum, and extreme apex of cuneus, bright red; remainder of antennae brown; eyes black; pronotum on pleural region and posterior quarter dorsally dark brown; clavus mesiad of row of puctures along claval suture dark brown; transverse macula on corium between apex of cuneus and base of membrane, and veins of membrane dull red brown; thoracic pleuron light reddish brown (except as above); abdomen lateroventrally and genital segments red, remainder white. Pronotum very obscurely rugulose, polished, shining; hemelytra hyaline, weakly shining. Frons slightly convexly produced between eyes; eyes weakly emarginate at antennal bases; labium just surpassing apex of meso- coxae. Measurements: Total length 3.20, maximum width 1.12, length head .26, width head .58, interocular space .26, length pro- notum .44, width pronotum .84, length scutellum .40, width scu- tellum .58, length corium 1.64, length clavus 1.10, length cuneus .66, width cuneus .30, length claval commissure .64, distance apex commissure-apex membrane 1.44, length metatibia 1.86; length antennal segments 1 — .32, 2 — 1.16, 3 — .66, 4 — .60; length labial segments 1 — .30, 2 — .30, 3 — .42, 4 — .40. 1974] Schuh: South African Orthotylinae and Phylinae 53 Figs. 13-18. Orthotylini. Fig. 13. Pseudopilophorus capeneri, male, holotype. Fig. 14. Zanchiella bowkeriae, male, holotype. Fig. 15. Zanchiella capensis, male, holotype. Fig. 16. Zanchiella ericae, male (Giants Castle, Natal). Fig. 17. Zanchiella natalensis, male, holotype. Fig. 18. Zanchiella sweeti, female (Fountains, Pretoria, Transvaal). Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous $, south Africa: Cape Province , Ysterhoutrug Picnic Site, 18 mi. NE Knysna, 10 Feb. 1968, T. Schuh, J. A. & S. Slater, M. Sweet (SANC). 54 Entomologica Americana [Vol. 47 Paratypes: Cape Province — 1 macropterous $, same data as holotype; 1 macropterous 2, Storms River Mouth, 14-15.X.1964 (Capener) (SANC, RTS). This species is named for its occurrence in the Cape Province. Zanchiella capensis is most colsely related to Z. natalensis and Z. bowkeriae but can be separated from them by the characters noted in the above key. No host or biological information is available for this species. Zanchiella ericae, new species Figure 16 Macropterous Male: Basic coloration greenish with strong yellow tinge, or nearly all yellow; cuneus red; membrane light smoky gray, veins slightly reddish; antennae and legs light brownish, an- tennal segments 3 and 4 and all tarsal segments slightly darker than remainder of appendages; labial segments 1 and 2 greenish yellow, segments 3 and 4 brown; eyes dark brown. Dorsum generally smooth and very finely granulose, weakly shining; hemelytra subhyaline; dorsum with scattered, fine, erect or semierect, moderately long, light brown hairs; antennae with a fine erect spine on interior surface of segment 1; eyes glabrous. Frons rather prominently convexly rounded between eyes, clypeus visible from above; anterior margins of eyes only weakly concave, not noticeably emarginate; labium slightly surpassing metacoxae; pronotal collar extremely narrow; anterior margin of pronotum weakly sinuate; posterior margin of pronotum broadly excavated across scutellum; row of punctures on clavus obsolete; tibial spines semierect. Measurements: Total length 3.92, maximum width .92, length head .30, width head .60, width vertex .26, length pronotum .30, width pronotum .76, length scutellum .50, width scutellum .62, length corium 1.96, length clavus 1.28, length cuneus .84, width cuneus .30, length claval commissure .76, distance apex commissure- apex membrane 2.00, length metatibia 2.06; length antennal seg- ments 1 — .38, 2 — 1.44, 3 — .98, 4 — .54; length labial segments 1 — .34, 2— .34, 3— .36, 4— .34. Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous S, south Africa: Natal , Giants Castle Park, 5800 ft. elevation, 6 Mar. 1968, T. Schuh, J. A. & S. Slater, M. Sweet (Adults and nymphs on Erica leucopelta Tausch.) (SANC). 1974] Schuh: South African Orthotylinae and Phylinae 55 Paratypes: 5 macropterous SS, 6 macropterous 2 2, same data as holotype (SANC, JAS, RTS). Additional Specimens: Cape Province — 3 macropterous 2$, Bainskloof Pass Summit, 21 Jan. 1968; 2 macropterous SS, 3 macropterous 9 9, Femkloof Nat. Res., Hermanus, 3 Feb. 1968; 1 macropterous 9, 2 mi. S. Goukamma, Knysna, 8 Feb. 1968; 1 macropterous S, 1 macropterous 9, Hermanus, 1 Feb. 1968; 4 macropterous $S, 15 macropterous 9 9 (in alcohol — 1 $,2 9 9, 4 nymphs), just W. of Knysna, 8 Feb. 1968 (Adults and nymphs on Erica floribunda Lodd.); 1 macropterous $, 1 macropterous 9, 6 mi. E. Plettenburg Bay, elevation 500 ft., 12-13 Feb. 1968; 2 macropterous $$, 4 macropterous 2 2 (in alcohol — 1 $), Yster- houtrug Picnic Site, 18 mi. NE Knysna, 10 Feb. 1968 (Adults and nymphs on Erica sp.). Natal — 9 $ $ , 12 22 (6 nymphs in alcohol), same data as holotype; 1 macropterous S, 1 macropterous 2, Sani Pass, 6200 ft., 10 Mar. 1968 (Adults and nymphs on Erica leuco- pelta Tausch.). Orange Free State — 7 macropterous $$, 2 mac- ropterous 2 2, Golden Gate, 12.X.66 (Capener) (Host plant — Erica maesta). Transvaal — 6 macropterous 2 2 (in alcohol — 1 $, 1 nymph), 22 mi. S. Barberton, 4900 ft. elevation, 24 Mar. 1968 (Adults and nymphs on Erica drakensbergensis Guth. & Bol.) (SANC, HM, BM[NH], JAS, RTS). This species is named for the host plant genus, Erica. I am placing this species in Zanchiella with some reservations. The other four species of the genus are very closely related based on the presence of claval punctures (obscure in sweeti and absent in ericae), the general body shape (much less oval in sweeti and ericae than in the other species), and the coloration pattern. Zanchi- ella ericae is isolated within the genus, but it seems unwise at the present time to erect a new genus for the reception of this species until the taxonomy of Zanchiella and related genera is better under- stood. The variation in coloration and relative proportions between specimens from different localities is in many cases extreme, and if an adequate series were not available many specimens would almost certainly be considered as representing distinct species. The most obvious variation is in the ratio of the total length to maximum width, the ratio of length to width of the cuneus, and the ratio of total width of the head to the interocular space. The Giants Castle specimens have a very elongate aspect with a long narrow cuneus and rather large eyes and a narrow vertex. Specimens from Fem- kloof Nature Reserve, Hermanus, are at the opposite extreme in all 56 Entomologica Americana [Vol. 47 of these characters. Specimens from other localities show inter- mediate conditions, but there is no obvious geographical pattern to the variation. Most specimens are light green with some yellowish suffusion, but others are almost totally yellow. Because of this vari- ation I have designated only those specimens from the type locality as paratypes. The structure of the male genitalia of all specimens examined is very similar. Z. ericae is apparently restricted to the genus Erica (Ericaceae), and seems to occur in nearly all regions of South Africa where Erica is present. Known host species include E. floribunda Lodd., E. maesta, E. leucopelta Tausch., and E. drakensbergensis Guth. and Bol. Zanchiella natalensis, new species Figure 17 Macropterous Male: Anterior portion of vertex and frons including clypeus brown; posterior half of vertex, head below eyes, and anterior third of pronotum very light brown; posterior two-thirds of pronotum and entire scutellum very dark brown; hemelytra hy- aline, nearly transparent; clavus along scutellum and commissure very dark brown; posterior third of clavus and macula on corium between apex of clavus and base of membrane brown; membrane, particularly veins, smoky brown; mesial margin of costal vein and lateral margin of clavus suffused with green; antennae dark brown; labium white; thoracic pleura generally brown; coxae and legs very light brown or yellowish; base of abdomen and genital segment dark brown; remainder of abdomen light greenish. Pronotum and scutellum polished, shining, with transverse ru- gosities. Head very weakly produced between eyes; anterior margins of eyes weakly emarginate; labium just surpassing apex of metacoxae. Measurements: Total length 3.64, maximum width 1.08, length head .28, width head .62, interocular space .20, length pro- notum .44, width pronotum .84, length scutellum .42, width scu- tellum .52, length corium 1.68, length clavus 1.28, length cuneus .66, width cuneus .30, length claval commissure .66, distance apex commissure-apex membrane 1.74, length metatibia 1.94; length antennal segments 1 — .40, 2 — 1.24, 3 — .81, 4 — .79; length labial segments 1 — .26, 2 — .30, 3 — .58, 4 — .20. Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous $, south Africa: Natal, Olivier- 1974] Schuh: South African Orthotylinae and Phylinae 57 shoek Pass Summit, 5400 ft. elevation, 25 mi. S. Harrismith, 4 Mar. 1968, T. Schuh, J. A. & S. Slater, M. Sweet (SANC). Paratypes: Natal — 1 macropterous 8, Cathedral Peak, Jan. 1964 (Capener); 1 macropterous 8 (?), Drakensberg, 1-22.1.1927 (Turner); 1 macropterous 8 , same data as holotype. Transvaal — 1 macropterous 9, Wylies Poort, 10.2.41 (Capener) (SANC, BM [NH], RTS). This species is named for its occurrence in Natal. Zanchiella natalensis is most closely related to Z. capensis and Z. bowkeriae. It can be recognized by the pronotum being light on the anterior third and dark on the posterior two-thirds, in combina- tion with the dark transverse macula on the posterior half of the corium. Zanchiella sweeti, new species Figure 18 Macropterous Male: Elongate, nearly parallel sided; head (excepting posterior margin of vertex and genae), pronotum, scu- tellum, clavus along posterior two-thirds of commissure, and trans- verse macula on corium at level of base of membrane orangish brown; genae, posterior half of vertex, hemelytra (excluding mem- brane), thoracic venter, legs, and labium very light green; abdom- inal venter green; proximal fifth of antennal segment 1 light green, distal four-fifths red; antennal segment 2 brown on proximal end, remainder with broad reddish brown bands alternating with light coloration; antennal segments 3 and 4 brown; membrane light smoky gray. Head polished, smooth, and shining; pronotum and scutellum obscurely transversely rugulose, weakly shining; hemelytra subhy- aline, dull; membrane rugulose; dorsum with semierect, moderately long, light hairs; anterolateral angles of pronotum with single, long, erect, fine seta; antennae with short, decumbent, light vestiture; abdomen with moderately long, reclining, light hairs. Frons weakly convexly produced between eyes; anterior mar- gins of eyes weakly emarginate; labium just surpassing metacoxae; calli rather widely separated, demarcated posteriorly by weak fur- row; posterior pronotal margin very slightly convex; punctures on clavus faint; tibiae with scattered, very fine, short, reclining light spines, hardly distinguishable from light tibial hairs; metatarsal segment 1 about half length of segment 2; segments 2 and 3 sub- equal in length. Measurements: Total length 3.24, maximum width .98, length 58 Entomologica Americana [Vol. 47 head .28, width head .60, interocular space .20, length pronotum .42, width pronotum .78, length scutellum .34, width scutellum .46, length corium 1.58, length clavus 1.04, length cuneus .58, width cuneus .30, length cl aval commissure .60, distance apex commissure- apex membrane 1.52, length metatibia 1.80; length antennal seg- ments 1 — .38, 2 — 1.14, 3 — .74, 4 — .64; length labial segments 1 — .30, 2— .28, 3— .38, 4— .34. Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous $, south Africa: Transvaal , Pre- toria, Fountains, 15 December 1967, M. Sweet, at light (SANC). Paratypes: Transvaal — 11 macropterous <$<$,8 macropterous $ 2, same data as holotype; 2 macropterous 2 2, Pretoria, Spring- bok Park, Jan. 1966 (Paliatseas) (SANC, BM[NH], JAS, RTS). This species is named for the collector, Dr. Merrill H. Sweet. Zanchiella sweeti can be recognized by the basic greenish col- oration of the hemelytra and the inconspicuous transverse macula on the posterior half of the corium. Zanchius Distant Zanchius Distant, 1904c, p. 477. — Carvalho, 1956b, p. 66. Zanchius can be characterized as follows — Macropterous Male: Body flattened, structure delicate; col- oration light; dorsal vestiture of moderately long, semierect, light hairs; head usually flattened and quadrate (or somewhat broader than long); eyes conspicuously set forward, small, granular, pro- tuberant, with short hairs present in some species; vertex flat or nearly so; frons convex, clypeus not or only barely visible from above; antennae inserted just above ventral margin of eyes, fossae nearly contiguous with eyes; antennal segment 1 long, cylindrical, moderately enlarged; antennal segment 2 slightly smaller in diameter than segment 1, segments 3 and 4 subequal in diameter, of slightly smaller diameter than segment 2; labium surpassing mesocoxae; calli distinct, set off by a transverse impression posteriorly; meso- scutum broadly exposed; scutellum nearly flat; hemelytra hyaline or subhyaline, very long; abdomen reaching at most to cuneal in- cisure; membrane with 2 cells; legs long and slender; tibiae with a few very thin, light colored spines about length of tibial diameter; metatibiae with several longitudinal rows of tiny closely spaced spines; metatarsal segment 1 shorter than segments 2 and 3; claws curved; parempodia fleshy, apically convergent, recurved; pul villi minute. 1974] Schuh: South African Orthotylinae and Phylinae 59 Male Genitalia: Vesica membranous, without spiculi, of the Orthotylini-type. Macropterous Female: Very similar to macropterous male. Female Genitalia: Posterior wall with well developed Re- structures. Zanchius is most closely related to Zanchiella in South Africa. The absence of the row of punctures paralleling the claval suture, the flattened head, and the absence of hemelytral maculae in Zanch- ius will help to separate it from Zanchiella in which there is a dis- tinct row of punctures on the clavus, the head is somewhat globose, and the hemelytra usually have a contrasting dark macula. The structure of the parempodia and male and female genitalia support placement of Zanchius in the Orthotylini. Distant (1904c) described Zanchius from India. The known range of the genus now includes the Southern Palearctic, South Africa, Southeast Asia, and the islands of the Southwest Pacific. Twelve species are currently placed in the genus, including those described as new below. Three female specimens from Roodeplaat, Transvaal, deposited in the South African National Collection of Insects, probably rep- resent a fifth new species in addition to those described as new in this paper. They differ from the other South African species of the genus in being somewhat smaller and generally orangish in coloration rather than green or white. Also available are several male and female specimens of what appears to be a new species of Zanchius; they are much smaller than any of the species of Zanch- ius described below and have the eyes only slightly set forward on the head, but agree closely with Zanchius in nearly all other char- acteristics. These specimens are deposited in the South African National Collection of Insects and the J. A. Slater Collection. Key to South African species of Zanchius 1. Antennal segment 1 with a black stripe on lateral surface; antennal segment 2 black proximally; general coloration light green or yellow-green nigrolineatus (Fig. 22) Antennal segment 1 without black stripe; coloration either very light greenish or white 2 2. Basic coloration white, hemelytra usually faintly suffused with light green; pronotum and hemelytra with elongate, longitudinal, yellow-orange markings buddleiae (Fig. 20) Basic coloration either white or very light green or greenish yellow without yellow-orange markings 3 3. Basic coloration dull white; large species, length 4.40 mm.; head 60 Entomologica Americana [Vol. 47 broad, ratio width across eyes to median length 37:14 alba (Fig. 19) Basic coloration very light green to nearly white; smaller species than above, length 3.48 mm.; head quadrate, ratio width across eyes to median length 2:1 leucosideae (Fig. 21) Zanchius alba, new species Figure 19 Macropterous Male: Entire body and appendages dull white or cream; hemelytra translucent but not hyaline, with only weak, scattered pigmentation; appendages and labium infuscate apically. Body surface smooth, dull; dorsum with rather long (about length of diameter of antennal segment 1), scattered, semierect, light hairs; antennae with short, fine, decumbent pubescence; ven- ter of abdomen with rather dense, semidecumbent, light hairs; fem- ora with semidecumbent, light hairs (about length of tibial diam- eter); tibiae with short, decumbent, light hairs and a few very light, fine, semierect spines about length of tibial diameter. Head short, much broader than long, vertical; eyes protuberant, noticeably granular, with some very short hairs; head constricted behind eyes; vertex nearly flat; frons weakly convex, not produced beyond anterior margin of eyes as viewed from above; clypeus not visible from above; antennae inserted just above ventral margin of eyes; antennal segment 1 moderately enlarged, nearly cylindrical, segment 2 about three-fourths diameter of segment 1, segments 3 and 4 subequal in diameter, about three-fourths diameter of seg- ment 2; head in frontal view triangular below eyes; bucculae small; labium just surpassing metacoxae; pronotum with anterior lobe somewhat swollen, demarcated by distinct transverse impression behind calli, anterolateral angles sharply rounded, anterior margin weakly sinuate, posterior lobe flattened, posterior margin shallowly excavated; mesoscutum about one-half length of scutellum; trans- verse impression separating scutellum and mesoscutum sinuate; scu- tellum weakly convex; lateral margins of hemelytra weakly convex, cuneal incisure shallow but distinct; fracture angled slightly antero- medially; metatarsal segment 1 half length of segment 3, segment 3 slightly shorter than segment 2. Measurements: Total length 4.40, maximum width 1.20, length head .28, width head .74, interoeular space .36, length pro- notum .36, width pronotum .82, length scutellum .48, width scu- tellum .74, length corium 2.24, length clavus 1.38, length cuneus .84, width cuneus .34, length claval commissure .68, distance apex 1974] Schuh: South African Orthotylinae and Phylinae 61 commissure-apex membrane 1.90, length metatibia 2.30; length antennal segments 1 — .48, 2 — 1.50, 3 — 1.00, 4 — .34; length labial segments 1 — .32, 2 — .38, 3 — .34, 4 — .40. Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous $, south Africa: Natal, Olivier- shoek Pass Summit, 5400 ft. elevation, 25 mi. S. Harrismith, 4 March 1968, T. Schuh, J. A. & S. Slater, M. Sweet (Adults and nymphs on Buddleia salviijolia [L.] Lam.) (SANC). Paratypes: Natal — 2 macropterous SS, 6 macropterous 2 2, Cathedral Peak, Jan. 1964 (Capener); 1 macropterous S, 1 mac- ropterous 2, Giants Castle Park, 5800 ft. elevation, 6 Mar. 1968 (Adults and nymphs on Buddleia salviijolia [L.] Lam.); 14 macrop- terous $ $ , 27 macropterous 2 2 , same data as holotype; 1 mac- ropterous S, 9 macropterous 2 2, Sani Pass, 6200 ft., 10 Mar. 1968 (SANC, TM, BM[NH], USNM, JAS, RTS). Additional Specimens: Natal — 7 macropterous SS, 5 mac- ropterous 2 2, 17 nymphs (in alcohol), same data as holotype. Transvaal — 3 males, 3 nymphs (in alcohol), 20 mi. NE Machado- dorp, Schoemannskloof, 4300 ft., 22 Mar. 1968 (ex: Buddleia salviijolia ) (RTS). This species is named for its very light coloration. Zanchius alba is the largest South African species of the genus. It is totally white or cream colored, becoming somewhat brownish in specimens preserved in alcohol. Z. alba is most easily separated from other South African species, particularly Z. leucosideae, by the length-width ratio of the head (see key). Zanchius alba was taken on Buddleia salviijolia (Loganiaceae) at all collection localities, but never at the same localities as Z. bud- dleiae, which apparently has the same host. Zanchius buddleiae, new species Figure 20 Macropterous Male: Basic coloration of body and append- ages dull, opaque white; hemelytra translucent, nearly devoid of white pigmentation, weakly and irregularly suffused with blue-green (this is absent in some specimens); dorsum with yellow-orange markings as follows — vertex medially at level of posterior margin of eyes with small round spot, posterior lobe of pronotum medially and on each side about one-third distance mesially from lateral margins with elongate markings, clavus with elongate streak, corium with elongate streaks along claval suture, claval commissure, and parallel to lateral margin of posterior half of clavus, cuneus with 62 Entomologica Americana [Vol. 47 round spot basomedially, and large cell of membrane with small spot medially. Body surface, pubescence, and structure very similar to Z. alba and Z. leucosideae, except as follows — head more or less quadrate, labium just attaining distal end of metacoxae, and posterior margin of pronotum very weakly sinuate. Measurements: Total length 2.92, maximum width .80, length head .28, width head .54, interocular space .24, length pronotum .28, width pronotum .60, length scutellum .38, width scutellum .50, length corium 1.50, length clavus 1.00, length cuneus .56, width cuneus .22, length claval commissure .58, distance apex commis- sure-apex membrane 1.24, length metatibia 1.60; length antennal segments 1 — .28, 2 — 1.10, 3 — .52, 4 — .46; length labial segments 1 — .24, 2— .26, 3— .34, 4— .26. Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous $, south Africa: Transvaal , Nat. Botanical Gardens, Pretoria, 22 November 1967, J. A. & S. Slater, T. Schuh (Adults and nymphs on Buddleia salviifolia [L.] Lam.) (SANC). Paratypes: 9 macropterous SS, 23 macropterous 2 2, same data as holotype (SANC, HM, JAS, RTS). Additional Specimens: 30 macropterous AS, 3 macropter- ous $2,4 nymphs (in alcohol), same data as holotype (RTS). This species is named for the host plant genus, Buddleia. Zanchius buddleiae most closely resembles Z. alba , but is much smaller, has orange markings on the pronotum and hemelytra and has a rather quadrate head compared to the broad head of alba (see key). Zanchius buddleiae is apparently host specific on Buddleia sal- viifolia ( Loganiaceae ) . Zanchius leucosideae, new species Figure 2 1 Macropterous Male: Basic coloration dull light green; legs, antennae, and labium infuscate apically. Entire body surface smooth, dull or weakly shining; hemelytra -> Figs. 19-22. Orthotylini. Fig. 19. Zanchius alba, male, holotype. Fig. 20. Zanchius buddleiae , male (Pretoria, Transvaal). Fig. 21. Zan- chius leucosideae , male, holotype. Fig. 22. Zanchius nigroleneatus, male, holotype. 1974] Schuh: South African Orthotylinae and Phylinae 63 64 Entomologica Americana [Vol. 47 subhyaline; vestiture essentially as in Z. buddleiae, except eyes glabrous. Structurally very similar to Z. alba and Z. buddleiae, except as follows — head more or less quadrate (see measurements); la- bium noticeably surpassing metacoxae; posterior margin of pro- notum broadly excavated across mesoscutum. Measurements: Total length 3.48, maximum width 1.10, length head .30, width head .60, interocular space .24, length pro- notum .28, width pronotum .70, length scutellum .52, width scu- tellum .64, length corium 1.84, length clavus 1.14, length cuneus .70, width cuneus .30, length claval commissure .70, distance apex commissure-apex membrane 1.80, length metatibia 2.34, length antennal segments 1 — .38, 2 — 1.36, 3 — .84, 4 — .42, length labial segments 1 — .26, 2 — .32, 3 — .58, 4 — .42. Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous S, south Africa: Natal, Sani Pass, 6200 ft., 10 Mar. 1968, T. Schuh, S. Slater, M. Sweet (SANC). Paratypes: Natal — 1 macropterous $, same data as holotype; 1 macropterous S , Sani Pass, 6000 ft., 10 Mar. 1968. Orange Free State — 2 macropterous 2$, 5 mi. N. Golden Gate Park, 17 Oct. 1967 (SANC, RTS). This species is named for the host plant genus, Leucosidea. Zanchius leucosideae resembles alba and buddleiae, but can be separated from the former by its more quadrate head and from the latter by its uniformly pale green coloration. Zanchius leucosideae is known to occur only on Leucosidea sericea Eckl. and Zeyh. (Rosaceae), which is endemic to South Africa (Phillips, 1951). Zanchius nigrolineatus, new species Figure 22 Macropterous Male: Basic coloration very light green or yellow green, hemelytra hyaline; veins of membrane greenish, mem- brane yellowish; antennae yellowish, segment 1 with broad black stripe laterally, extending entire length of segment, except for ex- treme distal end, segment 2 black proximally; tibiae yellowish; dull gray spot present at middle of inner vein of large cell and near apex of small cell; black spot present on vein at inner apical angle of large cell. Entire body and appendages smooth, shining; dorsum with light, moderately long (about length of greatest diameter of antennal seg- ment 1), semierect hairs; antennae with short decumbent pubes- 1974] Schuh: South African Orthotylinae and Phylinae 65 cence, segment 1 with some longer semierect hairs; abdominal ven- ter with moderately long, semierect hairs; legs with fine light hairs; tibiae with a few very fine, light spines about length of tibial diameter. Head quadrate, eyes set far forward, distance from posterior margin of eye to posterior margin of head slightly less than longitu- dinal diameter of an eye; vertex flat; frons convexly rounded be- tween antennal bases, clypeus just visible from above; eyes distinctly protuberant, granular, with short hairs (visible at 50X); antennae inserted just above ventral margin of eyes, fossae contiguous with eyes; antennal segment 1 moderately enlarged, greatest diameter near base, segment 2 about two-thirds diameter of segment 1, seg- ments 3 and 4 subequal in diameter, about two-thirds diameter of segment 2; bucculae moderately developed; labium reaching distal end of metacoxae; pronotum flat, depressed behind weakly elevated calli; anterior, lateral, and posterior margins of pronotum nearly straight; mesoscutum exposed, about one-third length of scutellum, separated from scutellum by medially interupted, weak, transverse impression; scutellum flat; lateral margins of hemelytra weakly con- vexly rounded; cuneal incisure obsolete, fracture at right angles to lateral corial margin; veins of membrane heavy, inner vein nearly parallel to mesial margin of cuneus; apex of abdomen not quite attaining cuneus; metatarsal segment 1 about one-half length of seg- ment 2, segment 2 about IV2, times length of segment 1. Measurements: Total length 3.56, maximum width 1.16, length head .30, width head .56, interocular space .24, length pro- notum .39, width pronotum .84, length scutellum .50, width scu- tellum .60, length corium 1.76, length clavus 1.24, length cuneus .64, width cuneus .28, length claval commissure .70, distance apex commissure- apex membrane 1.55, length metatibia 1.96; length antennal segments 1- — .36, 2 — 1.34, 3 — .64, 4 — approx. .70; length labial segments 1 — .22, 2 — .26, 3 — .36, 4 — .38. Male Genitalia: Not illustrated. See generic discussion. Holotype: Macropterous 8 , south Africa : Transvaal, Kruger Nat. Park, 3 mi. E. Skukuza Camp, 25 Apr. 1968, T. Schuh, J. A. & S. Slater, M. Sweet (SANC) . Paratypes: Transvaal — 3 macropterous 8 8, 6 macropterous 2 $, same data as holotype; 1 macropterous 8, Kruger Nat. Park, Oliphants River near Oliphants Camp, 30 Apr. 1968 (SANC, JAS, RTS). This species is named for the black stripe on the first antennal segment. Zanchius nigrolineatus is the most distinctive of the South Af- 66 Entomologica Americana [Vol. 47 rican species of the genus, especially in the structure of the head and pronotum. It can be easily separated from the other described species by the black stripe laterally on antennal segment 1. The specimens from the type locality were taken on Lantana sp. (Verbenaceae). SUBFAMILY PHYLINAE TRIBE HALLODAPINI Acrorrhinium Noualhier Acrorrhinium Noualhier, 1895, p. 176. Cinnamus Distant, 1909a, p. 441. New Synonymy. Acrorrhinium can be characterized as follows — Macropterous Male: Very elongate, nearly parallel sided; coloration pattern either mottled or with one or two contrasting hemelytral maculae. Body surface smooth, dull, or weakly shining; dorsum usually with short decumbent hairs, sometimes with erect peg-like hairs; antennae with short dense vestiture; abdominal venter with semi- decumbent shining hairs. Eyes protuberant, nearly hemispherical, removed from anterior margin of pronotum by at least one-third diameter of eye; head neck-like behind eyes; vertex horizontal; frons strongly convex, produced into more or less attenuated spine above clypeus, with five anteromedially directed transverse rugosities posterior to spine; clypeus compressed laterally, nearly vertical; antennae inserted at or just below middle of anterior margin of eyes, fossae contiguous with or only slightly removed from anterior margins of eyes; antennal segment 1 somewhat enlarged, about equal to length of head, seg- ment 2 about three-fourths diameter of segment 1, occasionally in- creasing in diameter distally, segments 3 and 4 subequal in diameter, about three-fourths diameter of segment 2; labium reaching or sur- passing metacoxae; pronotum with distinct transverse impression demarcating narrowed anterior lobe with flat collar about as wide as diameter of antennal segment 1, and steeply inclined, strongly swollen, broad posterior lobe; mesoscutum exposed, separated from scutellum by well defined transverse impression, inclined anteriorly; scutellum distinctly convex; clavus more or less inclined mesially to form ridge along claval commissure; cuneal incisure usually dis- tinct; membrane with two cells, the outer small, elongate, triangular, the inner large, rectangular, reaching to about apex of cuneus; legs long; tibiae with longitudinal rows of tiny, closely spaced, black 1974] Schuh: South African Orthotylinae and Phylinae 67 spines and scattered semierect spines about length of tibial diam- eter; tarsal claws long, smoothly curved; parempodia hair-like, par- allel; pulvilli minute. Male Genitalia: Figures 113-144. Genital capsule usually with a posteroventral spine; vesica strongly twisted, S-shaped; phal- lotheca L-shaped; left clasper trough -like, right clasper lanceolate. Brachypterous Female: See A. drakensbergensis and A. formicarium. Female Genitalia: Figures 145, 146. Sclerotized rings small, unusual in Hallodapini; posterior wall a simple sclerotized plate. Acrorrhinium can be separated from all other members of the Phylinae by the spiniform frons. Wagner (1970b) felt that Acrorrhinium was closely related to Aeolocoris and created a new tribe, the Aeolocorini, for these and other genera. I do not recognize this tribe, although as can be seen from the following discussion, Acrorrhinium and related genera do form a distinct group within the Hallodapini. Wagner based this relationship on the form of the male genitalia of Aeolocoris vidali (Wagner) and on external characters in A. vidali and Acrorrhinium conspersum, although he did not have access to the male genitalia of the latter species. The male genitalia of the Acrorrhinium spe- cies in South Africa show a specialized condition over those of Aeolocoris , at least as illustrated by Wagner (1970b), particu- larly in the apical region of the vesica which bears peculiar spine- like projections in Acrorrhinium and is simple in Aeolocoris. Other characters in Acrorrhinium , including the sclerotized rings in the female, and the spiniform frons, also suggest considerable special- ization. Even though the genus does possess a number of specialized characters, it does show its closest relationship to Aeolocoris and allied genera, including Azizus, Trichophorella, and Marmorodapus, based on the coloration pattern (which is often marmorate), the peculiar peg-like hairs (although these are much less common in Acrorrhinium than in other genera), and the virtual absence of hemelytral fasciae (present in some species of Acrorrhinium). Examination of the holotype female of Cinnamus rhinocerus Distant in the British Museum (Natural History), indicates that this species is not a member of the Cylapinae as indicated by Car- valho (1952a), but actually a species of Acrorrhinium , very closely related to A. pusae Ballard. The general body form, including the spiniform frons, and the pattern of coloration of rhinocerus are similar to A. pusae and A. lupa. I am therefore synonymizing Cinnamus Distant with Acrorrhinium Noualhier. 68 Entomologica Americana [Vol. 47 The South African species of Acrorrhinium can be divided into two rather distinct groups: the A. brincki group includes A. brincki, A. drakensbergensis , A. capensis, A. oudtshoornensis, and A. mon- ticola, all of which have castaneous markings on the corium as well as contrasting white hemelytral maculae (except in monticola, which is nearly unicolorous dull gray, although this may be a secondary loss of the more distinct color pattern found in the other species), dark unicolorous femora (see however brincki ), and a dark first antennal segment contrasting with the much lighter second segment; the A. muntingi group includes A. muntingi and A. incrassata (and possibly a third undescribed species represented by a single mac- ropterous specimen, lacking the abdomen, from Matjiesfontein, Cape Province, deposited in the British Museum [Natural History]), which are larger than the species of the brincki group, have the hemelytra rounded in transverse cross section, forming a humped appearance, and have femora and first antennal segments that are relatively light colored with numerous contrasting spots. Also the head is much smaller relative to the total body size in the muntingi group than in the brincki group. In South Africa, A. formicarium forms what is probably a third group, based on the rather anomalous structure of the female. Until males are known it will be difficult to assess the exact relationship of formicarium within the genus. Of the species of Acrorrhinium occurring outside South Africa, acutum Odhiambo, hebes Odhiambo, pauliana Carvalho, nilgiri- ensis (Distant), monoceros (Distant), lupa (Delattre), and spicatus (Distant) are all probably related to the brincki group; conspersus Noualhier, pusae (Ballard), and rhinocerus (Distant) seem to form another distinct group within the genus. Most of the known species of Acrorrhinium are from light traps; the females are unknown for four of the eight South African species and of the four species where females are known, only A. brincki includes macropterous specimens, and in this case no brachypterous specimens are known. Odhiambo (1959c) records macropterous females for A. acutum. The large series of A. muntingi taken at light with no females may indicate that the females are only rarely macropterous in this species, if at all, or that they are not attracted to lights. All species are probably ground living judging from those that have been observed in the field. The macropterous forms do not appear particularly ant mimetic when alive, but behavior of the brachypterous forms is very ant-like (see species discussion for A . oudtshoornensis) . 1974] Schuh: South African Orthotylinae and Phylinae 69 List of described species of Acrorrhinium acutum Odhiambo ( Acrorrhinium ), 1959c, p. 673. Kenya. brincki Carvalho and Becker ( Acrorrhinium ), 1960, p. 453. South Africa: Natal. capensis, new species. South Africa: Cape Province. conspersus Noualhier (Acrorrhinium) , 1895, p. 176. Ana- tolia; Syria. drakensbergensis, new species. South Africa: Natal; Transvaal; Lesotho. formicarium Poppius (Ectmetocranum) , 1914a, p. 36. South Africa: Cape Town. hebes Odhiambo ( Acrorrhinium ), 1959c, p. 676. Kenya. incrassata, new species. South Africa: Cape Province, Transvaal. lupa Delattre ( Seversyia ), 1950, p. 152. Ivory Coast: Bouake. monoceros Distant (Armachanus) , 1904c, p. 478. Ceylon. monticola, new species. South Africa: Cape Province. nilgiriensis Distant ( Armachanus ), 1909b, p. 60. South India: Nilgiri Hills. oudtshoornensis , new species. South Africa: Cape Province. pauliani Carvalho ( Acrorrhinium ), 1953a, p. 45. Madagascar. pusae Ballard ( Armachanus ), 1927, p. 67. North India: Bihar. rhinocerus Distant ( Cinnamus ), 1909a, p. 442. New Combination. Ceylon. spicatus Distant ( Armachanus ), 1904b, p. 203. North- western Australia. Key to South African species of Acrorrhinium Macropterous specimens 1. Metafemora with light ground color and numerous small brownish or reddish spots 2 Metafemora either entirely dark or dark distally and light proximally 3 2. Vertex and frons light with narrow, median, longitudinal reddish stripe; antennal segment 2 incrassate distally incrassata Vertex and frons mottled dorsally, without longitudinal reddish stripe; antennal segment 2 of uniform diameter muntingi 3. Metafemora light proximally and dark distally brincki Metafemora dark, unicolorous 4 70 Entomologica Americana [Vol. 47 4. Corium with reddish or castaneous areas strongly contrasting with white maculae; tibiae without dark dorsal stripe as below 5 Corium generally dull grayish brown with only weakly contrasting light maculae; tibiae with dark brown longitudinal dorsal stripe monticola 5. Corium with at least part of lateral margin dark (castaneous) (Fig. 23) 6 Corium with lateral margin (exocorium) light unicolorous drakensbergensis 6. Membrane with light halo-like areas between cunei; large species, length 6.40 mm.; ground color reddish brown capensis Membrane unicolorous, without halo-like area; small species, length 5.20 mm.; ground color dark castaneous oudtshoornensis (Fig. 23) Brachypterous specimens 1. Hemelytra strongly upturned apically; entire dorsum and legs cas- tanous, polished, covered with long, erect, light hairs formicarium Hemelytra flat; dorsum and legs dull, if castaneous or partly so, with only short, decumbent hairs 2 2. All tarsal segments dark brown oudtshoornensis All tarsal segments tan drakensbergensis (Fig. 24) Acrorrhinium brincki Carvalho and Becker Figures 113, 120-123 Acrorrhinium brincki Carvalho, Dutra, and Becker, 1960, pp. 453-454. Acrorrhinium brincki is very similar to A. drakensbergensis, described below, but differs from it in having the metafemora light proximally and dark distally; no other species in South Africa has this type of metafemoral coloration. This species was originally described from a macropterous female. Male specimens are now available and it is apparent that the structure of the two sexes is very similar. Measurements: Macropterous $ — Total length 5.92, greatest width 1.70. Male Genitalia: Figures 113, 120-123. This species is known from relatively high elevations along the Drakensberg Escarpment (circa 1875 meters; 6000 feet) and from the high veld of the Transvaal. No biological information is avail- able. Moderate numbers of male and female specimens were col- lected at an UV light in a light rain at Giants Castle Park, Natal, in early March, 1968. This is an area with a high proportion of macchia-related plants. 1974] Schuh: South African Orthotylinae and Phylinae 71 Specimens Examined: Natal — 1 macropterous 2, Royal Natal National Park, The Hostel, 3.IV.51, at light in evening (Brinck and Rudebeck) (holotype); 1 macropterous $, Royal Natal Na- tional Park, Tendele Camp, 5400 ft., 4-5 Mar. 1968, UV light; 1 macropterous 2, Natal National Park, iii.1932 (Ogilvie); 1 mac- ropterous S, Mont-aux-Sources, 4-6. IV. 1954 (Vari); 1 macrop- terous 11 macropterous 2 2, Giants Castle Park, 5800 ft. ele- vation, 6 Mar. 1968, UV light. Transvaal — 1 macropterous $, Lake Chrissie, 6 Nov. 1967; 1 macropterous 2, Pretoria, 6. IV. 1954, at light in evening (Rudebeck) (paratype) (SANC, TM, LU, BM [NH], JAS, RTS). Acrorrhinium capensis, new species Figures 116, 131-133 Macropterous Male: Basic coloration light mahogany; an- tennal segments 2 and 3 and all tibiae yellow; ostiolar peritreme, anterior half of corium, and quadrate macula at apex of corium adjacent to cuneal fracture white; elongate rectangular macula on endocorium just posterior to middle and contiguous with anterior white macula castaneous; membrane smoky brown with round, white, halo-like area between cunei; costal vein on corium at cuneal fracture orange; tarsi brown. Dorsal surface with scattered, short, decumbent, sericeous hairs; antennal segment 1 with decumbent dark hairs; antennal segments 2, 3, and 4 with dense, short vestiture; femora with scattered, short, dark, decumbent hairs. Eyes removed from anterior margin of pronotum by distance equal to one-third diameter of eye; vertex with weak transverse impression at level of posterior margin of eyes; spiniform frons not obscuring clypeus from above; antennal segment 1 moderately en- larged with several erect black spines on interior surface; antennal segments 3 and 4 slightly smaller in diameter than segment 2, seg- ment 2 about one-half diameter of segment 1; labium reaching to about abdominal sternite 3; posterior margin of pronotum evenly concave; tibiae with dark spines; metatarsal segments 1 and 2 sub- equal in length; segment 3 one-and-a-half times length of segment 2. Measurements: Total length 6.40, maximum width 1.84, length head .76, width head .82, interocular space .36, length pro- notum .62, width pronotum 1.32, length scutellum 1.04, width scutellum 1.08, length corium 3.04, length clavus 2.56, length cun- eus 1.00, width cuneus .62, length claval commissure 1.46, distance apex commissure-apex membrane 2.76, length metatibia 4.12; length 72 Entomologica Americana [Vol. 47 antennal segments 1 — .78, 2 — 2.76, 3 — 2.08, 4 — .96; length labial segments 1 — .74, 2 — .72, 3 — ?, 4 — ?. Male Genitalia: Figures 116, 131-133. Female unknown. Holotype: Macropterous S, south Africa: Cape Province, Knysna, Garden of Eden, 16-20.1.1955, A. J. T. Janse (TM). Paratype: Cape Province — 1 macropterous $, Cape Town, Kirstenbosch, 5-29.XII.1954 (Janse) (RTS). See key and A. oudtshoornensis discussion for separation of capensis from other South African species. Acrorrhinium drakensbergensis, new species Figures 24, 114, 124-127, 145, 146 Macropterous Male: General coloration dull brown; an- tennal segment 1, entire cuneus, procoxae and mesocoxae, and genital segment castaneous; elongate streak medially on clavus near claval suture and heavy quadrate macula submedially on endocorium deep mahogany; vertex between and behind eyes, anterior two-thirds of pronotum on either side of midline, antennal segment 1 proxi- mally, distal two-thirds of antennal segment 4, area around anten- nal fossae, streaked area above and below eyes posteriorly extend- ing onto pronotum, and distal margin of mesotrochanters and metatrochanters suffused with red; midline of pronotum, anterior half endocorium (anterior to large castaneous macula), exocorium generally, antennal segments 2 and 3 and proximal third of seg- ment 4, all tibiae and tarsi, and basal 2 segments of abdomen yel- lowish white; metacoxae gray to whitish; labium and profemora light brown; membrane with a round, halo-like, white suffused area between cunei. Entire body smooth, dull or only weakly shining; dorsum and thorax ventrally with scattered, short, decumbent, silvery hairs; an- tennae with very short, dense, shining hairs; coxae and femora with a few, scattered, semidecumbent hairs. Eyes removed from anterior margin of pronotum by distance equal to about one-third diameter of eye; vertex shallowly trans- versely sulcate at level of anterior margin of eyes; spiniform frons not obscuring clypeus from above; antennal segment 1 moderately enlarged, almost twice diameter of segment 2, segment 2 slightly greater in diameter than segments 3 and 4; labium reaching onto anterior third of abdomen; tibiae with a few very fine, light hairs and a few light spines mostly on ventral surfaces, about the length of tibial diameter; metatarsal segments subequal in length. 1974] Schuh: South African Orthotylinae and Phylinae 73 Measurements: Total length 4.96, maximum width ?, length head .66, width head .70, interocular space .30, length pronotum .56, width pronotum 1.20, length scutellum .72, width scutellum .90, length corium 2.34, length clavus 1.74, length cuneus .68, width cuneus .26, length claval commissure 1.00, distance apex commissure-apex membrane 2.18, length metatibia 3.40; length antennal segments 1 — .70, 2 — 2.32, 3 — 1.70, 4 — 1.00; length labial segments 1 — .58, 2 — .62, 3 — .56, 4 — .66. Male Genitalia: Figures 114, 124-127. Brachypterous Female: Head, thorax, and scutellum mostly yellowish; abdomen, procoxae and mesocoxae, all femora and la- bium deep brown with reddish suffusion; antennal segments 2 and 3 (and basal quarter of 4 — from paratype), metacoxae, and all tibiae and tarsi light yellowish; posterior portion of vertex, collar area of pronotum (and remainder of pronotum faintly), and most of scutellum on either side of light longitudinal midline (and distal three-fourths of antennal segment 4 — from paratype) reddish (also longitudinal reddish stripes at level of antennal fossae on genae, at dorsal and ventral margins of eyes on posterior portion of head and extending onto pronotum, and 2 longitudinal stripes on pro- epimeron); hemelytra mostly light transclucent, broadly suffused with brown along scutellum and claval commissure; small roundish mark at level of apex of scutellum on corium and large trapezoidal macula (with long extension from the anterior mesial corner) at level of claval commissure medially on corium, very dark brown; antennal segment 1 light brown. Body surface and vestiture similar to male; eyes with a few very short hairs. Structure of head similar to male, eyes smaller and vertex rela- tively wider than in male; labium reaching posterior margin of ab- dominal sternite 3; pronotum not strongly constricted anteriorly, nearly flat longitudinally; pronotal collar about as wide as diameter of antennal segment 1, well defined laterally, indistinct dorsally; posterior margin of pronotum shallowly and evenly concave; scutel- lum flattened longitudinally, weakly convex transversely; hemelytra greatly reduced, undifferentiated, reaching middle of abdominal tergite 4; posterior margin of hemelytra evenly rounded beginning at claval commissure, slightly upturned; abdomen broad medially (no gravid specimens examined but the abdomen presumably be- comes swollen and bulbous), pointed at anus; tibiae as in male. Measurements: Total length 4.40, maximum width ?, length head .76, width head .70, interocular space .40, length pronotum 74 Entomologica Americana [Vol. 47 .60, width pronotum .84, length scutellum .56, width scutellum .63, length hemelytron 1.40, length metatibia 3.68; length antennal seg- ments 1 — .80, 2 — 2.56, 3 — 2.00, 4 — 1.16; length labial segments 1 — .70, 2— .56, 3— .70, 4— .62. Female Genitalia: Figures 145, 146. Holotype: Macropterous S, south Africa: Natal, Royal Natal Nat. Pk., Tendele Camp, 5400 ft., 4-5 Mar. 1968, T. Schuh, J. A. & S. Slater, M. Sweet, UV Light Trap (SANC). Paratypes: Cape Province — 1 macropterous $ , Knysna, Keur- booms River, Jan. 1931 (Barnard). Natal — 1 macropterous $, same data as holotype; 1 bracypterous 9, idem, but not at UV light; 1 macropterous $, Howick; 1 macropterous S, P. Shepstone, 5.97. Transvaal — 1 macropterous S, Blouberg, Motlakeng, 5-6000 ft., 6-15.1.1955; 1 macropterous $, Pretoria, 22.12.1910 (Swierstra). Lesotho — 3 brachypterous 9 9, Sani Pass, 8000 ft., 10 Mar. 1968 (SANC, TM, BM[NH], JAS, RTS). This species is named for its occurrence on the Drakensberg. Acrorrhinium drakensbergensis appears to be most closely re- lated to A. brincki, A. capensis , and A. oudtshoornensis. The uni- colorous metafemora will separate it from brincki, the light uni- colorous corial margin from oudtshoornensis and capensis. The brachypterus females of drakensbergensis are very similar to those of oudtshoornensis but can be separated as in the key. In the series of males examined the length of the labium varies from just surpassing the metacoxae to reaching about one-third the length of the abdomen. The three brachypterous females from Lesotho, Sani Pass, were collected in association with Chrysocoma tenuifolia Berg. (Compositae). Acrorrhinium formicarium (Poppius) Ectmetocranum formicarium Poppius, 1914a, p. 37. Acrorrhinium formicarium is one of the most specialized spe- cies in the genus and to date is known only from the brachypterous female. The structure of the head is the only feature that obviously relates this species to the other members of the genus. The clypeus is strongly flattened and greatly produced and the frontal spine is proboscis-like. The entire body and all of the appendages are cas- taneous, highly polished, and covered with long, erect, light-colored hairs. The posterior margin of the pronotum is upturned. The scu- tellum is slightly elevated. The hemelytra are upturned just past the apex of the scutellum and form two points nearly as high as 1974] Schuh: South African Orthotylinae and Phylinae 75 the dorsum of the bulbous abdomen. The tibiae lack the longitu- dinal rows of tiny spines found in all other species of Acrorrhinium. The figure of A. formicarium in Poppius (1921) is basically ac- curate. When the male of this species is known it will be much easier to assess its relationship to other species in the genus. Poppius (1914a) stated that the two type females of A. formi- carium were deposited in the Paris Museum. In fact, none are in Paris, but at least one is in the Helsinki Museum, and I am desig- nating it as the lectotype. It bears the labels: “Museum Paris, Cape-Town, E. Simon, Coll. Noualhier 1898”; '‘Ectmetocranum formicarium n. gen. et sp. B. Poppius det.”; “Mus. Zool. H:fors, Spec. typ. No. 7786, Ectmetocranum formicarium Popp.”; and “LECTOTYPE Ectmetocranum formicarium Poppius, det. R. T. Schuh.” Acrorrhinium incrassata, new species Figures 119, 138-140 Macropterous Male: General coloration light brownish yel- low with the following dull red markings: narrow stripe on dorsal midline of head (interrupted at level of posterior margin of eyes), narrow stripe on either side of midline of anterior lobe of pronotum, posterior third of scutellum on either side of midline, two lines on head at level of dorsal margin of eyes and just below dorsal margin of eyes running anteriorly from eyes to antennal fossae and pos- teriorly to pronotum, line near ventral margin of eyes between eye and pronotum, lower margin of juga, two parallel lines on entire lateral margin of pronotum, numerous small spots on all femora (particularly mesofemora and metafemora), and suffused areas at apex of clavus and corium at cuneal fracture; posterior lobe of pronotum, most of scutellum, elongate area on corium along claval suture at level of midpoint of claval suture, venter of mesothorax, and most of genital segment black; antennal segment 2 distally, antennal segments 3 and 4, and all tarsi dark brown; irregular mark- ing on corium contiguous laterally with elongate black area de- scribed above and diffuse marking on clavus at same level white; much of corium and cuneus suffused with brown, veins lighter; membrane light yellow gray; pronotum and scutellum with distinct yellow midline dorsally. Entire body smooth, dull; dorsum with a few scattered, decum- bent, very short, silvery hairs antennal segments 2, 3, and 4 with dense, short, shining vestiture; tibiae and tarsi with short, dull hairs. Eyes removed from anterior margin of pronotum by distance 76 Entomologica Americana [Vol. 47 equal to about one-third diameter of eye; spine-like projection of frons very short; antennal segment 1 only very slightly enlarged, segment 2 increasing in diameter distally to same diameter as seg- ment 1, segments 3 and 4 about same diameter as segment 2 prox- imally, about two-thirds diameter of segment 1; labium attaining distal end of metacoxae; pronotal collar about equal in width to diameter of antennal segment 3; posterior margin of pronotum shal- lowly concave; femora with a few, very fine, erect spines; tibiae with scattered dark spines about equal in length to tibial diameter; meta- tarsal segments subequal in length. Measurements: Total length 7.52, maximum width 2.24, length head .90, width head .90, interocular space .40, length pro- notum .80, width pronotum 1.60, length scutellum 1.10, width scu- tellum 1.30, length corium 3.60, length clavus 2.68, length cuneus 1.24, width cuneus .56, length claval commissure 1.60, distance apex commissure-apex membrane 3.40, length metatibia 4.32; length antennal segments 1 — 1.02, 2 — 2.72, 3 — 1.96, 4 — 1.02; length labial segments 1 — .78, 2 — .84, 3 — .64, 4 — .74. Male Genitalia: Figures 119, 138-140. Female unknown. Holotype: Macropterous S, south Africa: Cape Province, Grootfontein, Middelburg, 4. XI. 65, E. Schoombee (SANC). Paratypes: 1 macropterous $, same data as holotype, except 3. XII. 1965. Transvaal — 1 macropterous S, Zomerkomst, Politzi, 23.X.64 (Johannsmeier) (SANC, RTS). This species is named for the incrassate second antennal segment. See discussion under A . muntingi for distinguishing characteristics. Acrorrhinium monticola, new species Figures 117, 134-137 Macropterous Male: General coloration dull gray-brown; anterior lobe of pronotum, scutellum (except as noted below), cu- neus, thoracic pleura and venter, abdominal venter, and labial seg- ments 2, 3, and 4 dark brown; medium brown as follows: dorsal surface of frons, much of vertex between eyes posteriorly, posterior lobe of pronotum (except as below), diffuse area medially on co- rium, antennal segment 1, and all femora; white as follows: clypeus, lora, metathoracic scent gland opening, longitudinal stripe on ex- treme posterior pronotal lobe medially and midlaterally (midlateral stripes not present in paratype), longitudinal stripe on posterior lobe of scutellum, ovoid spot just anterior to brown area medially on corium, and apex of corium at cuneus; antennal segments 2 and 1974] Schuh: South African Orthotylinae and Phylinae 77 3 yellow- white, segment 2 white basally; tibiae cream with contrast- ing longitudinal dorsal stripe; tibiae distally, and tarsi black. Entire body smooth, dull; dorsum with scattered (almost scale- like), short, decumbent, sericeous hairs; antennal segment 1 with short, dark, decumbent hairs; femora with scattered short hairs; tibiae with a few semidecumbent hairs distally. Eyes removed from anterior margin of pronotum by distance equal to about one-third diameter of eye; spine-like process of frons short, blunt; antennal segment 1 moderately enlarged, with several erect, black spines on interior surface, segments 2 and 3 of equal diameter, about half diameter of segment 1 (segment 4 missing); labium reaching posterior margin of abdominal sternite 5 (from paratype); posterior pronotal margin straight; femora with a few erect, thin, black spines mostly on dorsal surface; tibiae with scat- tered dark spines with dark bases; metatarsal segments subequal in length. Measurements: Total length 6.80, maximum width ?, length head .88, width head .90, interocular space .36, length pronotum .74, width pronotum 1.36, length scutellum 1.10, width scutellum .96, length corium 3.28, length clavus 2.52, length cuneus 1.16, width cuneus .80, length claval commissure 1.48, distance apex commissure-apex membrane 2.76, length metatibia 4.36; length antennal segments 1 — 1.00, 2 — 2.52, 3 — 2.08, 4 — ?; length labial segments 1 — .80, 2 — .84, 3 — .68, 4 — .80. Male Genitalia: Figures 117, 134-137. Female unknown. Holotype: Macropterous S, south Africa: Cape Province, Bainskloof Pass Summit, 21 Jan. 1968, J. A. & S. Slater, T. Schuh, M. Sweet (SANC). Paratype: 1 macropterous $, same data as holotype, but at UV light (RTS). This species is named for its occurrence in the mountains of the Southwest Cape. A. monticola is distinctive among the South African species of Acrorrhinium by virtue of its dull gray coloration and the dark brown stripe on the dorsal surface of the tibiae. Acrorrhinium muntingi, new species Figures 118, 141-144 Macropterous Male: General coloration light yellowish brown, with dark brown markings as follows: vertex between eyes irregularly, distinct transverse lines anteriorly on vertex (behind 78 Entomologica Americana [Vol. 47 spine), most of pronotum on either side of light midline, scutellum broadly around anterior and lateral margins, hemelytra suffused mostly on clavus and cuneus, labial segments 2, 3, and 4, thorax ventrally, abdomen ventrally (particularly anterior two-thirds of genital segment), antennal segment 1, and numerous small round spots on all femora; tibial spines with dark bases; tarsi black; mid- line of posterior lobe of scutellum suffused with red; pregenital ab- dominal segments lighter than most of venter and suffused with green. Body smooth, weakly shining; dorsum with short, scattered, decumbent, silvery hairs; antennal segment 1 with dark decumbent hairs; antennal segments 2, 3, and 4 with short, very dense, shining vestiture; femora with scattered short hairs. Eyes removed from anterior margin of pronotum by distance equal to about one-third diameter of eye; spine of frons very short; antennal segment 1 slightly greater in diameter than segment 2 (only segments 1 and 2 present in holotype); labium just surpassing meta- coxae; posterior margin of pronotum slightly concave; femora with a few moderately long very thin spines; tibiae with scattered dark spines about length of tibial diameter; metatarsal segments subequal in length. Measurements: Total length 7.68, maximum width 2.00, length head .78, width head .92, interocular space .36, length pro- notum .76, width pronotum 1.52, length scutellum 1.10, width scu- tellum 1.24, length corium 3.68, length clavus 2.84, length cuneus 1.40, width cuneus .54, length claval commissure 1.72, distance apex commissure-apex membrane 3.64, length metatibia 4.44; length antennal segments 1 — 1.06, 2 — 2.88, 3 — ?, 4 — ?; length labial seg- ments 1 — .82, 2 — .82, 3 plus 4 — 1.54. Male Genitalia: Figures 118, 141-144. Female unknown. Holotype: Macropterous S, south Africa: Cape Province , Grootfontein, Middelburg, 3. XII. 1965. E. Schoombee (SANC). Paratypes: 8 macropterous $ S, same data as holotype; 2 mac- ropterous $ $ , same data as holotype, but “October, M. Johanns- meier” (SANC, J AS, RTS) . This species is named for Mr. Jack Munting, formerly Curator of the South African National Collection of Insects, Pretoria. A. muntingi and A. incrassata can be separated from the brincki group as indicated in the generic discussion. A. muntingi has a spotted first antennal segment, a second antennal segment of uni- form diameter, and a mottled vertex, whereas incrassata has the 1974] Schuh: South African Orthotylinae and Phylinae 79 first antennal segment spotted but the second segment is enlarged distally; the vertex of incrassata is light, marked only with a median longitudinal red stripe. Acrorrhinium oudtshoornensis, new species Figures 23, 115, 128-130 Macropterous Male: General coloration dark gray-brown, suffused with red; antennal segments 2 and 3 (4 missing in holotype) and tibiae light yellowish gray; anterior third of corium and quad- rate macula at apex of corium along cuneal fracture white; small area on corium at apex claval commissure suffused with white; quad- rate macula on endocorium just posterior to and contiguous with white anterior portion of corium velvety castaneous; tarsi almost black; membrane dull gray-brown. Entire body smooth, dull; abdominal venter weakly shining; dorsum with scattered, short, decumbent hairs; antennal segment 1 and femora with scattered, dark, decumbent hairs; antennal seg- ments 2 and 3 with dense, short, shining vestiture. Eyes removed from anterior margin of pronotum by distance equal to one-half diameter of eye; vertex with weak, longitudinal, median sulcus between eyes; spiniform frons conical, not obscur- ing clypeus from above; antennal segment 1 moderately enlarged, almost twice diameter of segment 2, segment 2 of slightly greater diameter than segment 3; labium attaining distal end of metacoxae; posterior margin of pronotum very shallowly concave; tibiae and tarsi with scattered dark spines with dark bases; metatarsal seg- ments subequal in length. Measurements: Total length 5.20, maximum width 1.38, length head .66, width head .70, interocular space .36, length pro- notum .60, width pronotum 1.16, length scutellum .78, width scu- tellum .92, length corium 2.40, length clavus 1.86, length cuneus .72, width cuneus .44, length claval commissure 1.06, distance apex commissure-apex membrane 2.20, length metatibia 3.32; length antennal segments 1 — .58, 2 — 2.00, 3 — 1.54, 4 — ?; length labial segments 1 — .64, 2 — .64, 3 — .40, 4 — .60. Male Genitalia: Figures 115, 128-130. Brachypterous Female: Antennal segment 1, abdomen, pro- coxae and mesocoxae, and all femora dark gray brown suffused with red; most of head, pronotum, scutellum, and claval area of hemelytra yellowish gray; corial region of hemelytra, antennal segments 2 and 3, tibiae, and metacoxae yellow or yellow white; antennal segment 80 Entomologica Americana [Vol. 47 4 red; tarsi black; corium medially with heavy velvety black macula and an anteriorly contiguous oval white macula. Body surface texture and vestiture as in male; eyes with a few very short hairs. Structure very similar to that of brachypterous female of A. drakensbergensis; metatarsal segment 2 about half length of seg- ment 3, segment 1 subequal in length to segment 3. Measurements: Total length 4.48, maximum width ?, length head .88, width head .80, width vertex .42, length pronotum .64, width pronotum .88, length scutellum .64, width scutellum .70, length hemelytra 1.84; length antennal segments 1 — .80, 2 — 2.52, 3 — 1.80, 4 — .92; length labial segments 1 — .70, 2 — .76, 3 — .66, 4— .78. Female Genitalia: See generic discussion. Holotype: Macropterous $, south Africa: Cape Province , Swartberg Pass, elevation 5000 ft., 25 mi. N. of Oudtshoorn, Plat- berg, 19 Nov. 1967, M. H. Sweet (SANG). Paratypes: Cape Province — 1 macropterous $, same data as holotype; 1 macropterous S, 3 brachypterous $ $, just N. Outi- niqua Pass Summit S. of Oudtshoorn, 7 Feb. 1968 (SANC, JAS, RTS). Additional Specimens: 3 nymphs (in alcohol), just N. Outi- niqua Pass Summit S. of Oudtshoorn, 7 Feb. 1968 (RTS). This species is named for Oudtshoorn, the district in which all known specimens have been collected. A. oudtshoornensis is most closely related to A. capensis , but is much smaller, much darker in coloration, and lacks the light halo- like area on the membrane. The Outiniqua Pass specimens of oudtshoornensis were col- lected under Helichrysum orbiculare (Thunb.) Druce (Compositae) in association with workers of the ant Anoplolepis sp. The females of this species are very ant-like in their movements. Azizus Distant Azizus Distant, 1910a, p. 11. Azizus is probably most closely related to Aeolocoris. It can be recognized by the large eyes of the males, the well defined pro- notal collar, the mottled coloration, and the erect peg-like hairs on the dorsum and first antennal segment. The genus as presently constituted may bring together species that actually belong in dif- ferent genera. 1974] Schuh: South African Orthotylinae and Phylinae 81 The type species of the genus, A . basilicus Distant, is from India, whereas all other known species are from Africa. List of described species of Azizus basilicus Distant (Azizus), 1910a, p. 11. India: Bengal. basilewskyi Carvalho (Azizus), 1951b, p. 110. Congo. * dispar Odhiambo (Azizus), see Hallodapus dispar (Od- hiambo) New Combination. oculatus Poppius (Megacoeloides) , 1914a, p. 33. Togo; Swaziland. Azizus near oculatus (Poppius) Megacoeloides oculatus Poppius, 1914a, p. 33. A single male specimen with the data “Eranchi, Swaziland, XII- 1954, Capener” is in the J. A. Slater Collection. Comparison of this specimen with type material of A . oculatus (see below) in- dicates that it is congeneric, if not conspecific, with oculatus. Un- fortunately the male genitalia of the Swaziland specimen are very lightly sclerotized, probably because of its teneral condition, and therefore a critical comparison could not be made with other speci- mens of oculatus. A male and female of Azizus oculatus from Togo, apparently part of Poppius’ type series, are in the Helsinki Museum. Neither of these specimens appear to be the “type” of Poppius, but I am informed that it is also in Helsinki (personal communication, Martin Meinander). On subsequent examination of all specimens origi- nally studied by Poppius a lectotype will have to be designated. Carinogulus, new genus Macropterous Male: Elongate, ant mimetic; head, pronotum, and scutellum granular or finely rugulose, weakly shining; posterior third of corium, apex of clavus, cuneus, cell of membrane, and ad- jacent membrane at base of cell highly polished, shining; legs and venter polished, shining; dorsum with short, decumbent, light hairs and sometimes scutellum, corium, and clavus also with a few long erect hairs; eyes glabrous; antennae with very short, light, appressed pubescence; labium with a few, short, erect hairs; femora with short, decumbent, light hairs, sometimes metafemora with longer, erect, rather dense hairs on ventral surface; tibiae with short decumbent, light hairs, and semierect, fine, light spines about as long as tibial diameter; abdominal venter with reclining light hairs. Head strongly deflexed, either concave behind, or narrowed 82 Entomologica Americana [Vol. 47 1974] Schuh: South African Orthotylinae and Phylinae 83 behind eyes to form a short neck about as long as diameter of an- tennal segment 2 and as wide as interocular space; frons nearly vertical; eyes granular, large, reaching ventrally almost to gula; vertex raised above level of pronotal collar by at least diameter of antennal segment 2; vertex weakly convex or sometimes slightly depressed between eyes; frons transversely rugose; anterior margins of eyes sinuate; antennae inserted slightly above ventral margins of eyes; antennal fossae contiguous with anterior margins of eyes; an- tennal segment 1 slightly enlarged, segment 2 tapering proximally, distal diameter about equal to diameter of segment 1, segments 3 and 4 subequal to or slightly greater in diameter than distal end of segment 2; clypeus rounded transversely, prominent, separated from frons by distinct cleft; bucculae moderately expanded, crescent- shaped; gula moderately long, inclined, with distinct carina run- ning from posterior margin of buccal cavity to posterior margin of head; pronotum tumid and strongly inclined posteriorly, strongly narrowed anteriorly, lateral margins nearly straight, posterior mar- gin broadly rounded laterally, calli indistinct, defined only by slightly roughened surface texture; pronotal collar wide, flat; meso- scutum exposed, inclined anteriorly, steep-sided laterally; scutellum tumid, reaching dorsally to about height of pronotum; lateral corial margins reflexed ventrally on anterior half, distinctly sinuate, nar- rowest at level of midpoint of claval commissure; cuneal incisure obsolete; membrane with 2 cells; abdomen long, very narrow; femora nearly round, slightly enlarged in diameter apically, weakly bowed; tibiae with longitudinal rows of tiny, closely spaced spines; tarsal claws evenly curved, moderately long; parempodia hair-like, par- allel; pulvilli minute. Male Genitalia: Figures 147-159. Vesica twisted, S-shaped; apex of vesica weakly or strongly attenuated; phallotheca either with “thumb-like” structure near basal opening (Figures 156, 159) or without “thumb” and with apex of complex structure (Figures 148, 152); left clasper trough-like; right clasper lanceolate. Females unknown. Type Species: Carinogulus varii , new species. This genus is named for the distinctive carinate gula. Figs. 197-219. Leucophoropterini, Karoocapsus male genitalia. Fig. 197. Lateral view of vesica, K. bifasciatus. Fig. 198. Phallotheca, idem. Fig. 199. Left clasper, idem. Fig. 200. Lateral view of vesica, K. brunneus. Fig. 201. Phallotheca, idem. Fig. 202. Left clasper, idem. Fig. 203. Lateral view of vesica, K. middelburgensis. Fig. 204. Phallotheca, idem. Fig. 205. Left clasper, idem. Fig. 206. Lateral view of vesica, K. flavomaculatus. Fig. 207. Phallotheca, idem. Fig. 208. Lateral view of vesica, K. obscurus. Fig. 209. Phallotheca, idem. Fig. 210. Left clasper, idem. Fig. 211. Lateral view of vesica, K. occidentalis. Fig. 212. Phallotheca, idem. Fig. 213. Left clasper, idem. Fig. 214. Lateral view of vesica, K. pulchrus. Fig. 215. Phal- lotheca, idem. Fig. 216. Left clasper, idem. Fig. 217. Lateral view of vesica, K. trifasciatus. Fig. 218. Phallotheca, idem. Fig. 219. Left clasper, idem. 1974] Schuh: South African Orthotylinae and Phylinae 227 228 Entomologica Americana [Vol. 47 of the low and midaltitude regions of South Africa, although the Kalahari, Karoo, and South West Africa have not been well enough collected to know if it occurs there. Before we began our collecting in 1967-1968 only 52 specimens from 12 localities were to be found in all available collections. It is now known that australis is one of the most common and widespread species of Phylinae in South Africa. Known host plants for this species include Scirpus dioecus Boeck., Scirpus costatus Boeck., Cyperus rotundus L., Cyperus ladfolius Pair., and Cyperus distans L. (see also discussion under P. suturalis). Paramixia suturalis Reuter Figures 92, 335 Paramixia suturalis Reuter, 1900, p. 264. — Lindberg, 1958, p. 105. — Carvalho, 1958a, p. 86. — Linnavuori, 1961, p. 23. — Wagner, 1970a, pp. 4-5. Troitskiella minuta Poppius, 1914a, pp. 81-82. New Synonymy. Paramixia suturalis can be recognized by the following characters: basic coloration usually light yellow green, some specimens medium brown; antennal segment 1 entirely brownish black; metatibial spines without dark bases, tibiae unicolorous tan; and, vesica C-shaped, but rather sharply bent subapically, apical section “wavy” (Figure 335). The male genitalia of specimens from South Africa agree very closely in structure with those of specimens from the Cape Verde Islands (Lindberg, 1958) and the Nile Valley. Wagner (1970a) redescribed Paramixia suturalis and provided illustrations. As noted above, Wagner’s (1970a) interpretation of the structure of the parempodia in Paramixia was incorrect and therefore his illustrations of these structures are not accurate. Wag- ner examined 3 specimens of suturalis from the Helsinki Museum, from which Reuter (1900) apparently originally described the species, but did not designate a lectotype. I have selected a male bearing the following labels as the lectotype: “Vail, Nil.”; “spec, typ. Reuter”; “Mus. Zool. H:fors, Spec. typ. No. 3443, Paramixia Figs. 220-225. Phylini, Austropsallus male genitalia. Fig. 220. Lateral view of vesica, Austropsallus drakensbergensis. Fig. 221. Phallotheca, idem. Fig. 222. Left clasper, idem. Fig. 223. Lateral view of vesica, Austropsallus middelburgensis. Fig. 224. Phallotheca, idem. Fig. 225. Left clasper, idem. 1974] Schuh: South African Orthotylinae and Phylinae 229 230 Entomologica Americana [Vol. 47 suturalis Reut.”. I have also added the label “LECTOTYPE Para- mixia suturalis Reuter, det. R.T. Schuh.” Examination of the holotype of Troitskiella minuta Poppius indicates that the coloration of this species fits well within the range of variation found in P. suturalis and that the external morphology and structure of the male genitalia are identical with suturalis from the Nile Valley and from South Africa. I am therefore synonymizing Troitskiella minuta Poppius with Paramixia suturalis Reuter. Paramixia suturalis is a primarily tropical species occuring in East and West Africa and extending northward into the southern Mediterranean and southward into southern Africa. In South Africa this species is restricted to the coastal region of Natal, the lowveld of the eastern Transvaal, and from Pretoria north into the interior of the Transvaal. Lindberg (1958) recorded suturalis as occurring on Cyperaceae and Gramineae, and specifically on Cynodon dactylon. In South Africa it breeds on J uncus kraussii Hochst., Scirpus costatus Boeck., and Cyperus latifolius Pair. In nearly all situations where I collected suturalis in South Africa it was living in association with Paramixia australis, Cymodema basicornis (Motschulsky), Cymodema tabida Spinola, and less commonly with Cymus waelbroecki and an addi- tional undescribed species of Cymus. Specimens Examined: Natal — 1 2, Eshowe, 15 Nov. 1967; 1 8, 3 2 2, Mtunzini Plantation, 5 mi. N. Mtunzini, 15 Nov. 1967 (Adults and nymphs on Scirpus costatus Boeck.); 11 8 8, 17 2 2, St. Lucia Estuary, 14 Nov. 1967 (Adults and nymphs on J uncus kraussii Hochst. and Cyperus latifolius Pair. ). Transvaal — 1 2 , Hare- beespoort Dam, 20 mi. W. Pretoria, 30 Oct. 1967; 6 8 8, 2 2 2, Little Sabie River, Sabie, 29 Nov. 1967; 1 2, 17 mi. N. Louis Trichardt, 14 Dec. 1967; 1 8, Kruger Nat. Park, confluence of Limpopo and Luvuvho Rivers, Pafuri, 7 May 1968; 1 <3, 1 nymph, Kruger Nat. Park, 8 mi. W. Shingwidzi, Shingwidzi River, 4 May 1968; 1 2, top Magoebaskloof, 6000 ft., 12 Dec. 1967 (SANC, HM, BM[NH], JAS, RTS). Figs. 226-234. Phylini, Austropsallus male genitalia. Fig. 226. Lat- eral view of vesica, A. albonotum. Fig. 227. Phallotheca, idem. Fig. 228. Lateral view of vesica, A. senecionus. Fig. 229. Lateral view of vesica, A. saniensis. Fig. 230. Apex of vesica, idem. Fig. 231. Phal- lotheca, idem. Fig. 232. Lateral view of vesica, A. helichrysi. Fig. 233. Phallotheca, idem. Fig. 234. Right clasper, idem. 1974] Schuh: South African Orthotylinae and Phylinae 231 232 Entomologica Americana [Vol. 47 Pilophorus Hahn Pilophorus Hahn, 1826, p. 22. This very large genus is poorly represented in Africa, with only one known species. Most described members of the genus are Nearctic and Palearctic, but a large undescribed fauna is present in Southeast Asia. No species of Pilophorus are known from the Neotropical Region. Pilophorus pilosus Odhiambo Pilophorus pilosus Odhiambo, 1958b, pp. 326-330. Pilophorus pilosus can be separated from all other South African Miridae by the following combination of characters : the parempodia are fleshy, convergent apically, and recurved; and the dorsum is black with narrow, transverse bands of scale-like, sericeous hairs on the clavus and corium. Odhiambo (1958b) in the original description of this species, provided excellent illustration of the male genitalia and the hemelytra. The type specimens of pilosus from Uganda agree very closely with all specimens known from South Africa. No host or ecological information is available for pilosus. Specimens Examined: Natal — 4 macropterous macrop- terous 2, Port Shepstone, 5.97. Transvaal — 1 2, Letaba Valley, Tzaneen Dist., XII-20-1962 (Capener) (BM[NH], JAS, RTS). ZOOGEOGRAPHY No attempts have been made to analyze the zoogeography of the Miridae of southern Africa. In the following discussion I have tried to bring together those factors which seem to be most important in influencing the distributional patterns of the Orthotylinae and Phylinae in this region. Figs. 235-247. Phylini male genitalia. Fig. 235. Lateral view of vesica, Capecapsus tradouwensis. Fig. 236. Phallotheca, idem. Fig. 237. Left clasper, idem. Fig. 238. Lateral view of vesica, Coatonocapsus transvaalensis. Fig. 239. Left clasper, idem. Fig. 240. Right clasper, idem. Fig. 241. Lateral view of vesica, Coatonocapsus johannsmeieri. Fig. 242. Lateral view of vesica, Coatonocapsus pallidus. Fig. 243. Lateral view of vesica, Coatonocapsus sweeti. Fig. 244. Phallotheca, idem. Fig. 245. Lateral view of vesica, Denticulo phallus adenandrae. Fig. 246. Phallotheca, idem. Fig. 247. Left clasper, idem. 1974] Schuh: South African Orthotylinae and Phylinae 233 234 Entomologica Americana [Vol. 47 The Miridae as a group are largely phytophagous, or if predatory they are often host specific on a given species of phytophagous insect. Thus, one would expect the distributions of host plants to be im- portant in considering the distributions of mirids. Aside from edaphic factors, it seems that precipitation is the most important single factor affecting plant distributions (Stuckenberg, 1969) and there- fore it must be important in determining the distributions of the Miridae in at least an indirect way. As pointed out by Stuckenberg (1969), the South African flora can be divided into three basic components which are roughly cor- related with the amounts of rainfall and the rainfall regime. These are (1) the “Karoo” and (2) the “tropical and subtropical forest and grassland,” which receive most of their precipitation during the summer months, and (3) the “macchia” which receives most of its rainfall during the winter months. Sufficient data do not exist at present to allow for a careful correlation of the distribution of the mirid fauna with the floral types of South Africa, but certainly a strong affinity does exist at least between the endemic Orthotylinae and Phylinae and the almost totally endemic floras of the Southwest Cape (primarily macchia) and the Karoo (see following analyses). Recently Stuckenberg (1969) has applied H.P. Bailey’s (1960) concept of “effective temperature” (denoted as “ET” in the fol- lowing text) to South Africa and shown that the distributions of snakes and amphibians agree remarkably well with this measure of effective solar radiation. Stuckenberg (1969) points out that in using an effective temperature analysis the biologies of the animals under consideration must be taken into account, especially for those Figs. 248-268. Phylini male and female genitalia. Fig. 248. Lateral view of vesica, Ellenia obscuricornis. Fig. 249. Phallotheca, idem. Fig. 250. Left clasper, idem. Fig. 251. Lateral view of vesica, Emi- noculus drosanthemi. Fig. 252. Phallotheca, idem. Fig. 253. Left clasper, idem. Fig. 254. Sclerotized rings, idem. Fig. 255. Lateral view of vesica, Lamprosthenarus near sjostedti. Fig. 256. Phallo- theca, idem. Fig. 257. Left clasper, idem. Fig. 258. Lateral view of vesica, Lasiolabopella capeneri. Fig. 259. Phallotheca, idem. Fig. 260. Left clasper, idem. Fig. 261. Lateral view of vesica, Lepido- capsus rubrum. Fig. 262. Phallotheca, idem. Fig. 263. Left clasper, idem. Fig. 264. Lateral view of vesica, Macrotylus niger. Fig. 265. Left clasper, idem. Fig. 266. Lateral view of vesica, Macrotylus hemizygiae. Fig. 267. Phallotheca, idem. Fig. 268. Left clasper, idem. 1974] Schuh: South African Orthotylinae and Phylinae 235 236 Entomologica Americana [Vol. 47 animals that have an ability to control their environment. Certain taxa in the Orthotylinae and Phylinae (e.g. Paramixia and Pangania) are obviously restricted by factors other than host plant distributions in that the distributions of the mirid taxa are limited in South Africa, whereas the hosts are much more wide-ranging. This would suggest then, that distributions of at least certain of the Miridae are amenable to an “effective temperature” analysis. In those cases where adequate distributional data are available a positive correlation does seem to exist (see below). Consideration of the distributions of the Orthotylinae and Phylinae in relation to the long term evolution of the fauna in South Africa is difficult because no fossils exist by which it is possible to determine dates of origin. Even though it is not possible to determine the antiquity of the South African fauna, certain strong correlations appear to exist between the distributions of the Orthotylinae and Phylinae (especially the endemic elements) and the South African paleogenic element. Stuckenberg (1962) recognized two centers of paleogenic en- demicity in South Africa — the Cape Center and the Eastern High- lands Center. In the Orthotylinae and Phylinae group “la” (see below) in the endemic fauna fits closely with Stuckenberg’s Cape Center and group “lb” with his Eastern Highlands Center. Stucken- berg (1962) felt that the Eastern Highlands Center was the node from which many paleogenic groups have spread. He based this conclusion on the ancient nature of the region, which was formed by the pre-Cretaceous Stormberg lava series, and the fact that much of the Cape Center was drowned by Cretaceous seas. The greater Figs. 269-288. Phylini male genitalia. Fig. 269. Lateral view of ve- sica, Natalophylus heteromorphae. Fig. 270. Apex of vesica, idem. Fig. 271. Phallotheca, idem. Fig. 272. Left clasper, idem. Fig. 273. Lateral view of vesica, Odhiamboella solani. Fig. 274. Phallotheca, idem. Fig. 275. Left clasper, idem. Fig. 276. Lateral view of vesica, Parasciodema albicoxa. Fig. 277. Phallotheca, idem. Fig. 278. Left clasper, idem. Fig. 279. Lateral view of vesica, Parasciodema nigri- femur. Fig. 280. Lateral view of vesica, Stoebea barbertonensis. Fig. 281. Phallotheca, idem. Fig. 282. Left clasper, idem. Fig. 283. Lateral view of vesica, Stoebea elginensis. Fig. 284. Apex of vesica (obverse view), idem. Fig. 285. Lateral view of vesica, Stoebea plet- tenbergensis. Fig. 286. Lateral view of vesica, W iddringtoniola kir- stenboschiana. Fig. 287. Phallotheca, idem. Fig. 288. Left clasper, idem. 1974] Schuh: South African Orthotylinae and Phylinae 237 238 Entomologica Americana [Vol. 47 diversity and morphological specialization of the Orthotylinae and Phylinae in the Cape Center suggests that area as containing the most isolated elements in the South African fauna, and thus possibly being the evolutionary center for these taxa. Recent evidence from continental drift suggests that Africa has moved northward (Dietz and Holden, 1970) and therefore the region of Mediterranean climate has probably been reduced. Also, Pleistocene pluvials probably created an increased area of Mediter- ranean climate and allowed for northward expansion of the flora and fauna that is now compressed into the Southwest Cape (see van Zinderen Bakker, 1967). Both of these factors may thus be important in the evolutionary history of the Cape Center and help to explain the tremendous diversity that is found there as well as to eliminate, at least in part, the need for “explosive local evolution” as proposed by Stuckenberg (1962). Although certain groups of plants and animals have transantarctic distributions including South Africa, no evidence exists for such a pattern within the Orthotylinae and Phylinae except possibly in the Leucophoropterini (see tribal discussion). This apparent absence of circumaustral distributions may be the result of incomplete taxonomic knowledge, and certainly merits further investigation. The seasonal distribution of the Miridae in South Africa is at best only poorly understood. Knight (1941; 1968) has confirmed in North America that mirid reproduction generally takes place during the height of growth and flowering in the host. It is at this Figs. 289-309. Phylini, Parapseudosthenarus and Pseudosthenarus male genitalia. Fig. 289. Lateral view of vesica, Parapseudosthenarus buchenroederae. Fig. 290. Phallotheca, idem. Fig. 291. Lateral view of left clasper, idem. Fig. 292. Dorsal view of left clasper, idem. Fig. 293. Lateral view of vesica, Pseudosthenarus ater (Cape of Good Hope). Fig. 294. Phallotheca, idem. Fig. 295. Posterior view of left clasper, idem. Fig. 296. Dorsal view of left clasper, idem. Fig. 297. Right clasper, idem. Fig. 298. Lateral view of vesica, P. ater (Oudts- hoorn District). Fig. 299. Phallotheca, idem. Fig. 300. Lateral view of vesica, P. ater (Calvinia). Fig. 301. Phallotheca, idem. Fig. 302. Posterior view of left clasper, idem. Fig. 303. Lateral view of vesica, Pseudosthenarus grossus. Fig. 304. Phallotheca, idem. Fig. 305. Posterior view of left clasper, idem. Fig. 306. Posterior view of left clasper, P. rozeni. Fig. 307. Dorsal view of left clasper, idem. Fig. 308. Posterior view of left clasper, P. namaquaensis . Fig. 309. Dor- sal view of left clasper, idem. 1974] Schuh: South African Orthotylinae and Phylinae 239 240 Entomologica Americana [Vol. 47 time that the foliage is most succulent. The life cycles of mirids are thus rather short, terminating with the laying of eggs in plant tissue and diapause in the egg stage until the plant begins growing again the following season. Because a distinct summer-winter seasonality exists for most of South Africa and plant growth is correlated with this, it can be predicted that mirid abundance will be greatest in spring and early summer. The limited temporal samples that are available to some extent confirm this, particularly in the Southwest Cape. As noted by Stuckenberg (1962), the known distributions of many groups of animals in South Africa may reflect the activity patterns of entomologists. This should be borne in mind in the following analysis. Areas within South Africa for which the mirid fauna has been very poorly sampled include: the dry arid areas of the northwestern Transvaal; the Eastern Cape; the Orange Free State; the Northern Cape; the Great Karoo; Namaqualand; South West Africa; and, the Tsitsikama Forest area. Two areas of particular interest in South Africa, the Southwest Cape and the Drakensberg have received limited collecting, but deserve much greater attention than has been given in the past, particularly in view of the rapid shrinkage of the endemic flora. The Orthotylinae and Phylinae of South Africa can be divided into five components: 1) endemic; 2) tropical African; 3) paleo- tropical; 4) pantropical; and 5) cosmopolitan (including holarctic elements). 1 ) The following genera are endemic to South Africa (those in parentheses are known only from a limited number of localities and may prove to be more widely distributed): Austropsallus , Carino- gulus , Coatonocapsus, Denticulophallus , Eminoculus , ( Lasiolabo - pella), Karoocapsus , N amaquacapsus , Natalophylus , Neoambonea , Nichomachus , Parambonea , Parapseudosthenarus , Parasciodema, Pseudambonea , Pseudonichomachus , Pseudopilophorus , Pseudo- sthenarus , Stoebea, Widdringtoniola, and (Zanchiella) . They occur primarily in the Southwest Cape and the Drakensberg, are morpho- logically isolated, and have speciated in the region. -> Figs. 310-314. Phylini, Pseudosthenarus male and female genitalia. Fig. 310. Lateral view of vesica, P. namaquaensis. Fig. 311. Phal- lotheca, idem. Fig. 312. Ventral view of phallotheca, idem. Fig. 313. Sclerotized rings, idem. Fig. 314. Posterior wall, idem. 1974] Schuh: South African Orthotylinae and Phylinae 241 242 Entomologica Americana [Vol. 47 2) The tropical African element in South Africa is composed of the following genera: Aloea, Ambonea , Felisacodes , Formicopsella , Lamprosthenarus , Lepidocapsus , Myombea , Nanniella , Odhiambo- ella, Pangania , Plagiognathidea , Sknkuza , Systellonotopsis , Tricho- phorella, and Trichophthalmocapsus. Most of these genera are re- stricted to sub-Saharan Africa, although Aloea is known from North Africa and also from the Arabian Peninsula (Linnavuori, personal communication). 3) Genera occurring in South Africa which have paleotropical distributions are Acrorrhinium , Azizus , Hallodapus , Pseudoloxops , Systellonotus , and Zanchius. 4) Pantropical genera in South Africa are Ellenia and Paramixia. 5) Cosmopolitan genera in South Africa are Cyrtorhinus, Halti- cus, Macrotylus, Orthotylus, Pilophorus, Psallus, Sthenarus-Cam- pylomma, and Tytthus. Not all of these genera are truly cosmopolitan but all show very wide distributions; some are not presently well known from certain regions, particularly South America and Aus- tralia. Such genera as Orthotylus, Pilophorus, Psallus, and Sthenarus- Campylomma are very large and complex and in need of monographic revision; the presently recorded distributions for these genera may be inaccurate since some may not represent monophyletic units. The 54 genera of South African Orthotylinae and Phylinae show several more or less distinct distributional patterns within the region. They correlate well with those known for other groups, including the Lygaeidae (see Slater, 1964), other insect groups (see Stucken- Figs. 315-336. Pilophorini male and female genitalia. Fig. 315. An- terodorsal view of posterior wall, Aloea samueli. Fig. 316. Lateral view of posterior wall, idem. Fig. 317. Sclerotized rings, idem. Fig. 318. Lateral view of vesica, idem. Fig. 319. Phallotheca, idem. Fig. 320. Anterior view of left clasper, idem. Fig. 321. Apex of vesica, Ambonea munroi. Fig. 322. Phallotheca, idem. Fig. 323. Dorsal view of left clasper, idem. Fig. 324. Apex of vesica, Ambonea rusten- burgensis. Fig. 325. Phallotheca, idem. Fig. 326. Lateral view of left clasper, idem. Fig. 327. Lateral view of vesica, Parambonea trans- vaalensis. Fig. 328. Anterior view of left clasper, idem. Fig. 329. Lateral view of vesica, Neoambonea cynanchi. Fig. 330. Phallotheca, idem. Fig. 331. Anterior view of left clasper, idem. Fig. 332. Lat- eral view of vesica, Paramixia australis. Fig. 333. Phallotheca, idem. Fig. 334. Left clasper, idem. Fig. 335. Lateral view of vesica, Para- mixia suturalis. Fig. 336. Lateral view of vesica, Paramixia nigra. 1974] Schuh: South African Orthotylinae and Phylinae 243 244 Entomologica Americana [Vol. 47 Fig. 337. Distribution of six genera of the Miridae in South Africa (see text for explanation). berg, 1962) and non-insects including Amphibia and Reptilia (see Poynton, 1964; Stuckenberg, 1969). The five main patterns are discussed below, but must be considered as strictly tentative because of the infant state of the taxonomy of the South African Miridae. Distributions which are anomalous and cannot be analyzed with certainty are discussed under the most appropriate heading or in a general section at the end. 1 ) The most important pattern is that of endemic species distri- butions. This is the largest element of the South African fauna, representing 43% of the total genera. It can best be portrayed by division into several subgroups.. a) The most restricted pattern involves taxa confined to the Southwest Cape and Karoo (including portions of Namaqualand) (Figure 337) which are probably adapted to the unique flora of the area. Included genera are: Eminoculus, Denticulophallus , Widdring- toniola, Pseudosthenarus, and Ccipecapsus. The first three taxa are known to occur on endemic plant genera. The host of Pseudo- 1974] Schuh: South African Orthotylinae and Phylinae 245 16 20 22 24 26 28 30 32 34 Fig. 338. Distribution of seven genera of the Miridae in South Africa (see text for explanation). sthenarus is not known, but the most closely related genus, Para- pseudosthenarus, occurs at midaltitudes on the Drakensberg in Natal on Buchenroedera (Leguminosae), a plant genus which is endemic to Africa and almost completely restricted to South Africa (Phillips, 1951). The occurrence of Widdringtoniola and Eminoculus in the Transvaal is almost certainly the result of transplanting the hosts as ornamentals in Pretoria. The morphological distinctiveness of Eminoculus and Pseudosthenarus suggests long isolation in South Africa. b) A pattern of wider distribution is that of endemicity in the Southwest Cape but having strong affinities with the Drakensberg montane region and outlying Southwest Cape floral elements. Genera with this distribution are Austropsallus , Coatonocapsus , Stoebea , Parapseudosthenarus , the Acrorrhinium brincki group, Zanchius (except Z. nigrolineatus) , and Zanchiella. These genera, in large part, show a strong association with the endemic flora of South Africa and particularly with elements of the Southwest Cape Flora (Figure 246 Entomologica Americana [Vol. 47 Fig. 339. Distribution of five genera of the Miridae in South Africa (see text for explanation). 338). The occurrence of Zanchius buddleiae in the central Transvaal is almost certainly the result of introduction of the host into the National Botanical Gardens in Pretoria. Although the distributions are not well known for all species, at least Austropsallus drakensber- gensis is generally distributed in the northern Drakensberg, and further collecting will probably substantiate similar occurrences for other taxa. This is in marked contrast to both genera and species in “pattern a” which are not known to occur in the Drakensberg in Natal or the Transvaal. This agrees with many plants endemic to South Africa, where those in the Southwest Cape show very limited distributions and those of the Drakensberg and other outlying Cape Floral elements show much wider distributions. Zanchius and Zanchiella (Figure 338) are related to Felisacodes (see pattern 2), but I have placed them here because they appear to have radiated in South Africa and are primarily restricted to the endemic flora. Nichomachus and Pseudonichomachus (Figure 339) have been 1974] Schuh: South African Orthotylinae and Phylinae 247 16 20 22 24 26 28 30 32 34 Fig. 340. Distribution of three genera of the Miridae in South Africa (see text for explanation). collected from localities at relatively high elevations in the Transvaal where there are patches of Southwest Cape related vegetation. The species from the Cape Province are either from the Macchia region proper or from the Karoo. Natalophylus is known from a single locality on the Drakensberg at about 6000 feet (1875 meters) (Figure 339). Its morphological specialization in the male genitalia relative to the related Phylus from Europe, suggests isolation and possible endemism in South Africa. This genus may be dervied from a Phylus- like ancestor which arrived in South Africa from the north. The host genus Heteromorpha (Umbelliferae) is endemic to Africa (Phillips, 1951). Lepidocapsus and Lamprosthenarns are known from the South- west Cape and montane regions of South Africa, respectively (Figure 339). Both of these genera also occur in tropical Africa, but at the present time it is not possible to determine with which area they have their strongest affinities. An additional genus which may have a “type b” distribution is 248 Entomologica Americana [Vol. 47 Fig. 341. Distribution of two genera of the Miridae in South Africa (see text for explanation). Pseudambonea , which is known at present only from a single locality in the eastern Cape, north of Port Elizabeth. c) A third endemic pattern includes Namaqualand, the Great Karoo, and the dry Southwest Cape (with possible strong affinities with the Little Karoo). Genera having this distribution are Karoo- capsus , Parasciodema , the Acrorrhinium munting group, and to some extent Austropsallus and Coatonocapsus. Figure 340 shows the distribution of these taxa; many of the species are currently known only from Grootfontein, Middelburg, Cape Province (Note: In several cases the same species taken at Middelburg are also recorded from Zomerkomst, Politzi, Transvaal. According to Acocks (1951), the vegetation of these areas is unrelated. The latter localities may be in error.). The only close relatives of Karoocapsus are from Australia, which indicates that this unique South African genus is probably a relict. Namaquacapsus from Namaqualand (Figure 345) possibly has this type of distribution. It shows great morphological specialization, 1974] Schuh: South African Orthotylinae and Phylinae 249 Fig. 342. Distribution of two species of the Miridae in South Africa (see text for explanation). particularly in the type of vestiture. The distribution of the Halticini, to which Namaquacapsus belongs, is however primarily Mediter- ranean and therefore it is discussed under “pattern 4”. 2) A pattern showing primarily tropical affinities in South Africa and more or less confined to an area delimited by the 15° ET isoline includes Cyrtorhinus , Formicopsella, Hallodapus , Lcisiolabopella, Macrotylus niger, Odhiamboella, Pangania , Paramixia , Pseudo- loxops , Trichophorella , Trichophthalmocapsus , and Tytthus. These genera constitute 23% of the South African fauna. Most of the taxa with this distributional pattern belong to tropical African and paleotropical faunal elements and show greater diversity in tropical Africa than in South Africa (see Figure 343). Pangania fasciatipennis and Formicopsella regneri present well documented examples of this type of distribution (Figure 341). Both species are apparently associated with Acacia , particularly A. karroo . The distribution of these two mirid species is well defined by the 15° ET isoline despite the wider distribution of their host, 250 Entomologica Americana [Vol. 47 8 12 14 16 18 20 22 24 26 28 30 32 34 36 38 16 20 22 24 26 28 30 32 34 Fig. 343. Distribution of nine genera and one species of the Miridae in South Africa (see text for explanation) . especially in the highveld of the Transvaal and the Orange Free State. The one record of Pangania from the Southwest Cape, from the Little Karoo at Nooitgedacht near Oudtshoorn, corresponds with the 15° ET isoline and also with the distribution of Acacia in that area. Both of these species come to light and are relatively easily collected. The known distribution is therefore probably relatively accurate, although they will most likely be found in much of South Africa that is as yet very poorly collected. Paramixia has a similar distribution (Figure 342), although here one species is more strictly tropical than another, but nonetheless the 15° ET isoline defines well the distribution of the genus within South Africa. Felisacodes has what is apparently a “forest relict” distribution (Figure 343). It is known from Chirinda, Rhodesia, and Mt. d’Ambre, Madagascar, as well as South Africa. 3) A pattern of wide distribution within South Africa, and in some cases involving East Africa, includes Systellonotus and Ellenia. These two genera constitute 4% of the fauna. 1974] Schuh: South African Orthotylinae and Phylinae 251 Fig. 344. Distribution of a species of the Miridae in South Africa (see text for explanation) . Systellonotus is chiefly Mediterranean is distribution. In South Africa it is not restricted to the area of Mediterranean climate but is widely distributed, being known from Natal, the Orange Free State, and South West Africa (Figure 345). This may represent a disjunct Mediterranean distribution, but the widespread occurrence of the genus is South Africa suggests that it may eventually be found in East Africa as well. Ellenia obscuricornis is widely distributed in South Africa and East Africa (Figure 344); however, in South Africa it does not occur in tropical localities, such as the low veld in the Transvaal and the Natal tropical corridor. There is some indication that Ellenia , in South Africa, may be more or less restricted to Senecio, a very diverse and widely distributed genus. The absence of Ellenia from tropical South Africa, even though it occurs in East Africa, is not readily explicable. 4) A pattern of distribution associated with very dry areas and possibly showing some endemism in South West Africa includes 252 Entomologica Americana [Vol. 47 Fig. 345. Distribution of four genera of the Miridae in South Africa (see text for explanation). Carinogulus , Namaquacapsus , and Systellonotopsis, which constitute 6% of the fauna. Both Carinogulus and Namaquacapsus show rela- tionships with groups that occur primarily in the Horn of Africa or the Mediterranean. There are many examples of groups whose distributions show close affinities between these areas (van Zinderen Bakker, 1969; Moreau, 1966). Systellonotopsis is probably endemic to the dry areas of southern Africa, but its relationship to other groups is too poorly understood to speculate on the possible signif- icance of its distribution (see Figure 345). 5) About 15% of the South African genera have a strictly tropical distribution roughly delimited by the 16° ET isoline. They are Azizus , Halticus , Myombea, Nanniella , Pilophorus , Plagiog- nathidea , Pseudopilophorus , and Skukuza. These genera are all re- stricted to the Mozambique coastal plain and to the Natal tropical corridor (Figure 346). They primarily belong to the tropical African element, but Pilophorus and Halticus are cosmopolitan. Macrotylus hemizygiae and Zanchius nigrolineatus may also be 1974] Schuh: South African Orthotylinae and Phylinae 253 16 20 22 24 26 28 30 32 34 Fig. 346. Distribution of eight genera and two species of the Miridae in South Africa (see text for explanation). species with tropical affinities (Figure 346). They are presently known only from the Kruger National Park. Other South African members of these two genera have distributions that are not strictly tropical, as discussed above. Five genera, or 9% of the known fauna of South Africa, were not analyzed in the foregoing discussion. These include Brachycran- ella , Sthenarus-Campylomma , Leptoxanthus , Psallus , and Orthotylus. My inability to identify specimens of Brachycranella and Lepto- xanthus makes it impossible to discuss their distributions at the present time. Both genera are known only from South West Africa. The Sthenarus-Campylomma group is represented in South Africa by approximately eight species. These are known primarily from the area with the more tropical fauna rather than from the areas of ex- treme endemism. This distribution is in agreement with that of Campylomma and Sthenarus as a whole, which are rather widely dis- tributed in the Old World with only one introduced species of Campy- lomma presently recorded from the New World (Knight, 1941). 254 Entomologica Americana [Vol. 47 Approximately 10 Psallus-Yike species are present in collections from South Africa. They are known from all areas of the country. The extremely complex taxonomic problems involved with this genus make it impossible to determine at present the possible affinities of the South African fauna. Orthotylus in South Africa presents an even more complex picture than Psallus. Orthotylus is currently recorded from all zoo- geographic regions except South America. There are about 20 species occurring in South Africa, but they cannot be related to the faunas of other areas at present because of the paucity of knowledge on the genus as a whole except for those of the Pale arctic. PART 2. A PHYLOGENETIC ANALYSIS OF THE ANT- MIMETIC TRIBES OF THE ORTHOTYLINAE AND PHYLINAE FOR THE WORLD HISTORY OF THE CLASSIFICATION OF THE MIRIDAE The phylogeny of the Miridae has not received great attention since the exhaustive treatments of Reuter (1905a; 1910a). Van Duzee (1916), Knight (1923; 1941) and Carvalho (1952a) have reviewed or revised the classification; Slater (1950) and Kelton (1959b) examined the female and male genitalia, respectively, and made many valuable suggestions relative to the classification and phylogeny of the family; and, Wagner (1955) and Leston (1961) presented phylogenetic analyses of the Miridae with useful discus- sions, but unfortunately their works were based almost solely on Palearctic genera. Distant (1904c: 412-413) stated: “. . . at present the classifi- cation of the family is more reflective of personal opinion, and con- trived for purposes of entomological arrangement, than exhibiting an evolutionary or philosophical conception. The Capsidae are a very difficult group to study, their affinities are of the most difficult description, and for the present we must be satisfied with a somewhat artificial or cabinet arrangement.” Distant went on to say that he did not understand the system of Reuter and therefore did not follow it. This statement by Distant precipitated Reuter’s (1905a) pub- lication of the polemical “Hemipterologische Spekulationen. I. Die Classification der Capsiden” in which he accused Distant of making only superficial observations on the Miridae and therefore producing results of little or no value. Most mirid taxonomists since Reuter have more or less adopted his excellent classification. Even the 1974] Schuh: South African Orthotylinae and Phylinae 255 world classification of Carvalho (1952a) added little new morpho- logical information to that which Reuter (1905a; 1910a) presented. Reuter (1905a) included a dendrogram with his phytogeny of the Miridae. He considered the absence of a pronotal collar to be primitive. Reuter therefore placed the “Isometopidae,” which possess ocelli, but lack a pronotal collar, at the very base of his phylogenetic tree and grouped the Plagiognatharia, Oncotylaria, etc. (Phylinae) and the Cyllocoraria, Laboparia, etc. (Orthotylinae) together on one side of the dendrogram with the Phylinae nearer the base be- cause of the hair-like parempodia (considered to be “poorly devel- oped arolia” by Reuter). He derived the Dicypharia (Dicyphinae) and Cylaparia and Fulviaria (Cylapinae) from near the Orthotylinae. Reuter considered the Bryocoraria (Bryocorinae) and Clivenemaria (Deraeocorinae) to be somewhat isolated but relatively closely re- lated to one another and placed them by themselves near the base of the tree. He placed the Miraria, Capsaria, etc. (Mirinae) at the top of the dendrogram above the Orthotylinae. Reuter (1910a), in his later work, did not give a phyletic dendrogram for the Miridae but presented a scheme of classification similar to that discussed above. In this paper he grouped the Miridae into subfamilies whereas in the previous paper he recognized only divisions (tribes). Knight (1923; 1941) obviously modified his own dendrogram from that of Reuter but gave little supporting discussion regarding subfamily relationships. Both Reuter and Knight considered the Phylinae to be among the most primitive members of the Miridae, whereas I consider them to be among the most highly derived (see below). Wagner (1955) considered the Orthotylinae-type of male genitalia to be primitive in the family. He presented the Bryocorinae, Cyla- pinae, Pilophorini-Phylinae, and Halticini-Mirinae-Dicyphinae-Dera- eocorinae, in that order, as separate evolutionary lines branching off the main orthotyline stem at successively higher levels. This scheme was based on the male genitalia and did not take into account the parempodia, pulvilli, or pronotal collar, all of which would have to be evolved more than once to correspond with Wagner’s interpre- tation of the evolution of the male genitalia. Wagner considered the Orthotylinae and Phylinae to be closely related but also derived the Mirinae, Dicyphinae, Deraeocorinae, and Halticini in a single line of evolution from the main Orthotylinae stem. If the Orthotylinae male genitalia are considered as derived a much more parsimonious system can be devised. Leston (1961) presented a phylogenetic scheme based primarily 256 Entomologica Americana [Vol. 47 on the genitalic studies of Slater (1950) and Kelton (1959b) and on his own work on chromosomes, testis follicle numbers, and wing venation. He considered the Isometopinae to be the most primitive subfamily in the Miridae with the Bryocorinae somewhat more ad- vanced, but still near the base of the mirid stem. Leston (1961) proposed a relationship between the Deraeocorinae and Mirinae and also a possible link between the Cylapinae and Deraeocorinae. He considered the Orthotylinae, Phylinae, and Dicyphinae to be the most advanced groups in the Miridae. ANALYSIS OF CHARACTERS AND A REVISED CLASSIFICATION Reuter (1910a) used the parempodia as the primary characters in the classification of the Miridae and was followed by Carvalho (1952a). Both authors recognized three basic types of parempodia and considered them more or less invariable. I have also found that the parempodia are extremely valuable in the classification of the Miridae; however, some confusing variation exists in the different types, a situation which was first recognized by Wagner (1955). By interpretation of this variation and correlation with other char- acters, particularly those of the genitalia, the parempodia can be used to establish some primary divisions within the family. Kullen- berg (1947b) raised the most serious doubts about the value of the parempodia in the classification of the Miridae, suggesting that they are highly adaptive. The previous studies of Reuter (1910a) and Knight (1922; 1923) and the subsequent work of Carvalho (1952a) and Wagner (1952; 1955) and this study all show, however, that the parempodia are the most useful single character in the classification of the family. The other characters which are of greatest value in the systematics of the Miridae are the male and female genitalia, pronotal collar, wing dimorphism, pulvilli, feeding habits, presence or absence of ocelli, number of tarsal segments, ant-mimetic appearance, and possibly chromosome numbers and testis follicle numbers. Additional characters such as body form, punctation, pubescence, length of tarsal segments, and number of cells in the membrane are also important, but usually useful only below the subfamily and tribal levels. My use of characters in classifying the Miridae is more or less in agreement with previous authors except for differences in interpretation and increased emphasis on the male and female geni- talia. In addition to the parempodia, Reuter (1905a; 1910a) used the structure of the pronotum, including the collar, tarsi, membrane 1974] Schuh: South African Orthotylinae and Phylinae 257 cell, hamus, prosternal xyphus, and lora in his classification. Knight (1923) considered the parempodia, genitalia, biology, and modifi- cations of the thorax, in that order, to be of greatest importance in studying relationships within the family. I have not included the hamus, prosternal xyphus, or lora in developing the following classifi- cation of the Miridae, as I have found these structures to be of more limited value than the others listed above. The genitalia have received only limited acceptance in the study of mirid phylogeny. The attitude of many workers is typified by Carvalho (1952a: 34) . . the claws and arolia are still the best characters to be used in the subdivision of the family into subfamilies and tribes. . . . The genitalia alone have been found to be mis- leading in many respects and their acceptance as a primary character would certainly cause some changes in the present classification.” The first author to use the male genitalia in the separation of mirid species was Reuter in 1883 (Kelton, 1959b). Knight (1941) emphasized that the vesica in the Phylinae was a fundamental char- acter in classification. Wagner (1955) was the first author to use the male genitalia as an integrated unit in the classification of the family. The most comprehensive review of mirid male genitalia is that of Kelton (1959b); he suggested needed changes in the classifi- cation based on the vesica, but did not make them. I use the male genitalia extensively, including characters of the claspers, vesica, and phallotheca. Mirid female genitalia received very little attention until the studies of Kullenberg (1947a) and Slater (1950). The former author was concerned primarily with the functional aspects of the morphology. Slater, however, examined them from the point of view of higher classification and laid the groundwork for subsequent studies. Wagner (1956) was the first author to base taxonomic changes in the Miridae on the structure of the female genitalia. I use the structure of the posterior wall primarily, and the sclerotized rings secondarily, in the classification of the family. The possible value of wing dimorphism in classifying the Miridae was first alluded to by Reuter (1910a), but it has been used very little. In many Hemiptera (vis., Lygaeidae, semiaquatic families) reduction of the hemelytra occurs with more or less equal frequency in both sexes. In the Miridae two types of wing dimorphism exist, suggesting two independent mechanisms to account for the expression of the phenomenon. The Stenodemini and Halticini show the nonsex related type of wing dimorphism mentioned above. In the Phylinae and some Orthotylinae, however, wing dimorphism is almost always 258 Entomologica Americana [Vol. 47 sex related, the females being brachypterous, when short winged forms exist, and the males always being macropterous. A similar situation is thought to exist in the Schizopteridae (Emsley, 1969). Brinkhurst (1959) has established the mechanisms for determination of hemelytral types in the Gerridae, but this is not known for the Miridae. Presumably the mechanism of determination in the Steno- demini is hormonal and environmentally induced (Southwood, 1962), but in the Phylinae it is probably genetic. Southwood (1962) and Sweet (1964) have proposed that brachyptery in the Hemiptera is a development related to environ- mental stability. This probably applies to the Phylinae (and other groups with a similar type of wing dimorphism), with the male as the active agent of genetic interchange between populations. The females are only very rarely macropterous and therefore contribute less to gene flow between populations than the males (see Brink- hurst, 1963). The retention of the female in the habitat of the host must confer a selective advantage. In the Hallodapini (and some Leucophoropterini, Orthotylini, and Nichomachini) an additional selective advantage may be conferred on the females in that they are generally much more ant-like than the males. Carvalho (1952a) used ant-mimetic appearance as a primary character in the classification of the Miridae. An important aspect of this study has been to assess the value of mimicry in analyzing the phylogeny of the family. Within limits I have found that mimetic appearance is very useful, particularly at the tribal level (e.g. Hallo- dapini). In some cases, however, higher taxa cannot be defined on mimetic facies. Kelton (1959b) for the Mirinae, and other workers, have emphasized this point. In all groups where ant-mimics exist, they certainly represent a derived condition. However, in many cases (e.g. the Pilophorini) mimetic genera retain primitive characters. The morphological modifications most commonly found in the ant- mimetic Miridae are: the presence of one or more light hemelytral maculae or fasciae that contrast with the dark background coloration of the forewings and body; the sinuation of the lateral hemelytral margins; and the constriction of the pronotum anteriorly, with the tendency toward the development of a flattened pronotal collar. Certain tribes in the Phylinae and Deraeocorinae have the pronotum constricted anteriorly and bearing two erect, cone-like structures dor- sally. This structural modification also occurs in the genus Saldoida (Saldidae). Some genera (as yet undescribed) have the pronotum constricted medially so as to form an hour-glass shape. In certain undescribed genera in the Pilophorini and Leucophoropterini the 1974] Schuh: South African Orthotylinae and Phylinae 259 TABLE 1 Comparison of Nabidae and Miridae (p — primitive; d — derived) CHARACTER NABIDAE MIRIDAE Feeding habits predaceous (p) primarily phytophagous (d) Male genitalia (symmetry) nearly symmet- rical (p) strongly asymmet- rical (d) Male genitalia (vesica) membranous (p) membranous or sclerotized (p-d) Female genitalia (posterior wall) poorly developed (P) usually well devel- oped (d) Female genitalia (sclerotized rings) poorly developed (P) usually well devel- oped (d) Cuneus absent (p) present (d) Ocelli present (p) usually absent (d) Pronotal collar present or absent (p-d) present or absent (P-d) Parempodia hair-like (p) hair-like or fleshy (p-d) gula is carinate below the eye and gives the appearance of mandibles when viewed anteriorly. All of these structural characteristics have evolved more than once and are therefore indicative of the extreme adaptability of the Miridae to ant-mimic selection. The obvious convergence of several groups toward certain common mimetic facies for the most part precludes the definition of higher taxa within the Miridae on the basis of ant-like appearance alone. The zoogeography of the Miridae has never been analyzed or used as a tool in the classification of the family. With the Carvalho Catalogue as a source of information the assessment of distributional patterns becomes much easier than before and the study of the phylogeny of the family can be greatly enhanced, a position in opposition to that taken by Leston (1961). In this study I have not been able to consider the distributions of all subfamilies of Miridae in detail. The distributional patterns of the Orthotylinae and Phylinae, however, suggest that zoogeographic analyses within the family, especially at the tribal level, are very useful in understanding the evolution of the Miridae. Additional characters I have used in developing this classification are discussed below under the individual taxa. TABLE 2 260 Entomologica Americana [Vol. 47 C-W G 0 jU C/3 'G « s G 03 1 1 x> G CD G "»-c O Oh •a I G a Oh w a o O OjS Qj C/3 W PQg S 2<0 hHu Z w Q ffl £h 00 H O o £ W P4 h-1 u on — H G CD w > & 03 03 (D — , a 13 ° £ G CD w > ~ CD 13 ° £ o c G S-h G3 ^ S 5 CD 03 03 > o a 00 o o c 03 -O ^ a 3 CD g 03 I a c G « 03 03 in > o P /-N s 3 CD 03 03 CD — , Oh 00 o a ro a g CD w > r2n -V OD flj 133 > £ 03 03 qj _ a 13 ° £ G 03 I ' a 3 (D 133 > -G ^ 03 qj -h Oh 00 o C- c/3 G G O >> G 33 G G b G -O 33 G G CD a m a g CD w > >G ^ 03 H u HH P4 H 3* >H Oh w P4 X o PQ Q § Q O a 1974] Schuh: South African Orthotylinae and Phylinae 261 d o >b i-I GO 20 Pft Pu 2 i-4 Z d 9 Ie Q PQ 2 < 2 T 2 >% t-i o 3 T3 > O 3 T3 G o C CD > o 73 g >. E? -g « CO I> > o 2 2 £• i •s.g* (U CG w jp c» 3 ^ j2 O 3-5 *-• G ft O T3 G +-> CD . c g < H O 2 2 < g 2 2 (J 2 Pin H ot, 2 p4 2 U PQ 5 § Q O 2 262 Entomologica Americana [Vol. 47 In a phylogenetic or cladistic classification it is necessary to designate given states of a character as relatively primitive (plesio- morphic; ancestral) or derived (apomorphic) and monophyletic groups must be recognized only on the possession of shared derived characters (Hennig, 1966). The taxonomy of the Miridae is still so poorly known that it is difficult to determine for most characters what is the primitive and what is the derived state. By examining families related to the Miridae some comparative data can be as- sembled. Table 1 lists characters in the Nabidae and Miridae. The Nabidae have the greatest number of what are probably plesiomorphic characters in the Cimicoidea, including predatory feeding habits, ocelli, presence of a pronotal collar, membranous vesica, nearly symmetrical male genitalia, and absence of pulvilli. The Anthocoridae and Miridae, which are derived relative to the Nabidae, have in common the cuneus and therefore form a natural group within the Cimicoidea; however, these two families represent individually specialized lines of evolution. The Anthocoridae are entirely predatory, retain ocelli, and show a trend toward develop- ment of the highly specialized method of traumatic insemination. The Miridae, with the exception of the Isometopinae, have lost the ocelli, have a tendency toward phytophagy, and have specialized genitalia, although along a much different line than found in the Anthocoridae. Table 2 lists the subfamilies of Miridae and a number of char- acters that are important in the classification of the family. I am following Carayon (1958) and including the Isometopinae in the Miridae. If the individual mirid subfamilies are compared with the Nabidae, which seems justified, it appears that the Isometopinae possess the greatest number of plesiomorphic characters, namely the presence of ocelli, predatory feeding habits, and the absence of pulvilli. The Cylapinae also appear relatively primitive in that they are predaceous, possess a pronotal collar, and lack pulvilli. The view that these groups are not secondarily predaceous is supported by the uniformity of feeding habits within each subfamily. Support for the predatory nature of the ancestral cimicoid. stock can be found in the fact that the Nabidae, Microphysidae, Velocepedidae, and Anthocoridae and also Reduviidae (which are most closely related to the Cimicoidea [Cobben, 1968]), are all predatory. Only the Tingidae, Thaumasto- coridae and Miridae are phytophagous. In that the Miridae are specialized within the Cimicoidea, lacking ocelli (except Isometo- pinae) and possessing a cuneus and specialized male and female 1974] Schuh: South African Orthotylinae and Phylinae 263 genitalia, it is probable that the predaceous habit is ancestral within the family and that phytophagy is a derived condition. The feeding habits of the Miridae are, however, rather poorly understood. Many groups may be secondarily predaceous and some are probably oligo- phagous. The rounded pronotal collar is probably plesiomorphic in the Miridae, an hypothesis supported by the occurrence of the structure in most subfamilies, including the Cylapinae, Bryocorinae, Dicyphinae, Mirinae, and Deraeocorinae. It also occurs in the prostemmine Nabidae (although not in the Nabinae) and in the Anthocoridae, both of which possess a greater number of plesiomorphic characters than do the Miridae. The rounded pronotal collar is absent in the Isometopinae, Orthotylinae, and Phylinae, but this can probably best be attributed to secondary loss. If the rounded collar is con- sidered to be derived in the Miridae it must be evolved independently at least 4 times, whereas it is only necessary that it be lost twice when it is considered to be plesiomorphic. The genus Psallops Usinger should be considered here. Usinger (1946) placed this genus in the Phylinae, but subsequent examination of the male genitalia by Carvalho (1956b) revealed that Psallops is not a phyline, but has a membranous vesica more similar to all other Miridae than to the Phylinae. Carvalho (1956b) felt that someday Psallops would be placed in the Isometopinae, even though it does not possess ocelli. Psallops is in actuality probably most closely related to the Cylapinae. I have examined the female genitalia of an undescribed species of Psallops from South Africa. It has well developed sclerotized rings and a simple, sclerotized, plate-like posterior wall. Neither of these structures is highly specialized, but both are relatively primitive and similar to those found in the Cylapinae (personal investigation) and Isometopinae (Slater and Schuh, 1969). Psallops has only two tarsal segments. This condition is the rule in the Isometopinae, and also occurs in a few genera placed in other mirid subfamilies, including Vannius Distant and Peritropis Uhler in the Cylapinae and Hemisphaerodella Reuter in the Bryocorinae. The 2-segmented condition is almost certainly not ancestral in that the remainder of the Miridae, the Nabidae, and the Anthocoridae all have 3-segmented tarsi. Bergroth (1925) has noted that 2-segmented tarsi occur sporadically throughout the Heteroptera, and that those groups that possess them cannot be considered connecting links to other such groups. J. A. Slater (per- sonal communication) has suggested that they possibly represent a neotenic condition, because of the nymphs of the Geocorisae which 264 Entomologica Americana [Vol. 47 all have 2-segmented tarsi. The Isometopinae and Cylapinae there- fore cannot be related by the similar tarsal structure of some genera. Psallops also has only one membrane cell, a condition found in most, but not all Isometopinae (Bergroth, 1925), and lacks a pronotal collar, a structure which is also absent in all Isometopinae but present in most Cylapinae. Psallops looks much like many Isometopinae (see Isometopidea , in Slater and Schuh, 1969). This complex of characters suggests, but in no way confirms, a relationship between the Isometopinae and Cylapinae and may distinguish them as rela- tively primitive within the Miridae although they both possess many derived characters. The Dicyphinae have a rounded pronotal collar, a simple posterior wall in the female and a membranous vesica in the male, all of which are probably plesiomorphic characters and occur in varying combina- tions in other mirid subfamilies. Cobben (1968) has suggested a relationship between the Dicyphinae and the Helopeltis group (Monalionini) of the Bryocorinae based on the structure of the eggs. The pul villi of the Dicyphinae also relate them to the Bryocorinae. They are leaf-like and attached to the inner surface of the claw, whereas in all other Miridae, the pulvilli are minute and always attached to the ventral surface of the claw.1 The male and female genitalia of the Bryocorinae appear to be the most primitive in the Miridae although the condition may be secondarily derived (Kullenberg, 1947b). In this subfamily the phallus is very simple and in this sense resembles that of Nabis; it may not be divided into a vesica and conjunctiva as in other mirids (Kullenberg, 1947b). Also the posterior wall and ring glands of the female are very poorly developed in some tribes (Slater, 1950), as in Nabis (Kullenberg, 1947a), but more highly developed in others (see Schmitz, 1968). The Bryocorinae and most Isometopinae have only one membrane cell, whereas the rest of the Miridae have two, which also suggests a derived condition (see Leston, 1961). The genus Bunsua Carvalho, from Africa, which was originally placed in the Orthothylinae, is closely related to the Bryocorinae, and may have some characters which are intermediate between the Bryocorinae and other members of the Miridae, although there is no direct relationship to the Orthotylinae. 1 Carvalho (1952a) considered the pulvilli of the Phylinae (including Dicyphinae) to arise from the “base or inner surface of the claw” and those of the Bryocorinae to arise from the ventral surface of the claw. I have interpreted the Phylinae-type to arise from the ventral surface of the claw and the Bryocorinae-type (including Dicyphinae) from the inner surface. 1974] Schuh: South African Orthotylinae and Phylinae 265 The question of the monophyletic nature of the Bryocorinae and the relationship of the Dicyphinae to the Monalionini needs careful investigation. The Deraeocorinae lack pulvilli and have a rounded pronotal collar, both plesiomorphic conditions in the Miridae. They are, however, highly specialized in several respects and therefore are discussed below in relation to the Mirinae. The Mirinae, Orthotylinae, and Phylinae all possess modified fleshy parempodia and pulvilli that are attached to the ventral sur- face of the claws. They therefore form a group within the Mi- ridae. Leston (1961) considered this structural similarity to be a convergence. The Mirinae possess several derived characters. The vesica of the male is of a highly developed membranous type similar to that found in the Deraeocorinae (Kelton, 1959b). The posterior wall and sclerotized rings are specialized but do not show an extremely close relationship to other subfamilies in the Miridae (Slater, 1950). This subfamily has a rounded pronotal collar which I consider to be an ancestral condition relative to the Orthotylinae and Phylinae, both of which lack a collar. The Stenodemini do not have a collar, except for the genus Collaria Provancher. Knight (1941) felt that the Stenodemini were probably primitive because of their host plants (grasses) and distribution. I disagree with Knight on morphological grounds and consider the pronotal type in the Stenodemini to be derived. Also the grasses may not be so primitive as thought by Knight. Distant (1904c) considered the sulcation of the head in the Stenodemini to be of great importance, and assigned the group subfamily rank; he placed the Isometopinae in a second subfamily within the Miridae and lumped all of the remaining members of the family into a third subfamily. The monophyletic nature of the Mirinae is supported by the female genitalia (Slater, 1950) and the male genitalia (Kelton, 1959b). The Deraeocorinae, although possessing hair-like parempodia and lacking pulvilli, both of which are probably plesiomorphic characters in the Miridae, also possess a number of derived characters and probably show their closest relationship to the Mirinae, and may be derived from them. The vesica in the Deraeocorinae is similar to that found in the Mirinae and occurs nowhere else in the Miridae (Kelton, 1959b). The posterior wall is a simple sclerotized plate, which probably represents the plesiomorphic condition in the Miridae, but in the Deraeocorinae, when considered in light of other characters possessed by the subfamily, may represent a secondarily simplified 266 Entomologica Americana [Vol. 47 Fig. 347. Phylogeny of the Miridae. condition (Slater, 1950). The general facies of the Deraeocorinae, particularly Deraeocoris, are very much like that of many Mirinae. The type of pronotal collar and the punctate dorsum are common to the two subfamilies; the latter character occures only sporadically in all other mirids, but is relatively common in the Mirinae and almost universal in the Deraeocorinae. The Deraeocorinae are often very specialized predators, suggesting a specialization of feeding habits rather than what is probably a generalized predatory habit in many ground living Cylapinae (e.g. Fulvius). These facts then indicate secondary loss of the pulvilli, reevolution of the hair-like parempodia (see also Orthotylinae-Phylinae discussion), and simplification by reduction of the female genitalia in the Deraeocorinae from the type found in the Mirinae. A number of characters suggest that the Orthotylinae and Phylinae are closely related. These include 1) the absence of a rounded pronotal collar (a derived character relative to other Miridae, although it is secondarily evolved at least once in both subfamilies); 2) the presence of recurved, convergent parempodia in the Ortho- tylinae and some Phylinae (Pilophorini) ; 3) the tendency for in- folding of the lateral margins of the sclerotized rings in both sub- 1974] Schuh: South African Orthotylinae and Phylinae 267 families with a great accentuation of this condition in the Orthotylini; and 4) a tendency towards sclerotization of the vesica. Figure 347 presents a possible evolutionary scheme for the Mi- ridae. The Isometopinae probably diverged early as an isolated line which retained the ocelli. The Cylapinae also probably arose relatively early, in that they are predatory, primarily ground living, have relatively primitive male and female genitalia, lack pulvilli, and possess hair-like parempodia. The Cylapinae also possess several characters that are probably derived, including the form of the head, but none of these seems to relate them to other subfamilies. The Dicyphinae and Bryocorinae (Monalionini) may be related as suggested by Cobben (1968). If these two groups were placed in a single subfamily they would still be related to the remaining mem- bers of the Bryocorinae through the Monalionini by the pretarsal structures and the single cell in the membrane (which also occurs in some Isometopinae, but this is probably a convergence [Leston, 1961]). The male and female genitalia of the Bryocorinae may appear primitive as a result of reduction (Kullenberg, 1947b; Slater, 1950). If they are considered as primitive within the Miridae as a whole, it necessitates the loss of the ocelli twice or reevolving the ocelli in the Isometopinae. I have therefore derived the Bryocorinae- Dicyphinae above the level of the Isometopinae but below the level of the remaining subfamilies, based primarily on the structure of the male and female genitalia in the Dicyphinae. The Mirinae (including Deraeocorinae)-Orthotylinae-Phylinae line is probably the most advanced in the Miridae. The Mirinae are primarily phytophagous. The Deraeocorinae, although predatory, are often very highly specialized and in many cases resemble the Mirinae and are probably most closely related to them. I have there- fore derived them from a common stem. The Orthotylinae and Phylinae are related to the Mirinae by the pretarsal structures but differ from them in lacking a rounded pronotal collar. They are related to one another by the structure of the parempodia, the male and female genitalia, the absence of a rounded pronotal collar, and possibly by their mixed feeding habits (Leston, 1961). The tremen- dous diversity in the Mirinae, Orthotylinae, and Phylinae in most zoogeographic regions suggests an active evolution with little extinc- tion, and therefore probably an advanced position in the evolution of the family. The subfamily Palauocorinae (Carvalho, 1956b) was not discussed above because so little is known about it. Erected for a single genus and species from Micronesia, this unique insular subfamily has many 268 Entomologica Americana [Vol. 47 specialized features, but may not deserve such high taxonomic rank. It is probably specialized through great isolation in an island envi- ronment. Further information will be necessary to determine the correct placement for this taxon within the Miridae. THE RELATIONSHIPS OF THE ORTHOTYLINAE AND PHYLINAE Three types of parempodia exist within the Orthotylinae and Phylinae: 1) distinctly fleshy, convergent apically, recurved (lyre- shaped), and flattened laterally; 2) hair-like and parallel; and 3) fleshy, rod-like, of nearly uniform diameter, and weakly conver- gent apically. Types 1 and 3 were placed together as a single type in the Reuter and Carvalho systems of classification. Knight (1923) realized that there was classificatory confusion in groups with conver- gent parempodia and made some generic changes, but it was Wagner (1961) who first pointed out the distinctive nature of types 1 and 3. Excellent figures of all three types of parempodia are available in the current literature (see Carvalho, 1955a; Knight, 1923, 1941, 1968; Wagner, 1961). All genera of Orthotylinae and Phylinae with parempodia types 2 and 3 have male genitalia of the phyline-type, i.e., with a rigid, sclerotized vesica and characteristic left clasper and phallotheca. Miridae with type 1 parempodia have two types of male genitalia — most genera have a membranous vesica, which may or may not possess sclerotized spiculi (orthotyline-type) ; a much smaller number has the phyline-type. The phyline-type male genitalia are structurally distinct from all others found in the Miridae. They also possess a unique and com- plex functional relationship. It is almost inconceivable that such a combination of structure and function could have evolved indepen- dently more than once and therefore all taxa possessing it must be placed in a single, derived group. The phyline-type of male geni- talia probably evolved from the less specialized orthotyline-type (Singh-Pruthi, 1925) which resembles that of the other subfamilies of mirids more closely than it does the phyline-type. Since type 1 recurved convergent parempodia occur in taxa with and without phyline-type male genitalia, it seems logical to believe that of the parempodial types discussed above, type 1 is the ancestral condition and types 2 and 3 represent derived states which arose from an- cestors with type 1 parempodia and phyline-type male genitalia. All previous authors have defined the Orthotylinae as those Mi- ridae with apically convergent parempodia. As can be seen from the 1974] Schuh: South African Orthotylinae and Phylinae 269 above discussion, this definition does not recognize the different types of convergent parempodia (types 1 and 3) and therefore brings to- gether mirids with two types of male genitalia. Because the phyline- type male genitalia are derived, all of those genera possessing them must be placed in one higher category to form a monophyletic group. I am therefore redefining the Orthotylinae to include only those gen- era with type 1 parempodia and orthotyline-type male genitalia. This definition excludes the Pilophorini, which were placed in the Orthotylinae by all previous authors, and also several genera which have type 3 parempodia; all of these excluded taxa belong to the Phylinae. As redefined, the Phylinae now include all genera with the derived phyline-type of male genitalia but with all three types of parempodia discussed above. SUBFAMILY ORTHOTYLINAE Diagnosis: Facies, coloration, and vestiture variable; sometimes ant mimetic; females occasionally and males less often brachypter- ous; pronotum sometimes with a flattened or rounded collar; par- empodia fleshy, convergent apically, recurved (lyre-shaped), and flattened laterally; pulvilli minute, attached to ventral surface of claws; vesica membranous, inflatable to at least a limited degree, sometimes with long sclerotized spiculi apically; phallotheca fixed to phallobase, claspers variable, left usually larger than right; female with posterior wall varying from a simple sclerotized plate to a highly modified form with K-structures (Figure 109); sclerotized rings ranging from nearly flat to highly infolded on lateral margins. Discussion: All Orthotylinae have apically convergent, re- curved parempodia. The male genitalia have a membranous vesica with or without sclerotized spiculi. The claspers show some tribal characters, but in general the male genitalia are not indicative of distinct phyletic trends within the subfamily. The female genitalia in the Orthotylinae present a somewhat different evolutionary picture than those of the male and are very useful in recognizing phyletic lines within the subfamily. There are two basic types: 1) those with the posterior wall usually relatively simple and plate-like and with the sclerotized rings varying from flat to somewhat upturned laterally; and 2) those with a highly specialized posterior wall possessing K-structures and with the scle- rotized rings strongly upturned laterally. The simple plate-like posterior wall (type 1 above) probably represents the plesiomorphic condition in the Miridae because it is present in some of the Orthotylinae and in at least five other sub- 270 Entomologica Americana [Vol. 47 TABLE 3 Characters used in tribal classification of Orthotylinae CHARACTER PRIMITIVE DERIVED Parempodia fleshy, convergent apically, recurved all genera Posterior wall without K-struc- tures : Halticini, Nichomachini with K-structures: Orthotylini Vesica membranous : Halti- cini, Nichomachini often with sclero- tized spiculi: Orthotylini Pronotal collar absent : many Orthotylini present: Falconia, Nanniella Lateral corial margin convex or straight: most genera sinuate: Nichomachini, Pseudopilophorus, Sericophanes- group Punctations on dorsum absent in most genera present: Nanniella, Falconia Body form robust: most genera very elongate: Aetorrhinella, Felisacodes Vestiture setiform hairs: most genera scale-like hairs: Melanotrichus Wing dimorphism both sexes macrop- terous: most Orthotylini females brachyp- terous : Serico- phanes, Nichomachus; both sexes brachyp- terous: many Halti- cini, Laurinia families. The posterior wall with K-structures is without question a derived condition as it occurs in only a limited group of mirids (Orthotylini), all of which possess type 1 parempodia. The Pilo- phorini, which I am placing in the Phylinae (see above and also tribal discussion), have type 1 parempodia which relate them to the Orthotylinae; they also have a relatively simple posterior wall, as in some Orthotylinae, but have highly specialized male genitalia. Therefore, the ancestral orthotyline stock must have had a simple posterior wall and type 1 parempodia. The Phylinae must have di- verged from this line relatively early and the posterior wall became specialized within the Orthotylinae subsequently. 1974] Schuh: South African Orthotylinae and Phylinae 271 o & o &' W D5 n If n P3 CD Fig. 348. Phylogeny of the Orthotylinae. Variation in the posterior wall in the Orthotylinae is greater than in the remainder of the Miridae. The great dissimilarity in type of female genitalia in those mirids with type 1 parempodia, particu- larly in the structure of the posterior wall, suggests that a case can be made for placing the genera with K-structures ( Ortho tylini) in a distinct subfamily apart from the remaining genera. Based on the stability of the parempodia and male genitalia, however, I prefer to treat the Orthotylinae as a monophyletic group of subfamilial rank. I recognize three tribes within the Orthotylinae — Halticini, Nicho- machini, and Orthotylini. All of the Nichomachini and some Hal- ticini and Orthotylini are ant mimetic. Table 3 lists characters important in the tribal classification of the Orthotylinae. They are categorized as primitive and derived. Where a character state has evolved independently more than once, several unrelated genera that posses the character are noted. A pro- posed phylogeny of the subfamily is given in Figure 348. Zoogeography: The Orthotylinae appear to be nearly cosmo- politan in distribution in the Carvalho system of classification. With the revised tribal classification presented below, certain definite dis- 272 Entomologica Americana [Vol. 47 Fig. 349. Distribution of the Halticini, Orthotylini, and Nichomachini (see text for explanation). 1974] Schuh: South African Orthotylinae and Phylinae 273 tributional patterns emerge. The removal of the Pilophorini to the Phylinae is significant in this respect. The most distinctive feature of the zoogeography of the Ortho- tylinae is the preponderance of advanced genera (Orthotylini) in the New World, including nearly all of the ant-mimetic genera in the subfamily (Figure 349c). A much smaller number of advanced orthotylines occurs in the Old World, but the relatively primitive groups in the subfamily (Halticini and Nichomachini) are virtually restricted to the Eastern Hemisphere. Australia has a number of rather interesting genera, the most notable of which is Myrmecor- oides Gross, 1963, which by virtue of its bizarre morphology, espe- cially of the head, shows a probable long isolation in Australia. When better known Myrmecoroides will probably be placed in a new tribe. The occurrence of Myrmecoroides and other isolated genera in Aus- tralia indicates that that faunal region may be of particular interest in understanding the evolution of the Orthotylinae. TRIBE HALTICINI Diagnosis: Usually black or dark colored, sometimes with lighter markings; body usually robust, sometimes elongate; dorsum usually smooth, often polished and shining, seldom with heavy punc- tures; vestiture variable, occasionally dense and very long; head usu- ally dorsoventrally elongated, height of gena greater than height of eye; pronotum and scutellum seldom highly modified (except in Myrmecophyes)’, often brachypterous (sometimes males as well as females); metafemora often greatly enlarged; vesica membranous, without spiculi; left clasper usually elongate with small hook apically; right clasper flattened and expanded apically, usually spoon-shaped or club-shaped; posterior wall of female without K-structures, al- though sometimes specialized ( Labops)\ sclerotized rings variable; other structural features as in Orthotylinae. Discussion: The most useful structures for recognizing the Hal- ticini are: 1) the recurved convergent parempodia; 2) an apically enlarged right clasper; 3) a posterior wall lacking K-structures; 4) the elongated genae; and 5) the enlarged metafemora. Although the Halticini are often difficult to separate from the Orthotylini on external characters, the posterior wall will always distinguish the two groups (Wagner, 1956), as will usually the right clasper. Reuter (1910a) and Carvalho (1952a) did not include these genital char- acters in their definition of the tribe and consequently certain genera were either incorrectly excluded from (e.g. Myrmecophyes ) or in- cluded in (e.g. Slaterocoris Wagner) the Halticini. 274 Entomologica Americana [Vol. 47 Much recent discussion has centered around the possible poly- phyletic character of the Orthotylinae and particularly the Halticini (Slater, 1950; Leston, 1961). The variability of the posterior wall has been the major cause for this concern. I consider the relative uniformity of the head, with its elongate genae, and the peculiar club-shaped right clasper as apomorphic characters that bind the Halticini together. The posterior wall can be viewed as an inher- ently variable structure within the Halticini which gave rise to the highly derived K-structures in the Orthotylini. The Halticini are morphologically somewhat isolated in the Or- thotylinae with their peculiar head structure, greatly enlarged meta- femora, and generally jet black coloration. Even though they pos- sess these specialized features, they are probably still primitive within the subfamily as judged primarily on the simplicity of the posterior wall and sclerotized rings in most genera. The Halticini are most closely related to the Nichomachini, which have a simple posterior wall (Figure 98), but very peculiar sclerotized rings (Figure 99). The ancestral stock of the tribe probably originated early in the evolution of the Orthotylinae and became specialized subsequently. Only the genus Myrmecophyes Fieber is ant mimetic. Zoogeography: The Halticini are most diverse in the Pale- arctic (Figure 349a), particularly in the Mediterranean region, but lack of modern generic definitions may misrepresent the actual num- ber of genera involved. The cosmopolitan Halticus occurs in both tropical and temperate regions, including areas of high rainfall, and is the only genus that is widespread. Most other halticines are adapted to areas of Mediterranean climate which are relatively dry. The tribe is not well represented in many areas of the world with a Mediter- ranean climate, which suggests low dispersal ability and limited adaptibility or possible replacement by more advanced groups. Discussion Of Individual Genera. Myrmecophyes Fieber, 1870. Carvalho (1952a; 1958b) placed Myrmecophyes in the Pilo- phorini because of its ant-mimetic facies and convergent parem- podia. Wagner (1952; 1955) recognized the halticine character of the head and male genitalia and moved the genus to the Halticini. Approximately 16 species are described, primarily from the Eastern Mediterranean. * Strongylocoris Blanchard, 1840 (in part), see Slaterocoris Wagner (Orthotylini). 1974] Schuh: South African Orthotylinae and Phylinae 275 NICHOMACHINI, NEW TRIBE Diagnosis: Ant mimetic; dark colored; males usually macrop- terous, elongate with very slender abdomen, in brachypterous forms abdomen similar to females; hemelytra with two or three partial, light, transverse fasciae; females brachypterous, abdomen strongly constricted basally and swollen medially, hemelytra greatly reduced, covering only two basal abdominal segments; head nearly vertical, concave behind, including eyes, posterior margin of vertex usually carinate; antennal segments 3 and 4 equal to or slightly greater in diameter than segment 2; pronotum usually with distinct anterior and posterior lobes or with anterior lobe only poorly differentiated, posterior lobe in macropterous forms tumid; scutellum elevated, bluntly conical or only convex; hemelytra in macropterous forms exceeding apex of abdomen; lateral corial margins weakly to rather strongly sinuate; membrane with two cells; vesica membranous (Fig- ure 94) or with sclerotized bands (see Wagner, 1957b); left clasper with spine-like group of stiff hairs on basal lobe, shaft slender, apex barbed (Figure 95); right clasper greatly reduced; posterior wall of female simple, lacking K-structures (Figure 98); sclerotized rings very small, contorted (Figure 99). Discussion: The African genera Nichomachus Distant, Pseudo- nichomachus Schuh, and Laurinia Reuter, form a distinct group, based on the structure of the claspers, the simple posterior wall, the peculiar sclerotized rings, general mimetic facies, and profound sex- ual dimorphism. I am therefore placing them in a new tribe. Eu- compsella Poppius and Kuomocoris Odhiambo, both from Mada- gascar, probably belong in the Nichomachini, but the genitalia have not been examined for either genus. Wagner (1957b) placed Laurinia in the Orthotylini because of the parempodial structure and the form of the male genitalia; he related the genus to Globiceps LePeletier and Serville. Additional characters provided by examination of the female genitalia of Nicho- machus, viz., the simple posterior wall and the very small, anoma- lous sclerotized rings, do not confirm this relationship, because Glo- biceps is a typical member of the Orthotylini and has well developed K-structures. The Nichomachini are probably most closely related to the Hal- ticini, where the posterior wall also lacks K-structures and is generally much simpler than in the Orthotylini (although see Slater, 1950; Wagner, 1955, 1956). The male genitalia of the Nichomachini can be derived from those of the Halticini, and are a specialization of the 276 Entomologica Americana [Vol. 47 halticine-type. The right clasper in the Nichomachini is greatly reduced, a situation found in very few Orthotylinae. The Nichoma- chini have become very highly specialized in many features but rep- resent an old (and possibly relict) stock within the Orthotylinae that diverged from the main line of evolution before the development of K-structures. Zoogeography: At present the Nichomachini are known only from the Ethiopian Region, including Africa and Madagascar (Fig- ure 349b). Their greatest diversity is in South Africa. All known species are ground living and probably adapted to dry areas. Discussion Of Individual Genera. Eucompsella Poppius, 1914a. Eucomp sella is related to the Nichomachini by 1) the structure of the pulvilli; 2) the structure of the head and pronotum; 3) the number of hemelytral fasciae; and 4) the distribution. Poppius (1914a) stated that he examined three male specimens of Eucomp- sella elegantula Poppius and that they were deposited in the Paris Museum. I was unable to locate these specimens in Paris, but at least one is in the Helsinki Museum and I am designating it as the lectotype of the species. It bears the following labels: “Museum Paris, Madagascar, Tananarive, Coll. Noualhier 1898”; uEucomp- sella elegantula n. gen. et sp., B. Poppius det.”; “Mus. Zool. H: fors, Spec. typ. No. 7777, Eucompsella elegantula Popp.”; and “LECTO- TYPE, Eucompsella elegantula Poppius, det R. T. Schuh”. Poppius’ (1921) dorsal view drawing is not accurate (see also discussion under Kuomocoris) . Kuomocoris Odhiambo, 1967, pp. 1683-1687. Kuomocoris was placed in the Pilophorini with considerable reservation by Odhiambo (1967). He noted that the male genitalia were not phyline, but did not illustrate or describe them. After ex- amining the holotypes of K. rabalus Odhiambo and K. rubellus Od- hiambo, I believe they are closely related to the Nichomachini. This placement is strengthened by the structure of the head, pronotum, and hemelytra, including the transverse fasciae, the narrow, basally constricted abdomen, and the form of the parempodia. The scu- tellum is only convex and not conical as in Nichomachus and Pseudo- nichomachus. The male genitalia are missing from the holotype of K. rubellus and those of K. rabalus have not been dissected. Kuo- mocoris is known only from Madagascar and is very closely related to Eucompsella. 1974] Schuh: South African Orthotylinae and Phylinae 277 Laurinia Reuter, 1884. Carvalho (1952a) placed Laurinia Reuter in the Herdoniini (Mirinae) and Formicocoris Lindberg in the Pilophorini (Ortho- tylinae). Lindberg (1956) synonymized the two genera. Wagner (1957b) reviewed the systematic position of Laurinia and on the structure of the parempodia and male genitalia related it to Globiceps in the Orthotylini. The male and female genitalia in the Nichoma- chini are unique and do not show the close relationship to the Ortho- tylini that was suggested by Wagner (1957b). Only a single species is presently included in Laurinia , L. fugax Reuter, from North Africa (see Wagner, 1957b). Nichomachus Distant, 1904a, see page 29. Pseudonichomachus Schuh, new genus, see page 35. TRIBE ORTHOTYLINI Diagnosis: Facies, coloration, and vestiture variable; sometimes ant mimetic; females occasionally brachypterous; dorsum seldom punctured heavily; pronotum occasionally with flattened collar, less often with rounded collar; cuneal fracture very rarely absent ( Sula - mita Kirkaldy); male genitalia with an inflatable membranous ves- ica, with or without long sclerotized spiculi; left clasper usually larger than right; female genitalia with K-structures (Figure 109); lateral margins of sclerotized rings usually strongly infolded (Figure 112); other structural features as in Orthotylinae. Discussion: The single most distinctive feature uniting the Or- thotylini is the presence of K-structures on the posterior wall of the female genitalia. The strong infolding of the lateral margins of the sclerotized rings is also useful, but this condition occurs in an al- most equally advanced state of development in some Phylinae. Char- acters helpful in separating generic groups within the tribe are: 1 ) the presence or absence of spiculi on the vesica; 2) the presence or absence of a pronotal collar; 3) the ant-mimic facies; 4) the length of the ovipositor; 5) the presence or absence of punctation on the dorsum; and 6) the hyaline or opaque hemelytra. The variabil- ity of all of these characters is very poorly understood. Therefore, a generic revision of the tribe including the use of those characters listed above, as well as a search for new characters, is badly needed. This becomes obvious in the following generic group analysis. There have probably been several independent evolutions of ant mimicry within the Orthotylini, but the phenomenon is still very 278 Entomologica Americana [Vol. 47 poorly understood from a phylogenetic viewpoint, primarily because of the lack of material from the New World tropics where the great- est number of Orthotylini ant-mimic genera occur. The Sericophanes group (see below) is the only evolutionary line in the tribe in which all members are ant mimetic. The Orthotylus group. This group, the largest in the Orthotylini, includes genera that can be most easily distinguished by well devel- oped, long, heavily sclerotized spiculi on the vesica. Other characters, including the claspers, labial length, pubescence, and color, are all extremely variable. Most of the genera have an Orthotylus- type facies, but a few, primarily from the Palearctic, are ant mimetic. Included genera are (genera in parentheses have not been dissected, but appear very close to genera that have been) : Aetorhinella Noual- hier, Aserymus Distant, Bifidungulus Knight, Blepharidopterous Kolenati, Brachynotocoris Reuter, Canariocoris Lindberg, Cyllecoris Hahn, Cyrtorhinus Fieber, Cyrtotylus Bergroth, Diaphnidia Kelton, Dryophilocoris Reuter, Erythrocorista Lindberg, Excentricus Reuter, Ficinus Distant, Fieberocapsus Carvalho and Southwood, Globiceps LePeletier and Serville, Hadronema Uhler, Heterocordylus Fieber, Heterotoma LePeletier and Serville, Hyoidea Reuter, Ilnacora Reu- ter, Ilnacorella Knight, ( Kalania Kirkaldy), ( Kamehameha Kirk- aldy), ( Koanoa Kirkaldy), Labopidea Uhler, Lopidea Uhler, Mara- lauda Distant, Mecomma Fieber, Orthotylus Fieber, Pachylops Fieber, Parthenicus Reuter, Platycranus Fieber, Pseudambonea Schuh, Pseudoclerada Kirkaldy, Pseudoloxops Kirkaldy, Pseudopilo- phorus Schuh, Pseudopsallus Van Duzee, Reuteria Puton, and ( Thermus Distant). A number of tribes have been proposed for groups of genera that I am including in the Orthotylus group. The most important from a phylogenetic viewpoint are discussed below. Zimmerman (1948) proposed the tribe Pseudocleradini for the endemic Hawaiian genus Pseudoclerada. Although Pseudoclerada has a peculiar facies, the male and female genitalia are clearly of the type found in the Orthotylus group. This superficially unique island genus certainly represents only a morphologically specialized segment of the main Orthotylini stem. Although the head and body shape are unlike that of most other members of the tribe, but re- semble closely the predatory lygaeid Clerada , the K-structures of the female and the vesical spiculi relate Pseudoclerada closely to Orthotylus and its congeners. Pseudoclerada is very similar to Mara- lauda from the Seychelles, and may be closely related to it. Zimmerman (1948) also proposed the tribe Kalaniini for the 1974] Schuh: South African Orthotylinae and Phylinae 279 endemic Hawaiian genus Kalania. He placed it in the Bryocorinae. I have followed Carvalho (1952a) in considering Kalania an ortho- tyline. Examination of the type female suggests that the morpho- logical attributes of Kalania hawaiiensis Kirkaldy, the only species in the genus, are the result of extreme isolation in the Hawaiian Is- lands and that the genus is certainly derived from the main Ortho- tylini stem. Ant mimicry within the Orthotylus group is limited to a few gen- era, such as: Cyllecoris, Dryophilocoris , and Globiceps from the Palearctic; Pseudoxenetus from the Nearctic; Ficinus from Mexico; and Pseudopilophorus from South Africa. The disjunct distribution of these genera suggests multiple independent evolutions of ant mimicry, but verification of this must await a thorough morpholog- ical study of the Orthotylus group. The Falconia group. This small assemblage of genera can be recognized by the rounded pronotal collar, heavily punctured dor- sum, vesica without spiculi, and extremely short ovipositor. Included genera are: Adfalconia Carvalho, Falconia Distant, Falconiodes Reuter, and Solanocoris Carvalho from the Neotropical Region, and Sulamita from the Hawaiian Islands. It is probably most closely related to the Zanchius group, the most obvious difference being that all Falconia group genera are heavily punctured and those of the Zanchius group are not. Sulamita was placed in the tribe Sulami- tini (Sulamitaria Kirkaldy) by Zimmerman (1948) in the subfamily Bryocorinae; Carvalho (1952a) later moved the genus to the Ortho- tylini. The basic structure of members of the genus is very similar to Falconia. I have dissected the females of both Falconia and Sula- mita and confirmed the presence of K-structures. The cuneus is fused with the corium in Sulamita , a secondary development, which accentuates the coleopteroid appearance found in all members of the Falconia group. The Zanchius group. Diagnostic features of the group are: 1) the flattened appearance; 2) the very delicate body structure; 3) the usually hyaline hemelytra; and 4) the vesica without spiculi. In- cluded genera are: Brasiliomiris Carvalho, Felisacodes Bergroth, Hyalochloria Reuter, ltacoris Carvalho, Jobertus Distant, Malaco- coris Fieber, Parachius Distant, Paraproba Distant, Pliniella Berg- roth, Zanchius Poppius, and Zanchiella Schuh (also probably Zo- nodorellus Poppius and Zonodoropsis Poppius). The distribution of the Zanchius group is more or less pantropical with the greatest diversity occurring in the Neotropics. Generic limits appear to be 280 Entomologica Americana [Vol. 47 poorly understood. It is very difficult to find definitive characters on which to base genera, a situation which results mainly from the very delicate body structure and small size of the members of the group. The Sericophanes group. This group forms the major ant-mimic complex within the Orthotylinae. It can be recognized by the pres- ence of a more or less well developed pronotal collar, at least some degree of ant resemblance (females often brachypterous and much more ant-like than macropterous males), spiculi usually absent from the vesica, and ordinarily some type of hemelytral maculae or fascia. Included genera are: Borgmeierea Carvalho, Ceratocapsus Reuter, Eucerella Poppius, Hallodapoides Carvalho, Laemocoridea Poppius, Lepidotaenia Poppius, Pamilia Uhler, Pilophoropsis Poppius, Reno- daeus Distant, Schaffneria Knight, Sericophanoides Carvalho, Seri- cophanes Reuter, and Tuxenella Carvalho. The genera Ceratocapsus , Pamilia , Pilophoropsis, and Schaff- neria, were placed in the tribe Ceratocapsini by Knight (1968). Carvalho (1952a) placed Ceratocapsus and Pamilia in the Ortho- tylini and Pilophoropsis in the Pilophorini. Kelton (1959b) showed that Ceratocapsus lacks vesical spiculi and is therefore very similar to Sericophanes, which Carvalho (1952a) and Knight (1968) placed in the Pilophorini. Kelton (1959b) also showed that the vesica of Pamilia has spiculi and that it is therefore closely related to Hadro- nema, Slaterocoris, and other members of the Orthotylus group, and thus its resemblance to Sericophanes may be one of convergence. I have not examined the male genitalia of Pilophoropsis (and I have not seen Schaffneria), but externally Pilophoropsis appears very closely related to Sericophanes. The females of Sericophanes are brachypterous and ant-like whereas those of Ceratocapsus are mac- ropterous and much less ant-like. The placement of Pamilia and Ceratocapsus in the Orthotylini by Carvalho (1952a) is evidence of the subjective character of tribes defined on ant-mimetic facies alone, because Pamilia behrensi Uhler (and certain Ceratocapsus species) are nearly as ant-like as some species of Sericophanes, which Carvalho (1952a) placed in the Pilophorini. The type of ant mimicry found in the Sericophanes group is not as morphologically sophisticated as in many hallodapine genera (Phylinae), particularly in that no Orthotylini genera have the head convex behind (as in the Formicopsella group of the Hallodapini), but always concave and contiguous with the anterior margin of the pronotum. In some genera, e.g. Renodaeus, the general body form 1974] Schuh: South African Orthotylinae and Phylinae 281 is extremely similar to that of Pilophorus , showing a remarkable con- vergence between the Orthotylinae and Phylinae. The Sericophanes group is the only unit in the Orthotylini which exhibits pronounced brachyptery similar to the type found in the Phylinae, where the males are always macropterous and the females brachypterous (if short winged individuals exist). Most genera are not well known, but in Sericophanes, for example, it appears that the females are always brachypterous and much more ant-like than the males. This group is primarily Neotropical with a limited representation in the eastern and southwestern United States. Carvalho (1952a) synonymized Xenofulvius Bergroth from Luzon, Philippines, with Ceratocapsus. This action needs verification as no others members of the group are known from outside the Western Hemisphere. Genera placed in the Orthotylini by Carvalho and subsequent authors that I have not examined or for which adequate information is lacking to determine subfamily placement include: Acroderrhis Bergroth, Campylotropis Reuter, Compsoscytus Reuter, Cysteorracha Kirkaldy, Deleapidea Knight, Dichaetocoris Knight, Druthmarus Distant, Hadronemidea Reuter, Hyporhinocoris Reuter, Lopidella Knight, Macrotyloides Van Duzee, Melanostictus Reuter, Mesotropis Reuter, Noctuocoris Knight, Pseudoneoborus Knight, Rhinocapsi- dea Reuter, Squamocoris Knight, Sthenaridea Reuter, and Uleana Carvalho. Zoogeography: The Orthotylini show a marked concentration in the New World (Figure 349c). The Falconia group and the Seri- cophanes group are primarily restricted to the neotropics. The Zan- chius group is pantropical and widely distributed in the Pacific Is- lands, suggesting great dispersal ability. The Orthotylus group is the most generally distributed in the Orthotylini and it is also prob- ably the least specialized morphologically. Although the Orthotylini have radiated as ant mimics in the neo- tropics, and to a lesser extent in the Nearctic, mimetic forms are virtually absent in the Old World tropics. The opposite situation obtains in the Pilophorini (except for North America), Leucophor- opterini, and Hallodapini, which are essentially absent from the New World, but diverse in the Old World. Based on the mimics in par- ticular, it appears that the Orthotylini have undergone long isolation in the neotropics and that even though the tribe is represented in the Old World (primarily by nonmimetic forms) the earlier mimetic radiation of the Phylinae in the Old World precluded radiation of 282 Entomologica Americana [Vol. 47 the Orthotylini as ant mimics in the Eastern Hemisphere (see also discussion under Phylinae). Discussion Of Individual Genera. Many genera included in the Orthotylini by Carvalho do not belong there and must be moved to other tribes and subfamilies. Also many genera placed in the Pilophorini by Carvalho are cor- rectly placed in the Orthotylini. Borgmeierea Carvalho, 1956c, pp. 235-237. When he described Borgmeierea from Natal, Brazil, Carvalho (1956c) related it to Lepidotaenia, Renodaeus , and Pilophoropsis. He placed the genus in the Pilophorini. Although Carvalho did not illustrate the genitalia, his dorsal view drawings indicate a relation- ship of Borgmeierea to Sericophanes , as well as to the above men- tioned genera. The type of parempodia, general facies, and occur- rence in South America strengthen the probable affinities of this genus even though the genitalic information is not available. Under my redefinition of the tribes of the Orthotylinae, Borgmeierea is a member of the Orthotylini. The genus is known only from a single species from Brazil. * Bunsua Carvalho, 1951b, Bryocorinae, see misplaced genera. Coriodromus Signoret, 1862. This genus closely resembles N esidorchestes Kirkaldy from Ha- waii, which was placed in the Halticini by Carvalho (1952a). Study of the male and female genitalia is needed to determine of these two genera are closely related and to which tribe Coriodromus actually belongs. Coriodromus occurs only in the Southwest Pacific and Australia. * Ellenia Reuter, 1910a, Phylini, see page 157. Erythrocorista Lindberg, 1958, pp. 107-109. Erythrocorista Lindberg was incorrectly placed in the Phylinae by Lindberg (1958). The parempodia are plainly fleshy, convergent, and recurved and the male genitalia are not of the phyline-type but of the orthotyline-type. Lindberg (1958) designated E. echii Lind- berg as the type species of the genus. I have examined specimens from the Helsinki Museum labeled as holotype (Type No. 11109) and allotype (Type No. 11110) of echii. Each pin bears three spec- imens with no indication as to which specimen is the type. It is there- fore necessary to designate a lectotype. The situation is additionally 1974] Schuh: South African Orthotylinae and Phylinae 283 confused because the pin bearing the holotype label has two males and one female on it; Lindberg (1958) indicated that the holotype was a male. Therefore I have placed a male specimen with the or- iginal locality label of Lindberg, a “holotypus” label, and the iden- tification label on a separate pin and labeled it “LECTOTYPE Erythrocorista echii Lindberg, det. R.T. Schuh” and relabeled the remaining specimens. Eucerella Poppius, 1921. The structure of the parempodia and its occurrence in South America strengthen the placement of this genus in the Orthotylini, rather than in the Pilophorini as by Carvalho (1952a). The struc- ture of the head (concave behind), narrow flattened pronotal collar, and mimetic facies ally Eucerella at least provisionally, with the Seri- cophanes group. Eucerella is known only from Bolivia. The holotype of Eucerella hirtipes Poppius, the only available specimen for the genus, is not in the Paris Museum, as stated by Poppius (1921), but in the Helsinki Museum (Type No. 7781). Hallodapoides Carvalho, 1951a. In his original description, Carvalho (1951a) referred Hallodap- oides to the Pilophorini. Subsequently he moved it to the Hallo- dapini (Carvalho, 1958a), but gave no explanation for this action. Carvalho’s (1951a) illustrations of the male genitalia indicate that Hallodapoides is most closely related to Sericophanes and allied gen- era. This relationship is confirmed by the structure of the parem- podia, the general facies, and the distribution. Hallodapoides con- tains only a single species, H. guaraniensis Carvalho, from Paraquay. * Hypseloecus Reuter, 1891, see Pilophorini. * ldiomiris China, 1963, see genera incertae sedis. Kirkaldyella Poppius, 1921. I have examined the male genitalia and parempodia of Kirkaldy- ella rugosa Poppius, and place Kirkaldyella in the Orthotylini, based on these characters. Carvalho (1952a) considered the genus to be in the Pilophorini. The general appearance is not particularly ant- like. A male of K. rugosa from Sydney, New South Wales, Australia, is deposited in the Helsinki Museum (Type No. 12106). Poppius (1921) indicated that there is also a male in the Hungarian Museum. This latter specimen will have to be examined before a lectotype can 284 Entomologica Americana [Vol. 47 be designated. I have also seen an undescribed species of Kirkaldy- ella from Borneo. Laemocoridea Poppius, 1921. Laemocoridea is most closely related to the Sericophanes group based on the flattened pronotal collar, male genitalia, and general facies. I have examined the holotype male of L. quadrimaculata Poppius, which is deposited in the Helsinki Museum (Type No. 7784), rather than in the Paris Museum, as indicated by Poppius (1921). Lepidotaenia Poppius, 1921. Lepidotaenia is probably most closely related to Tuxenella , based on the upturned, carinate anterior margin of the pronotum, and the male genitalia of L. bergrothi Poppius. This species has two trans- verse bands of lepidote hairs on the hemelytra, similar to the type found in Pilophorus. The pronotum in bergrothi is constricted med- ially, forming a distinct anterior and posterior lobe. Two species of Lepidotaenia are known from Bolivia. I have examined the holotype male of L. bergrothi, which is de- posited in the Helsinki Museum (Type No. 7779), rather than in the Paris Museum as stated by Poppius ( 1921 ) . * Millerimiris Carvalho, 1951b, see Phylini. * Nanniella Reuter, 1904, Halticini, see page 28, 511 Orthotylellus Knight, 1935, see Paramixia Reuter, Pilophorini, see page 210. Pamilia Uhler, 1887, see discussion under Sericophanes group. Five species of Pamilia are known from the eastern and south- western United States. * Parasthenaridea Miller, 1937, see Pilophorini. Pilophoropsis Poppius, 1914c, see discussion under Sericophanes group. Three species of Pilophoropsis are known from Arizona. * Platyscytus Reuter, 1907a, see Phylini. Pseudoxenetus Reuter, 1909. The female genitalic studies of Slater (1950) and male genitalic studies of Kelton (1959b) correctly established the position of Pseudoxenetus in the Orthotylini, rather than in the Pilophorini, as 1974] Schuh: South African Orthotylinae and Phylinae 285 placed by Carvalho (1952a). Two species of Pseudoxenetus are known from the eastern United States. Renodaeus Distant, 1893. Although the general facies of Renodaeus are very much like those of Pilophorus, the male genitalia (see Carvalho and Becker, 1959) confirm that the genus belongs to the Orthotylini and is a member of the Sericophanes group. Three species are known from Texas, Guatemala, and Guyana. Distant (1893) described Renodaeus ficarius from two female specimens. I have designated as the lectotype a specimen in the British Museum (Natural History) bearing the labels: “Cerro Zunil, 4-5,000 ft., Champion”; “sp. figured”; “ Renodaeus ficarius Dist.”; and “LECTOTYPE Renodaeus ficarius Distant, det. R.T. Schuh.” Schaffneria Knight, 1966, see discussion under Sericophanes group. One species is known from Texas. * Semium Reuter, 1876a, see Phylini. Sericophanes Reuter, 1876a. Kelton (1959b) confirmed the relationship of Sericophanes to the Orthotylinae on the basis of the male genitalia which he con- sidered as related to Ceratocapsus. I have examined the female genitalia of S. heidemanni Poppius, which has well developed K- structures; therefore the genus must be placed in the Orthotylini (see also discussion under Sericophanes group). Sericophanes pres- ently includes 20 species, all from the New World, and shows its greatest radiation in the tropics (Maldonado, 1970). Sericophanoides Carvalho and Fonseca, 1965, pp. 53-57. Although placed in the Pilophorini by Carvalho and Fonseca (1965), Sericophanoides is closely related to Sericophanes by the general facies, and the form of the male genitalia and belongs to the Orthotylini. Two species are known from South America. Slaterocoris Wagner, 1956, pp. 277-281. Primarily on characters of the female genitalia, Wagner (1956) recognized the distinctive nature of the North American species pre- viously placed in Strongylocoris. He erected for them the new genus Slaterocoris , belonging to the Orthotylini. This was the first taxo- nomic use of the K-structure, the importance of which was pointed out by Slater (1950). Sulamita Kirkaldy, 1902a, see Falconia group discussion. 286 Entomologica Americana [Vol. 47 Tuxenella Carvalho, 195 2d. Carvalho and Dutra (1959) illustrated the male genitalia of Tuxenella which confirm the placement of the genus in the Ortho- tylini, although Carvalho (1952a, etc.) placed the genus in the Pilo- phorini. This genus probably belongs to the Sericophanes group, but it does not have the well developed pronotal collar of most gen- era in that group and has a more complex vesica with spiculi. Nine species are known from Chile. SUBFAMILY PHYLINAE Diagnosis: Facies, coloration, and vestiture variable; sometimes ant mimetic; males always macropterous, females often brachypter- ous; pronotum sometimes with a flattened collar; parempodia either 1) fleshy, convergent apically, recurved (lyre-shaped) and flattened laterally, 2) fleshy, rod-like, of nearly uniform diameter, and weakly convergent apically, or 3) hair-like and parallel; pulvilli usually minute, always attached to ventral surface of claw, sometimes en- larged and either attached only at base or over entire ventral sur- face of claw; male genitalia distinctive; vesica elongate, sclerotized, rigid (Figure 228); gonopore variable, phallotheca not fixed to phal- lobase; left clasper always larger than right, trough-like and receiving apex of phallotheca in repose (Figure 222); right clasper flattened, leaf-like (Figure 234); female genitalia with simple undifferentiated posterior wall, sometimes with posterior margin evaginated (Figure 316); sclerotized rings usually slightly to rather strongly infolded laterally. Discussion: Carvalho (1952a) defined the Phylinae as those mirids with hair-like parallel parempodia and with the pulvilli at- tached to the inner surface of the claws (see discussion on page 264). He recognized three tribes within the subfamily— Phylini, Hallo- dapini, and Dicyphini. Kelton (1959b) showed that on the basis of the male genitalia the Dicyphinae are unrelated to the Phylinae and that the Pilophorini are much more closely related to the Phy- linae than to the Orthotylinae, where they were placed by Carvalho (1952a; 1958b). The hair-like parempodia of the Phylinae are derived from the convergent parempodia of the Orthotylinae (see above). The par- empodia of the Dicyphinae, although similar to those of the Phy- linae, are of an independent origin and may be the ancestral type in the Miridae. This view is supported by the dicyphine male gen- italia which have a membranous, inflatable vesica, more similar to 1974] Schuh: South African Orthotylinae and Phylinae 287 that of other mirids than to the Phylinae. Considering the Dicy- phinae as closely related to the Phylinae requires the derviation of of the dicyphine-type male genitalia from the phyline-type or vice versa. The former situation requires a reevolution of the general- ized dicyphine-type from the highly specialized phyline-type; the latter requires the independent evolution of convergent recurved par- empodia in both the Pilophorini and Orthotylinae sensu novo and the evolution of the phyline-type male genitalia from the dicyphine- type. Both of these alternatives are less parsimonious and require more unlikely evolutionary events than does acceptance of the dicy- phine and phyline parempodia as independently evolved. Kelton (1959b) indicated no relationship between the Dicyphinae and Phylinae on the basis of the vesica, but proposed an affinity of the Dicyphinae with the Deraeocorinae and Cylapinae. Evidence of this relationship is further strengthened by the presence of a rounded pronotal collar (which is absent in the Phylinae) and hair-like par- empodia; the Deraeocorinae and Cylapinae lack pulvilli, which the Dicyphinae have, however. Slater (1950) related the Dicyphinae to the Phylinae on the structure of the female genitalia. This rela- tionship, however, is based on what I consider to be primitive char- acters, since the simple plate-like posterior wall of the Dicyphinae also occurs in the Isometopinae, Cylapinae, Deraeocorinae, and Phylinae. Therefore, the Dicyphinae are not closely related to the Phylinae and must be placed in a separate subfamily. This is the status given the group by Knight (1941; 1968) and other authors. The Dicyphinae are possibly related to the Monalionini (Bryocori- nae) as noted above. The convergent recurved parempodia of the Pilophorini relate them to the Orthotylinae, but, as discussed above, the Pilophorini have the unique phyline-type male genitalia. I therefore place them in the Phylinae and consider them to be among the most prim- itive members of the subfamily. The female genitalia of the pilo- phorines are specialized relative to the rest of the Phylinae, an evo- lutionary event which must have taken place subsequent to the divergence of the nonpilophorine Phylinae. The type 3 rod-like convergent parempodia (see above) have apparently evolved independently several times in the Phylinae and represent a specialized condition within the subfamily. The derived nature of the type 3 parempodia relative to type 2 (hair-like), can be established with some certainty because they occur in all tribes and often in genera that possess many derived characters. The type 2 parempodia could also be considered intermediate between types 288 Entomologica Americana [Vol. 47 TABLE 4 Characters used in tribal classification of Phylinae CHARACTER PRIMITIVE DERIVED Head behind Antennae Labium Frons Eyes Anterior margin of pronotum Pronotum Scutellum Lateral corial margins Wing dimorphism Vestiture Rows of minute tibial spines Length tarsal segment 1 concave : all Pilo- phorini, most Leucophoropterini segments 3 and 4 slender: most genera long: most genera with transverse rugosities: most genera glabrous: genera in all tribes finely carinate: Phylini, Pilophor- ini, some Leuco- phoropterini constricted ante- riorly : most genera in all tribes flat: all tribes except Hallodapini straight or convex: all Phylini, most Pilophorini, some Leucophoropterini both sexes macrop- terous: all(?) Pilo- phorini, most Phylini setiform hairs: all Hallodapini, some genera in all tribes present: most genera in all tribes shorter than seg- ments 2 and 3: nearly all genera convex: many Phylini, some Hallodapini segments 3 and 4 en- larged: some Hallo- dapini and Leuco- phoropterini short: some genera in all tribes without transverse rugosities : some genera, all tribes hairy: genera in all tribes flattened collar: all Hallodapini, some Leucophoropterini hour-glass shaped: some genera in Pilophorini and Leucophoropterini protuberant or spini- form: some Hallo- dapini sinuate : most Hallo- dapini and Leuco- phoropterini, some Pilophorini females brachypter- ous: many Hallodap- ini and Leucophorop- terini, some Phylini wooly or scale-like hairs: some Phylini, Leucophoropterini, and Pilophorini absent : some genera in all tribes longer than segments 2 and 3: Cremno- cephalus and Myrmicomimus 1974] Schuh: South African Orthotylinae and Phylinae 289 TABLE 4 (continued) CHARACTER PRIMITIVE DERIVED Pulvilli minute : most genera in all tribes enlarged : Macrotylus, Eminoculus, Coquil- lettia, etc. Parempodia fleshy, convergent apically, recurved: Pilophorini hair-like: most genera of Phylini, Hallodapini, and Leucophoropterini ; sometimes weakly fleshy and conver- gent apically Abdomen broad: all Phylini, most Pilophorini slender, constricted basally: most Hallo- dapini and Leuco- phoropterini Vesica U-shaped, not twisted: all Pilophorini, some Leucophorop- terini S-shaped, twisted: some Leucophorop- terini, all Phylini and Hallodapini Phallotheca straight, opening terminal: all Pilo- phorini except Para- mixia L-shaped : all Leuco- phoropterini, Phy- lini, and Hallo- dapini Posterior wall simple plate, pos- terior margin not evaginated: all tribes except Pilophorini simple plate, pos- terior margin evagi- nated: Pilophorini Sclerotized rings more or less ellip- tical: most genera in all tribes shaped otherwise: Acrorrhinium 1 and 3, but this does not appear to be the case in the majority of the genera possessing them. I am recognizing four tribes within the Phylinae — Pilophorini, Phylini, Leucophoropterini and Hallodapini. Some Pilophorini, and all Leucophoropterini and Hallodapini, are ant mimetic. Table 4 lists characters that are important in the tribal classifi- cation of the Phylinae. For those characters in which the derived state has evolved more than once, several genera are listed. Figure 350 is a proposed phylogeny for the subfamily. Zoogeography: The most distinctive feature of the phyline 290 Entomologica Americana [Vol. 47 Fig. 350. Phylogeny of the Phylinae. distribution is the concentration of tribes in the Old World (exclu- sive of Australia) (Figures 351a, b, c, and d). Of the four tribes, only the Phylini are at all well represented in the New World. The Hallodapini and Pilophorini are poorly represented (except for Pilo- phorus, which has speciated in the Nearctic) and the Leucophorop- terini are absent from the Western Hemisphere. The Phylinae form a limited component of the ant-mimic mirid fauna in North America and are absent from the Neotropical Region, whereas they are the dominant mimetic mirids in the Old World. The Orthotylinae and Mirinae (Herdoniini) are the major mimetic mirid taxa in the New World. The Phylinae appear to be extremely numerous and diverse in the Palearctic, but this may result from the advanced state of tax- onomy in that region. In addition to being well collected and de- scribed, the Palearctic Phylinae are rather finely divided at the gen- eric level, so that the actual number of genera might be somewhat reduced if the same generic' concepts were applied in the Palearctic as currently exist in less well known regions. The Phylinae are most diverse in temperate areas, including the Nearctic, Palearctic, and southern Africa, and much less so in trop- ical regions. This may be an adaptation to temperate floras, but 1974] Schuh: South African Orthotylinae and Phylinae 291 Fig. 351. Distribution of the Leucophoropterini, Hallodapini, Pilo- phorini, and Phylini (see text for explanation). 292 Entomologica Americana [Vol. 47 the evidence for this is not available. The pattern is particularly evident in the Phylini (Figure 351c), but is probably important in the Leucophoropterini also (Figure 351b). The Hallodapini and Pilophorini also have conspicuous temperate elements (see tribal discussions). TRIBE HALLODAPINI Diagnosis: Ant mimetic; usually dark with contrasting light hemelytral maculae; pronotum always with well developed flattened collar (rounded in Cremnocephalus and Myrmicomimus)\ head us- ually convex behind, sometimes concave, eyes either contiguous with anterior margin of pronotum or removed from it by distance equal to nearly twice diameter of eye; pronotum always constricted an- teriorly, sometimes forming a long neck; scutellum often tuberculate or occasionally spiniform; hemelytra either straight or sinuate later- ally; abdomen usually constricted basally; parempodia hair-like, par- allel (in Cyrtopeltocoris weakly fleshy and convergent apically); pulvilli usually minute, occasionally long and free or fused to claws; vesica always S-shaped or otherwise bent, sometimes very long (Fig- ure 175); gonopore well developed, only rarely indistinct; left clasper always wedge shaped, trough-like; right clasper flattened and leaf-like; phallotheca always L-shaped; posterior wall a simple plate, never with evaginated posterior margin. Discussion: The Hallodapini are the largest ant-mimic group in the Miridae, consisting of about 41 genera. The morphological adaptations of the Hallodapini for ant mimicry are unexcelled in the Miridae, and probably the Insecta, and are equalled in the Mir- idae only by the Leucophoropterini and Nichomachini. Wagner (1952; 1970b), and other authors have given the hallodapines sub- family rank, a position that has little merit, because it is based only on the superficial uniqueness of the group. The Hallodapini ( Systellonotini and Cremnocephalaria of older authors) have generally included all Phylinae that are ant mimetic, have a flattened pronotal collar, and usually have light maculae on the hemelytra. To this definition must be added characters of the male genitalia and distribution, otherwise genera belonging to the Leucophoropterini will be included in the tribe. The Hallodapini as a tribe possess several characters that are derived relative to other members of the Phylinae and therefore appear to represent a monophyletic unit within the subfamily. These characters include: 1) the ant-mimic facies; 2) the flattened pro- notal collar; 3) the hemelytral maculae; and 4) the structure of the male genitalia. 1974] Schuh: South African Orthotylinae and Phylinae 293 The strongest indication that the Hallodapini are monophyletic is found in the flattened pronotal collar. Although the rounded pro- notal collar, as found in the Mirinae and other subfamilies, probably represents the ancestral condition in the Miridae, the flattened collar in the Hallodapini is almost certainly derived. This idea is supported by the fact that the Pilophorini, which are probably the most prim- itive members of the Phylinae, do not possess a pronotal collar. Also, the mode of evolution of the flattened collar from a pronotum with a finely carinate upturned anterior margin (as found in the Pilo- phorini, some Leucophoropterini, and most Phylini) can be traced in the Leucophoropterini (see tribal discussion), and suggests a possible mechanism for the evolution of the flattened collar in the Hallodapini from a more primitive phyline in which the anterior margin of the pronotum was finely carinate and upturned. The flattened collar is known to occur in only two genera of Phylinae in addition to the Hallodapini. They are Eminoculus from South Africa and an undescribed genus from Southeast Asia belonging to the Leucophoropterini; both of these taxa are specialized and, on the basis of additional derived characters, are more closely related to other evolutionary lines within the subfamily than to the Hallodapini. All hallodapine genera either possess hemelytral maculae or are obviously related to genera that do. Although maculae have evolved more than once in the Phylinae, they are of very limited occurrence outside the Hallodapini and Leucophoropterini (occurring, e.g. in Auchenocrepis in the Phylini; in Pilophorus in the Pilophorini a transverse fasica is formed by sericeous scale-like hairs), and those non-hallodapine stocks that possess them are related to other evo- lutionary lines in the subfamily by additional derived characters. The male genitalia of the Hallodapini are very similar struc- turally to those of the Phylini and much less so to those of the Pilo- phorini and Leucophoropterini. The tribe retains its monophyletic unity when the male genitalia are considered in conduction with the pronotal collar and hemelytral maculae. As outlined below, several lines of evolution can be traced in the Hallodapini. Each of the generic groups shows a strong affinity with at least one other generic group within the tribe, and thus all genera can be related to one another through other genera. The geographic distribution of the Hallodapini also strengthens the ar- gument for a monophyletic tribe, with the greatest diversity in Africa and the Palearctic and with only limited but obviously related ele- ments occurring in Southeast Asia and North America. The phylogenetic affinities of the Hallodapini within the Phylinae 294 Entomologica Americana [Vol. 47 are obscure because no intermediate types exist through which the tribe can be derived from other members of the subfamily. Now, the Leucophoropterini at least suggest a logical progression toward development of the flattened pronotal collar from a type that prim- itively had no collar; however, the Hallodapini are probably not evolved from the Leucophoropterini. They appear to be most closely related to the Phylini, but no known genus suggests itself as a pos- sible ancestral type. Six groups can be recognized within the Hallodapini. The Aeolocoris group. Wagner (1970b) erected the tribe Aelo- corini [sic] for the genera Aeolocoris Reuter and Acrorrhinium Noualhier. He based the tribe on what he felt was a significant dif- ference in the structure of the male genitalia between these genera and the rest of the Hallodapini, namely that the phallobase was heavily sclerotized apically and lightly sclerotized basally and that it was not firmly attached to the genital capsule. Wagner (1959) first described this condition for Saharocylapus vidali Wagner from Morocco. At that time he proposed a relationship between the Cy- lapinae and the Hallodapini based on the apparent absence of pul- villi in Saharocylapus and the similarity between the male genitalia of the Hallodapini and Cylapinae, viz. Saharocylapus and Para- fulvius Carvalho. Later Wagner (1970b) synonymized Saharocy- lapus with Aeolocoris , giving scant attention to his previous place- ment of the former genus in the Cylapinae. If the pulvilli are actually absent in Aeolocoris vidali this is a derived condition, in that all other known phylines have them. Wagner’s idea that the Hallodapini are related to the Cylapinae by the structure of the male genitalia is based on the incorrect subfamily placement of Parafulvius by Car- valho (1954) (see Phylini discussion of individual genera). Genera belonging to the Aeolocoris group are: Acrorrhinium Noualhier, Aeolocoris Reuter, Azizus Distant, Kapoetius Schmitz, Marmorodapus Schmitz, Syngonus Bergroth, and Trichophorella Reuter (and probably Bibundiella Poppius). These genera consti- tute the most primitive element in the Hallodapini based on the broad abdomen (narrowed in Trichophorella and Marmorodapus ), the absence of hemelytral fasciae (except in some species of Acrorrhin- ium and an undescribed species of Syngonus ), the relatively non- ant-like appearance, and the absence of brachyptery (except in some species of Acrorrhinium) . Although the group as a whole appears to have many primitive features, some included genera possess several derived characters, e.g. the spiniform frons and specialized vesica in Acrorrhinium. 1974] Schuh: South African Orthotylinae and Phylinae 295 The Aeolocoris group is held together by the presence of pecu- liar peg-like hairs on the dorsum and first antennal segment of most genera and the generally marmorate color pattern. Also, in certain genera, including Aeolocoris, Azizus, Marmorodapus (?), and Trich- ophorella, the eyes of the males are very much larger than those of the females. Although male Miridae generally have slightly larger eyes than the females, in Aeolocoris and the other genera, and also in some members of the Hallodapus group (see below), this dimor- phism is extreme. The Aeolocoris group is most diverse in tropical Africa, with all of the known genera occurring there. Only Azizus and Acrorrhinium are represented in Southeast Asia (the latter also occurring in north- ern Australia). The distributional pattern of the group agrees closely with that of the Hallodapini as a whole. The Hallodapus group. The following genera form a distinct group based on uniform habitus, type of hemelytral fasciae, and other characters discussed below: Boopidella Reuter, Eremachrus Lind- berg, Hallodapus Fieber, Laemocoris Reuter, Paralaemocoris Lin- navuori, Ribautocapsus Wagner, and Trichophthalmocapsus Pop- pius (and probably Omphalonotus Reuter). All species are relatively small and ground living in those cases where the habits are known. This group possesses the first stridulatory mechanism recorded in the Miridae, although they are well known in other families of Heteroptera (see Leston, 1957; Ashlock and Lattin, 1963). It exists in Trichophthalmocapsus , Laemocoris , and some species of Hallo- dapus (I have not examined specimens of Ribautocapsus , Eremach- rus, and Paralaemocoris for this structure). The stridulatory device consists of a stridulitrum of fine teeth on the lateral corial margin and a plectrum of varying structure on the inner surface of the meta- femur. In all species known to have the stridulatory mechanism both sexes possess it. In Trichophthalmocapsus and Laemocoris all of the species I have examined are apparently capable of stridulation. These highly derived genera probably arose from the more gener- alized genus Hallodapus in which only a limited group of species possess the structure. The stridulatory mechanism is absent in Boopidella, which otherwise appears most closely related to Tricho- phthalmocapsus. Trichophthalmocapsus (and probably Boopidella, although no females have been available for examination) has strongly sexually dimorphic eyes. The eyes of the males are much larger than those of the females, a situation also found in certain genera of the Aeolo- coris group. 296 Entomologica Americana [Vol. 47 All genera of the Hallodapus group occur in Africa and are us- ually most diverse there. Hallodapus is well represented in the Pale- arctic and Southeast Asia, but it is not known from New Guinea or Australia. This distribution agrees with the Hallodapini as a group, showing a major radiation in Africa with a more limited represen- tation in the Palearctic and Southeast Asia. Hallodapus albiceps (Lethierry) from Guadaloupe Island in the Lesser Antilles (Car- valho, 1958a) is not a phyline (see page 92). The Systellonotus group. This group includes a relatively large number of genera. Although the basic facies are quite variable, cer- tain structures of the head, scutellum, and male genitalia suggest close relationships. Included genera are Carinogulus Schuh, Cyrtopel- tocoris Reuter, Diocoris Kirkaldy, Gampsodema Odhiambo, Glaph- yrocoris Reuter, Hypomimus Lindberg, Mimocoris Scott, Pangania Poppius, Systellonotopsis Poppius, and Systellonotus Fieber. Carin- ogulus, Diocoris, Gampsodema, and Pangania all possess projections either dorsally or laterally on the phallotheca (the male genitalia of Systellonotopsis, Mimocoris, and Hypomimus are not known). The phallothecal projection in Pangania is much more highly developed than in the other three genera. The dorsal phallothecal projection also occurs in Hallodapus similis, the only species in the Hallodapus group known to possess the structure. This suggests a close relation- ship between the genera placed in the Hallodapus group and some members of the Systellonotus group, if in fact the dorsal phallothecal projection is homologous in all species in which it occurs. Further study of the male genitalia of the Hallodapini will be necessary to determine the phylogenetic significance of this structure. Glaphy- rocoris, Hypomimus, Carinogulus, and Cyrtopeltocoris have affinities with one another based on the tuberculate scutellum and also the head, which is concave behind (except in some species of Carino- gulus). Most of the genera in this group are ground living. Pangania, however, is arboreal, but may be secondarily adapted to this habit. The habits of Diocoris are unknown. Pangania, Diocoris, and Gampsodema are more or less restricted to tropical Africa. The remaining genera are distributed primarily in the Mediterranean Region with a limited representation in South Africa. Cyrtopeltocoris is endemic to North America, and most di- verse in the arid southwestern United States. The Systellonotus group appears to be adapted to relatively dry areas with its greatest evolu- tion in the Mediterranean, with a possibly wider distribution across Africa in the past and an invasion of North America by the ancestor of Cyrtopeltocoris (see also Coquillettia group). 1974] Schuh: South African Orthotylinae and Phylinae 297 The Formicopsella group. This group contains the morpholog- ically most ant-like genera in the Hallodapini. Included genera are Alloeomimus Reuter, Aspidacanthus Reuter, Formicopsella Poppius, Malgacheocoris Carvalho, Myombea China and Carvalho, Skukuza Schuh, and Sohenus Distant. The most distinctive structural feature of the Formicopsella group is the head, which is dorsoventrally elongated and narrowed poste- riorly, forming a neck; the eyes are far removed from the anterior margin of the pronotum. Hemelytral markings consist of a white fascia at the base of the cuneus as well as medially on the corium. The vesica is long, strap-like, and usually bent several times (Fig- ures 160, 182) (the male genitalia are imperfectly known for Al- loemimus, Malgacheocoris , and Aspidacanthus). An interesting morphological specialization in Aspidacanthus and Myombea is the presence of a thin, erect spine on the scutellum. Laemocoris in the Hallodapus group shows a similar but presumably independent development. The Formicopsella group is probably primarily ground living, although at least one genus, Formicopsella , is arboreal. The greatest diversity is in tropical Africa, but the range includes the southern Mediterranean, Madagascar, and India. The Cremnocephalus group. Wagner (1970b) erected the tribe Cremnocephalini sensu Wagner to receive the genera Cremnoceph- alus Fieber, which contains two European species, and Closterocoris Uhler, which contains a single North American species (see discus- sion under misplaced genera). He substantiated the creation of this new taxon on the possession of long first tarsal segments and peculiar hemelytral markings in the included genera, and also the form of the right clasper in Cremnocephalus. Wagner specifically excluded Myrmicomimus Reuter, from southern Europe, the only other hal- lodapine genus with a long first tarsal segment, because it has a com- plete, narrow, light, transverse fascia on the anterior third of the corium and clavus, rather than the series of light lines parallel to the claval suture found in Cremnocephalus. Both Cremnocephalus and Myrmicomimus have a rounded pronotal collar that is unlike the flattened structure found in all other Hallodapini. Wagner (1970b) stated that Cremnocephalus is not ant mimetic, citing Kul- lenberg (1944), and that it is arboreal, whereas the Hallodapini sensu Wagner are ground living. He felt this supported a more dis- tant relationship from the remaining hallodapines. However, several genera in the Hallodapini are arboreal (see Pangania and Formi- 298 Entomologica Americana [Vol. 47 copsella ), and some are not less convincingly ant-mimetic than Cremnocephalus (e.g. Orectoderus) . The long first tarsal segment and peculiar hemelytral markings of Cremnocephalus occur nowhere else in the Phylinae and are of only very limited occurrence in the Miridae as a whole. This sug- gests that they are probably apomorphic characters. These charac- ters set off Cremnocephalus as (part of) a distinct evolutionary unit, but one not significantly different from the remainder of the Hallo- dapini to merit tribal status. Furthermore, hemelytral markings are variable in the Hallodapini, at least much more so than the length of the first tarsal segment. Therefore, if the Cremnocephalini were to be recognized, Myrmicomimus would have to be included with Cremnocephalus because they are very closely related by the general body form, the rounded pronotal collar, type of male genitalia, and long first tarsal segment. I consider these two genera to represent nothing more than a specialized evolutionary line within the Hallo- dapini sensu lato. The Coquillettia group. The genera Coquillettia Uhler, Orecto- derus Uhler, and Teleorhinus Uhler (and Cyrtopeltocoris , discussed under Systellonotus group) are the only Hallodapini that occur in the New World. Reuter (1910a) placed Coquillettia and Orectoderus in the Cremnocephalini and Teleorhinus in his genera incerta. Knight (1923) and Carvalho (1952a; 1958a) placed all three genera in the Hallodapini. Knight (1941; 1968) considered these genera to be related to Macrotylus (and other genera with long pulvilli) and placed them in the Phylini. Although Colquillettia and Orectoderus-Teleorhinus form two rather distinct groups they can be related on a number of derived characters. These are: 1) the presence of a long carinate gula in the males (the gula is ecarinate in the females); 2) the type of bra- chyptery in the females; 3) the enlarged pulvilli; and 4) the absence of minute rows of spines on the tibiae (present in almost all other Phylinae). In the Coquillettia group, as in the Leucophoropterini, there is what appears to be an independent evolution of two types of gono- pores in the male genitalia. This is suggested by the fact that Coquil- lettia has an obscure poorly developed gonopore situated at the apex of the vesica, whereas in Orectoderus the gonopore is large, well developed, and situated subapically. The type found in Coquillettia is possibly primitive because the flattened pronotal collar is of the form found in most other Hallodapini, whereas in Orectoderus and Teleorhinus , it is greatly modified. 1974] Schuh: South African Orthotylinae and Phylinae 299 Coquillettia is related to Systellonotus by the general body form and structure of the vesica. The carinate gula found in the males of all Coquillettia group genera is not present in Systellonotus, although it does occur in both sexes of Mimocoris and at least in the males of Carinogulus. Systellonotus does not have the greatly enlarged pulvilli of this group, although at least in S. triguttatus they are dis- tinctly larger than in most hallodapine genera. Coquillettia is the least specialized of the three genera in the group. Orectoderus and Teleorhinus have diverged greatly from Coquillettia, losing the hemelytral fascia (in most species), having the pulvilli fused to the ventral surface of the claws (free in Coquil- lettia), having the pronotal collar greatly modified, and in Teleo- rhinus having a strongly clavate second antennal segment. Based on morphological and distributional evidence it seems reasonable to assume that the ancestor of this group was similar to Systellonotus in many respects and that it invaded North America from the Palearctic. The genera belonging to the Coquillettia group are widely distributed in North America, but show their greatest diversity in the Western mountains. Van Duzee (1921) described Coquillettia uhleri from Pasadena, California. This species was in- correctly recorded from “Austria; n.m. Europe” by Carvalho (1958a). Zoogeography: The distribution of the Hallodapini as por- trayed by the Carvalho Catalogue is cosmopolitan, with the greatest diversity in the Palearctic and Ethiopian Regions. I have excluded the genera Amazonocoris Carvalho, Closterocoris Uhler, Glossopeltis Reuter, Hallodapoides Carvalho, Heidemanniella Poppius, Nico- stratus Distant, and Tylopeltis Reuter, which were included in the Hallodapini by Carvalho (1958a), and moved them to other sub- families. The tribe is thus predominantly Old World, with only two very limited groups occurring in the Nearctic, and is totally absent from the Neotropics. The Hallodapini are most diverse in Africa and the Palearctic (Figure 351a) and the faunas of these two areas are very closely related. Hallodapines are distinctive for their extensive adaptation to relatively dry areas of the Old World. This is exemplified by the large number of genera in the Mediterranean and Africa. At present the only genera known from Southeast Asia and Australia are Hal- lodapus, Azizus, and Acrorrhinium. In these regions the Leuco- phoropterini and Pilophorini are most abundant and probably re- place the Hallodapini ecologically. 300 Entomologica Americana [Vol. 47 Discussion Of Individual Genera. Acrorrhinium Noualhier, 1895, see page 66. Aeolocoris Reuter, 1903, see page 121 and discussion under Aeolo- coris group. A lloeomimus Reuter, 1910b. The male genitalia (as illustrated by Hoberlandt, 1953, for Al- loeomimus kurdus), parempodia, pronotal collar, and facies confirm the placement of this genus in the Hallodapini. Two species are known from the Mediterranean. * Amazonocoris Carvalho, 1952c, see genera incertae sedis. * Anapsallus Odhiambo, 1959c, see Phylini. Aspidacanthus Reuter, 1901. The structure of the head and pronotum, hemelytral coloration, and general facies place this genus in the Hallodapini. The scutellar spine shows a very close relationship to Myombea. Two species are known from Senegal and Turkestan. Azizus Distant, 1910a, see page 80. Bibundiella Poppius, 1914a. I have not examined specimens of this genus, but based on the original description, it belongs to the Hallodapini. The type speci- men of Bibundiella obscura Poppius, is in the Helsinki Museum (personal communication, Martin Meinander, Helsinki Museum), although Poppius’ (1914a) original description indicates that it was deposited in the Berlin-Humboldt Museum. Boopidella Reuter, 1907b, see page 121. Carinogulus Schuh, new genus, see page 81. Chaetocapsus Poppius, 1914a. I have not examined Chaetocapsus binotatus Poppius, the type species of the genus, but the original description indicates that it belongs to the Hallodapini. The holotype of C. binotatus is in the Helsinki Museum (personal communication, Martin Meinander, Helsinki Museum), although Poppius (1914a) indicated that it was deposited in the Berlin-Humboldt Museum. * Closterocoris Uhler, 1 890, Mirinae, see misplaced genera. Coquillettia Uhler, 1890, see discussion under Coquillettia group. 1974] Schuh: South African Orthotylinae and Phylinae 301 Cremnocephalus Fieber, 1860, see discussion under Cremnocephalus group. Cyrtopeltocoris Reuter, 1876a. Reuter (1910a) placed Cyrtopeltocoris in the Cremnocephalaria and was followed by Van Duzee (1917) who placed the genus in the Hallodapini (Cremnocephalaria = Hallodapini); Knight (1968) later placed it in the Pilophorini, the position given the genus by Carvalho (1952a; 1958b). Kelton (1959b) noted that the male genitalia are of the phyline-type. The parempodia, which are weakly fleshy and slightly convergent apically, have created confusion as to proper tribal placement. The male and female genitalia, the flat- tened pronotal collar, the white transverse fascia on the hemelytra, and the general body form all confirm placement in the Hallodapini (see also discussion under Systellonotus group). At least 13 species are presently placed in Cyrtopeltocoris , most from the Southwestern United States (see Knight, 1968). Diocoris Kirkaldy, 1902c, see page 122. Eremachrus Lindberg, 1958, pp. 105-106. Described from brachypterous specimens, and placed in the Hal- lodapini by Lindberg (1958), Eremachrus is extremely closely re- lated to Hallodapus. Only a single species is known from the Cape Verde Islands. * Eucerella Poppius, 1921, see Orthotylini. F ormicopsella Poppius, 1914a, see page 89. Gampsodema Odhiambo, 1959c, pp. 648-649, see Diocoris , page 122. Glaphyrocoris Reuter, 1903, see page 84. * Glossopeltis Reuter, 1903, Deraeocorinae, see misplaced genera. * Hallodapoides Carvalho, 1951a, see Orthotylini. Hallodapus Fieber, 1858, see page 91. * Heidemanniella Poppius, 1914c, Mirinae, see misplaced genera. Hypomimus Lindberg, 1940, see discussion under Carinogulus, page 84. Kapoetius Schmitz, 1969, pp. 72-81. Kapoetius belongs to the Aeolocoris group. One species is known from the Sudan. 302 Entomologica Americana [Vol. 47 Laemocoris Reuter, 1879, see page 103. * Lissocapsus Bergroth, 1903, see genera incertae sedis. * Makakix Odhiambo, 1967, Deraeocorinae, see misplaced genera. Malgacheocoris Carvalho, 1952b. Malgacheocoris is probably most closely related to Formicopsella and Myombea from Africa. Only a single species is known from Madagascar. Marmorodapus Schmitz, 1970, pp. 512-520. Marmorodapus was described with a single included species, M. spinulatus Schmitz. Unfortunately, by what must have been an inadvertent error, no locality data or holotype designation was in- cluded with the original description. The genus is from Africa (Congo?) and belongs to the Aeolocoris group. Mimocapsus Wagner, 1953. Mimocapsus may be closely related to Systellonotus. One species is known from Egypt. Mimocoris Scott, 1872. See discussion under Systellonotus group. Two species are de- scribed from southern Europe and the Mediterranean. A third spe- cies, Mimocoris scotti Berg, from Argentina, is almost certainly placed in the wrong genus and subfamily. Myombea China and Carvalho, 1951, see page 104. Myrmicomimus Reuter, 1881, see discussion under Cremnocephalus group. * Myrmicopsella Poppius, 1914a, see Leucophoropterini. * Nicostratus Distant, 1904a, Deraeocorinae, see misplaced genera. Omphalonotus Reuter, 1876b. Omphalonotus is probably most closely related to Hallodapus and allied genera. Two species are known from Europe and North Africa. Orectoderus Uhler, 1876, see discussion under Coquillettia group. Pangania Poppius, 1914a, see page 104. Paralaemocoris Linnavuori, 1964, pp. 326-328. This genus is most closely related to Laemocoris and Hallodapus. Three species are known from the Middle East. 1974] Schuh: South African Orthotylinae and Phylinae 303 Ribautocapsus Wagner, 1962, p. 83. This genus is most closely allied to Laemocoris and Hallodapus. One species is known from Spain and Algeria. Sohenus Distant, 1910a. Sohenus appears to be very closely related to Formicopsella, from Africa, by the structure of the head, pronotum, and hemelytra, and also the color pattern. Further study may reveal that the two are synonymous. The male genitalia of S. uvarovi Ballard are typical of the Hallodapini, the vesica being long, with several bends, and having a well developed subapical gonopore. Two species are known from India. Syngonus Bergroth, 1926. Originally described under the preoccupied name Bibundia (Pop- pius 1914a), this genus was renamed by Bergroth (1926). Poppius (1914a) stated that the holotype of Syngonus nigra (Poppius), the only described species in the genus, was deposited in the Berlin- Humboldt Museum. In fact it is in the Helsinki Museum (Type No. 11958). Syngonus is probably most closely related to Acror- rhinium and Trichophorella. It is peculiar in the Hallodapini in being black. An undescribed species from Ghana has a very broad white fasica medially on the hemelytra, whereas nigra has only a faint light marking on the corium. The former condition is not found in other members of the Acrorrhinium group. The head is missing from the holotype of S. nigra, from Cameroon. Systellonotidea Poppius, 1914a, see Diocoris Kirkaldy, page 122. Systellonotus Fieber, 1858, see page 112. Teleorhinus Uhler, 1890, see discussion under Coquillettia group. Trichophorella Reuter, 1905b, see page 114. Trichophthalmocapsus Poppius, 1914a, see page 117. * Tylopeltis Reuter, 1904, Deraeocorinae, see misplaced genera. LEUCOPHOROPTERINI, NEW TRIBE Diagnosis: Usually ant mimetic; generally dark, often with con- trasting light hemelytral maculae; head usually concave behind, eyes usually contiguous with anterior margin of pronotum; head some- times convex behind, eyes well removed from pronotum; genae oc- casionally extremely hairy; pronotum usually with finely carinate 304 Entomologica Americana [Vol. 47 upturned anterior margin or with more or less well developed flat- tened collar; pronotum slightly to strongly constricted anteriorly or rarely constricted medially (hour glass shaped); scutellum always flat; hemelytra straight or weakly or strongly sinuate laterally; ab- domen narrow; parempodia usually hair-like, parallel, very seldom fleshy, rod-like, and weakly convergent apically; pulvilli always minute; vesica occasionally U-shaped, weakly twisted (Figure 200), usually S-shaped, strongly twisted; male gonopore undeveloped, poorly developed, or rarely well developed; left clasper always wedge- shaped, trough-like; right clasper flat, leaf-like; posterior wall simple, posterior margin not evaginated. Note: Many genera in the Leucophoropterini are undescribed and therefore cannot be given names in the following discussion. Discussion: Many genera in the Leucophoropterini are super- ficially very similar to members of the Hallodapini and Pilophorini. Fortunately, a distinct evolutionary sequence can be traced in the Leucophoropterini, for if this were not possible, at least some of the genera would be placed in the other two tribes. Two of the most primitive genera in this tribe ( Leucophoroptera and Karoocapsus) have well developed light hemelytral maculae. The anterior margin of the pronotum in these genera is finely carinate and upturned and the head is concave behind, resembling structurally the situation found in the Pilophorini. Tytthus , which also appears to be rela- tively primitive, usually has a dark head and pronotum and light hemelytra; the head is convex behind and the anterior margin of the pronotum is similar to that of Leucophoroptera. Most of the derived members of the tribe have rather poorly developed hemely- tral maculae and have the head and prontum variously modified from the structure found in Leucophoroptera and Karoocapsus (see discussion below) . The male genitalia of the Leucophoropterini are distinct from those of the Hallodapini, Phylini, and Pilophorini, although the dif- ferences are often small and difficult to categorize. The gonopore is either apical and not developed ( Karoocapsus and Tytthus) or subapical and poorly developed. In an undescribed genus from the Philippines, the gonopore is very well developed, but this is almost certainly a convergence toward the form found in most Hallodapini, because all other characters of this genus agree closely with the Leucophoropterini. In Karoocapsus and Tytthus the vesica is U- shaped and only very slightly twisted, resembling the form found in the Pilophorini and emphasizing the plesiomorphic character of these genera. In all other genera the vesica is distinctly twisted 1974] Schuh: South African Orthotylinae and Phylinae 305 and S-shaped. The male genitalia are small relative to the total body size. In contrast to the Leucophoropterini, the Hallodapini almost always have a well developed subapical gonopore (exceptions in- clude Coquillettia) , a strongly twisted S-shaped or more elaborately bent vesica and genitalia that are relatively large in comparison to the total size of the insect. The female genitalia of the Leucophorop- terini show only very minor differences from those of the Hallodapini and Phylini, and differ from those of the Pilophorini in not having the evaginated posterior margin of the posterior wall. Two lines of evolution toward effective ant mimicry can be rec- ognized in the Leucophoropterini. One involves the development of a head-pronotum combination similar to that found in Formicop- sella in the Hallodapini. In this line there is a definite transition from the type of head and anterior pronotal margin found in Karoocapsus and Leucophoroptera to an anteriorly constricted pronotum with a flat collar and head “necked” behind the eyes (genera of this type are as yet undescribed). The other line of evolution involves ac- centuation of the head which is concave behind, again as in Karoo- capsus, and at the same time specialization of the pronotum to con- form to the outline of the posterior margin of the head. At its highest degree of specialization the pronotum is constricted medially into an hourglass shape; this type has also evolved independently in the Pilophorini (genera in neither tribe are as yet described, however) . Two convergences, in addition to those mentioned above, occur between the Leucophoropterini and other phyline tribes. In certain undescribed genera the gena is “carinate”, forming a broad ridge below the eye. In frontal view this resembles the outline of mandi- bles, and its most advanced condition, includes the buccula as the apex of the mandible. A similar development occurs in at least one undescribed species of pilophorine, related to Pilophorus , from the Philippines. In another undescribed genus of Leucophoropterini from the Philippines the parempodia are fleshy, rod-like, and weakly convergent apically, a condition very similar to that found in some Phylini and Hallodapini. Zoogeography: Two of the most primitive genera in the Leuco- phoropterini, Karoocapsus and Leucophoroptera , occur in South Africa and Australia (and New Guinea) respectively. The more derived genera are found in New Guinea, New Ireland, the Solomon Islands, Borneo, and the Philippine Islands. Therefore the group probably evolved in temperate and subtropical areas of Australia (and South Africa) and subsequently spread to nearby tropical is- lands and became more highly specialized there. The paucity of 306 Entomologica Americana [Vol. 47 leucophoropterine genera in Africa (Figure 351b) suggest that either Karoocapsus is a relict or that the environmental conditions in trop- ical Africa were unsuitable for the evolution of the group. Also, com- petition from the Hallodapini may have been important in limiting the evolution of the Leucophoropterini in Africa. Tytthus appears to be in a somewhat distinct evolutionary position in the tribe. It is nonmimetic, has the head convex behind, and is cosmopolitan. Discussion Of Individual Genera. Bilirania Carvalho, 1956a, pp. 215-216. Bilirania myrmecoides Carvalho, from the Philippines, was placed in the Pilophorini and related to Leucophoroptera by its author. Carvalho (1956a) did not illustrate the male genitalia, but the facies of Bilirania indicate that it probably belongs to the Leucophorop- terini. The complex distribution of the Leucophoropterini and the Pilophorini in Southeast Asia and the great external similarity of the two groups makes it necessary to only tentatively assign Bilirania to the Leucophoropterini, until specimens can be examined and the male genitalia dissected. Karoocapsus Schuh, see page 123. Leucophoroptera Poppius, 1921. Leucophoroptera was originally described from New South Wales, Australia, and New Guinea. I have examined an undescribed species from Queensland, Australia, in which the male genitalia are similar to those of Karoocapsus , but the vesica is more strongly S-shaped. The female genitalia of L. quadrimaculatus have a posterior wall consisting of a simple sclerotized plate. Poppius (1921) described L. quadrimaculatus , the type species of the genus, from specimens from New South Wales, Australia, and New Guinea (“Ins. Deslacs”). I have examined a female from the Helsinki Museum from the latter locality. The other specimens are apparently in the Hungarian Museum and they must be studied be- fore a lectotype can be designated. Myrmicopsella Poppius, 1914a. The holotype female of Myrmicopsella nitidipenne Poppius from Tananarive, Madagascar, is the only known representative of the genus. It almost certainly belongs to the Leucophoropterini, and is probably most closely related to Karoocapsus. Poppius (1914a) noted that this specimen was deposited in the Paris Museum, but in fact, it is in the Helsinki Museum (Type No. 7788). 1974] Schuh: South African Orthotylinae and Phylinae 307 Tytthus Fieber, 1864, see page 135. TRIBE PHYLINI Diagnosis: Facies, coloration, and vestiture variable, never ant mimetic; females occasionally brachypterous; flattened pronotal collar absent (except in Eminoculus)\ dorsum very seldom heavily punctate; parempodia usually hair-like, parallel, occasionally fleshy, rod-like, of nearly uniform diameter, weakly convergent apically; pulvilli minute or enlarged, free or fused to ventral surface of claws; vesica always twisted (only very slightly in Pseudosthenarus and Parapseudosthenarus) , gonopore usually subapical, well developed; left clasper always wedge shaped, trough-like (somewhat modified in Pseudosthenarus ); posterior wall never with evaginate posterior margin. Discussion: The Phylini have traditionally been the tribe in the Phylinae which contained all nonmimetic and/or collarless gen- era. Until a thorough analysis of this large cosmopolitan group can be undertaken, I am defining the tribe as consisting of all nonmimetic, collarless genera without convergent recurved parempodia. Tribes recognized by certain authors that I include in the Phy- lini are Cremnorrhini, Harpocerini, Camptotylini, Exaeretini, Tu- poniini, and Semiini. A few additional tribes have been recognized, but these are of mostly historical interest. The Oncotylini have been recognized by Reuter (1883; etc.), Van Duzee (1916), Knight (1923), and other authors to include genera with enlarged pulvilli (e.g. Lopus Hahn and Macrotylus Fie- ber). Although many genera with this tarsal condition are probably closely related, they do not appear to merit tribal status, when con- sidered relative to the total variation of the pulvilli in the Phylini. Also, the condition has evolved independently in the Hallodapini (see Coquillettia group). The Cremnorrhini have recently been recognized (Wagner and Weber, 1964) to include the single Eastern Mediterranean genus Cremnorrhinus with the single included species C. basalis Reuter; the Harpocerini (Wagner, 1952) to include Harpocera Curtis, from Western Europe and the Mediterranean; the Camptotylini and Ex- aeretini (Wagner, 1952; Wagner and Weber, 1964) to include the Mediterranean genera Camptotylus Fieber and Exaeretus Fieber, respectively; and the Tuponiini (Wagner, 1952, as a subtribe; Wag- ner and Weber, 1964, as a tribe) to include Tuponia Reuter and closely related genera. None of the above tribes is based on a com- 308 Entomologica Americana [Vol. 47 parative analysis of the world fauna, and for the most part they are founded on structures that vary throughout the Phylini sensu lato. The genitalia in Camptotylus (see Wagner and Weber, 1964) (and probably Exaeretus) , are somewhat peculiar in the Phylinae, but in the form of the right clasper, a parallel to Camptotylus is found in Pseudosthenarus from South Africa, although the two are probably unrelated, based on the structure of the vesica. The Harpocerini and Cremnorrhini are defined on color characters (Wagner, 1952). The Tuponiini are separated from the Exaeretini (= Camptotylini) (Wagner and Weber, 1964) on the length and shape of the labium, a character I have found to be extremely variable (e.g., compare Pseudosthenarus , Capecapsus , and Coatonocapsus with one another and with other phylines). The tribe Semiini (Knight, 1923) was erected within the Orthotylini, but the only included genus, Semium Reuter, has since been transferred to the Phylini (Kelton, 1959a). Several genera, in addition to those discussed above, are some- what anomalous within the Phylini. Reuteroscopus Kirkaldy, from North and Central America, has a vesica quite distinct from all other known Phylinae (see Kelton, 1964). The coleopteroid females and stylate eyes of Eminoculus, from South Africa, are unique in the Phylinae, and resemble those of Pachytomella Reuter in the Halti- cini. Also Eminoculus is the only known Phylini genus with a flat- tened pronotal collar. This character might relate the genus to the Hallodapini, but otherwise Eminoculus bears no obvious relationship to that tribe, and the collar is probably independently evolved rela- tive to the Hallodapini. The male genitalia of Pseudosthenarus and Parapseudosthenarus from South Africa, show no close relationship to any other known genera, although externally Pseudosthenarus closely resembles some species of Sthenarus. Until the Phylini as a whole can be carefully studied, I do not consider it advisable to subdivide the tribe. Although the other tribes recognized within the subfamily may in some cases represent derivatives of the Phylini (especially the Hallodapini), it seems de- sirable to recognize individual phyletic lines of specialization, e.g. ant mimicry, within the Phylinae, rather than conceal them within an omnibus tribe. Subdivision of the Phylini along phyletic lines at the present time would, however, be nearly impossible. I have not examined the following genera placed in the Phylini by Carvalho (1958a) and have not found references which will allow confirmation of subfamily or tribal placement: Alloeotarsus Reuter, Boopidocoris Reuter, Capellanus Distant, Cephalocapsidea Poppius, Decomia Poppius, Demoplesia Poppius, Ectagela Schmidt, Ectenellus 1974] Schuh: South African Orthotylinae and Phylinae 309 Reuter, Ephippiocoris Poppius, Eucharicoris Reuter, Euderon Puton, Exaeretus Fieber, Hadrophyes Puton, Homolaner Kiritschenko, Ibi- aris Horvath and Reuter, Leucodellus Reuter, Litoxenus Reuter, Myochroocoris Reuter, Nicholia Knight, Nyctidea Reuter, Oligobiella Reuter, Opisthotaenia Reuter, Pararagmus Poppius, Phoenicocapsus Reuter, Pleuroxonotus Reuter, Pronotocrepis Knight, Sceodamia Poppius, Sthenaropsis Poppius, Taeniophorus Linnavuori, Trevessa China, and Utopnia Reuter. Zoogeography: The Phylini are the only tribe in the Phylinae that occur in the Neotropical Region (with the exception of one spe- cies of Pilophorini). Some records for South America are old and pertain to species placed in large, widely distributed genera, which may in fact be incorrectly assigned. The actual amount of endemism in the Neotropics cannot therefore be accurately determined. Data for the Nearctic indicate that the fauna consists of two basic elements: 1) an endemic fauna; and 2) a fauna closely related to the Palearctic at the generic level (Figure 351c). The Palearctic phyline fauna is extremely large, but probably not as diverse as the data would indicate. As discussed above, Wagner (1952) and Wag- ner and Weber (1964) have divided the Palearctic Phylini into five tribes, but the majority of the genera are placed in one tribe, the Phylini. The one element of the Palearctic fauna that has received attention recently, particularly from Wagner (1957b; 1959; 1961; etc.) and Linnavuori (1961; 1964; etc.) is the Mediterranean. This region is very interesting and seems to hold most of the anomalous types in the Palearctic fauna as a whole. With regard to the Ethiopian Region, my investigations on the Phylini of South Africa reveal some interesting facts. Several generic groups, based on the structure of the male genitalia, appear to exist within the Ethiopian fauna, even though the relationship of the gen- era is not obvious from general facies. For example, heavy punc- tations on the dorsum are extremely uncommon in the Phylini. Lamprosthenarus, which is heavily punctate, has a vesica very sim- ilar to that of Coatonocapsus, Austropsallus , Odhiamboella , and others, which are impunctate. The genera of Phylini presently recorded from the Oriental region are mostly of wide distribution, occurring in several faunal regions (Fig. 351c). At present no endemic Phylini are known from Aus- tralia. The Phylini are adapted primarily to temperate regions and prob- ably to floras of those areas. This is suggested by the abundance of genera in the Palearctic and Nearctic and the paucity of genera in 310 Entomologica Americana [Vol. 47 tropical areas, a phenomenon that may be to some extent the result of inadequate collecting in the tropics. However, I have examined large collections of Miridae from Africa, and have found very few Phylini from areas other than the Mediterranean and South Africa. Ex- amination of the known distributions of the Bryocorinae and Der- aeocorinae, which are no more or less well known than the Phylini, reveals that they are primarily tropical, with only a very few rep- resentatives in temperate regions. This confirms that although ab- solute faunal compositions are not known, the relative diversity of mirid taxa in temperate and tropical areas is well enough known to make useful comparisons. Discussion Of Individual Genera. Anapsallus Odhiambo, 1959c, pp. 680-681. Odhiambo (1959c) placed Anapsallus in the Hallodapini be- cause of its wide pronotal collar. My examination of the holotype of A. marmoratus Odhiambo reveals that in fact there is no pronotal collar and that the genus belongs in the Phylini. Ellenia Reuter, 1910a, see page 157. Erythrocorista Lindberg, 1958, see Orthotylini. Millerimiris Carvalho, 1951b. Carvalho (1951b) placed Millerimiris in the Orthotyini, but his illustrations of the male genitalia indicate that it is actually a member of the Phylini. My examination of the holotype of M. punctatus Car- valho reveals that the parempodia are only weakly fleshy and sim- ilar to the type found in Ellenia and Capecapsus. Parafulvius Carvalho, 1954. This genus was placed in the Fulviini (Cylapinae) by Carvalho (1954), on the basis of the type of claws and male genitalia. Car- valho (1954) stated that Parafulvius resembles Amblytylus Fieber. In fact it is probably closely related to that genus. The genitalia as illustrated by Carvalho are definitely phyline, and the claws, although they may be long and slender, fit into the range of variation found in the Phylinae. Paramixia Reuter, 1900, Pilophorini, see page 210. Platyscytus Reuter, 1907a. This genus has been assigned to the Orthotylini by Carvalho (1952a; 1958b). Examination of illustrations of the male genitalia of species of Platyscytus described by Carvalho (1953b) and Car- 1974] Schuh: South African Orthotylinae and Phylinae 311 valho and Fonseca (1965) and examination of specimens of the genus, indicate that it belongs to the Phylini. Also, the parempodia are hair-like and not of the type found in the Orthotylini. Psallops Usinger, 1946, Cylapinae ?, see pages 263-264. Semium Reuter, 1876a. Placed in the Orthotylini by Carvalho (1958b), this genus was correctly moved to the Phylini by Kelton (1959a). TRIBE PILOPHORINI Diagnosis: Elongate or robust, sometimes ant mimetic; seldom if ever strongly brachypterous or sexually dimorphic; head declivous to nearly vertical, concave behind, posterior margin of vertex usually carinate; pronotum usually broad and nearly flat, although some- times highly modified with tubercles or strongly constricted me- dially; hemelytra usually without defined fasciae contrasting with background coloration; often with light transverse band on hemelytra formed by aggregations of sericeous scale-like hairs; parempodia fleshy, recurved, convergent apically; pulvilli minute; vesica simply curved, U-shaped, not twisted, without enlarged apical or subapical gonopore (Figure 318); phallotheca usually nearly straight, without right-angle bend (L-shaped); opening usually terminal (Figure 325); left clasper sometimes distinctly trough-like ( Paramixia , Figure 334), usually splayed out, wing-like (Figure 320); right clasper small and leaf-like, typical of subfamily; female genitalia with sclerotized rings usually with moderate lateral infolding (Figure 317); posterior wall simple, lacking K-structures (Figure 315), but with evagination dorsally along posterior margin (Figure 316). Discussion : Most authors have defined the Pilophorini as those ant-mimetic mirids with convergent parempodia. Wagner (1952; 1955) was the first author to realize that the tribe, as so defined, was composed of unrelated genera and he redefined the group as those mirids with convergent recurved parempodia and Phylinae- type male genitalia. In analyzing the Orthotylinae and Phylinae I have concluded that the convergent recurved parempodia are ancestral and that the hair- like parempodia found in the Phylinae are derived from them. I have reached this conclusion because, when convergent recurved parempodia are regarded as derived, as can be inferred from most classifications, it becomes necessary to evolve the phyline-type male genitalia twice. I am following Knight (1941) who regarded the complex structure of the phyline male genitalia as a fundamental 312 Entomologica Americana [Vol. 47 character in classification; I consider them to be derived and there- fore (as previously discussed) place the Pilophorini in the Phylinae, rather than the Orthotylinae as all previous authors have done. I interpret the convergent recurved parempodia as primitive; thus the phyline-type male genitalia need to be evolved only once. As members of the most primitive phyline tribe, Pilophorus and closely related genera (e.g. Aloea) have the simplest male genitalia in the subfamily. The vesica is a U-shaped, untwisted tube with a gonopore that is little more than a subapical opening in the wall of the vesica. The most complex pilophorine vesica is that of Para- sthenaridea Miller. Although U-shaped and untwisted, the vesica is no longer a simple tube, but consists of what resemble two par- tially concentric sclerotized bands. Peculiar projections on the inner surface of the vesica are a unique feature of many pilophorine genera (e.g. Pilophorus , Parasthenaridea) . The pilophorine phallotheca is also plesiomorphic in the Phylinae. It is not bent at a right angle (except in Paramixia ) as in all other Phylinae, and is structurally closest to the orthotyline-type. As discussed above, the pilophorine posterior wall is advanced in the Phylinae and must have evolved subsequent to the split of the Phylinae into the Pilophorini and non- Pilophorini lines of evolution. The Pilophorini apparently do not possess brachypterous forms, although they are common in all other phyline tribes. The greatest degree of wing modification or reduction in this tribe is what might best be called submacroptery, as found in females of Aloea. Even in the ant-mimetic genus Pilophorus , brachypterous females are un- known. Wing polymorphism is an adaptation to a specialized or stable environment (see Sweet, 1964) and therefore must be de- rived. The apparent absence of brachyptery in the Pilophorini is a further suggestion of their primitiveness in the Phylinae. Zoogeography: My redefinition of the Pilophorini on a world basis changes the zoogeographic picture from one of more or less equal distribution in all zoogeographic regions in the Carvalho clas- sification to one of greatest diversity in the Old World tropics and virtual absence from the neotropics (Figure 35 Id). Two relatively distinct groups of genera can be recognized in analyzing the distribution of the Pilophorini. The Ambonea group is not ant mimetic. It is restricted primarily to Africa, constituting almost the entire fauna there; the only nonmember of the Ambonea group found in Africa is a single species of Pilophorus. The Pilo- phorus group, which is distinctly ant mimetic, is most diverse in Southeast Asia, with limited representation in the Nearctic and Pale- 1974] Schuh: South African Orthotylinae and Phylinae 313 arctic. In North America Pilophorus has radiated extensively, but no studies have been undertaken to determine if the 50 or so de- scribed species from that region are all closely related or if they represent separate elements in the genus and therefore independent invasions of the continent. The limited generic representation of the tribe in North America indicates invasion of the area from Southeast Asia, possibly via the Bering Land Bridge, rather than migration from North America to Southeast Asia. Paramixia , a morphologically somewhat anomalous genus in the Pilophorini , is pantropical, P. carmelitana (Carvalho) being the soli- tary neotropical pilophorine. Parasthenaridea Miller, a pilophorine with specialized male genitalia, but resembling the Ambonea group, is known only from Malaya. Discussion Of Individual Genera The incorrect subfamily placement and ant-mimic definition of the Pilophorini by Carvalho (1952a; 1958b) obscured the true re- lationships of the group as a whole and also of many included genera. Alepidea Reuter, 1909. Alepidea is very closely related to Pilophorus by the structure of the male genitalia (Kelton, 1959b) and by the structure of the female genitalia, in which the posterior margin of the posterior wall is evaginated. Two species are known, both from the Eastern United States. Alepidiella Poppius, 1914b. This genus contains only a single species from the Eastern United States. It is very closely related to Alepidea. Aloea Linnavuori (in press), see page 196. Ambonea Odhiambo, 1960b, see page 201. * Anthropophagiotes Kirkaldy, 1908, see genera incertae sedis. * Borgmeierea Carvalho, 1956c, see Orthotylini. * Cyphopelta Van Duzee, 1910, Mirinae, see misplaced genera. * Cyrtopeltocoris Reuter, 1876a, see Hallodapini. * Dolichostenia Poppius, 1921, see genera incertae sedis. * Eucerella Poppius, 1921, see Orthotylini. * Eucompsella Poppius, 1914a, see Nichomachini. 314 Entomologica Americana [Vol. 47 * Hallodapoides Carvalho, 1951a, see Orthotylini. Hyseloecus Reuter, 1891. The placement of Hypseloecus in the Pilophorini (Wagner, 1952) is verified by the structure of the parempodia and male genitalia. A single species is known from the Palearctic. * Kirkaldyella Poppius, 1921, see Orthotylini. * Laemocoridea Poppius, 1921, see Orthotylini. * Lasiomimus Poppius, 1914a, see genera incertae sedis. * Lepidotaenia Poppius, 1921, see Orthotylini. * Leucophoroptera Poppius, 1921, see Leucophoropterini. * Lutheriella Poppius, 1913, see genera incertae sedis. * Myrmecophyes Fieber, 1870, see Halticini. * Myrmecoridea Poppius, 1921, see genera incertae sedis. * Myrmecozelotes Berg, 1883, see genera incertae sedis. Neoambonea Schuh, new genus, see page 204. * Nichomachus Distant, 1904a, see Nichomachini. * Opistocyclus Poppius, 1914a, Deraeocorinae, see misplaced genera. Parambonea Schuh, see page 207. Paramixia Reuter, 1900, see page 210. Parasthenaridea Miller, 1937. See Pilophorini tribal discussion. The female genitalia will help to confirm tribal placement of Parasthenaridea. Only a single species is known from Malaya. * Pilophoropsis Poppius, 1914c, see Orthotylini. Pilophorus Hahn, 1826. Reuter (1910a) placed Pilophorus in the division Heterotomaria in the subfamily Heterotomina; Carvalho (1952a) placed the genus in the Pilophorini, as have most other modern authors. Slater (1950) noted the marked differences in the female genitalia between Pilo- phorus and Pseudoxenetus , the two genera he studied in the Pilo- phorini, particularly the lack of K-structures in Pilophorus and their presence in Pseudoxenetus. Kelton (1959b) correctly noted that 1974] Schuh: South African Orthotylinae and Phylinae 315 the male genitalia of Pilophorus appeared much more closely allied to the Phylinae than to the Orthotylinae. Pilophorus may be com- posite as presently constituted, but the basic body plan is the same in all species, and the male genitalia are very similar from species to species for those representatives that have been examined. The female genitalia have the distinct evagination along the posterior margin of the posterior wall. Pilophorus is well represented in the Palearctic, Nearctic, and Oriental regions. Only one species is known from sub-Saharan Africa and none are as yet recorded from the neotropics or from Australia. * Pseudoxenetus Reuter, 1909, see Orthotylini. * Renodaeus Distant, 1893, see Orthotylini. * Sericophanes Reuter, 1876a, see Orthotylini. * Tuxenella Carvalho, 195 2d, see Orthotylini. * Zanchisme Kirkaldy, 1 904, see genera incertae sedis. Zaratus Distant, 1909b. Zaratus is known only from the holotype female of Z. repandus Distant, from India. The genus was placed in the Pilophoraria by Distant (1910b), genera incerta by Reuter (1910a), and the Pilo- phorini by Carvalho (1952a). My examination of the holotype in- dicates that the genus is very closely related to Pilophorus , although I have not examined the genitalia. This conclusion is supported by its occurrence in Southeast Asia. MISPLACED GENERA1 Bunsua Carvalho, 1951b. This African genus was placed in the Orthotylini by Carvalho (1952a). Examination of a paratype of Bunsua bryocoroides Car- valho reveals that the genus has the pulvilli attached to the interior surface of the claws and that the posterior wall lacks K-structures. Bunsua must therefore be removed from the Orthotylinae and placed in the Bryocorinae, at least tentatively. Careful examination of the type material of the genus Petasma Odhiambo, 1960 (pp. 343-348), reveals that it is synonymous with Bunsua Carvalho (New Synonymy) . 1The genera listed were placed in the Orthotylinae or Phylinae by Car- valho (1952a; 1958a, b) or subsequent authors but actually belong in other subfamilies. 316 Entomologica Americana [Vol. 47 Closterocoris Uhler, 1890. The tribal position of Closterocoris has been in dispute for some time. Carvalho (1952a) placed the genus in the Hallodapini, al- though Knight (1922) had shown rather conclusively that it belongs to the Mirinae. Kelton (1959b) confirmed the placement in the Mirinae on the basis of the male genitalia. Wagner (1970b) placed Closterocoris in his Cremnocephalini (Phylinae), even though he had access to Kelton’s work on the male genitalia and routinely used the vesica in the Phylinae as a diagnostic feature of the subfamily. Cyphopelta Van Duzee, 1910. Kelton (1959b) confirmed the placement of Cyphopelta in the Mirinae on the basis of the male genitalia, although previous workers showed great disagreement on the proper subfamily position. Car- valho ( 1952a) placed Cyphopelta in the Pilophorini. Glossopeltis Reuter, 1903. This African genus was placed in the Hallodapini by Carvalho (1952a). The strongly toothed claws, hair-like parempodia, punc- tate dorsum, rounded pronotal collar, male genitalia, and claws with- out pulvilli all confirm a position in the Deraeocorinae, Surinamellini (Carvalho and Fonseca, 1962), however. Specimens of G. coutierei Reuter, the type species of the genus, are present in both the Helsinki and Paris Museums. The single fe- male specimen in Paris bears no locality labels, but has a determina- tion label reading “ Glossopeltis coutierei Reuter n.g. et n. sp., spec, typ.”. Single male and female specimens from Helsinki bear “Obock” labels; the female also bears a determination label of Poppius. Reu- ter (1903) did not indicate that specimens from the type series which he examined were placed in Helsinki, but Poppius (1914a) cited the same locality data as Reuter (1903) and noted that spec- imens did exist in Helsinki. The locality data of the two specimens in Helsinki do not agree exactly with that given in Reuter’s original description (they read “Museum Paris, OBOCK, Maindron 871- 93”), but the specimens are probably those examined by him. I have therefore labeled the female specimen in the Paris Museum as the lectotype — “LECTOTYPE Glossopeltis coutierei Reuter, det R. T. Schuh.” Heidemanniella Poppius, 1914c. The North American genus has long been placed in the Hallo- dapini (Carvalho, 1952a). My examination of the holotype of H. 1974] Schuh: South African Orthotylinae and Phylinae 317 scutellaris Poppius, reveals that Heidemanniella is probably most closely related to Cyphopelta and Closterocoris in the Mirinae. The only specimen is the holotype female and at the time of my exam- ination it was glued to a card so that the parempodia were not visible. Additional specimens and further study will almost certainly con- firm placement of Heidemanniella in the Mirinae rather than the Phylinae. Makakix Odhiambo, 1967, pp. 1673-1676. This African genus is very closely related to Opistocyclus Pop- pius. Odhiambo (1967) placed Makakix in the Hallodapini, with reservation. He gave excellent illustrations of the tarsal claws, which are strongly toothed at the base, but did not mention a possible re- lationship to the Deraeocorinae. The male genitalia of Makakix are not available. Examination of related genera, including Nicostratus Distant, reveals that the form of the tarsal claws is a valid subfamily character for recognizing mimetic as well as nonmimetic Deraeo- corinae. Makakix belongs to the Deraeocorinae, Surinamellini. Nicostratus Distant, 1904c. Carvalho (1952a) assigned this peculiar Southeast Asian genus to the Hallodapini. The strongly toothed tarsal claws and the male genitalia, however, unequivocally place it in the Deraeocorinae, Surinamellini. Opistocyclus Poppius, 1914a. This African genus is most closely related to Makakix and Glos- sopeltis , as confirmed by the strongly toothed tarsal claws, conical scutellum, and punctate dorsum. Therefore it must be placed in the Deraeocorinae, Surinamellini. Poppius (1914a) stated that the type of O. myrmecoides Pop- pius, the only species in the genus, was deposited in the Berlin- Humboldt Museum; however, it is in the Helsinki Museum (Type No. 7775). Tylopeltis Reuter, 1904. My examination of the holotype of Tylopeltis albosignata Reuter, the only species in the genus, in the Brussels Museum, indicates that Tylopeltis does not belong to the Hallodapini (Carvalho, 1952a), but to the Deraeocorinae, Surinamellini. This position is supported by the structure of the male genitalia, the conical scutellum, the punc- tate dorsum, and the rounded pronotal collar. 318 Entomologica Americana [Vol. 47 Genera incertae sedis1 Amazonocoris C arvalho, 1952c. Amazonocoris longipilosus Carvalho was placed in the Hallo- dapini by Carvalho (1952a). On the basis of the male genitalia as illustrated by Carvalho (1952c), this Brazilian genus appears to be much more closely related to the Dicyphinae than to the Phylinae. Carvalho does not mention the structure of the parempodia in the original description, but says only that the pulvilli are minute, which would suggest that Amazonocoris does not belong to the Dicyphinae. I have not been able to find the holotype of A. longipilosus in the British Museum (Natural History) and therefore cannot determine the correct subfamilial placement. Anthropophagiotes Kirkaldy, 1908. The whereabouts of the holotype of the only species included in this Fijian genus is unknown, and the original description is in- adequate for placing the genus in the correct subfamily. Carvalho (1952a) placed Anthropophagiotes in the Pilophorini. Dolichostenia Poppius, 1921. I have not seen specimens of this genus erected by Poppius (1921) for a species from Chile, and therefore its subfamily place- ment must remain uncertain. Carvalho (1952a) placed Dolicho- stenia in the Pilophorini. This is almost certainly incorrect, and it will probably prove to be a member of the Orthotylini. Idiomiris China, 1963, pp. 709-711. China described this peculiar genus from Chile, and on the basis of the pretarsal structures and male genitalia placed it in the Ortho- tylini. This is certainly incorrect. Idiomiris is much more closely related to either the Mirinae or the Deraeocorinae than to the Ortho- tylinae. I was not able to find the male genitalia in the British Mu- seum (Natural History). All of the characters enumerated by China, however, indicate the closest relationship with the Deraeocorinae, as does the basic facies, even though the claws are not toothed ba- sally, as is usually the case in that subfamily. Lasiomimus Poppius, 1914a. I have not examined specimens of this African genus, although they are probably present in the Leningrad Museum. Carvalho (1952a) placed Lasiomimus in the Pilophorini. xThe genera listed were placed in the Orthotylinae and Phylinae by Car- valho (1952a; 1958a, b) or by subsequent authors, but are unknown to me or are otherwise of uncertain systematic position. 1974] Schuh: South African Orthotylinae and Phylinae 319 Lissoca psus B ergroth , 1903. Bergroth (1903) commented that the type specimen of L. was- manni Bergroth was received from E. Wasmann of Luxembourg, but he did not say where it was deposited. Until specimens of this species can be located and carefully examined, the subfamily place- ment must remain in question. Carvalho (1952a) placed this Mada- gascan genus in the Hallodapini. Lutheriella Poppius, 1913. I have not seen specimens of this genus from Ceylon, and there- fore cannot confirm its placement in the Pilophorini (Carvalho, 1952a). Myrmecoridea Poppius, 1921. The type specimens of this Australian genus are probably de- posited in the Hungarian Museum (Poppius, 1921) and will have to be examined before its subfamily placement in the Pilophorini (Carvalho, 1952a) can be confirmed. Myrmecoroides Gross, 1963, pp. 7-10. This very peculiar genus from Australia has apically convergent recurved parempodia, but the bizarre structure of the head and the strong ant-mimetic facies require that the male and female genitalia be examined before subfamily placement can be confirmed. Myrmecozelotes Berg, 1883. This Argentinian genus was placed in the Pilophorini by Car- valho (1952a). I have not examined specimens and have not found adequate information in the literature to determine its correct sub- family placement. Zanchisme Kirkaldy, 1904. This Neotropical genus probably belongs to the Orthotylini. Car- valho (1952a) placed it in the Pilophorini. I have not examined specimens or found adequate information in the literature to con- firm the subfamily placement of Zanchisme. ACKNOWLEDGMENTS I would especially like to thank Dr. James A. Slater of the Uni- versity of Connecticut, who took a genuine personal interest in this project. Dr. Slater’s willingness to discuss problems and to lend new ideas, and the unrestricted use of his fine library of hemipterological literature, helped immensely in bringing this work into its final form. 320 Entomologica Americana [Vol. 47 Dr. W. G. H. Coaton, Director, Systematic Entomology, Plant Protection Research Institute, Pretoria, provided untiring assistance during our stay in South Africa during 1967-1968 and was instru- mental in the success of the field work and local arrangements. The staff of the South African National Herbarium, Pretoria, graciously identified all of the host plant specimens. My wife, Janet C. Schuh, constantly stimulated my thinking and typed the original manuscript. Others who have read portions of the manuscript or otherwise helped in ways too diverse to enumerate here are Drs. M. H. Sweet, C. W. Schaefer, and H. W. Pfeifer, and Mrs. D. B. Wilcox. The following persons have supplied specimens or identifications and many also provided museum space and a pleasant working en- vironment: Dr. P. Arnaud, California Academy of Sciences; Dr. M. Beier, Naturhistorisches Museum, Vienna; Drs. R. C. Froesch- ner and J. L. Herring, United States National Museum; Mr. Wayne Gagne, Bishop Museum, Honolulu; Dr. U. Gollner-Scheiding, Hum- boldt University Zoological Museum, Berlin; Dr. A. J. Hesse, South African Museum, Cape Town; Dr. W. J. Knight and Mr. W. R. Dolling, British Museum (Natural History); Dr. Carl Lindroth, Lund University Zoological Institute; Mr. M. Meinander, Helsinki Uni- versity Zoological Museum; Dr. J. Munting, former curator, South African National Collection of Insects; Dr. P. I. Persson, Naturhis- toriska Riksmuseum, Stockholm; Drs. P. Basilewsky and G. Schmitz, Musee Royal de l’Afrique Centrale, Tervuren; Dr. D. R. Smith, Systematic Entomology Laboratory, United States Department of Agriculture; Dr. Henri Synave, Institut Royal des Sciences Naturelles de Belgique, Brussels; Dr. L. Vari, Transvaal Museum, Pretoria; Drs. Andre Villiers and J. Carayon, Museum National d’Histoire Naturelle, Paris; and Dr. P. Wygodzinsky, American Museum of Natural History, New York. The National Science Foundation, through a grant to the Uni- versity of Connecticut for graduate studies in evolutionary biology, and the University of Connecticut Research Foundation have pro- vided generous financial support. This has allowed me to travel to South Africa, to visit museums in the United States and Europe, and devote an uninterrupted year to the completion of this work. REFERENCES Acocks, J. P. H. 1953. Veld types of South Africa. Bot. Surv. s. Afr., Mem. 28: 192 pp. Ashlock, P. D. and J. D. Lattin. 1963. Stridulatory mechanisms in the Lygaeidae, with a new Amer- 1974] Schuh: South African Orthotylinae and Phylinae 321 ican genus of Orsillinae (Hemiptera: Heteroptera) . Ann. Entomol. Soc. Arner. 56: 693-703. Bailey, H. P. 1960. A method of determining the warmth and temperateness of a climate. Geogr. Annaler. 42: 1-16. Ballard, E. 1927. Some new Indian Miridae (Capsidae). Mem. Dept. Agric. India, Ent. 10(4) : 61-68. Berg, C. 1883. Addenda et emendanda ad Hemiptera Argentinae (2). Ann. Soc. Cien. Argen. 16: 5—32. Bergroth, E. 1903. Neue myrmecophile Hemipteren. Wiener Entomol. Z. 29: 235-238. 1917. On the types of the exotic Hemiptera Heteroptera described by V. Motschulsky. Revue Russe d’Entomologie 17: 96-109. 1925. On the “annectant bugs” of Messrs. McAtee and Malloch. Bull. Brooklyn Entomol. Soc. 20: 159-164. 1926. Uber einige athiopische Miridae (Hem. Het.). Deut. Entomol. Z. pp. 60-64. Brinkhurst, R. O. 1959. Alary polymorphism in the Gerroidea. J. Animal Ecol. 28: 211-230. 1963. Observations on wing-polymorphism in the Heteroptera. Proc. Roy. Entomol. Soc. London , Ser. A , Gen. Entomol. 38: 15- 22. Carayon, J. 1958. Etudes sur les Hemipteres Cimicoidea. -1. Mem. Mus. Natl. Hist. Natur., Paris, Ser. A, Zool. 16: 141-172. Carvalho, J. C. M. 1948. Mirideos neotropicais, XXX: Generos Ellenia Reuter, Eury- chilopterella Reuter, e Rhinocloa Reuter, com descrigoes de especies novas (Hemiptera). Bol. Mus. Nac., Rio de Janeiro, Zool. 85: 13 pp., 24 figs. 1951a. Neotropical Miridae (Heteroptera), XLVI. A new genus and two new species in the collection of the Natural History Museum in Vienna. Ann. Naturhist. Mus. Wien. 58: 103-107. 1951b. Five new genera and eleven new species of African Miridae (Hemiptera). Rev. Zool. Bot. Afr. 45: 100-115. 1952a. On the major classification of the Miridae (Hemiptera). (With keys to subfamilies and tribes and a catalog of the world gen- era.) An. Acad. Bras. Cienc. 24: 31-110. 1952b. Trois nouveaux genres de Miridae de Madagascar (Hemip- tera). Mem. Inst. Rech. Sci. Madagascar, Ser. E, Entomologie 1: 93-100. 1952c. Neotropical Miridae. LVI. Description of three new genera 322 Entomologica Americana [Vol. 47 and five new species from Brasil and British Honduras (He- miptera). Rev. Brasil. Biol. 12: 265-272. 1952d. Neotropical Miridae, 47: Notes on the Blanchard, Spinola, and Signoret types in the Paris Museum. Rev. Fr. Entomol. 19: 181-188. 1953a. Sur les Miridae Madagascar (Hemipteres). Mem. Inst. Rech. Sci. Madagascar, Ser. E, Entomologie 3: 41—51. 1953b. Neotropical “Miridae”, LXIII: A new genus and new species from South America (Hemiptera). Rev. Brasil. Biol. 13: 33- 40. 1954. Neotropical Miridae, LXXIV: Two new genera of Cylapinae from Brazil (Hemiptera). Proc. Iowa Acad. Sci. 61: 504- 510. 1955a. Keys to the genera of Miridae of the World (Hemiptera). Bol. Mus. Paraense Emilio Goeldi, Nova Ser. Zool. 11(2): 151 pp., 263 figs. 1955b. Analecta Miridologica: Miscellaneous observations on some American museums and bibliography. Rev. Chilena Entomol. 4: 221-226. 1956a. On a new genus of mirid from the Biliran Island (Hemiptera). An. Acad. Brasil. Cienc. 28: 215-216. 1956b. Insects of Micronesia. Heteroptera: Miridae. Honolulu, Bernice P. Bishop Museum, Vol. 7(1): 100 pp. 1956c. Neotropical Miridae, LXXVIII: A peculiar new genus of “Orthotylinae” (Hemiptera). Rev. Brasil. Biol. 16: 235-237. 1958a. Catalogue of the Miridae of the World. Part II. Subfamily Phyline. Arq. Mus. Nac., Rio de Janeiro 45: 216 pp. 1958b. Catalogue of the Miridae of the World. Part III. Subfamily Orthotylinae. Arq. Mus. Nac., Rio de Janeiro 47: 161 pp. 1960. Catalogue of the Miridae of the World. Part V. Bibliography and general index. Arq. Mus. Nac., Rio de Janeiro 51: 194 pp. 1965. A new species of “Melanotrichus” Reuter from Australia (Hemiptera, Miridae). Rev. Brasil. Biol. 25: 265-267. Carvalho, J. C. M. and J. Becker. 1959. Neotropical Miridae, LXXXV: New species of Orthotylinae in the collection of the United States National Museum (He- miptera, Heteroptera). An. Acad. Brasil. Cienc. 31: 113-117. Carvalho, J. C. M. and J. A. P. Dutra. 1959. Mirideos Neotropicais, LXXXVI: Revisao do genero Tuxen- ella Carvalho (Hemiptera). Arq. Mus. Nac., Rio de Janeiro 49: 139-153. Carvalho, J. C. M., J. A. P. Dutra, and J. Becker. 1960. Hemiptera Heteroptera: Miridae. South African Animal Life. Stockholm, Swedish National Research Council, Vol. 7: 446- 477. 1974] Schuh: South African Orthotylinae and Phylinae 323 Carvalho, J. C. M. and A. Fonseca. 1962. Mirideos Neotropicais, XCI: Uma tribo e dois generos novos (Hemiptera). Rev. Brasil. Biol. 22: 427-432. 1965. Mirideos Neotropicais, XCI: Sobre um genero e duas es- pecies novas de Orthotylinae (Hemiptera). Rev. Brasil. Biol. 25: 53-57. Carvalho, J. C. M. and D. Leston. 1952. The classification of the British Miridae (Hem.), with keys to the genera. Entomol. Mon. Mag. 88: 231-251. Carvalho, J. C. M. and T. R. E. Southwood. 1955. Revisao do complexo Cyrtorhinus Fieber-Mecomma Fieber (Hemiptera-Heteroptera, Miridae). Bol. Mus. Paraense Emilio Goeldi, Nova Ser. Zool. 11(1): 72 pp., 25 figs. China, W. E. 1932. A new species of Trichophthalmocapsus Poppius (Hemiptera, Heteroptera, Capsidae) from Kenya. Ann. Mag. Natur. Hist. (10)10: 594-597. 1938. Die Arthropodenfauna von Madeira nach den ergebnissen der Reise von Prof. O. Lundblad Juli-August 1935. Ill — Ter- restrial Hemiptera. Ark. Zool. 30A(2) : 1-68. 1945. New and little known West African Miridae (Capsidae) (Hemiptera-Heteroptera), Bull. Entomol. Res. 35: 171-191. 1963. Hemiptera-Heteroptera collected by the Royal Society Ex- pedition to South Chile 1958-1959. Ann. Mag. Natur. Hist. (13)5: 705-723. China, W. E. and J. C. M. Carvalho. 1951. A new genus of Hallodapini and two new species of the genus Nicostratus Distant (Miridae, Phylinae). Ann. Mag. Natur. Hist. (12)4: 1120-1125. COBBEN, R. H. 1968. Evolutionary trends in Heteroptera. Part I. Eggs, architecture of the shell, gross embryology and eclosion. Wageningen, Centre for Agricultural Publishing and Documentation, 475 pp. Davis, N. T. 1955. Morphology of the female organs of reproduction in the Mi- ridae (Hemiptera). Ann. Entomol. Soc. Amer. 48: 132-150. Delattre, R. 1950. Description de nouveaux Miridae africains (Hem.). Bull. Soc. Entomol. Fr. 55: 151-153. Dietz, R. S. and J. C. Holden. 1970. The breakup of Pangea. Sci. Amer. 223(4): 30-41. Distant, W. L. 1893. Biologia Centrali Americana. Insecta. Rhynchota. Hemiptera- Heteroptera. London, 1 (Supplement): 304-462, 1884-1893, 39 pis. 324 Entomologica Americana [Vol. 47 1904a. XI. — Rhynchotal Notes. — XX. Ann. Mag. Natur. Hist. (7) 13: 103-114. 1904b. XXI. — Rhynchotal Notes. — XXI. Ann. Mag. Natur. Hist. (7) 13: 194-206. 1904c. The fauna of British India including Ceylon and Burma. Rhyn- chota. (Heteroptera). London, Taylor and Francis, Vol. II, 503 pp. 1909a. Descriptions of Oriental Capsidae. Ann. Mag. Natur. Hist. (8)4: 440-454. 1909b. Oriental Capsidae. Entomologist. 42(550): 58-60. 1910a. Descriptions of Oriental Capsidae. Ann. Mag. Natur. Hist. (8)5: 10-22. 1910b. The fauna of British India including Ceylon and Burma. Rhyn- chota. Vol. V. Heteroptera: Appendix. London, Taylor and Francis, 362 pp. Emsley, M. 1969. The Schizopteridae (Hemiptera: Heteroptera) with descrip- tion of new species from Trinidad. Mem. Amer. Entomol. Soc. 25: 154 pp. Fieber, F. X. 1858. Criterien zur generischen Teilung der Phytocoriden (Cap- sini auct.). Wiener Entomol. Monatschrift 2: 289-327; 329- 347, 1 pi. 1860-61. Die europaischen Hemipteren. Halbflugler. (Rhynchota Heteroptera). Wien, Carl Gerold’s Sohn, 2 pts. I: 1-112; II: 113-444. 1864. Neuere Entdeckungen in europaischen Hemipteren. Wiener Entomol. Monatschrift 8(3) : 65-86. 1870. Dodecas neuer Gattungen und neuer Arten europaischer Hem- ipteren. Verhandl. K. K. Zool. Bot. Ges. Wien. 20: 243-264, pis. 5 & 6. Goel, S. C. and C. W. Schaefer. 1970. The structure of the pulvillus and its taxonomic value in the land Heteroptera (Hemiptera). Ann. Entomol. Soc. Amer. 63: 307-313. Gross, G. F. 1963. A new ant-mimicking mirid bug (Hemiptera-Heteroptera) from Victoria. Mem. Nat. Mus. Victoria 26: 7-10. Hahn, C. W. 1826. leones ad Monigraphium Cimicum. Lechner, Nurnberg. Hahn, C. W. and G. A. W. Herrich-Schaeffer. 1831-1853. Die wanzenartigen Insecten. Nurnberg, Vol. I-IX. Hennig, W. 1966. Phylogenetic systematics. Urbana, University of Illinois Press, 263 pp. 1974] Schuh: South African Orthotylinae and Phylinae 325 Hoberlandt, L. 1953a. Hemipteren-Heteropteren von Shaqlawa in Sudkurdistan. Beitr. Entomol. 3: 377-384. 1953b. Results of the Armstrong College Expedition to Siwa Oasis (Libyan Desert), 1935, under the leadership of Prof. Omer- Cooper. Bull. Soc. Fouad. Entomol. 37: 359-370. Hutchinson, J. 1946. A botanist in southern Africa. London, P. R. Gawthorn, 686 pp. International Code of Zoological Nomenclature. International Trust for Zoological Nomenclature, London, 1964, 176 pp. Kelton, L. A. 1959a. Synopsis of the genus Semium, and description of a new spe- cies from Mexico (Hemiptera: Miridae). Can. Entomol. 91: 242-246. 1959b. Male genitalia as taxonomic characters in the Miridae (He- miptera). Can. Entomol. Supplement 11: 72 pp. 1964. Revision of the genus Reuteroscopus Kirkaldy 1905 with de- scriptions of eleven new species (Hemiptera: Miridae). Can. Entomol. 96: 1421-1433. Kerzhner, I. M. AND T. L. Yaczewski. 1964. Order Hemiptera (Heteroptera) (In: Keys to the insects of the European USSR, Vol. 1, Apterygota, Palaeoptera, Hemi- metabola). 851-1118. (Translated from Russian.) Kirkaldy, G. W. 1902a. Fauna Hawaiiensis. Vol. Ill, Part II. Hemiptera. Cambridge Univ. Press. 1902b. Miscellanea Rhynchotalia No. 5. Entomologist. 35(475): 315-316. 1902c. Memoir upon the rhynchotal family Capsidae auctt. Trans. Roy. Entomol. Soc. London. 2: 243—272, pis. V and VI. 1904. Bibliographical and nomenclatorial notes on the Hemiptera — No. 3. Entomologist. 37: 279-283. 1905. Neue und wenig bekannte Hemiptera, Wiener Entomol. Z. 24: 266-268. 1908. A catalog of the Hemiptera of Fiji. Proc. Linnaean Soc. N. S.W. 33: 345-391, pi. IV. Knight, H. H. 1918. Synoptic key to the subfamilies of Miridae (Hemiptera-Heter- optera). J. N. Y. Entomol. Soc. 26: 40-44. 1922. The genus Cyrtopeltis in North America (Heteroptera-Miri- dae). Bull. Brooklyn Entomol. Soc. 17: 65-67. 1923. The Hemiptera or sucking insects of Connecticut. Hartford, State Geological and Natural History Survey, Bull. 34, pp. 422-658. 326 Entomologica Americana [Vol. 47 1935. Insects of Samoa. Part II, Hemiptera. Miridae and Antho- coridae. British Museum (Natural History), Fasc. 5, pp. 193— 228. 1941. The plant bugs, or Miridae of Illinois. Bull. III. Nat. Hist. Surv. 22(1): 234 pp. 1966. Schaffneria, a new genus of ground dwelling plant bugs (He- miptera, Miridae). Iowa State J. Sci. 41: 1-6. 1968. Taxonomic review: Miridae of the Nevada Test Site and the Western United States. Brigham Young Univ. Sci. Bull., Biol. Ser. 9(3): 282 pp. Kullenberg, B. 1944. Studien liber die Biologie der Capsiden. Zool. Bidr. Uppsala. 23: 522 pp., 25 pis. 1947a. fiber Morphologie und Funktion des Kopulations-Apparats der Capsiden und Nabiden. Zool. Bidr. Uppsala. 24: 218 — 418, 23 pis. 1947b. Der Kopulations-Apparat der Insekten aus phylogenetischen Gesichtspunkt. Zool. Bidr. Uppsala. 25: 74-90. Leston, D. 1957. The stridulatory mechanisms in terrestrial Hemiptera Heter- optera. Proc. Zool. Soc. London 128: 369-386. 1961. Testis follicle number and the higher systematics of Miridae (Hemiptera-Heteroptera) . Proc. Zool. Soc. London. 137: 89- 106. Lethierry, L. 1881. Liste des Hemipteres recueillis par M. Delauney a la Guade- loupe, La Martinique, et Saint-Barthelemy. Ann. Soc. Entomol. Belg. 25: 1-12. Lindberg, H. 1940. Inventa entomologica itineris Hispanici et Maroccani, quod a. 1926 fecerunt Harald et Hakan Lindberg. XXVI. Die Capsidenfauna von Marokko. Commentat. Biol., Soc. Sci. Fenn. 7(14): 1-55. 1951. Canariocoris, eine neue kanarische Miridengattung. Commen- tat. Biol., Soc. Sci. Fenn. 12(9) : 1-12. 1953. Hemiptera Insularum Canariensium. Commentat. Biol., Soc. Sci. Fenn. 14(1): 114-153. 1956. Uber einige Miriden in E. de Bergevins Sammlung. Notulae Entomol. 36: 53-64. 1958. Hemiptera Insularum Caboverdensium. Commentat. Biol., Soc. Sci. Fenn. 19: 246 pp. Linnavuori, R. 1961. Hemiptera of Israel, II. Acta Zool. Soc. ‘Vanamo’. 22(7): 51 pp. 1964. Hemiptera of Egypt, with remarks on some species of the ad- jacent Eremian region. Ann. Zool. Fenn. 1: 306-356. 1974] Schuh: South African Orthotylinae and Phylinae 327 1965. Some new and interesting Hemiptera from the Middle East. Ann. Ent. Fenn. 31(4) : 264-269. Maldonado, J. 1969. The Miridae of Puerto Rico (Insecta, Hemiptera). Univ. of Puerto Rico Agri. Expt. Sta., Tech. Pap. 45: 133 pp. 1970. New species in the genus Sericophanes Reuter, (Hemiptera: Miridae). Proc. Entomol. Soc. Wash. 72: 98-106, 20 figs. Miller, N. C. E. 1937. A new genus of Malayan Capsidae (Rhynchota) from palm. Bull. Entomol. Res. 28: 535. 1956. The biology of the Heteroptera. London, Leonard Hill Books, Limited, 162 pp. Moreau, R. E. 1966. The bird faunas of Africa and its islands. New York-London, Academic Press, 424 pp. Motschulsky, V. 1863. Essai d’un catalogue des insectes de File Ceylon. Bull. Soc. Nat. Moscou. 36: 82-88. Noualhier, M. J. M. 1895. Supplement a la liste des Hemipteres d’Akbes. Rev. Entomol. 14: 176. Odhiambo, T. R. 1958a. Notes on East African Miridae (Hemiptera). IV. A new species of Pseudosthenarus Poppius from Uganda. Ann. Mag. Natur. Hist. (13)1: 241-246. 1958b. Notes on the East African Miridae (Hemiptera). VI: A new species of Pilophorus Hahn from Uganda. Ann. Mag. Natur. Hist. (13)1: 326-330. 1958c. Notes on the East African Miridae (Hemiptera). IX: De- scriptions of a new species of Lasiolabops Poppius and of Psallus Fieber. Ann. Mag. Natur. Hist. (13)1: 758-764. 1959a. Notes on East African Miridae (Hemiptera). XII: New species of the genera Campylomma Reuter and Sthenarus Fieber. Ann. Mag. Natur. Hist. (13)1: 421-438. 1959b. Notes on the East African Miridae (Hemiptera). XIII: Twelve new species of Psallus Fieber. Ann. Mag. Natur. Hist. (13)2: 513-544. 1959c. Notes on the East African Miridae (Hemiptera). XIV: New genera and species of the tribe Hallodapini. Ann. Mag. Natur. Hist. (13)2: 641-687. 1960a. Notes on the East African Miridae (Hemiptera). XVII: A new genus of Dicyphini. Ann. Mag. Natur. Hist. (13)3: 343- 348. 1960b. Notes on the East African Miridae (Hemiptera). XVIII: A new genus of Pilophorini. Ann. Mag. Natur. Hist. (13)3: 393-400. 328 Entomologica Americana [Vol. 47 1960c. Notes on the East African Miridae (Hemiptera). XX: One new genus and two new species of Phylinae. Ann. Mag. Natur. Hist. (13)2: 513-520. 1963. Two new species of myrmecophilous Miridae (Hemiptera) from Tanganyika. Proc. Roy. Entomol. Soc. London, Ser. B, Taxon. 32: 111-113. 1967. A taxonomic study of some genera of the Ethiopian Miridae (Hemiptera) (Part 1). Bull. Inst. Fondam. Afr. Noire, Ser. A. 29: 1655-1687. Phillips, E. P. 1951. The genera of South African flowering plants. Bot. Sur. S. Afr., Mem. 25. Poppius, B. 1912. Die Miriden der Athiopischen Region. I. Mirina, Cylapina, Bryocorina. Acta Soc. Sci. Fenn. 41(3): 203 pp. 1913. Zur Kenntnis der Miriden, Isometopiden, Anthocoriden, Na- biden und Schizopteriden Ceylon’s. Entomol. Tidskr. 34: 239-260. 1914a. Miriden der Athiopischen Region. II. Macrolophinae, Hetero- tominae, Phylinae. Acta Soc. Sci. Fenn. 44(3): 136 pp. 1914b. Ubersicht der Pilophorus — arten nebst Beschreibung verwand- ter Gattungen (Hem. Het.). Ann. Entomol. Soc. Belg. 58(8): 237-254. 1914c. Einige neue Miriden — Gattungen und Arten aus Nord-America und Cuba. Ann. Soc. Entomol. Belg. 58: 255-261. 1921. Beitrage zur Kenntnis der myrmecoiden Heteropteren. Miridae. Ann. Mus. Nat. Hungarici. 18: 31-69, 2 pis. POYNTON, J. C. 1965. The Amphibia of Southern Africa. Ann. Natal Mus. 17: 1- 334. Reuter, O. M. 1876a. Capsinae ex America boreali in Museo Holmiensi asservatae, descripta. Ofv. K. Vet. Akad. Fork. 32: 59-92. 1876b. Sur le Capsus 4-guttatus Kirschbaum. Petrop, Nouv. Entomol. 2(145): 26. 1879. Till kannedomen om mimiska Hemiptera och deras lefnads- historia. Ofv. Finska Vet.-Soc. Fork. 21: 141-198. 1881. Analecta Hemipterologica, zur Artkenntnis, Synonymie und geographischen Verbreitung der Palearktischen Heteropteren. Berliner Entomol. Z. 25: 155-196. 1882. Ad cognitionem Heteropterorum Africae occidentalis. Ofv. Finska Vet.-Soc. Fork. 25: 1-43. 1883. Hemiptera Gymnocerata Europae. Hemipteres Gymnocerates d’Europe, du bassin de la Mediterranee et de l’Asie Russe. III. Acta Soc. Sci. Fenn. 13: 313-496, 5 pis. 1884. [Diagnosen neuer arten] in P. M. Ferrari: Materiali per lo 1974] Schuh: South African Orthotylinae and Phylinae 329 studio della fauna Tunisia raccolti de G. e L. Doria V. Rincoti. Ann. Mus. Civ. Stor. Nat. Genov. 2: 482. 1890. Capsidae novae ex Africa boreali, descripsit. Rev. Entomol. 9: 255-260. 1891. Ein falscher und einer echter Sthenarus (Capsidae). Wiener Entomol. Z. 10: 49-51. 1900. Capsidae novae mediterraneae descripsit. II. Ofv. Finska Vet.- Soc.Forh.42 : 259-267. 1901. Capsidae rossicae descriptae. Ofv. Finska Vet.-Soc. Fork. 43: 161-194. 1903. Capsidae ex Abessinia et regionibus confinibus enumeratae novaeque species descriptae. Ofv. Finska Vet.-Soc. Forh. 45 (6): 1-18. 1904. Ad cognitum Capsidarum aethiopicarum. I. Ofv. Finska Vet.- Soc. Forh. 46(10) : 1-8. 1905a. Hemipterologische Spekulationen. I. Die Classification der Capsiden. Festschr. Palmen 1: 58 pp. 1905b. Ad cognitum Capsidarum aethiopicarum. II. Ofv. Finska Vet.- Soc. Forh. 47(10): 1-22. 1905c. Capsidae stalianae secundum specimena typica redescriptae, I. II. Ofv. Finska Vet.-Soc. Forh. 47(12): 1-20. 1905d. Ad cognitum Capsidarum aethiopicarum. III. Ofv. Finska Vet.- Soc. Forh. 47 (22) : 1-9. 1907a. Capsidae novae in insula Jamaica mense Aprilis 1906 a D. E. P. van Duzee collectae. Ofv. Finska Vet.-Soc. Forh. 49(5) : 1-27. 1907b. Ad cognitum Capsidarum aethiopicarum. IV. Ofv. Finska Vet.- Soc. Forh. 49(7) : 1-27. 1909. Bermerkungen fiber nearktische Capsiden nebst Beschreibung neuer Arten. Acta Soc. Sci. Fenn. 36(2): 86 pp., 3 pis. 1910a. Neue Beitrage zur Phylogenie und Systematik der Miriden nebst einleitenden Bermerkungen fiber die Phylogenie der Heteropteren-familien. Mit einer Stammbaums-tafel. Acta Soc. Sci. Fenn. 37 (3) : 169 pp. 1910b. Mitteilgungen uber einige Hemipteren des Russischen Reiches. Horae Soc. Entomol. Rossicae. 39: 73-88. Schmitz, G. 1968. Monographie des especes Africaines du genre Helopeltis Sig- noret (Heteroptera, Miridae) avec un expose des problemes relatives aux structures genitales. Ann. Mus. Roy. Afr. Cent., Ser. Quarto Zool. 168: 247 pp., 80 figs. 1969. Description d’un nouveau genre et d’une nouvelle espece d’Hal- lodapini (Miridae, Phylinae) du Sudan. Ann. Entomol. Fenn. 35: 72-81. 1970. Contribution a la fauna du Congo (Brazzaville). Mission A. Villiers et A. Descarpentries. XCVIII. Hemipteres. Miridae 330 Entomologica Americana [Vol. 47 et Isometopidae (lre partie). Bull. Inst. Fondam. Afr. Noire, Ser. A. 32: 501-530. Scott, J. 1872. Notes on some Corsican Insects. Entomol. Mon. Mag. 8: 193-195. SlGNORET, V. 1862. Coriodromus n.n.f. Ocypus Montruzier. Bull. Soc. Entomol. Fr. 4: 5. Singh-Pruthi, H. 1925. The morphology of the male genitalia in Rhynchota. Trans. Roy. Entomol. Soc. London. 5: 127-267, 32 pis. Slater, J. A. 1950. An investigation of the female genitalia as taxonomic characters in the Miridae (Hemiptera). Iowa State J. Sci. 25: 1-81, 7 pis. 1964. Hemiptera (Heteroptera) : Lygaeidae. South African Animal Life. Stockholm, Swedish National Research Council, 10: 15- 228. Slater, J. A. and T. Schuh. 1969. New species of Isometopinae from South Africa (Hemiptera: Miridae). J. Entomol. Soc. S. Afr. 32: 351-366. Southwood, T. R. E. 1953. The morphology and taxonomy of the genus Orthotylus Fieber (Hem., Miridae) with special reference to the British species. Trans. Roy. Entomol. Soc. London. 104(11): 415-449, 152 figs. 1962. Migration of terrestrial arthropods in relation to habitat. Biol. Rev. (Cambridge) 37: 171-214. Southwood, T. R. E. and D. Leston. 1959. Land and water bugs of the British Isles. London and New York, Frederick Warne and Co. 436 pp. Stal, C. 1858. Orthoptera och Hemiptera fran Sodra Afrika. Ofv. Svenska K.Vet-Akad. Fork. 15: 307-320. Stuckenberg, B. 1962. The distribution of the montane paleogenic element in the South African invertebrate fauna. Ann. Cape Prov. Mus. Natur. Hist. 2: 190-205. 1969. Effective temperature as an ecological factor in Southern Af- rica. Zool. Afr. 4: 145-197. Sweet, M. H. 1964. The biology and ecology of the Rhyparochrominae of New England (Heteroptera: Lygaeidae). Part I. Entomol. Amer. 43: 124 pp. Uhler, P. R. 1876. List of the Hemiptera of the region west of the Mississippi 1974] Schuh: South African Orthotylinae and Phylinae 331 River, including those collected during the Hayden explorations of 1873. U. S. Geol. and Geog. Surv. 1: 269-361, pis. 19-21. 1887. Observations on some Capsidae with descriptions of a few new species. Entomol. Amer. 3: 29—35. 1890. Observations on North American Capsidae, with descriptions of new species (No. 5). Trans. Maryland Acad. Sci. 1: 73- 88. USINGER, R. L. 1946. Hemiptera Heteroptera of Guam. In: Insects of Guam. — II. Bull. Bernice P. Bishop Mus. 189: 58-91. Van Duzee, E. P. 1910. Descriptions of some new and unfamiliar North American Hemiptera. Trans. Amer. Entomol. Soc. 36: 73-88. 1916. Check list of the Hemiptera (excluding Aphididae, Aleurod- idae, and Coccidae) of America North of Mexico. New York, New York Entomological Society, 111 pp. 1917. Catalog of the Hemiptera of America North of Mexico (ex- cepting Aphididae, Coccidae, and Aleurodidae). Univ. Calif. Pubs. Entomol. 2: 902 pp. 1921. Characters of some new species of North American hemipter- ous insects, with one new genus. Proc. Calif. Acad. Sci. 11: 111-134. Wagner, E. 1952. Die Tierwelt Deutschlands und der angrenzenden Meeresteile nach ihren Merkmalen und nach ihrer Lebensweise. 41-teil- Blindwanzen oder Miriden. Jena, Gustav Fischer, 218 pp. 1953. Nachtrag zu “Einige neue Miriden-Arten aus Agypten”. Bull. Soc. Fouad. Entomol. 37 : 477. 1955. Bermerkungen zum System der Miridae (Hem. Het.). Deut. Entomol. Z. 2(5) : 230-242. 1956. On the genus Strongylocoris Blanchard, 1840 (Hemiptera, Het- eroptera, Miridae). Proc. Entomol. Soc. Wash. 58: 277-281. 1957a. Die systematische Stellung der Gattung Amixia Reuter, 1883. (Hem. Het. Miridae). Deut. Entomol. Z. 4: 22-25. 1957b. Laurinia Reuter, 1884, und Formicocoris Lindberg, 1940 (Hem. Het. Miridae). Deut. Entomol. Z. 4: 263-267. 1959. Bermerkungen zum System der Miridae. II (Hem. Het.). Deut. Entomol. Z. 6: 1-7. 1961. Hemiptera Insularum Madeirensium. 2. Der Cephalocapsus- komplex auf den Makaronesischen Inseln (Hem. Het. Miri- dae). Commentat. Biol., Soc. Sci. Fenn. 24: 85-110. 1962. Berichtigung zu einigen neueren Arbeiten iiber Miridae (Hem. Het.). Notulae Entomol. 42: 83-84. 1965. Die Gattung Atomoscelis Reuter, 1875, und ihre Verwandten in der Palaarktis (Hem. Het. Miridae). Notulae Entomol. 45: 74-92. 332 Entomologica Americana [Vol. 47 1970a. tiber Paramixia Reuter, 1900 (Hem. Het. Miridae). Nachr. Bayer. Entomol. 19: 1-5. 1970b. Zur Systematik der Hallodapinae van Duz. (Heteroptera, Mi- ridae). Reichenbachia. 13: 149-155. Wagner, E. and H. H. Weber. 1964. Faune de France. 67. Heteropteres, Miridae. Paris, Federa- tion Frangaise des Societes de Sciences Naturelles, Office Cen- tral de Faunistique, 589 pp. Woodroffe, G. E. 1957. A preliminary revision of the British Psallus Fieber (Hem., Miridae), with a description of a new species. Entomol. Mon. Mag. 93: 258-271. Wygodzinsky, P. 1966. A monograph of the Emesinae (Reduviidae, Hemiptera). Bull. Amer. Mus. Natur. Hist. 133: 614 pp. Zimmerman, E. C. 1948. Insects of Hawaii. Vol. 3. Heteroptera. Honolulu, University of Hawaii Press, 255 pp. Zinderen Barker, E. M. van. 1967. Upper Pleistocene and Holocene stratigraphy and ecology on the basis of vegetation changes in the sub-Saharan Africa, pp. 125-147. In: Background to evolution in Africa, Walter W. Bishop and J. Desmond Clark, Editors, Chicago, University of Chicago Press. Zinderen Barker, E. 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