—— i, r i i te eon etyal Gh taO) tu 3 * ‘anpve Soeke ' ett, Vee vices iy pots Ry Ne Ms Re aus thie aymatanantonh Ata Aare ae 4 b 4 s i ea 4 Rh Wa De 2, ROMs ee a RT sTs Oe HS Pe Heine 2 te ‘ : - " ; DUM ANGI so err eS na Mahan Agta MLB de PATA 9 RAM - ; : , " e " Sub adm hr Bane Rag Rei PALM LerreN NST PAB 9 BA ERA MB ons ee r > - , : : . : 0 nt eth Mi 9 an ‘ a far hin Ma Hs NAN ANNI Mic ATER MAH AYPS NIM Zire tiah ait WPM, Hie ONES Hala aden eal Ped oi ae a8 ae Me IE wee Me tae awit e sepata Name tenctiadon taeiAe Foam tm oeetima Ten he May, Maa thns mts we 0 A a? ee a Ral a ee oat wpe ee PS nian a taeda OF, ey ALi Bima ett DR NAMEN A EM iret DO Le ia ad A tele ont mla ge me g tee ARORA On BO ak al 0 on oS BoM AAT ome hint 9 pt teaR ual ot minadin® “at HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology ; 44 : , ie — i f] S if : 7 ae a 7 4 a, : iv : | j . i - ) i ’ or i j ; " is 2 ‘ w.~.~.©.©.©_©. 2, 0,=,.°.".9,-5"5"9"s797s707070,8.0.0.0 9 © 0 8-0'0 0'0 0°00 9.0.0.02.000 0.000.000 0000200 0 e706 0705007070 070"6"0-0- 6.0.6.0 0000008 e eee ata ate ote eee O60 0 000 00 08 060806 '0 060s" s 0's e's" o' e's"! Not ot0100e7 01070. 0.0 0-0-0 -0-0.0.0.0.9.9, 95 "a nQe a e000 0's O60 2006 0.00000 '0'0-0-0-0-0-0-0-0-0-0'0-0'0-0'0'0-0°0'0' 0000's 0's" e: 00066 6 060000088 0900000008898 00 00950055808 F Ose ee es eee 2 2 2 oo ss oe oo = =~. -5-5-5 es 68” aa et ot oe Bi BN a i SCIENCE BULLETIN Mc LIBRARY DEC 24 1996 HARVA UNIVERS ry Se oo es RRR OD ei ocesesesecetesccsatereterecotecetece A SYNOPSIS OF THE KANSAS MOSSES WITH KEYS AND DISTRIBUTION MAPS RRS SRR RS nn Ee DR Sg *otee oe etatatetetatetetatetetatetetetatetetetetocecen 0.0.0 0.0.0.0 0 0-90 By Steven P. Churchill I ag tg" a a OCH IR IR SR SR Se eraPeraPeaateretas PR RR es Vol. 53, No. 1, pp. 1-64 October 22, 1985 SESE eee eeeeeeenneeeeneeaee eatetetetetece ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas University Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its inception, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now exhausted. However, most volumes of the University of Kansas Science Bulletin are still available and are offered, in exchange for similar publications, to learned societies, colleges and universities and other institutions, or may be purchased at $20.00 per volume. Where some of these volumes were issued in parts, individual parts are priced at the rate of 2 cents per page. Current policy, initiated with volume 46, is to issue individual papers as published. Such separata may be purchased individually at the rate of 3 cents per page, with a minimum charge of $1.00 per separate. Subscriptions for forthcoming volumes may be entered at the rate of $20.00 per volume. All communications regarding exchanges, sales and subscriptions should be addressed to the EXCHANGE LIBRARIAN, UNIVERSITY OF KANSAS LIBRARIES, LAWRENCE, KANSAS 66045. Reprints of indivudal papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletzn. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor William L. Bloom Editorial Board Philip W. Hedrick Rudolf Jander Charles D. Michener David Paretsky Norman A. Slade George W. Byers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 53, No. 1, pp. 1-64 October 22, 1985 A Synopsis of the Kansas Mosses with Keys and Distribution Maps! STEVEN P. CHURCHILL? Division of Biological Sciences, University of Kansas, Lawrence, Kansas 66045 CONTENTS ABSTRACT 5 Il INTRODUCTION. Eo Were sae History oF Moss @srpecrines AND MD RIGA TONE Stor Ine coves aoe ee ee ee Kansas Moss Types AND NOMENCLATURAL COMBINATIONS . 5 DIVERSirY. DISTRIBUTION AND: BIOGEOGRAPHY ...4-.4.-24.05+.)08 0s 5se ogee REPRODUCTIVE | PIOLOGY: tan etcans beeen 5 ae Ga We See oe eR aera IGEY TOITHE GENERA = 2 25 esos Se ee eres Rm Bee Lr GT eae wae et La eRe as capeinsc Witt KEVe TO TE SREGIES”,.. on se es Uae ar ee oe Se as Ria ee. SUMMARY OF CORRECTIONS FOR THE KANnsAs Moss Rion tee Clete eee eA en 80) DISTRIBUTION NUAR Sey ake Coe enue ome a, tee ae ear iee ee aes | ae a eo gen ee eee ei AGCKNOWEE DEMENTS) cee. cirulanouet ie. helen 2 ose SEEN SRE Ras ep me ee RM ore ee OZ GER RATUIRENGinED eta oe 8 eth ROR Meee nO) |S nee na ets eg ie eee ee ee ee” SEE NDS pes ca beeue Re whee gels Se A BN Se ole on OE eatced Jet) Ai ce Rite oct Re ear ie OAL ABSTRACT There are 151 species of mosses distributed among 77 genera and 31 families in Kansas. The average number of mosses encountered in counties in the west, central and east is 7, 15, and 28, respectively; the percent of species unique to each of these regions is 4, 12, and 32. The Sorensen Similarity Index shows a 83% similarity between species of the east and central, 42% between west and central and only 20% between west and east. The reproductive biology is expressed in types of sexuality (i.e., monoicous or dioicous and success of sporophyte production) and asexuality. The number of species that are dioicous and monoicous is about equal throughout the state. Overall, sporophyte production is greater in monoicous than in dioicous species. A higher number of dioicous species produce sporophytes in the east than in the west. Monoicous species produce sporophytes about equally state-wide. The percent of asexually reproducing species is relatively low for the state, but slightly greater in the west than in the east. Ecological parameters can be expressed as a moisture gradient for the mosses: hydric-9% , mesic-56%, and xeric-34%. Habitat substrates for mosses are: soil and Se ease vegetation 70%, saxicolous (rock) 40%, and corticolous (bark) 17%. The North American affinities of the Kansas mosses are: eastern 54%, widespread 29%, western 7%, central 5%, northern 4%, and endemic 1%. Many of the distributions can be attributed to the late-Wisconsinan, and early and mid-Holocene events as well as to the existing physiographic features found in Kansas. ! This study is dedicated to a remarkable naturalist and phycologist, the late Dr. Rufus H. Thompson, a University of Kansas faculty member from 1942-1979. ? Present address: New York Botanical Garden, Bronx, N.Y. 10458. 2 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN INTRODUCTION The Kansas moss flora exhibits an interesting sample of two major biotic regions, the grasslands and the eastern deciduous forests. The position of Kan- sas offers a nearly uniform east-west transition from the oak-hickory and post-blackjack oak forests in the east with intercalated prairie, to the tall- grass bluestem prairie, the mixed prairie, and the shortgrass prairie plains in the west (see Fig. 4). The transition is due in part to the interac- tion of physiographic and climatic con- ditions shaped and modified by the past glacial events. This biotic diversity is correlated with the recorded richness of the Kansas moss flora having a total of 151 species distributed among 77 genera and 31 families. The first significant account of Kan- sas mosses was published by Minnie Reed 90 years ago. Studies at that time and later provided a wealth of data, both in the form of collections and in publications that have made possible this present work. This study is the first modern floristic treatment for mosses of any Great Plains state. The entire region of the Great Plains is one of several areas in North America that have been cited as in need of intensive field work (Schofield, 1980). My own field work conducted throughout Kan- sas resulted in some 2000 collections (see last page of distribution maps for the counties collected in by the author). Particular emphasis was placed on east- ern Kansas where more detailed work was required to abut an important work already completed in western Kansas by Smith (1966). I have also examined about 4000 — specimens mainly from the New York Botanical Garden (NY), the Missouri Botanical Garden (MO), the Nebraska State Herbarium (NEB), Kansas State Uni- versity Herbarium (KSC), University of Kansas Herbarium (KANU), and Iowa University Herbarium (IA). The intent of this synopsis is to provide a guide for the identification of the Kansas mosses, and also to intro- duce several historical and biological aspects that should be of interest to biologists and naturalists alike. The keys were developed to identify sterile material, i.e., haploid gametophytic leafy plants, but in a few cases some genera or species can only be positively identified with fully developed sporo- phytes, e.g., Bruchia or Bryum. While the majority of Kansas mosses can be identified, it is always best when possi- ble to collect material with capsules. The keys are original though much use has been made of existing keys and descriptions. In the preparation of this treatment I have made use of the fol- lowing texts: Crum, 1976; Grout, 1928-1940; Redfearn, 1972; and Weber. 1973: The annotated species list includes information on the vegetation type and substrate that the moss is generally associated with. Reproductive infor- mation includes the presence of spo- rophytes (C.fr.) and any specialized asexual features that are present (e.g., gemmae etc.). A subjective estimate is given as to the frequency that a partic- ular moss was observed in Kansas and the distribution is indicated based on the dot maps and extrapolation as to where any particular moss is likely to be found. The county dot maps are based entirely on specimens that I have actually examined in the course of this study. There are three exceptions based on literature records that I have not examined but have accepted: Atrichum undulatum based in part on Ireland (1969a), Funaria americana based in part on Smith (1980), and Physcomitrium col- lenchymatum based on Crum and Ander- son (1960). Nomenclature generally A SYNOPSIS OF THE KANnsAs MOSSES 3 follows Crum et al. (1973), except for the Grimmiaceae (Churchill, 1981) and the Mniaceae (Koponen, 1968). This paper represents the seventh in a series on the mosses of the Great Plains. History oF Moss COLLECTING AND PUBLICATIONS Kansas has been fortunate to have had a significant number of both natu- ralists and botanists who collected mosses over a long period of time. Moss collecting in Kansas dates back at least to the 1870’s with the collections of Elihu Hall. The decades of 1880 and 1890 were a very active time. Much of the diversity of Kansas mosses became -» a. established by several important local collectors, in particular: Joseph Henry (1816-1887), Saline County (very little is known about Henry although he was important in the early botanical history of Kansas); Grace Meeker (?-1948), Franklin County; and Minnie Reed (1867-1959), Pottawatomie, Riley and Wyandotte counties. Minnie Reed contributed more to our early knowl- edge of Kansas mosses than any other individual. She was born in St. Clere, Pottawatomie County, Kansas in 1867; her parents were farmers. At the age of 15 she entered what is now Kansas State University at Manhattan, earned a B.S. in 1886, anda M.S. in 1893 (see Fig. 1). Little more is known except Fic. 1. Minnie Reed, seated center. Photograph taken by W. A. Kellerman at Kansas State College, Manhattan, Summer of 1891. Individuals are from left to right: standing, Geo. K. Thompson and C. H. Thompson; seated, Kate (Oldham) Sisson, Minnie Reed, and Elsie (Crump) Ames. Photograph courtesy of Rufus Thompson. 4 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN that she was a teacher and died on September 12, 1959 in Los Angeles, California. The publication by Minnie Reed (1893-1894) on the Kansas mosses surely must be one of the very first state moss floras in the United States. Her work is based on collections she made, supplemented by collections made by others (Bartholomew, Beede, Bennett, Knerr, Smyth, McClung, Meeker). Included also were species reported by others such as Renauld and Cardot. A total of 165 species distributed among 50 genera were re- ported for Kansas by Reed. In addition to keys and illustrations, she provided an introduction to the morphology and history of mosses. Reed obtained assis- tance on the identification or verifica- tion mainly from Elizabeth Britton of the New York Botanical Garden. Reed’s study suffered only by the natu- ral inclusion of ‘‘species of reports,’’ especially from the publications of Re- nauld and Cardot (see below), who where less than meticulous in the nam- ing or identification of Kansas mosses sent to them by Joseph Henry (see also section on Kansas types). While collections were made after the activity of two decades prior to the twentieth century, it was not until shortly before 1950 that further activity was initiated and continued off-and-on until the present time. Recent collec- tors such as Ronald L. McGregor, Harold L. Smith and the present au- thor have collected widely throughout the state and contributed to an under- standing of distribution patterns. In addition to the significant work by Minnie Reed, another notable work on the Kansas mosses is that by Harold L. Smith (1966). This study included the Great Plains proper and the Arkansas River Lowlands (about the western half of Kansas). Smith’s study gives a fine account of the observational as- pects of moss ecology and species asso- ciations encountered in that region. There is also an account of the minute ephemeral mosses and their possible adaptations to the Plains environment. This is followed by an annotated list of 117 species and varieties, which in- cludes descriptions of habitat and gen- eral physiographic areas. The serious deficiency of this work lies mainly with misidentifications, particularly for spe- cies reported as new state records. I have indicated these in the synopsis and in the summary of corrections at the end of the synopsis. Also, in Smith’s study no locality data were provided for most species, so that dis- tributional information could not be documented. Other primary publications based on collections are Renauld and Cardot (1892), Gier (1949), McGregor (1950), and Churchill (1979, 1980). Minor publications are Rau (1884, 1885a, 18856, 1886), Renauld and Cardot (1888, 1889, 1890a, 18906), Smyth (1890, 1891-1892, 1893-1894), Gier (1940), Hartman (1956), McGregor and Hartman (1956), Cridland (1959, 1960), Waller and Bass (1967), Magill and Kirkpatrick (1976), McGregor (1976, 1977), and Churchill (1982). Major catalogues providing lists of the Kansas mosses, usually based on litera- ture reports, are Renauld and Cardot (1892), Smyth (1892), Smyth and Smyth (1911), and Churchill (1976). Unpublished works concerning Kansas mosses are Duncan (1959), Anony- mous (1966), and Smith (1961). The unpublished work by Smith (1961) is an important contribution and con- cerns the mosses of the Chautauqua Hills in southeastern Kansas. Gruger (1973) reported the only known subfossil mosses for Kansas from the late Wisconsinan that in- cluded Mnium affine Bland s.1. (= Plagi- A SYNOPSIS OF THE KANSAS MOSSES 5 omnium), Mnium seligert Jur. (ques- tionable determination; = Plagiomnium elatum), Mnium affine f. integrifolium (Lindb.) Mild. (= Plagiomnium affine s.l.), Calliergon cf. ~— sarmentosum (Wahlenb.) Kindb., and Bryum pseudo- triquetrum (Hedw.) Gaertn., Meyer & Scherb. between the dates of 24,500 + 800 BP to 15,880 +600 BP. In the studies by Hartman (1956), Duncan (1959), Anonymous (1966), and McGregor (1976), as far as I am aware, no specimens were collected to verify the study or at least no indication was given that specimens were depos- ited in any herbaria. For example, the ecological study on the Kansas prairie bryophytes by Duncan (1959), a poten- tially important study, cites such taxa from prairie sites as Ditrichum pusillum, Fissidens osmundioides, Helodium pal- udosum and Tortula mucronifolia—all of which are very likely incorrectly deter- mined! These studies should be used with caution. The earlier moss collections are now found in scattered herbaria, mostly outside Kansas. The Henry collections are most likely deposited with the Re- nauld and Cardot collections at Paris, France (PC). I have examined some of these collections, mostly types, as well as some apparent duplicate type collec- tions at the New York Botanical Gar- den (NY). The majority of the Meeker and Reed collections are deposited at the New York Botanical Garden (NY), although a number of the Reed collec- tions can also be found at Kansas State University (KSC). The entire bryo- phyte collection of the University of Kansas Herbarium containing 7,224 specimen packets (4,402 mosses and 2,822 liverworts) was donated to the New York Botanical Garden on May 5, 1969. These collections contained all of the specimens collected by both McGregor and Smith as well as a few other individuals. My own work on the Great Plains region, including Kansas, started in the early 1970’s with most of my first set of specimens being depos- ited with the Missouri Botanical Gar- den (MO). Beginning in 1978 and continuing through mid-1980, the Bio- logical Survey of Kansas initiated a survey of the bryophytes. In addition to a complete set deposited at MO of my collections, a number of duplicates can be found at the University of Kansas Herbarium (KANU); there are also a few collections made by various other individuals contained in KANU. Fur- ther representive duplicates of my Kansas collections are deposited at the following herbaria: ALTA, B, CANM, COLO. DUKE. F Hie GIRO aE HSC, KRAM, KSC, MICH, MEXU, NEB, NY, SMU, SMS, TENN, UBC, UK, and US. A Great Plains exsiccatt, ‘‘Musci Planitiebus Incolae,’’ is in preparation by the author and will contain a number of Kansas mosses. This will be distributed by the New York Botanical Garden to 25 institu- tions (ALTA, B, BA, BM, C, CANM, CBG, ‘COLO, DURE. FEL-G. Er KSC). LE, MEXU, MICH, Me; NIGH. NY, P@2.PE. RM) sUmand UBC). Kansas Moss Types AND NOMENCLATURAL COMBINATIONS The following list gives the names of various moss taxa named as new to science from collections made in Kan- sas. These names constitute an impor- tant historical and nomenclatural record in the taxonomy of North American mosses. The original name with author and place of publication is listed. I have included the location and habitat of the type if provided with the original publication. Finally the pres- ent status of each name is noted. 6 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Amblystegium riparitum BS var. serratum Ren. & Card. var. nov. Renauld, F. & J. Cardot. 1889. New mosses of North America. III. Bot. Gaz. 14:98. ‘‘Kansas; Saline County, roots of tree (Joseph Henry).’’ =Am- blystegium riparium (Hedw.) B.S.G. Aschisma kansanum Andr. sp. nov. Andrews, A.L. 1915. Bryological notes I. Aschisma kansanum, a new species with remarks upon the genus. Torreya 15:63-67. ‘‘Prairies of western Kansas, silicious soil.’’ This species is still ac- cepted; the present author is studying the genus. Holotype: NY! Atrichum undulatum Beauy. var. al- tecristatum Ren. & Card. var. nov. Renauld, F. & J. Cardot. 1890. New mosses of North America. IV. Bot. Gaz. 15:58. ‘‘Kansas: Saline County (Joseph Henry).’’ =Atrichum_ altecris- tatum (Ren. & Card.) Smyth & Smyth. See Ireland (1969a, 1970). Barbula Henrict Rau. sp. nov. Rau, E.A. 1886. Fourth contribution to the knowledge of Kansas mosses. 5th Report on the Progress of the Wash- burn College Biological Survey of Kan- sas. Page 172. ‘‘Hab. On rocks, Saline county, (Joseph MHenry).’’ = Ptery- goneuron subsessile (Brid.) Jur. var. henrici (Rau) Wareh. This variety is still rec- ognized. I consider this only a minor form not worthy of taxonomic status. Campylopus Henrict Ren. & Card. sp. nov. Renauld, F. & J. Cardot. 1888. New mosses of North America. I. Bot. Gaz. 13:197-198, with Plate 14. “‘Kansas: Saline County, on sandy ground, where it was discovered by the late Joseph Henry.’’ = Dicranella heteromalla (Hedw.) Schimp. Coscinodon Renauldi Card. sp. nov. Renauld, F. & J. Cardot. 1890. New mosses of North America. III. Bot. Gaz. 15:41-42, with Plate 6b. ‘‘Kan- sas: Saline county (Joseph Henry).’’ = Jaffueliobryum rau (Aust.) Thér. Lec- totype PC! Britton (1890) correctly pointed out that Coscinodon Ranauldi was a synonym of the then Coscinodon raui (Aust.) Lesq. & James. This spe- cies is now considered a member of the genus Jaffueliobryum (Churchill, unpub. data). Fissidens obtusifolius Wils. var. Kan- sanus Ren. & Card. var. nov. Renauld, F. & J. Cardot. 1890. New mosses of North America. III. Bot. Gaz. 15:40. ‘‘Kansas: Saline county (Joseph Henry).’’ This variety is still recognized, however the status is under investigation by the present author. Microbyrum Floerkeanum Sch. var. Henrict Ren. & Card. var. nov. Renauld, F. & J. Cardot. 1889. New mosses of North America. III. Bot. Gaz. 14:91. ‘‘Kansas: Saline county, on sandy ground (Joseph Henry).’’ = Phascum cuspidatum Hedw. Physcomitrium kellermani Britt. sp. nov. Britton, E.G. 1894. A revision of the genus Physcomitrium, with descriptions of five new species. Bull. Torrey Bot. Club 21:204-205, with Plate’ 200. ‘‘Manhattan, Kansas, Kellerman (1889).”’ =P. pyriforme Hedw. sensu Crum et al. (1973), but still recognized by Lawton (1971). Further studies on this genus are warranted to determine the range of phenotypic variation. This moss was named in honor of William Kellerman (1850-1907), a noted my- cologist and a member of the faculty of Kansas State Agriculture College at Manhattan (Willard, 1909). Ptychomitrium pygmaeum Lesq. & James sp. nov. Lesquereux, Iu; é& IP, James! 1678- 1879. Description of some new species of North American mosses. Proc. Amer. Acad. of Art & Science 14:136. ‘‘Hab. Near the Neosho River, Kan- sas, and at Bolivar, Missouri (E. A SYNOPSIS OF THE KANSAS MOSSES Vl Hall).”’ =P. incurvum (Schwaegr.) Sull. A number of nomenclatural com- binations for mosses were made _ by Smyth and Smyth (1911) in a catalogue of the Kansas flora. These combina- tions were brought to my attention by Dr. Ronald McGregor, University of Kansas, and their validity was verified by Dr. Marshall Crosby, Missouri Bo- tanical Garden. The names are listed as they originally appeared in the cata- logue by Smyth and Smyth (op. cit.). Each name is annotated as to its pres- ent status. Fissidens Kansanus Ren. & Card. This appears to be a new combination based on the variety originally recog- nized by Renauld and Cardot (1890). The proper citation should be Fissedens kansanus (Ren. & Card.) Smyth & Smyth. =F”. obtusifolius Wils. var. Kan- sanus Ren. & Card. Mnium elatum (B. & S.) Apparently based on Mnium affine Funck var. elatum B.S.G. = Plagi- omnium elatum (B.S.G.) Kop. Atrichum altecristatum (R. & C.) Based on Atrichum undulatum (Hedw.) P. Beauv. var. alte-cristatum Ren. et Card. The: proper citation should be A. al- tecristatum (Ren. & Card.) Smyth & Smyth. The combination by Smyth and Smyth (op. cit.) supersedes that of Ireland (1970). Brachythectum acuminatum (Hedw.) This combination was made earlier by Austin. Brachythecium setosum (S. & L.) The combination apparently based on Hedwig’s Leskea setosa Hedw. = Brachythectum acuminatum (Hedw.) Aust. Eurhynchium hians (Hedw. ) Combination made earlier by Sande- Lacoste in 1866. Rhynchostegium serrulatum (Hedw.) The combination by Smyth and Smyth is superseded by Jaeger in 1878. Amblystegium orthocladon (Beauv. ) This combination was made earlier in 1892 by Macoun. This taxon is pres- ently not known for Kansas. Probably can be referred to A. tenax. (2?) Amblystegium irriguum (Hook. & Bruch, Schimper and Guembel made this combination in Bryologia Eu- ropeae in 1855. Amblystegium adnatum (Hedw.) This combination was made earlier by Austin in 1870. = Homomallium adnatum (Hedw.) Broth. Amblystegium cariosum (Sull.) Based on Hypnum riparium Hedw. var. cariosum Sull. in Rau. =Amblystegium riparium (Hedw.) B.S.G. Amblystegium serratum (R. & C.) Smyth and Smyth combination based on A. ripartum (Hedw.) B.S.G. var. serratum Ren. & Card. = Brachythecium acutum (Mitt.) Sull. This Brachytheciwm is not known for Kansas: if at asea species of Brachythecium, then it is likely to be B. acuminatum (Hedw.) Aust. since the Renauld and Cardot type was collected in tree roots. Campylium chrysophyllum (Brid.) Combination made earlier by J. Lange in 1887. Harpidium aduncum (Hedw.) =Dre- - panocladus aduncus (Hedw.) Warnst. Diversity, DistRIBUTION AND BIOGEOGRAPHY The accumulated data on the mosses of Kansas are sufficient to permit anal- yses of species diversity and patterns. The pattern of diversity of the Kansas mosses exhibits an expected decrease in species richness from east to west. ‘To gain some quantitative expression of this, the two vertical tiers of counties both in the east and west, plus three central tiers of counties were examined for the number of species present in each county (Fig. 2). The mean aver- age calculation exhibited a_ halving 8 ‘THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Fic. 2. Diversity of the Kansas mosses. Means based on numbers of species per county. Percentages of similarity for any two areas based on the Sorensen Similarity Index! (see text for further discussion). WIS, = C/!/2 (A+ B) X 100) where IS, = Sorensen’s Index of Similarity, c= number of species held in common by 2 areas, A = total number of taxa in first region, B= total number of taxa in second region. from the east to the central zone, and again from the central to the west zone. This appears to be a characteristic pat- tern and probably true of various groups of taxa. The number of species unique to each zone shows a similar pattern: the east zone with 32%, the central zone with 12%, and the west zone with 4%. ‘The Sorensen Similar- ity Index was used to examine relation- ships between the three zones (Fig. 2). The similarities between the east and central zones were highest (83%), while the similarity between west and central zones were nearly half of the first (42%), and finally 20% similarity existed between the east and west zones. The methods used to examine the distribution patterns and_ the _bio- geography involve consideration of only those species that have been col- lected well enough to permit analysis. Those species that are found through- out (i1.e., showing a ubiquitous dis- tribution) are not considered (ca. 20 species). Taxa known only from a few localities are used but with caution. In any biogeographical study the main process is the search for recurring pat- terns, whether the emphasis is ecologi- cal (MacArthur, 1972) or historical (Croizat, 1964). Once the patterns are established, the second phase involves hypothesis testing. The 151 species can be divided into several broad distributional categories, each of which have several subcatego- ries. To obtain a better impression of the patterns in the distribution of Kan- sas mosses, 52 distributions were su- perimposed on the same map—this produced a number of recurrent pat- terns. These patterns have been re- duced to five major patterns and are characterized by the following repre- sentative species: (1) widespread west- ern distribution—Aloina rigida, Barbula acuta, Didymodon rigidulus, Hypnum vauchert, Jaffueliobryum rau, J. wrightit, Pterygoneurum subsessile, and Tortula ruralis; (2) broadly eastern distribu- tion—Anomodon attenatuas, A. minor, A. rostratus, Atrichum angustatum, Brachythe- cium oxycladon, Campylium chrysophyllum, Entodon compressus, E. seductrix, Hedwigia A SYNOPSIS OF THE KANSAS MOSSES 9 ciliata, Homomallium adnatum, Leskea gra- cilescens, and Lindbergia brachyptera. ‘The remaining southeastern or easternmost distributions represent varying degrees of the eastern distribution: (3) extreme eastern—Aulacomnium heterostichum, Bartramia pomiformis, Bryum pseudotrique- trum, Climactum americanum, Dicranum scopartium, Plagiomnium ciliare, Pohlia nutans, and Thutdium delicatulum; (4) mostly southeastern—Drummondia pro- repens and Leucodon julaceous; and (35) sandstone sites in extreme east central —Atrichum oerstedianum, Bryhnia gramint- color, Bryoandersonia tllecebra, and Diphys- cum foliosum. What remains is to develop a hy- pothesis which will allow for the most parsimonious explanation of the extant moss distributions in Kansas. To do so, we must also consider the North Amer- ican affinities of the Kansas mosses and the historical events that led up to the patterns we see today. The broad North American affinities for the Kansas mosses are as follows: 54% of the Kansas mosses are eastern, 29% are widespread, 7% are western, 5% are central, 4% are northern, and 1% are endemic. These percentages compare roughly to that given for the Kansas vascular plants (Table 1). ‘This correla- tion suggests an important point about the nature of moss distributions. First of all, the distribution of the mosses is most often the same as those of vascular plants (Anderson, 1971; Crum, 1972). This distribution has been influenced by primarily historical events, particularly of the late Quater- nary period. Mosses are not haphazard in their distribution, and the chance of long-distant dispersal, not unlike the insects discussed by Udvardy (1969), are probably less than 1%. While it is difficult for the dispersalist to under- stand, the chance of long-distance dis- persal is the exception (but verified by TaBLeE 1. Percentage of the North American affinities of the Kansas mosses and vascular plants. Vascular plant data modified from Bare and McGregor (1970); based on 479 spe- cles. Region Mosses_ Vascular plants Eastern 54 49 Widespread 29 14 Western 7 17 Central 5 8 Northern 4 13 Endemic 1 0 Southern 0 9 a few islands), while continental land- forms provide evidence for the rule! The controlling factors of distribution (up to the level of biotic formation) are climate and_ geographical/geological factors. The influence of the Pleistocene pe- riod on extant distributions is best doc- umented by Wisconsinan and _post- Wisconsinan studies. Earlier periods of the Quatenary are less well known. Kapp (1965) does however present evi- dence concerning the Illinoian glacial and Sangamon interglacial of north- west Oklahoma and southwest Kansas. The evidence is meager but suggests the existence of Picea and Pinus during the Illinoian in this region, while in the Sangamon there was marked decrease of Picea and some decline of Pinus. This, together with the earlier work by Hibbard (1955) on fossil vertebrates, suggests a rather mesic environment for this area compared to the present interglacial. The Wisconsinan and Holocene are better documented for Kansas, at least the eastern region. Gruger (1973) stud- ied two sites in northeastern Kansas. Dates on the strata for which micro- and macro-fossil material were col- lected indicate Picea in the area from 10 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN about 23,000 yr. BP to at least 15,000 yr. BP. So dates are available for the Wisconsinan maximum and the early Holocene times. From the studies by Gruger (1973), Stewart (1979), and others, a hypothesized Wisconsinan glacial maximum vegetation for Kan- sas and adjoining states has been pro- posed by P.V. Wells (unpubl. map). This includes Rocky Mountain Con- ifer in central and most of the west in Kansas, while northwest and eastern Kansas was composed of Taiga (Fig. 3). Areas further south and southeast of Kansas were thought to be composed of transitional Taiga-Deciduous forests. It was some time after 15,000 yr. BP that prairie-deciduous elements entered northeastern Kansas replacing the Taiga vegetation. To the west, the Rocky Mountain Conifer contracted and the grasslands expanded. During this period (10,000 to 5,000 yr. BP) the climate apparently became arid, marked by increased grass and com- posite pollen in the studies by Gruger. Northeastern Kansas then assumed an appearance that one generally sees to- O | it TRANSITION Fic. 3. Past vegetation of Kansas and surrounding states, ca. 14,000 yrs BP (courtesy of P. V. Wells, unpubl. map). A SYNOPSIS OF THE Kansas MosseEs 11 day with transitional prairie and oak- hickory forest (Fig. 4; see Kuchler, 1974, for a detailed description of Kan- sas vegetation). A similar sequence of events was recorded by King (1973) for the western Missouri Ozarks. The distribution patterns of the Kansas mosses based on the evidence of the Quaternary events can be sum- marized. Western and central (more xeric types) elements could have been derived from the south or southwest in one or two periods of aridity at the close of the Wisconsinan or during the Hypsithermal. Although it is possible that these elements were already associ- ated with the Rocky Mountain Conifer as they are today in the Pine Ridge of grama-buffalograss wheatgrass-bluestem - needlegrass sandsage-bluestem TT bluestem-grama prairie bluestem prairie western Nebraska. Associated western elements include such genera as Barbula, Didymodon, Hypnum, Jaffuelio- bryum, Pterygoneurum and Tortula. East- ern and northern elements were likely associated in part with the Taiga in eastern Kansas during the maximum Wisconsinan. Several of the Kansas mosses are today found commonly in the southern Taiga forests of Canada as well as the eastern deciduous forests farther east; examples include Aulacom- nium, Bartramia, Bryum, Dicranum, Leu- cobryum, Pohlia, and Thuidium. Other taxa more typical of the rich eastern deciduous forests include Anomodon, Campylium, Drummondia, Entodon, Ho- momallium, Leskea, Leucodon and a — ———_ — = —_ Rea, ly U/L Wie Sas RUZ wh Win wl, ) Wi wy wey, S N 2 Wa, o. tet? whi Wy, Wi a Wy er - re ’ . =f i. us =: we fea! 33S, ee SS a 3. 43a Wy, - Breer ye ve 5 7 oe Fens 2 Wr OD S nd ° Willy wally ° Min}o win Mr, wily 3 Peers PO) & Ue ory P\4 2° ra Wty wir lyfe She Soo % wh wlll Wy Wy wa °° s Po 0°o,0 po 2 co Wh SMM Wig Wt Wy aa | a Pa AS I =~ | Wily wy, Mr, oh BEA WW, ly wit Ze Pos a Sg) Or . Cy > 4 neh Ra Wn eee Me wes § Gtaskier aires Ure errant g aks eames oe 6 6. Alar cells conspicuously differentiated, enlarged, often yellow to golden vyellow;Ooperculum. lomg OStrale «ain «oss so ae pe lee Sematophyllum 6, Alay-cells meonspicuous, not, cnlargeds. 24.4 sae Sut a eae 7 7. Leaf tip hyaline serrate; cells incrassate (thick-walled), with 1-2 papillae (oltem branched. sot 5 one ¥ dys gay ee een PS ae Hedwigia 7. Leaf tip not hyaline; cells incrassate or not, smooth not papillose ....... 8 8. Branch leaves large, usually 1-2 mm or more long; on trees in eastern IGanisast. 926, Feit ee ss Bea a ud Be ec etheee aes cemmreeeen mone oe oem Leucodon 8. Branch leaves small, usually 0.5 mm long; costa often present or appearing absent; on soil or rock in western Kansas ..... Pseudoleskeella PLEUROCARPOUS; LEAVES COSTATE 9, ueat.cells smooth, amot.papillose:: On trees* 224 5. 5.5 = = ee ee Drummondia Plants lacking above combinations; leaf cells not rounded, usually short to long rhombodial, linear or oblong; on various substrate ............ 22 22. Leaves lanceolate, very small (up to 1.2 mm), costa nearly precurrent; leaf margins slightly to distinctly serrate, especially at base; brood- bodies‘usually present onsleat tips 4-0 Rhynchostegiella 22. Leaves lacking above combinations, brood-bodies never present... . . 3) Leaves ovate-lanceolate, tip long acuminate; margins distinctly irregularly dentate;usually foundvabove base of trees. . 2-0... eee Fabronia Leaves usually lanceolate to ovate-lanceolate; regularly serrate if toothed; vVanousssubstrates, but usually motitrees = +) =] ee Dek 24. Leaves julaceous, deeply concave, ovate to round-ovate . . . Bryoandersonia 24. Leaves not julaceous, slightly or not at all concave, lanceolate... .. . 29 Leaves plicate; alar cells distinctly quadrate, median cells long rhombodial OT MMe AT ia Oe eae... Menon ee tains arn ene ean ee Brachythecium Leaves smooth, not plicate; alar cells usually not distinct, median cells shortly: rhombodialirs.% 225 eae Re epee ieee 26 26. Apex cells shorter than median cells; margins distinctly serrate ... . og He Wonks GUE ae oe ae ae olen oe aa ee Oe tea a ar Sates Pe ea Eurhynchium 16 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN 26. Apex cells as long as median cells; margins smooth to slightly dentate See he St ag a a Se cea ear ag dt ogee ces eae ote s Oe ene ne Amblystegium PLEUROCARPOUS; LEAF CELLS PAPILLOSE OR MAMMILLOSE 27. Leaf cells papillose on back only (lower or abaxil surface) ........ Bryhnia 27. eat-cells: papillose or mammiullose on both suriaces. 252. 2. 6 2 2 oe 28 28. Leaf cells pleuripapillose (several papillae per cell)......... Anomodon Jose discellssumipapllosea i 4 seals aaa ae cee a at ee ee 29 29. Leaf tip hyaline; plants loose and scattered, usually on upper tree trunks Se Pe cm Orne Ae NA s. Leh eA os Reet Ae ree Bg vee ee Lindbergia 29. Leaf tip not hyaline (if hyaline then leaves broadly ovate, i.e. Thelia); if on PREC SAGE Miaty DAS Cir thm AS Uh at She ale cece ge a ge hee ee eee a 30 30. Plants lacking or with only a few scattered paraphyllia; papillae low, Shrerhielye lon ning aoe ao Sarge arent ee nae. = hee aI et Leskea 30. Plants with abundant paraphyllia; papillae usually very distinct... . . oil 31. Leaf apex cell with 2 (3) papillae; plants distinctly 1- several pinnately ramehedneet:, Pe Sati ch etn Lh Re ote de ceeeeen 205 oon siete meee Thuidium 31. Leaf apex cell with 1 papillae; plants indistinctly pinnate or once pinnate . . 32 32. Plants julaceous; leaves broadly ovate, often with cilia on margins; papillae Inehvanduottem branched! 2.0 2c « i.2 + ete pe Gea eens Thelia 32. Plants not julaceous, leaves generally narrowly ovate, margins lacking cilia? papillae low and not branched’... .. =... = ss Benue sce: Haplocladium ACROCARPOUS (SPOROPHYTE USUALLY PRODUCED AT TIP OF STEM OR BRANCH) 33 Leayesilackime lamellae onadaxial (upper) surlace s . 2s pegs eos ee oS) 33> eaves with lamellae;presention adaxial-surtac€ 4.4 36%, chesnut tees 2 a 34. Lamellae filamentous, simple or branched, not in rows; leaf tip cucul- late leaves.shont. 1-2/9 mini lone. 2 8 2 et ee o> iene Aloina 34. Lamellae in rows, of few to several cells high; leaf tip acuminate to acute-rounded; leaves mostly longer than 2.S:mim: .2252 243. 4 30 35. Leaves ovate to spatulate, with hyaline hair point; lamellae 2-4....... Sle Ae, ao aa MONEE Me argc abate, aise sete ied einen ie IS See ees Pterygoneurum 35. Leaves lanceolate, with hyaline hair point generally lacking; lamellae mostly PTC ALET UNA kam. 5 Washes cae apse: Cagt ohaytaso id, vag, ae ty BEN on en el eer 36 36. Lamellae forming 4-6 (8) rows; leaf base not distinctly expanded nor Sheathing meats soe foo By cetyl tee tt ALS eee eae” Qe ee es Atrichum 36. Lamellae forming more than 10 rows; leaf base expanded and sheathing ee ae ee ee eS ee er ne. OFC ates 37 37. eaves) 125=35:.0: (4) smmclong-capsule:cylandrical™ 3. one ee Pogonatum on Leaves (4) 4°9-3° (10) mim tone, capsule-ancled 1. 2.0018 eaters ape eee 38 Jon lamellae terminating with aysquare cell; 320. oh Polytrichastrum 38. Lamellae terminating with a distinct notched or rounded cell... .. . a EO ak ees re ena enya oes cay ee eae rere Polytrichum oF igeal :CelistsmmOOU 2.5 sys a eco ee Oe OS ce aa eer ee 40 39, eat icells mammiullosesor-papilose’.5--.5e4ene veer were ee ea eee eee 60 40. Plants ephemeral, small, very inconspicuous; capsules cleistocarpous (lacking a defined capsule mouth and operculum—lid); and seta equal or shorter than pericheatial leaves (except Anomobryum)........... al a gel. 425) APO) POR Os 4 28 fe 49: an8)2 one ils Dos o3:. A SYNOPSIS OF THE KANSAS MOsSES 17 40. Plants larger, conspicuous; capsules stegocarpous (well defined capsule mouth and operculum); and seta usually longer than perichaetial leaves Paes ear sce Ria Seley SRN ted Sch aac tie tal a ee crag males DEEL ee Tat ae 46* eavieshihoOau lyr Vale meet for teenie ee cs 2.9 ce aii aan roe aes may (ean ae 42 [eaviestlanccolate tovlime aig east rah tek: nese ys home ae ae ee ran a ae aS 42. Leaves julaceous, shiny golden brown, apex acute to ovate . . Anomobryum 72 wlicaves Not julaccous. Greens acCUiMiM Ale ry. 3) s.csa. esos Soy ae Pyramidula Leaves setaceous; costa extending well beyond lamina; capsule round-ovoid, ANGUS ERIC E MMe CK FASE IN ei eed, tree ed overs) meee Sara Pleuridium Leaves linear to lanceolate; costa ending at tip or just beyond ........ 44 44. Costa excurrent; capsule pyriforme, neck region distinct....... Bruchia 44. Costa precurrent; capsule ovoid-rounded, neck region lacking. .... . 45 Leaf margins irregularly serrated; spores less than 100 um ..... Ephemerum Leaf margins smooth to evenly dentated; spores about 200 um _ .. . Archidium 46. Leaf margins differentiated, usually a border of linear cells longer or Gifierent trom. macdiarmCellls * 2: ots ets ca eo eee eer core kes ee ee ee 7 46. Leaf margins similar to median cells, or very indistinctly differenti- 2 LeXe | eee er eee See er ae a en ee Re PMR Ty aN lee nl SS 50 Leaves linear-lanceolate; costa long excurrent; capsule pyriforme..... . nae a eee diy tA Leas eis Seen a a tet ate Rte a a oe ca REN Leptobryum Leaves lanceolate-ovate to oblong; costa not long excurrent.......... 48 48. Leaves distinctly serrate from base or middle to apex... .. . Plagiomnium 46; heaves smooth-or only serrate:imcupper. third, 45.4% 2a) eee 49 Leaves large, up to 8-10 mm long; forming a distinct rosette of leaves Bae PY NO a OS rd at na en nee tig Se ete aan, ie ae Rhodobryum Leaves medium up to 5 mm long; not forming a rosette of leaves... . Bryum 50. Basal leaf cells differentiated, usually + inflated and rusty golden yellow ig eager set aii to th ewe emer Re hunaes Rane Ten Ota ceawpaee tess ee eee eee Dicranum 50: Basal leaticells.notidiflerentiateds 2c nt & cee ee ete oil eaves. selacous: tO: limCarss.: % 1. cus pase geet Seno fae ee kee eee a2 eaves lanceolate tonovate;, ovate-oblong uaa. .< Jerse oe ee a9 52. Lamina extending only a fifth or less along the long excurrent costa; capsule not constricted below mouth; annulus large and compound. . BSP SoMa Sas? yi By rnin lS cnc cpa cre Cevigge eae carat oy WE Sata Peers coe SN aa Ditrichum 52. Lamina extending half or more along the costa length; costa excurrent or precurrent; capsule constricted below mouth; annulus small or absent SMe Oph tnds mor Beis iyy ses CaN eic ect Roe gala je cadees ten ee Dicranella eat imedian::cells:isodiamietnie s.. Faceod, core SSsec sea bene eee ea ee 54 Leat-median, cellssrhomboidal to linear... ¢ 5. = 2-.s. + a ee 58 54. Leaves oblong to oblong-ovate, margin sharply serrate from middle to apex; costa ending:several cells below apex:. 22 2... am) aus Aulacomnium 54. Leaves oblong-lanceolate to lanceolate, margins smooth to dentate or bluntly serrate; costa precurrent or if ending several cells below apex then AGUStiNG Ely sSOuc—m 22 bt cawit pnw lovesae: nae op us oaecete ee a ee 55 * Couplet inadvertently left out. Leaves several layers thick, outer layers composed of empty hyaline cells surrounding a central layer of green cells; plants; palestor white; forming sctshionsi a eyerei-icias ie ene yeas ei caste enn ae ae nee Leucobryum Leaves lacking the above combinations of characters, plants various in color and habit................ 46 18 D0. D0. OW Oye Jo), og. 61. 61 63. 63. 69: 65: 67. O72 69. 69. Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN Weaf-apex hyalme:;-eithera short‘or long awit ees ae 56 ealrapex: lacking. auliyalime stipes ete he asta teh PELL re eine ee 57 56. Leaves ovate, apex acute to rounded; unistratose; calyptra campanulate Pe a eee een et Ae eNE IE NES t oe Cee Conc eee eee Jaffueliobryum 56. Leaves lanceolate to oblong-lanceolate, apex acute to acuminate; partially bistratose--caly ptraseuculllatere’ U9. tae ee ec ete Grimmia Leaf base + ovate, upper half becoming narrow and acuminate, apex not MOUGIE Gls ta Hecrrame? Semaine AES Wei sn emo a i Sema eeirane some Canoe Gane Barbula Leaf base and upper half continuous, not notably differentiated, gradually acuminate; apex notched; capsule curved and grooved ¥: .... 42 2: Ceratodon 58. Leaves lanceolate to ovate-lanceolate; either cells long rhomboidal or plantsapalevoreeiics sire. os ee ee et eee a Ren Gk ee ee Pohlva 58. Leaves oblong-ovate to obovate; plants neither with long cells or pale CRECIE eee es Gee he Sik ec) aaah dw Be eae Roe ea aReee eee te an meee 2 a9 Capsule inclined, mouth small, peristome present; costa usually ending in ADE XO SHOU CXCUITEME. G's) 2 2 oi 5 ¢ ac eacte me Pea te eg ene We eee Funaria Capsule erect, mouth large, peristome absent; costa usually ending several CEM SHClOW GH ADEX: cscgacait 2 oh a ane Goh kage HERS ae aut Oe em ee Physcomitrium LEAF CELLS PAPILLOSE OR MAMMILLOSE 60: Leal marginsancuryed throughout 3.50... 5-4. ess 0s 4 essere eeeeos 61 60: "Leat mareimsaplane orerecurved <7. G05 Gina 2 ajah ao eo Ne nese oe eee 62 Capsule immersed’ among leaves, cleistocarpous.....' 22 22s ae 2. Astomum : Capsule exserted above leaves, stegocarpous:;...4.2.454 54 Bieta are Werssia 62. Leaves ecostate, silver-gray green color; on sandstone rock . . . . Hedwigia 62> Leaves costate, variously colored; substrate various. :<-.. <>... 3. 63 eat margin distinctly serrate (occasionally serrulate) :. <2 % acess) use 64 Leaf margin smooth (rarely dentate, if so usually toward apex). ....... 70 64. Margin serrate at juncture of hyaline and papillose cells .... . Eucladium 04. Mar ein: serration mot-comtined: to jumcttive. 0 %..2 2 28: te oe ee 65 Leaf sheath distinct, of hyaline cells, upper lamina cells mammillose; calyptra often remaining attached to mature capsule neck ........ Timmia Leaf lacking a differentiated sheath and upper lamina; calyptra never attached to, mmattire Capsule. 21a fe ete a oe SM sues oe bo bo aon teae weee ee meme 66 66. Leaf median cells oblong-linear, papillose at lower and/or upper ends of Cel ee A iG Ae ee Regs oe. ania ea ie ae Rl ni ony Cook cre ee Philonotis 66. Leaf median cells quadrate-rounded, papillae not confined to ends of CES, -s5.5 wane og OR ee Sal ris ee, ee ea eet 67 Leaf oblong to spatulate; brood-bodies present in leaf axil ....... Hyophila Leaf-lanceolate to: linear: brood-bodiés absents— "2. Nels > ie ee 68. Plants very small, up to 2.0-2.5 mm tall; capsule cleistocarpous; ProlLoOnemia; PErsistemirs ss ce a ete ee cee eg eet Ephemerum 68. Plants larger, usually over 4 mm tall; capsule stegocarpous; protonema MOL PELSISCEME Sn, fe =e Sorc Sica Shoe teste a ie megs es ee 69 Leaves oblong-lanceolate, serrate only at apex; stalked propagula usually PEESENt; marshes amd Swamps. 295% a. Gacy i ae Aulacomnium Leaves linear-lanceolate, serrate except at base; propagula absent . . Bartramia Hal 7A hes): Tek 13: Use dei: Tile (oO TS): 81. 81. A SYNOPSIS OF THE Kansas MosseEs 19 70. Leaf hyaline basal cells extending up along margin, forming a V-shape: leavesnoblone-lanceolatey 66 “oy i.e ae ae ee eee ee ee Tortella 70. Leaf hyaline basal cells not forming a V-shape; leaves variously shaped Boe MEE ae nae eee ee a ee am aa eS eR eras er rs en ie Pea 71 Leaf cells with mammillae or papillae rounded, not C-shaped........ . qe Eeatscelisiwithtpapillaei@-shaped a 7..5 = ass. ee 78 72. Plants small, up to 2 mm tall; confined to margin of clear or white Guantz;pepbles-*cleistocanpous...9. 4. 2 oss on ee Aschisma 72. Plants larger, 4 mm or more tall; not confined to quartz pebbles; SIGS OCAMBOUS as tee oad si it Oa es See seth ap ae ce ee 13 Upper leaf cells partly bistratose; basal cell walls thick; costa shortly excurrent to precurrent; capsule immersed among leaves; columella remain- ingrattached!tooperculunr ees. oa ele ee eel ee eee © Schistidium Plants lacking above combination of characters; never with columella kemaimine, attached; Go, OPerculuim « pit toes keene ene eee eee We (oP lantssproducinsapropaculave sie (Wien yo ok, eee eee 1K) (Ae Plants mot producing propagulars tech) Gunso meni tha ae eee 76 Propagula present in leaf axils, round, not stalked; leaves lanceolate, costa PORE CUTE 162s faire ha" Seed so ace ce NE ie Hearne cart cea ait Serge. so eae Didymodon Propagula present on leaf, clavate, stalked; leaves ovate-lanceolate. . . Barbula 76. Leaves strongly decurrent; apex obtuse to rounded; costa ending below apex, plants Of wet, calcaneous-sites. 4 0. sete es ee Didymodon 76. Leaves not or very short decurrent; apex various; costa usually precur- FEN COVER CULLEME: plants) MOL Ol Wet sIteS. 2.2) 2 229, =) ee ee Ei} Costa ending just below apex or excurrent in a short hyaline awn; peristome teeth in 8 pairs; calyptra plicate, often hairy; plants on trees or limestone ROCKY Hasyn nas Sad dant Scag ftps a) Ee hae catia, vege eee Tee ae ot Orthotrichum Costa precurrent or excurrent but not hyaline; peristome of 16 divided teeth; calyptra smooth, not hairy; plants on soil, rarely on sandstone or NIMES COME SHOCKS. Co tA ah, Mae Phe ee ence ecg oa 2 ahaha A Barbula 78. Leaves oblong-lanceolate; apex acuminate, ending in a short awn; cleistocarpous; seta shorter than capsule::on soil. 92.2 Phascum 78. Leaves oblong to oblong-ovate, rarely oblong-lanceolate; apex acute or often abruptly rounded to obtuse; stegocarpous; seta longer than capsuleon: varioussulostrates... sy. 7. eee ee 79 Fropaculaipresentam leat axils: onetrees. ©)... oc ee Tortula Propagulatabsent:mot-onvtrees* = \2 5.22 6 2 2. ee eee 80 80. Leaves apiculate or mucronate, not ending in a hyaline awn. ...... 81 80" Leaves long excurrent, ending in a: hyaline: awn.) =.= 82 Leaf tip reflexed and pinched; costa ending below apex.......... Tortula Leaf tip not reflexed nor pinched; costa precurrent or shortly excurrent . cred he Ghee eet glce 2 ebh 3 DN Si castle Net grace tein deen ke Desmatodon 82. Excurrent awn serrate, awn a third or more the length of lamina; leaves When MmMOISt squarcrose-recurved:) 4 .)..6 4.) eaee ee Tortula 82. Excurrent awn smooth to + dentate, awn a fourth or less the length of lamina; leaves erect spreading when moist...) oe Desmatodon 20 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN ANNOTATED LIST WITH KEYS TO THE SPECIES Acaulon C. Muell. Pottiaceae . Costa present from tip to base of leaf; leaves only slightly papillose at back to completely smooth; spores smooth to finely granulate........ 1. A. muticum . Costa present from tip to mid region but lacking at leaf base; leaves papillose on both sides; spores smooth to finely papillose........ 2. A. schimperianum . Acaulon muticum (Hedw.) C. Muell. In open grassy areas; on soil. Uncommon, but probably overlooked. Presently known only from the central portion of the state. . Acaulon schimpertanum (Sull.) Sull. ex Sull. G Lesq. In prairies; on partly disturbed soil. Only a single collection known, made by Smith 2681 (NY!) from Reno County. Rare. Aloina (C. Muell.) Kindb. Pottiaceae. . Aloina rigida (Hedw.) Limpr. In High Plains grassland; on calcareous, gypsum and sandstone rock and soil. Based on the collections that I have made both here in Kansas and elsewhere (Churchill, 1979a, 19796, 1983) it appears that this species is probably not an uncommon component of the High Plains moss flora. Western half of Kansas. Amblystegium B.S.G. Amblystegiaceae . Median leaf cells linear, often vermicular, averaging 7-15:1.... 1. A. ripartum . Median leaf cells oblong to rhomboidal, averaging less than 7:1 ........ 2. 2. Costa noticeably thick, averaging 40 um or more in width at leaf base . Se ea eee ONT Rea eine? WRN ir Oey er Be pee eee we rots Ss 3. A. tenax 2 “Costa. =. thim, Jlessthan 40 umm: width at leaf base... 2 oS ee ee 3. Costa. extending into the acumimate leaf+tip; cells 1-2 (3):1 < .. 2. 5. A. varium . Costa extending 1'/2-°/+ way up the leaf, but not into the acuminate leaf tip; (tes | SOS) ca eee co te eee are ee ee a Pee er poe os a2, fF lieatca. 12-2: 2 mmm lone= cells:Cas Oil nu. cane tty occ ke 4. A. trichopodium A eaicau0.0-1 mrmelone, cells ca... 32 icon se cose, Waar acne ese 2. A. serpens . Amblystegium riparium (Hedw.) B.S.G. (Leptodictyum) In woods or marshy areas; most often found on decaying logs or over decaying marsh vegetation, occasionally on soil. C.fr. Occasional. Probably scattered throughout Kansas, particularly the eastern half. . Amblystegium serpens (Hedw.) B.S.G. (A. juratzkanum) In open to closed woods and prairies; found on trees (Fraxinus pennsylvanica and Ulmus sp.), on decaying logs, on soil, soil covered sandstone rocks, and on decaying marsh vegetation. C.fr. Frequent. Scattered throughout the state. . Amblystegium tenax (Hedw.) C. Jens. (Hygroamblystegium tenax and A. fluviatile) Along and in creeks both in woods and prairies; on marsh banks, creek beds, decaying logs but most often found on either limestone or sandstone rocks. C.fr. Common. Throughout Kansas. . Amblystegium trichopodium (Schultz) Hartm. (Amblystegium kochi and Leptodic- tyum trichopodium) In open to closed woods, less common in prairies; on soil, decaying logs and over sandstone rock. C.fr. Occasional. Throughout Kansas. . Amlystegium varium (Hedw.) Lindb. In open to closed woods, occasional in A SYNOPSIS OF THE KANSAS MOSSES 21 prairies; on soil, sandstone rock, decaying logs and trees (Carya sp. and Quercus muhlenbergi). C.fr. Occasional. Throughout Kansas. NOTE: Some authors still recognize the ecological genera Hygroamblystegium and Leptodictyum but these have no phyletic meaning at the generic level. The Am- blystegium species that one finds in the Plains region exhibit a wide range of phenotypic variation of the gametophyte. Experimental laboratory work and biometrical analysis of this genus would be most worthwhile. The many species, varieties and forms that have been named in the past are certainly only environmen- tal variants not worthy of nomenclatural status. Anomobryum Schimp. Bryaceae 1. Anomobryum filiforme (Dicks.) Solms. In open woods; on sandstone boulder. Rare. Known only from Woodson County (Churchill, 19796). ‘This minute species should be sought in other open Quercus wooded sandstone sites. Anomodon Hook. @ Tayl. Thuidiaceae i leeat tiprwith hyaline point.atipacumminates {e120 2) a5 se 3. A rostratus flteattipalackine a hyaline.pointstip.ObtuSe:Omacute 4 2 tee eee Ds Zeleatstiprouncdedytoro biUsewen eee etens Cac eee Bae as ce 2. A. minor Jeera laolngaCUle™ a5 Gk auat Catena units eels 6 oo eh Caos) Cth eos bee 1. A. attenuatus 1. Anomodon attenuatus (Hedw.) Hueb. In woods; on moist to dry soil, limestone and sandstone rock. C.fr. Common. Eastern half of Kansas. 2. Anomodon minor (Hedw.) Fuernr. In woods; on moist to dry soil, limestone rock, and base of trees (including Carya sp., Quercus macrocarpa and Ulmus americana). C.fr. Very common. Eastern half of Kansas. 3. Anomodon rostratus (Hedw.) Schimp. In woods; on moist to dry soil, on limestone and sandstone rocks, and base of trees. Rare c.fr. Common. Eastern half of Kansas. Archidium Brid. Archidiaceae 1. Archidium ohioense Schimp. ex C. Meull. Exposed in open woods and prairies; on sandy and alluvial soils. C.fr. Rare? Probably more common but overlooked. South-central Kansas. Aschisma Lindb. Pottiaceae 1. Aschisma kansanum Andr. In High Plains grassland; on exposed knolls, confined to edge of clear or white quartz pebbles that are usually 2-5 cm in diameter. C.fr. Rare. Endemic to the Plains, at present known only from south central Kansas but should be sought in Oklahoma and Colorado. See discussion under Kansas types. If any moss should be placed on a ‘‘threat- ened or endangered’’ list for mosses of the United States, this species surely qualifies. Astomum Hampe Pottiaceae ie Setasmuchashorter than’ the, capsule 12.) ai.snc eee 1. A. muhlenbergianum ie seta,as) long or longer than, the capsules» ieeemees ene eee 2. A. phascoides 1. Astomum muhlenbergianum (Sw.) Grout Exposed to shaded, most often in prairies, occasionally in open woods; on soil or sandstone rock. C.fr. Common. Throughout Kansas. Often associated with Werssza controversa with which it is known to hybridize freely; I have seen a few collections from mixed 99 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN populations where some of the capsules were poorly developed suggesting hybrid forms. An excellent summary and study is presented by Anderson and Lemmon (1972) concerning hybridization among these taxa. 2. Astomum phascoides (Hook. ex Drumm.) Grout In open disturbed areas and prairies; on soil. C. fr. Rare? NOTE: One other Astomum has been previously reported: A. ludovicianum Sull. (Smith, 1966). Atrichum P.-Beauv. Polytrichaceae 1. Upper leaf cells averaging 12-17 um in longest dimension; middle stem leaves mostly deameomless: wide dioicousy a6 ceAcuaneas 2 a te 1. A. angustatum 1. Upper leaf cells averaging 17 um or more in longest dimension; stem leaves usually wider than 1 mm; autoicous or dioicous.......... 2. A. undulatum 1. Atrichum angustatum (Brid.) B.S.G. In woods, occasionally in prairies; on moist to dry soil, sandy soil, and thin soil over sandstone rock. C.fr. Common. Eastern half of Kansas. 2. Atrichum undulatum (Hedw.) P.-Beauv. In Kansas there are two recognized species of this complex: A. altecristatum (Ren. G Card.) Smyth & Smyth. Lamellae averaging 4-6 cells high; plants autoicous; stem lacking whitish rhizoids. Reported by Ireland (19692). Uncommon. Probably to be found in the eastern third of Kansas. A. oerstedianum (C. Muell.) Mitt. Lamellae 2-4 cells high (up to 8); plants dioicous; stems clothed with whitish rhizoids. Reported by Churchill (1980) for Douglas County. NOTE: If the appropriate characters are present, one can readily identify these various taxa in the Atrichum undulatum complex. However, if plants are sterile or have a variable number of lamellae cells, then it may be best to simply recognize this as a complex for floristic purposes while realizing that these are very likely both good evolutionary and biological species. Certainly detailed cytological, morphological and chemical work is needed for this interesting genus. Reference: Ireland (1969a). Aulacomnium Schwaegr. Aulacomniaceae 1. Leaf tip rounded to obtuse; margins plane; cells smooth ... 1. A. heterostichum I. Leaf tip acute; margins revolute; cells unipapillose.. .. 2.42... 2. A. palustre 1. Aulacomnium heterostichum (Hedw.) B.S.G. In woods; on moist soil, and thin soil over sandstone rock, often encountered on bank slopes. C.fr. Occasional. Eastern third of Kansas. Some of our plants exhibit the small deciduous leaves at the stem tip which may function as propagula analogous to the propagula commonly associated with A. palustre (see discussion by Lowry and Steere, 1946). 2. Aulacomnium palustre (Hedw.) Schwaegr. In marshes, around seepage areas or border of ponds. Propagula present. Uncommon. Eastern half of Kansas. The report by Smith (1966) of Dichodontium pellucidum is this taxon. Barbula Hedw. Pottiaceae 1. Leaves oblong-lanceolate; obtuse-acute and apiculate ....... 2.2.4.5: Ze 1. Leaves lanceolate to subulate, from a broad to ovate base; acuminate... . . 3. 2 sbropagula, Presemts; xian of nae spe Rane heer eae ee ereraes ee 3. B. indica 2 ~hropacula. absent... Oba s.5 cee a ee ee 4. B. unguiculata A SYNOPSIS OF THE KANSAS MOSSES Ds) . Leaves with a costa precurrent, costa ending below the apex. ... . 2. B. fallax . Leaves with a costa excurrent, extending distinctly bevomelstne wean -ttuke aah tev, aden nets ok soe Oha ge ce ert et nae ae 1. B. acuta . Barbula acuta (Brid.) Brid. In prairies and plains; on sandy soil and calcareous rock. Common. Western half of Kansas. See notes under Didymodon rigidulus. . Barbula fallax Hedw. In prairies and open woods; on clay or sandy soil, often of disturbed sites. Uncommon. Eastern third of Kansas. . Barbula indica (Hook.) Spreng. (B. cancellata and B. cruegeri) In open woods; on sandstone rock. Propagula present. Uncommon. Eastern third of Kansas. . Barbula unguiculata Hedw. In open woods, prairies and disturbed areas; on soil, sand dunes, and limestone rock. C.fr. Common. Throughout Kansas. Bartramia Hedw. Bartramiaceae . Bartramia pomiformis Hedw. In woods; on moist soil and soil over sandstone rock, often on hill slopes. C.fr. Occasional. Eastern third of Kansas. Brachythecium B.S.G. Brachytheciaceae . Stem leaves ca. 2 mm long, distinctly plicate; alar cells subquadrate, not forming a) distinct compact group of smallveells’) 442%. 2 oe 3. B. oxycladon . Stem leaves ca. 1.5 mm long or shorter, plication indistinct; alar cells quadrate, forming a smiallidistmetyeroupsof cellsis a. 2: ee De 2. Leaves subjulaceous; plants dioicous; often found on tree trunks; eastern IAMS aS ta tena ey ee es Ls ae aes 1. B. acuminatum 2. Leaves + spreading, or leaf tips + curved; plants autoicous; on calcareous Solleancd erocksewestern daAnsase 2 6.0 yes sceck ae bs see en ogee 2. B. collinum . Brachythecium acuminatum (Hedw.) Aust. (Chamberlainia) In open to closed woods; occasionally on soil or thin soil over limestone, but most often encountered on base trunk of trees and shrubs (including Quercus muehlenbergii and Ribes missouriense!), also on decaying tree stumps. Occasional. Eastern half of Kansas. . Brachythecium collinum (Schleich. ex C. Muell.) B.S.G. In High Plains; on calcareous soil and rock. Rare? Originally this was determined as Brachythe- cium salebrosum by Smith (1966) based on Smith 1944 (NY) from Cheyenne County. The distribution reported for Nebraska (Churchill, 1979a) would suggest that this species may be found in additional sites in western Kansas. . Brachythecium oxycladon (Brid.) Jaeg. G Sauerb. (Chamberlainia) In open to closed woods, occasional in prairies; on moist to dry soil, on thin soil over limestone and sandstone rock, and on decaying logs. C.fr. Very common. Eastern three-fourths of Kansas. The reports by McGregor (1950) and Smith (1966) of Brachythecium salebrosum (Web. & Mohr) B.S.G. and/or B. calcareum Kindb. are B. oxycladon. Bruchia Schwaegr. Dicranaceae . Spores pitted, up to 27 um wide; seta mostly immersed, shorter than the Cap sulesccine. cer Se eee eta eer ees eee 1. B. brevifolia . Spores spinuose or reticulate, up to 40 um wide; seta often as long or longer thanzthe capsules 5 oe ae acs, taaganee othe ge Sd ceo oe nee en 2 2, Spores spinose, up to 40 unv wide; awn smooth’... 2. B. flexuosa 2. Spores reticulate, up to 45 um wide; awn rough........... 3. B. texana 2a 3 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN . Bruchia brevifolia Sull. In prairies; on sand dunes. C.fr. Rare? Known only from Reno County. . Bruchia flexuosa (Sw. ex Schwaegr.) C. Meull. Old crop fields; on bare exposed soil. C.fr. Rare? Known only from Douglas County. Bruchia texana Aust. Open areas; on dry sandy soil. C.fr. Rare? Known only from Wilson County. NOTE: It is essential that spores be used for identification of the Bruchia species. The distribution maps suggest a rare occurrence for all Bruchia taxa but it is more likely that these minute species, never over a few millimeters tall, are simply overlooked. Reference: Britton, 1913. i: Bryhnia Kaur. Brachytheciaceae Bryhnia graminicolor (Brid.) Grout. In woods; on moist soil bank and soil over sandstone rock. Occasional. Eastern third of Kansas. Bryoandersonia Robins. Brachytheciaceae . Bryoandersonia illecebra (Hedw.) Robins. In woods; soil over sandstone and on shaded bank. This species is frequently found in southern Missouri (Red- fearn, 1972) and eastern Oklahoma (Churchill et al., 1981). Recently reported as a new state record for Kansas (Churchill, 1982), this species should be sought elsewhere in eastern Kansas, particularly the wooded sandstone sites. Bryum Hedw. Bryaceae . Plants julaceous, appearing whitish-silver; lacking a distinct marginal border BI leg I ye sok Re ANT a ol he oe, Sh Se nih fly EL SE aA OR 1. B. argenteum . Plants not julaceous, light to dark green occasionally appearing bleached; Inanatnal border ustallyodistineh a2. cxssenc.c soap veg Sieg tachometer a Ze 2. Plants with abundant filiform brood-bodies in leaf axil; costa precurrent Sete ue: tee Aa. ia Rie sine Tb Lea, spas Miata, yack Gee eM alae aeeae 3. B. capillare 2. Plants lacking filiform brood-bodies; costa precurrent or excurrent..... Be oe CO Stal GistinCthy4eExCUNNe Miles ia. 6 .f ss ct cn nisi. Aso tpl ok ceacettn sects cK ge ee “Costasprecurrenton indistinctly excunnemt «ik ce lecre ea ra en 6. eer MI SeGIOLCOUS AM Sitat Tey i Shy ucent we heel eee ik oe eee A ek eee 4. B. caespiticium He ALAIN SES Va @OLCOUS 5 1 hag ates aPC AUR, Ba aa En as ad ae seme Di. . Cilia lacking, endostome poorly developed, lower half adhering to exostome Ss Fae Yeh Aen Soles AN i ae ee a de OD eR de Yaa tet 1. B. algovicum . Cilia appendiculate, endostome well developed, and not adhering to ex- OSHOTINE yp io 5a Bore Re orto 2 gouil coe Bates) > Ts rs Cae ere eek 6. B. creberrimum 6. Leaf apex obtuse-rounded; marginal border indistinct; distinct serrulation BAG Goris pte cold INS ne, pak dei I. at a Sk eae 5. B. gemmiparum 6. Leaf apex acute-acuminate; marginal border distinct; serrulation confined LOAAPERO © os fn bo PARI ce we Be i cada ich ee nea eee ee qT. . Leaf ca. 1 mm long, base not decurrent (axillary bulbiform gemmae SOMME MICS PTESEME)) Act cia Sad t TAN, Pea Oy ee ee eee ee 3. B. bicolor eat ea, 2-2-9 mm. long, basevdecurrent:(gemmaciabsent)- =. 22 eee Re aera ea eed nee tra ear ay am dae eR ae SA To 8. B. pseudotriquetrum . Bryum algovicum Sendtn. ex C. Muell. (B. angustirete and B. pendulum) In open woods and prairies; on sandstone rock. C.fr. Uncommon. Scattered through- out Kansas. No (Sy A SyNopsiIs OF THE Kansas MOsSEs 2. Bryum argenteum Hedw. In natural and disturbed areas in most habitats, usually uncommon in shaded woods; on soil. C.fr. Common. Throughout Kansas. 3. Bryum bicolor Dicks. In prairies; on soil and thin soil over sandstone. Bulbiform gemmae often present. Uncommon. Probably scattered through- out Kansas, particularly the eastern half. 4. Bryum caespiticum Hedw. In natural and disturbed areas of prairies, open to closed woods, and less common in the plains; on soil, on thin soil over sandstone and limestone. C.fr. Frequent. Scattered throughout Kansas. 5. Bryum capillare Hedw. In prairies; on sandstone rock in ravines. Known only from Ellsworth County based on Smith 1297 (NY!). Brood bodies present. 6. Bryum creberrimum ‘Tayl. (B. cuspidatum) In woods, prairies and plains; on soil and soil over rock. C.fr. Occasional. Scattered throughout Kansas. 7. Bryum gemmiparum De Not. In High Plains; on calcareous soil and rock of seepage sites. Uncommon. Western half of Kansas. 8. Bryum pseudotriquetrum (Hedw.) Gaertn., Meyer & Scherb. Open woods, upland prairie springs and disturbed sites; most often found on thin soil over sandstone rock, occasionally on sandy soil. Occasional. Eastern half of Kansas. NOTE: Bryum is a difficult genus and if sterile, identification is quite impossible for some species—particularly B. algovicium, B. caespitictum and B. creberrimum. For mapping the Kansas Bryum’s, sexual condition and gametophytic features were used in combination when sporophytic capsules were lacking. Specimens lacking capsules or sexual features were not mapped for the taxa mentioned above. Campylium (Sull.) Mitt. Amblystegiaceae 1. Leaf with single costa extending half or somewhat farther to apex... .. . ST ge ee one rR trae OR cg SF eT hea 1. C. chrysophyllum 1. Leaf with very short, usually double costa extending a fourth to fifth way from LY Pn ae ee ge OR ER eRe: OR Dae rer Saas ii 2. C. hispidulum 1. Campylium chrysophyllum (Brid.) J. Lange. In prairies and woods; on moist to dry soil, on loam and sandy soil, thin soil over sandstone rock. Frequent. Eastern half of Kansas. 2. Campylium hispidulum (Brid.) Mitt. In prairies, plains and woods; moist to damp soil, often in calcareous and sandstone areas under rock ledges and crevices, and at base of trees. Frequent. Throughout Kansas. Ceratodon Brid. Ditrichaceae 1. Ceratodon purpureus (Hedw.) Brid. In prairies and open woods, especially disturbed or exposed sites; on sandy to loam soil. C.fr. Occasional. Through- out Kansas. Climacium Web. & Mohr Climaciaceae 1. Climacium americanum Brid. Open to closed woods; marsh areas along tributaries and wooded slopes, most often over sandstone rock. Uncommon. Eastern third of Kansas. Desmatodon Brid. Pottiaceae 1. Leaves with a distinct light border of cells, papillae lacking or nearly so along margin wlealoapex "aACuLer -uc, mye eR eset ee a eee ee ee 3. D. portert 26 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN 1. Leaves without a distinct border of differentiated cells, all cells appearing papillose along margin; leaf apex obtuse-rounded or with a long Papell e™ AVANT Dee cree cocker Rah a fie, AOC Ree ae ge I I are rt arc el 2. 2. Leaves with a hyaline awn, greater than 0.5 mm and usually up to 1 mm oe i eR CME ORS a SPR oo Beale Bk er ea 2. D. plinthobius 2. Leaves lacking a distinct awn, occasionally with a short apiculate or hyaline miucro-tip. lesssthans0;) mmelong. 5 a= sp see ae 2 1. D. obtuszfolius 1. Desmatodon obtusifolius (Schwaegr.) Schimp. In woods and prairies; usually found on sandstone rock, less common on calcareous rock. C.fr. Common. Throughout Kansas. 2. Desmatodon plinthobius Sull. G Lesq. ex Sull. In open woods, less common in prairies; on calcareous and sandstone rocks. C.fr. Common. Throughout Kansas. 3. Desmatodon portert James. In woods; on sandstone. C.fr. Rare? Eastern third of Kansas. Many of the collections made by Smith (1961) and identified under this name are in fact Desmatodon obtuszfolius. NOTE: Conard (1951) noted the occasional occurrence in which Desmatodon obtusifolius has a short awn; a few collections from Kansas also exhibit this feature, but can be separated by the length of the awn and the collenchymatus submedian cells that are lacking in D. plinthobius. Dicranella (C. Muell.) Schimp. Dicranaceae 1. Seta yellow; capsule plicate (when empty); leaves usually falcate-secund; margins serrulate from the tip to beyond the leaf middle; margins plane. . AE RRS, SRR Ree 5 ee ek eae AMO EEE inter OF My ae ep RCE 1. D. heteromalla 1. Seta red-brown; capsule smooth (when empty); leaves erect to slightly spreadine + mostly entire: margins recurved 5 ys coe ale os oe 2. D: varia 1. Dicranella heteromalla (Hedw.) Schimp. In open to closed woods; on sandy soil, soil over sandstone rock, and riparian banks. C.fr. Occasional. Eastern half of Kansas. 2. Dicranella varia (Hedw.) Schimp. In open to closed woods, wooded margins, often of disturbed sites; sandy or clay soils, occasionally found on sandy soil over standstone rock. C.fr. Occasional. Eastern third of Kansas. Dicranum Hedw. Dicranaceae 12) pper leat cells: quadrate tommectameular, 1-2) 13s cee 1. D. condensatum i Wipper leaticells elongate, (2 pOroses Sel. r a. ee ee as wpe os 2. D. scoparium 1. Dicranum condensatum Hedw. (D. sabuletorum) In woods; on sandy soil, and soil over sandstone rock. C.fr. Uncommon. Eastern third of Kansas. Given the recent discovery of this taxon by the author in Douglas County and the reports given by Smith (1961), this should be sought in other slightly xerophytic open wooded sites with sandstone substrate in eastern Kansas. 2. Dicranum scoparium Hedw. In woods; often in cushions over sandstone rock or on sandy soil banks. C.fr. Frequent. Eastern third of Kansas. Didymodon Hedw. Pottiaceae 1. Leaves strongly decurrent; axillary gemmae absent; mesic or aquatic sites. SED Sc eae Aw PO IER eee eee a ete EEE ee 2. D. tophaceus A SyNoPSIS OF THE Kansas MOsseEs No “I . Leaves not noticeably decurrent; axillary gemmae present; xeric sites. . SRR eee ae a A teres, OK LP ASAIN EY. cee TAO Mee tt 1. D. rigidulus . Didymodon rigidulus Hedw. In plains and prairies; on gravel or calcareous soil and on calcareous or sandstone rock. Axiallary gemmae present. Common. Western half of Kansas. Both Didymodon rigidulus and Barbula acuta are equally encountered in western Kansas, and may be confused. Barbula acuta has leaves that are squarrose-spreading when wet, the costa typically extends well beyond the lamina, and lacks gemmae; Didymodon rigidulus has erect-patent leaves when wet, precurrent costa, and axillary gemmae. Many of the collections made and reported on by Smith (1966) of Didymodon rigidulus are in fact Barbula acuta (NY!). . Didymodon tophaceus (Brid.) Lisa. In spring and marsh sites; often on calcareous substrates of soil and rock. Occasional. Scattered throughout Kansas. The distinctive long decurrent leaves is a diagnostic feature found in no other Pottiaceae moss in Kansas. Diphyscium Mohr Buxbaumiaceae . Diphyscium foliosum (Hedw.) Mohr. In open to closed woods; on sandstone rock. C.fr. Rare. Known only from Douglas and Woodson counties. Ditrichum Hampe Ditrichaceae . Ditrichum pallidum (Hedw.) Hampe. In open to closed woods, disturbed prairies and roadsides; on soil, often on sandy soil or thin soil over sandstone rock. C.fr. Common. Eastern half of Kansas. The long yellow seta and subulate leaves are diagnostic. Drepanocladus (C. Muell.) Roth. Amblystegiaceae . Drepanocladus aduncus (Hedw.) Warnst. In exposed wet areas, ponds, stream banks, marshes etc. Uncommon. North central Kansas. Drummondia Hook. Orthotrichaceae . Drummondia prorepens (Hedw.) Britt. In woods; usually on upper trunk of trees (including Quercus). C.fr. Occasional. Southeastern Kansas. Entodon C. Muell. Entodontaceae . Leaves julaceous; quadrate alar cells more than 10 along the margin; PeristommessmmOOth Teo.) cy Ag.c 8s se A ee ee a 3. EF. seductrix . Leaves complanate; quadrate alar cells 10 or fewer along the margin; PELISCOMECVANIOUS 2% sa) 4 los. ee eee es 6, oe ee eee De 2. Leaf length 1(1.5) mm; width of leafy branch ca. 1(1.5) mm; peristome Papulosewec my ate © wie eA er 8 an Cee ee, eee 1. E. compressus 2. Leaf length (1.5)2 mm; width of leafy branch ca. 2 mm; peristome smooth ss SES vol wap eas cok fas OS eR Loa Oa, os ee ONE ete retary: 2. E. cladorrhizans . Entodon compressus C. Muell. (E. challengeri) In woods; on base trunk of trees (including Celtis occidentalis, Quercus macrocarpa and Q. muehlenbergii), occasion- ally decaying logs, rarely on soil. C.fr. Frequent. Eastern half of Kansas. . Entodon cladorrhizans (Hedw.) C. Muell. In woods; on trees, decaying logs, and over limestone. Uncommon. Eastern third of Kansas. . Entodon seductrix (Hedw.) C. Muell. In open to closed woods, rarely in sandstone prairies; on moist to dry soil, sandstone and limestone, decaying THE UNIVERSITY OF KANSAS SCIENCE BULLETIN logs, base trunk of trees (including Fraxinus sp. and Quercus muehlenbergit). C.fr. Common. Eastern half of Kansas. Ephemerum Hampe Ephemeraceae . Cells in the middle third of leaf smooth, in diagonal rows from the costa to EVAR CUNT Ie ea atte Ue eee et oe et See eRe Rete Mee hae irc Mea a 1. E. cohaerens . Cells of the leaf papillose above or if smooth then cells not in diagonal rows 2. Upper leaves spinose, teeth mostly recurved at an angle of 45° or more; leaves usually narrowly linear-lanceolate; median cells 4-8:1........ SB a Ma iNet Lea sates AS oe we eeey. oe Oey a Rte 3. E. spinulosum 2. Upper leaves entire to strongly toothed, teeth at an angle of less than 45°; median? cellsmiessethanvacl ee eats eS a: eee 2. E. crassinervium . Ephemerum cohaerens (Hedw.) Hampe. In woods and prairies; on soil, often on exposed or disturbed soil. C.fr. Occasional. Eastern half of Kansas. . Ephemerum crassinervium (Schwaegr.) Hampe. In prairies; on sandy soil. C.fr. Rare. Known only from Reno County. . Ephemerum spinulosum Bruch G Schimp. ex Schimp. In prairies and sand dune areas; on sandy soil, also soil over limestone rock. C.fr. Uncommon. Central; expected in eastern Kansas. Reference: Bryan & Anderson (1957). Eucladium B.S.G. Pottiaceae . Eucladium verticillatum (Brid.) B.S.G. Shaded in woods; on wet to moist limestone rock; often associated with Fisszdens. Uncommon. Southeast Kan- sas. Eurhynchium B.S.G. Brachytheciaceae branchtleaves- ovate, acute;-seta TOUCH 2 2% G4 See hee wae See 1. Be hrans . Branch leaves narrowly ovate to lanceolate, acutely blunt to obtuse; seta SINIOO CE pur eee Fo hassc eRe S Secuat et ata Mets ck) ear at Cae ke 2. E. pulchellum . Eurhynchium hians (Hedw.) Sande Lac. In woods or margin of woods, often in disturbed sites; on moist soil. C.fr. Frequent. Eastern half of Kansas. . Eurhynchium pulchellum (Hedw.) Jenn. In open woods; on moist soil, and thin soil over sandstone rock. Occasional. Eastern half of Kansas. Fabronia Raddi Fabroniaceae . Fabronia ciliaris (Brid.) Brid. (F. imperfecta) In woods; lower and more often upper tree trunks (including Celtzs occidentalis, Juglans nigra, Quercus spp., and Ulmus sp.). C.fr. Frequent. Eastern half of Kansas. Past reports (McGregor, 1950) of F. wright Sull. are F. cilzarts. Fissidens Hedw. Fissidentaceae . Plants aquatic; leaves linear-lanceolate; vaginant lamina extending only a fourth: tosthird the way-up. the daminay. 2 aces renee Teer 3. F. fontanus . Plants not aquatic, but often on moist or wet substrates; leaves various shapes but not linear; vaginant lamina extending to about half the way up the lamina ict 5! 4, ee act a Some eh Lee aA ee ee ee eae oe ee Ze 2: dneayes distinetly obtuse-rounided* 273 seat eee eee 5. F. obtusifolius 2 PICAVES ACULEG c's Sida vee loge ube aU anaes es aren ee er 3: A SYNOPSIS OF THE KANSAS MossEs 99 . Leaves with a distinct border of linear cells extending beyond the middle to Meatauherapexi assets eos hel ah er er ee a ae mS 1. F. bryovdes . Leaves lacking a border made up of linear cells extending beyond the middle 4. Leaves with a border of linear cells at the base, plants less than 5 mm tall Ries Ree ered airman Me ahs UN es Cea ero eS germ ra a oof a 4. F. garberi 4. Leaves lacking border of linear cells; plants mostly greater than 5 mm tall et Re OO hte aan tre hey ata RNG tks eA ee MLE lt Nee ha eae oO. eaves zeoular, finely;crenulate-serrulate’ 2... =. =12, 5/084 2 ao 7. F. taxifolius , lbeaves arresularly dentate; sernulate-or:crenulate-serrate : 9: 4)5 05. eects 6. 6. Costa ending several cells below the apex, costa cells mammillose ... . desea ih att Tn AR. SET ba te ICRA, Dae. eeacee ene aan A 6. F. subbasilarts 6. Costa percurrent (extending into the apex), costa cells not mammillose . ENOE ES eOohs oy REL os EL PRACT a, SMR ORL hoa: god eae ah eee 2. F. cristatus . Fissidens bryowdes Hedw. (F. minutulus, F. sublimbatus and F. viridulus) In woods and plains; on calcareous and sandstone rocks, usually in crevices or over- hanging ledges, occasionally found on sandy or loam soil especially on stream banks. C.fr. Frequent. Scattered throughout Kansas. . Fissidens cristatus Wils. ex Mitt. Open to closed woods and in prairies; on sandstone rock or sandy soil over sandstone, occasionally on loam soil banks. C.fr. Occasional. Eastern half of Kansas. . Fisstdens fontanus (B.-Pyl.) Steud. (F. debilis and F. julianus) In woods or sandstone prairies; aquatic, attached to rocks or occasionally to twigs and branches of trees in streams. Uncommon. Eastern half of Kansas. . Fissidens garberr Lesq. G James. In woods; on moist sandstone rock. Rare. Known only from Woodson County. . Fissidens obtusifolius Wils. Open woods and prairies; on moist calcareous or sandstone rocks, usually found in crevices or under over-hanging ledges. C.fr. Occasional. Scattered throughout Kansas. Two forms of this taxon occur in Kansas: var. obtusifolius has quadrate cells at base of leaf, whereas var. kansanus Ren. & Card. has linear cells along the leaf base margin. This latter taxon is confined to central and southwestern North America whereas the typical form is found in eastern and western North America. The report by Smith (1966) of F. osmundioides Hedw. belongs to F. obtusifolius. . Fisstdens subbasilaris Hedw. In woods; on rock, soil or base trunk of trees. Uncommon. Eastern third of Kansas. The report by McGregor (1950) of F. bushi (Card. & Ther.) Card. & Ther. is F. subbasilaris. . Fissidens taxifolius Hedw. In woods; on soil banks, often on clay soil. Occasional. Eastern third of Kansas. Fontinalis Hedw. Fontinalaceae . Fontinalis missourica Card. Submerged in streams; attached to rocks. Rare. The last known collection of this taxon was made by Grace Meeker in 1900 from Middle Creek in Franklin County. Several other collections from that county are dated 1892 and 1894. The type locality for this taxon is in nearby Benton County, Missouri. See Welch (1960) for further discussion. Funaria Hedw. Funariaceae . Capsule only slightly bent, smooth, constricted just below mouth; annulus 30 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN AIDSE Rae eT ea tke ek A ede el ie las ey ee ee oe p 1. F. americana . Capsule bent or inclined, usually grooved or ribbed, lacking a constriction below mouth annulus presents... ens es ee ye kee seh ee ee Ze 2. Leaves acuminate with the costa excurrent; inner peristome blunt, ca. half the length of outer peristome; spores rough, 20-50 um wide. . 2. F. flavicans 2. Leaves acute (or shortly acuminate) with the costa percurrent; inner peristome nearly equal to the outer; spores smooth, 14-17 um wide. . . SORA eters ey ey ee ne MO Care gee a a, ea 3. F. hygrometrica . Funaria ameriwana Lindb. In prairies; on sandstone rock, or thin soil over sandstone. C.fr. Uncommon. Central Kansas. Smith (1980) has charac- terized this species as a strict calciphile with a Pleistocene or post-Pleistocene distribution. The distribution map is based in part on Smith (1980). . Funaria flavicans Michx. In open woods and prairies, often in disturbed sites; on soil, often sandy. C.fr. Common. Eastern half of Kansas. . Funarta hygrometrica Hedw. In woods and prairies, often in disturbed and burnt areas, e.g. roadsides etc.; on soil, occasionally rocky or sandy substrates. C.fr. Frequent. Throughout Kansas. Grimmia Hedw. Grimmiaceae . Leaves flat to slightly concave in section, not keeled; margins plane, not HECUNC OM Me AU RS FRc se wi nag 408 GR d wae ew ginger Duer eee oes Ze . Leaves keeled in cross-section (in upper half); margins + recurved...... oy 2. Leaves unistratose in section; hyaline awn denticulate; seta sigmoid; capsule gibbous, immersed among leaves .:.2....2. 2. 2. G. plagiopodia 2. Leaves bistratose; hyaline awn spinulose; seta straight; capsule symmetri- caly exserted:slichtly beyond the-leaves . 2. a 1%. 222%, 202. 1. G. laevigata . Upper leaves with a long hyaline awn; central strand of stem well defined; Capsule Olen Presents go tes 2 toh eh we Rae ke 2 Oe ee Ne oe 3. G. pulvinata . Upper leaves with a muticous tip or a very short spinulose awn; central strand of stem poorly defined; capsule unknown .=...:.. =<: .+¢ 4%. 4. G. teretinervis . Grimmia laevigata (Brid.) Brid. In open to closed woods, occasionally in prairie transition from woods, or in prairies; most often on sandstone, found once on limestone. Rare c.fr. Frequent. Eastern two-thirds of Kansas. . Grimmia plagiopodia Hedw. In prairies; exposed on sandstone. C.fr. Rare? Known only from Ellsworth County. This taxon should be sought in other sandstone or calcareous sites in western Kansas. . Grimmia pulvinata (Hedw.) Sm. In open woods and prairies; on sandstone. C.fr. Occasional. The plants reported by McGregor and Hartman (1956) of G. olneyi Sull. is this taxon. Central and southeast Kansas. . Grimmuia teretinervis Limpr. In plains; on exposed calcareous rocks. Uncom- mon. West central Kansas. Haplocladium (C. Muell.) C. Muell. Thuidiaceae . Stem leaves broadly ovate to rounded, tip short acuminate; branch leaves acute rounded; leaves not plicate (or if so, indistinct); upper margin + Gentatetoventiren re eee ek en ae eee ee 2. H. virginianum . Stem leaves lanceolate, broad at base, tip + long acuminate; branch leaves also acuminate; leaves usually plicate; upper margins sinuate-serrulate. . . 1. H. microphyllum A SYNOPSIS OF THE KANSAS MOSSES Sil . Haplocladium microphyllum (Hedw.) Broth. (Thuzdium) In woods and adjoining prairies; on soil, often sandy, trunk of trees, and decaying logs. Occasional. Eastern half of Kansas. . Haplocladium virginianum (Brid.) Lindb. (Thuidium) In woods; on soil and bark of trees (including Ulmus sp.). Uncommon. Eastern third of Kansas. Hedwigia P.-Beauv. Hedwigiaceae . Hedwigia ciliata (Hedw.) P.-Beauv. In woods and prairies; on limestone and sandstone. C.fr. Frequent. Eastern half of Kansas. Homomallium (Schimp.) Loeske Hypnaceae . Homomallium adnatum (Hedw.) Broth. In woods and prairies; on limestone and sandstone, rarely on bark of trees. C.fr. Common. Eastern half of Kansas. The report of Homomallium mexicanum Card. by Smith (1966) is Hypnum vauchert Lesq. Hyophila Brid. Pottiaceae . Hyophila involuta (Hook.) Jaeg. G Sauerb. (H. tortula) In woods; on limestone. Brood bodies present. Rare. Probably to be found in the eastern half of Kansas. This species is often associated with Fissedens, especially F. bryoides, and may be overlooked among such collections. Cridland (1960) reported collections from Douglas and Woodson counties. Hypnum Hedw. Hypnaceae . Leaves with many small quadrate alar cells; median cells short (ca. 5-7:1); WIESE RMNICATISAS Gi. 5 Wat oe) acuity ah as eRe ty a ay fe ge eR 3. H. vauchern . Leaves with few to several large inflated alar cells; median leaf cells longer (greater than al0:1);“easterm Kansas: 2 4.35752 55) eee i cence ere Me 2. Stem cortex with an outer hyaline layer of cells, larger and thinner than the NEXCIMMNEL: SEMLES 4. ag tdec, da We eee ed ee ae Ee ee 2. H. lindbergi 2. Stem cortical cells small, thick and usually golden or dark yellow in color, MSOs yale a cere cs tale fe een a CACORE Ieee ee 1. H. curvifolium . Hypnum curvifollum Hedw. In woods; on thin soil over sandstone rock. Occasional. Eastern third of Kansas. The report of Brotherella recurvans (Michx.) Fleisch. by McGregor & Hartman (1956) in this taxon. . Hypnum lindbergi Mitt. (H. patientiae) In marshy areas; on wet soil or decaying vegetation. Uncommon. Eastern third of Kansas. . Hypnum vauchert Lesq. In plains, usually associated with rock outcrops; on soil or rock, mostly calcareous substrate. Occasional. Western half of Kansas. The report of H. cupressiforme Hedw. by Smith (1966) belongs to this taxon. With the recent work of Ando (1976) it now is clear that Hypnum vaucheri is the common Hypnum of the arid plains, whereas H. cupressiforme is known only from the mesic Black Hills of South Dakota within the Plains states (Churchill, unpubl. data). Jaffueliobryum Thér. Grimmiaceae . Costa inconspicuous; leaves broadly ovate with upper lamina rounded; lower leaves on.plantjappressed!... <¢4..5. 25.560 2s Sho ee ee 2. J. wright . Costa conspicuous, raised above the lamina surface; leaves ovate-lanceolate, upper lamina acute-acuminate; lower leaves spreading. ......... Lee/earait 32 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN 1. Jaffueliobryum raui (Aust.) Ther. (Grimmia, Coscinodon) In plains and prairies; on calcareous and sandstone rock. C.fr. Frequent. Western two-thirds of Kansas. 2. Jaffueliobryum wrightii (Sull. in Gray) Thér. (Grimmia, Coscinodon) Location and habitat the same as for Jaffueliobryum raui. C.fr. Frequent. NOTE: These two species occasionally occur together, but can be easily separated in the field by the distinctly julaceous branches of /. wrightit as opposed to the patent or spreading leafy branches of /. rauwz. Reference: Churchill (unpubl. data). Leptobryum (B.S.G.) Wils. Bryaceae 1. Leptobryum pyriforme (Hedw.) Wils. In woods; on moist soil, sandstone and at base of trees (Salix). C.fr. Occasional. Eastern half of Kansas. Leskea Hedw. Leskeaceae 1. Leskea gracilescens Hedw. In open to closed woods; on soil, limestone and sandstone, decaying logs, but most often found on trees (including Acer negundo, Carya sp., Celtrs occidentalis, Fraxinus pennsylvanica, Gleditsia triacanthos, Juglans nigra, Juniperus virginiana, Maclura pomifera, Quercus macrocarpa, Q. marilandica, Q. muehlenbergit, Q. stellata, Populus deltoides, Ulmus americana, and U. rubra). C.fr. Common. Eastern two-thirds of Kansas. NOTE: The variability of this taxon often approaches that of two other taxa: L. obscura Hedw. and L. polycarpa Hedw. Until further work is carried out with this group, in particular a biometric analysis, it seems best to regard this as a complex for the Great Plains region. Nearly all our material from this region is L. gracilescens in the strict sense. In Kansas, only L. obscura has been reported in addition to L. gractlescens (McGregor, 1950). Those wishing to separate their material based on traditional characters may use the following key: We heat length: ereaterstham twice the width 5%... 22 .7.Ps 2.2 L. polycarp Pe Eeatleneth less, than twice the width: 221-2 95 6 cea ee eee Ze 2. beavessasymimetrical, not: plicate*s,.2 5-2: 5252 = tere: L. obscura 2. Leaves symmetrical, biplicate, margins with one or both margins RECUIVEG! S552 fon Crp et the a: Sen tO aR, Oe, pees oC Ais L. gracilescens Leucobryum Hampe Leucobryaceae 1. Leucobryum glaucum (Hedw.) Angstr. ex Fr. Most often found in woods, rarely in moist uplands of open woods and prairies; in cushions on soil over sandstone. Occasional. Eastern third of Kansas. NOTE: Leucobryum albidium (Brid. ex P.-Beauv.) Lindb. was reported by McGregor (1950) but in a later study by Smith (1961) this taxon was not included. A specimen of this taxon could not be located at NY, and is excluded from the Kansas moss flora. Leucodon Schwaegr. Leucodontaceae 1. Leucodon julaceus (Hedw.) Sull. In open to closed woods; on various trees (including Celtis sp., Juglans nigra, Quercus marilandica, and Q. stellata). C.tr. Frequent. Eastern third of Kansas. NOTE: Leucodon brachypus Brid. was listed by Grout (1928-1940; vol. III, part 4: 217-218) for Kansas. There is a specimen at NY!, but I believe that the label information may be incorrect, and therefore the species is tentatively excluded from the flora of Kansas. a Oo SX) A SYNOPSIS OF THE Kansas MOSSES Lindbergia Kindb. Leskeaceae . Lindbergia brachyptera (Mitt.) Kindb. In open to closed woods; on trees (including Acer saccharum, Quercus muehlenbergii, Quercus sp., and Ulmus sp.), occasionally on bark of fallen trees or decaying logs. This moss is usually found on upper tree trunks in thin, loosely scattered strands. Rare c.fr. Common. Eastern half of Kansas. Orthotrichum Hedw. Orthotrichaceae leaves With-a distinct ayalime-awmmn.s «eG. e426) ee 1. O. diaphnum . Leaves lacking a hyaline awn, tip occasionally hyaline-apiculate ........ Zs 2. Capsule smooth, + lacking ribs; stomata superficial; calyptra naked, STOOP Me ry. ae tae ee tate Settee RCA hey IG Te Ae iNT ee, 3. O. pusillum 2. Capsule ribbed; stomata immersed; calyptra with scattered hairs. .... . Be . Upper leaf cells averaging 7-12 um wide; leaves bistratose on margin in upper halt; plantsstound: ongrocks...¢ sus yee ieah eas O58. va eee 4. O. strangulatum . Upper leaf cells averaging 14-20 um wide; leaves unistratose; plants found on GRE CAUIUUIT Sear thot rics vate sum te One qe ene eer re On et gigy oan 2. O. pumilum . Orthotrichum diaphanum Brid. In open woods; on trees (including Juniperus virginiana and Ulmus sp.). C.fr. Occasional. Central Kansas. . Orthotrichum pumilum Sw. In woods; on trees (including Juglans nigra, Maclura pomifera, Quercus spp. and Ulmus spp.), occasionally found on decaying logs. C.fr. Common. Eastern three-fourths of Kansas. . Orthotrichum pusillum Mitt. In woods; on trees (including Celtis tenuifolia, Cercis canadensis, Quercus marilandica, and Q. velutina). C.fr. Common. Eastern half of Kansas. Orthotrichum strangulatum P.-Beauv. Mostly in open woods; on limestone rock. C.fr. Occasional. Eastern third of Kansas. NOTE: Orthotrichum sordidum Sull. G. Lesq. in Aust. previously reported by McGregor (1950) for Kansas in O. pusillum according to Vitt (1973). A duplicate collection of that cited by McGregor (1950) (Holland 93 in KSC!) contains both O. pumilum and O. pusillum. Reference: Vitt (1973). Phascum Hedw. Pottiaceae . Phascum cuspidatum Hedw. In open woods, prairies and plains; on soil, often sandy. C.fr. Frequent. Throughout Kansas. Philonotis Brid. Bartramiaceae . Leaf cells papillose at the lower end; margin with partly double serration, cell Inargins concave with extended, pomted ends 2.4... . ns 1. P. fontana . Leaf cells papillose at upper ends; cell margins pointed only on upper ends, WOLTCONCAVE Tyce on as ss ead a, Soe eee cep ts Cae ae ee 2e 2. Leaves long triangular-lanceolate, straight or falcate and flat, cells long and narrow (9-15:1), papillae pointed at the extreme upper cell ends; FNOMOICOUS «a cera teehee, coe erat het ge ee cee ee ee 2. P. longiseta 2. Leaves triangular to slightly ovate-lanceolate, carinate; cells rectangular (less than 9:1); papillae rounded near the upper cell ends; dioicous . . . 3. P. marchica . Philonotis fontana (Hedw.) Brid. In springs and marshes; on wet soil over rock. Known only from Leavenworth County. 34 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN 2. Philonotis longiseta (Michx.) Britt. In open to closed woods; on moist to wet soil over sandstone. C.fr. Occasional. Scattered throughout Kansas. 3. Philonotis marchica (Hedw.) Brid. In woods; on moist to wet soil on sandstone. C.fr. Occasional. Eastern half of Kansas. NOTE: The report by Grout (1928-1940) of Philonotis gracillima Angstr. probably refers to Holzinger’s Musci Acrocarpii Boreali-Americani No. 71, a collection made by Meeker at Princeton, Kansas; this collection is P. marchica. The taxon listed by Grout as P. glaucescens (Hornsch.) Broth. is not supported by any known specimen, and is therefore excluded from the flora. Reference: Zales (1976) and correspond- ence with present author (1979). Physcomitrium (Brid.) Fuernr. Funariaceae 1. Leaves smooth or slightly dentate in upper half; annulus of 3 layers, revoluble Orapersistent,.sporeslessuthan 30h mm diameter: 2.22 20-5 ao: 2. P. hookert 1. Leaves distinctly dentate or serrate in upper half; annulus 1-2 layers, usually persistent; spores of various sizes, mostly greater than 30 um in diameter .. . 2. Upper leaves 2 mm or less long; capsule on drying becoming shallow with avery wide, flared.imouth; spores 24-37 wm inidiameter... s.04) 06 De ReneS tenia eeits Te Aneta ce. Pater ts racine Fe, eee, sche Re 1. P. collenchymatum 2. Upper leaves usually greater than 2.5 mm long; capsule mouth turbinate when dry, not flared; spores ca. 25-50 um in diameter..... 3. P. pyriforme 1. Physcomitrium collenchymatum Gier. On exposed mud bank along creek. First reported for Kansas by Cridland (1960), this particular taxon was discussed in detail by Crum & Anderson (1964) who included a citation of a specimen at CANM from Douglas County, the only known Kansas locality. Specimen not examined. Distribution map based on Crum & Anderson (1964). 2. Physcomitrium hooker’ Hampe. In plains and prairies; often in somewhat exposed marshy places, on soil. C.fr. Uncommon. Western two-thirds of Kansas. 3. Physcomitrium pyriforme (Hedw.) Hampe. (P. acuminatum, P. kellermanu, and P. turbinatum) In woods and prairies, often found in disturbed sites including recently burnt areas, roadside ditches and foot paths; on moist soil, occasion- ally on sandy soil, or over sandstone rock, usually exposed. C.fr. Common. Eastern two-thirds of state. Plagiomnium 'T. Kop. Mniaceae 1. Leaf toothed. on the margin from near the base to the tip... .- 1, P.coliare i. Leaf toothed on the margin from about the middle tothe tip ... ~-.5 0... 1. Plagiomnium ciliare (C. Muell.) T. Kop. (Mnium affine of past reports, and Mnium ciliare) In woods; on moist soil and thin soil over sandstone rock. C. fr. Occasional. Eastern third of Kansas. 2. Plagiomnium cuspidatum (Hedw.) T. Kop. (Mnium) In open to closed woods, rarely in prairies; on moist soil, thin soil over sandstone and limestone rock, on base trunk of trees and over decaying logs. C.fr. Common. Eastern half of Kansas. NOTE: These two species often occur together, particularly in rich deciduous woods. In addition to the characters listed above, P. cuspidatum has obovate leaves A SYNOPSIS OF THE Kansas MOsSEs Bo whereas P. ciliare has oblong to elliptic leaves. The previous reports of Mnium affine (McGregor, 1950, and Magill & Kirkpatrick 1976) are all Plagieomnium ciliare (MO! SINY1), Platygyrium B.S.G. Hypnaceae 1. Platygyrium repens (Brid.) B.S.G. In woods; most often found on decaying logs, occasionally on trees (including Quercus sp.). C.fr. and axillary brood- branchlets present. Common. Eastern half of Kansas. Pleurrdium Brid. Ditrichaceae 1. Pleurtdium subulatum (Hedw.) Rabenh. In prairies and margin of woods; exposed on bare soil and on soil among grasses. C.fr. Uncommon? Eastern half of Kansas. NOTE: Pleuridium sullivantt Aust. has been reported from Kansas (McGregor, 1950). This taxon has leaves erect-appressed, imbricate, ovate or mucronate; P. subulatum has leaves spreading or secund, lanceolate-subulate (Britton, 1913). No specimen of Pleuridium sullivanti: could be located at NY and is therefore excluded from the flora until verified. Pogonatum P.-Beauv. Polytrichaceae 1. Leaves acute to acuminate; 10-15 lamellae on adaxil surface of leaf; margins SenrateIMatlpper taal: olle ale ea ren cone icc ay ener ee csi 2. P. pensilvanicum 1. Leaves + obtuse; 20-70 lamellae on adaxil surface of leaf; margins entire in Mpperaalt: Ofleate #0 shen aie ek Ae Cen eee ee 1. P. brachyphyllum 1. Pogonatum brachyphyllum (Michx.) P.-Beauv. In open woods and prairies; on wet to moist soil over sandstone rock, often found at spring sites. C.fr. Occasional. Eastern half of Kansas. These gregarious plants occur in very thick protonematal sods, a feature that no other Kansas moss shares. 2. Pogonatum pensilvanicum (Hedw.) P.-Beauv. In woods; on soil over sandstone. C.fr. Uncommon. Eastern third of state. The report by McGregor & Hartman (1956) of this taxon from Chautauqua County belongs to P. brachyphyllum (NY!). Pohlia Hedw. Bryaceae 1. Median leaf cells 3-5:1; leaves ovate-lanceolate; plant leaves distant, appear- ing whitish-green with a-distimct*red stem. 222 22. >=): 2. P. wahlenbergi 1. Median leaf cells greater than 7:1; leaves lanceolate-acuminate; plant leaves Ambricate sshiny-colden-ereenis 43°5 Aoe1 an) A. ele ee 1. P. nutans 1. Pohlia nutans (Hedw.) Lindb. In open to closed woods; on soil over sandstone, often among cushions of Leucobryum, Dicranum, and Polytrichum. C.fr. Fre- quent. Eastern half of Kansas. 2. Pohlia wahlenbergii (Web. G Mohr) Andr. In marshes, seepages and spring sites of open woods, prairies, and plains; on decaying marsh vegetation, exposed mud banks, and on thin soil over sandstone rock. Occasional. Scattered throughout Kansas. The leaf shape of Pohlia wahlenbergiit resembles Bryum gemmiparum De Not., but the latter like Pohlia nutans, has leaves that are imbricated. Polytrichastrum G. L. Sm. Polytrichaceae 1. Polytrichastrum ohionense (Ren. G Card.) G. L. Sm. (Polytrichum) In woods; on THE UNIVERSITY OF KANSAS SCIENCE BULLETIN sandy soil or thin soil over sandstone. Uncommon. Eastern third of Kansas. The square marginal cells of the lamellae differentiate this taxon from members of Polytrichum. Polytrichum Hedw. Polytrichaceae > Leaves withea distinct» hyalime: awn. 2s. 2-4 555 32) shee ons 3. P. piliferum sieavesswithout 2a: distinct lyalimes await ipso eee tn eek. bee cee roe 2 2. Leaves with marginal teeth; margins not folding over and reaching the middle of the leaf; terminal lamellae cells notched........ 1. P. commune 2. Leaves without teeth; margins folded over and reaching the middle of the leat--termuinal lamellacccellls Conic 1. fa 2s 6 es ss 2. P. juniperinum . Polytrichum commune Hedw. In woods; in loose to compact cushions over moist sandstone rock. Occasional. Eastern third of Kansas. . Polytrichum juniperinum Hedw. In woods and occasionally found in upland prairies; usually over sandstone, rarely over limestone, often forming cushions in association with P. commune. C.fr. Occasional. Eastern half of Kansas. . Polytrichum piliferum Hedw. In prairie; on sandy soil. Known from a single collection from Ellsworth County. Pseudoleskeella Kindb. Leskeaceae . Pseudoleskeella tectorum (Funck ex Brid.) Kindb. ex Broth. On calcareous soil in High Plains. Known only from Cheyenne County in extreme northwest Kansas. The report by Smith (1966) of Amblystegiella subtilis (Hedw.) Loeske 1s this taxon (Smith 1981, 1982, NY!). The very small leaves and variable costa length or absence of costa are diagnostic. Rare. Pterygoneurum Jur. Pottiaceae . Sporophytes exserted beyond the leaves; seta longer than the capsule; leaves apparent and not obscured by the’ hyaline awns % 92%. 722922 1. P. ovatum . Sporophytes immersed to occasionally emergent among the leaves; seta shorter or equal in length to the capsule; leaves obscured by the long hyaline AWA AAS oe Sake re Seer Ne Wal Lea, A eae eras Teer aes 2. P. subsessile . Pterygoneurum ovatum (Hedw.) Dix. In plains; on exposed soil in short grassland. C.fr. Rare. Known only from southwest Kansas in Stanton County. . Pterygoneurum subsessile (Brid.) Jur. In plains and prairies; on calcareous soil among grasses, occasionally found on disturbed soil. C.fr. Frequent. Western half of Kansas. Ptychomitrium Fuernr. Grimmiaceae . Ptychomitrium incurvum (Schwaegr.) Sull. In prairies and open Quercus upland woods; on sandstone. C.fr. Occasional. Southeast and central Kansas. This species is somewhat similar in aspect to Weissza and Astomum, but the lobed mitrate calyptra and dark green to black appearance of the gametophyte that lack a mucronate leaf tip help to distinguish this species from the Pottiaceae taxa. Pylaisiella Kindb. Hypnaceae . Pylaisiella selwynii (Kindb.) Crum, Steere @ Anderson. In woods; on trunks of re -—s A SYNOPSIS OF THE KAnsAs MOossEs trees (including Celtis occidentalis, Quercus stellata, Populus deltoides, and Ulmus sp.), rarely found on bark of fallen trees. C.fr. Occasional. Eastern half of Kansas. Pyramidula Brid. Funariaceae . Pyramidula tetragona (Brid.) Brid. Margin of woods, usually in disturbed sites; on exposed soil, and thin soil over sandstone. C.fr. Uncommon. Eastern third of Kansas. Rhodobryum (Schimp.) Hampe Bryaceae . Rhodobryum ontariense (Kindb.) Kindb. (R. roseum of past authors) In woods; on soil, usually over sandstone rock. Uncommon. Northeast Kansas. Rhynchostegiella (B.S.G.) Limpr. Brachytheciaceae . Rhynchostegiella compacta (C. Muell.) Loeske. (Amblystegium americanum) In plains; exposed in marshy springs. Brood bodies present. Uncommon. Western half of Kansas. The report by Smith (1966) of Brachythecium velutinum can be referred here (NY!). Rhynchostegium B.S.G. Brachytheciaceae . Rhynchostegium serrulatum (Hedw.) Jaeg. G Sauerb. (Eurhynchium) In open to closed woods; most often on moist soil, occasionally on base trunk of trees (including Quercus sp.), or on decaying logs. C.fr. Frequent. Eastern half of Kansas. The diagnostic features of serrulate margins, twisted leaf tip, and smooth seta aid to distinguish this taxon from Brachythecium. The complanate nature of leaves is also found in Taxiphyllum, but Taxiphyllum lacks the single long leaf costa found in RAynchostegium. Schistidium Brid. Grimmiaceae . Leaves. with, a. distinct hyaline tip; spores about 10 um.or less “3 2 a2 ae | Sao Ra aged a mee ee tee eer ane Caan Seen cotinine os 2 2. S. apocarpum . Leaves lacking a hyaline tip (ocasionally the perichaetial leaves have a very short hyaline tip); spores usually greater than10 um......... 1. S. revulare . Schistedium rivulare (Brid.) Podp. (Grimmia alpicola and Schistidium alpicola, of recent authors) In open to closed woods, prairies and along transitional tributaries; usually on limestone rock, but also on sandstone. C.fr. Common. Eastern half of Kansas. . Schistidium apocarpum (Hedw.) B. GS. in B.S.G. (Grimmia apocarpa) In woods or prairies; on sandstone or limestone rock. C.fr. Uncommon. Eastern half of Kansas. Many of the reports given by McGregor (1950) of Grimmia apocarpum are S. rivulare. Reference: Bremer, 1980a & 1980b. Sematophyllum Mitt. Sematophyllaceae . Sematophyllum demissum (Wils.) Mitt. (S. carolinianum) In woods; over sloping or vertical sandstone rock. C.fr. Occasional. Eastern third of Kansas. In aspect Sematophyllum demissum may be confused with Homomallium adnatum, but differs markedly by having a long rostrate operculum and golden-yellow leaves that have slightly inflated alar cells. Sphagnum L. Sphagnaceae . Green cells of branch leaves exposed only or more broadly on the inner THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Sunlace ofleaf lik ChOSS-SECHON = 95,24 Gs ope eee 1. S. capillifolium . Green cells of branch leaves exposed only or more broadly on the outer SULA CCNOMMC Al faut hy cache pei Bia ceed wen ce ere meee eee eae Fe 2. S. cuspidatum . Sphagnum capillifollum (Ehrh.) Hedw. (S. capillaceum & S. nemoreum) Marsh and seepage areas in sandstone prairies and upland woods; wet places among aquatic vascular vegetation and occasionally in seepage over sandstone. Rare. Southeast and central Kansas. The dot map reported herein of S. nemoreum should be referred to as S. capillifolium. . Sphagnum cuspidatum Hoftm. (S. trinitense) Along margin of old strip mines; in shallow water and wet shore line, among plants of Scirpus and Juncus. Known only from Cherokee County in southeast Kansas. Both experimental and biometrical studies by Lane (1981) have shown that the taxon S. trinitense C. Muell. is only a form of S. cuspidatum; the expression of serrulated leaf margins denoting S. trinitense appears to be only an environmental modifica- tion. Therefore the report of S. trinztense (Churchill, 19796) is placed under S. cuspidatum. The dot map reported herein of S. trinitense should be referred to S. cuspidatum. Taxiphyllum Fleisch. Hypnaceae . Leaves imbricate, margins plane or slightly recurved at base; quadrate alar Cells-o-Ordlone Marciidal TOW seas ult + Sa ees saat eek as 1. T. deplanatum . Leaves not imbricate (usually distant), margins recurved to at least the middle; -cellsiusually rectangular at basei:%.. : Fhe ow. se 2. T. taxirameum . Taxiphyllum deplanatum (Bruch & Schimp. ex Sull.) Fleisch. (Plagiothectum) In woods; on forest soil, often on hill slopes and occasionally found on sandstone, rarely on decaying wood or base of trees (including /Juglans nigra). Occasional. Eastern third of Kansas. Plants of this species generally form a thick distinctive green shiny mat. . Taxiphyllum taxirameum (Mitt.) Fleisch. (Plagiothecium geophilum) In open to closed woods; on forest soil or more often on thin soil over sandstone rock. Occasional. Eastern half of Kansas. Reference: Ireland (19690). Thelia Sull. Theliaceae , eat cell papillae simple..net branched. © 25 .2)..06 a5) Sefer oe 2. T. hirtella > eat cell: papillae branched 25 ik 2g. Defies 5 ts ate reas ose Ao eae eee 2 2. Stems creeping, radiculose; leaves ciliate-papillose above, long-ciliate below along margin; acute or obtuse, long apiculate or piliform-apiculate; MOst often" on tree trunks 22. feat a sk ae, See ee eee 1. 7. asprella 2. Stems crowded and ascending; not or only slightly radiculose; leaves not or somewhat ciliate-papillose along margin, not long-ciliate below; obtuse, short-apiculate: “most eitenvon solljor rock ia). 2 495 ae 3. T. lescunst . Thelia asprella Sull. In woods; on base trunk of trees (including Quercus muehlenbergu and Ulmus americana), and sometimes on decaying wood as well. C.fr. Occasional. Eastern third of Kansas. . Thelia hirtella (Hedw.) Sull. In woods; on limestone rock. Uncommon. Eastern third of Kansas. . Thelia lescurtt Sull. In open to closed woods, occasionally found in prairies; A SYNOPSIS OF THE KANSAS MOSSES 39 most often on sandstone rock or sandy soil, and occasionally found on limestone. Occasional. Eastern half of Kansas. NOTE: Experimental studies by Gier & Kennedy (1955) suggested that Thelza lescurtt is only an environmental form of 7. asprella. This hypothesis needs further detailed testing. I have followed the taxonomic study by Crum in the present treatment. Evidence from field work in Kansas tends to support the hypothesis that these are two distinct taxa. Reference: Crum (1966). Thuidium B.S.G. Thuidiaceae 1. Paraphyllia with papillae at the upper ends of the cells; costa filling the ELGG UITUNG Utecary Pre teat ee cea utes aks einen Nye te re Stee alias cae ME ea 2. T. recognitum 1. Paraphyllia with papillae mostly in the middle of the cells; costa ending below the apex,andumot filling the acumen <.02 4s eres eee ree ee 1. 7. delicatulum 1. Thuidium delicatulum (Hedw.) B.S.G. In woods; on moist soil, often bank slopes, and soil over sandstone. Uncommon c.fr. Frequent. Eastern third of Kansas. 2. Thuidium recognitum (Hedw.) Lindb. In woods; on limestone, and sandy soil. Uncommon. Southeastern Kansas. Timmia Hedw. Timmiaceae 1. Timmia megapolitana Hedw. In woods; on moist shaded bank slopes. C.fr. Occasional. Northeast Kansas. Tortella (Lindb.) Limpr. Pottiaceae 1. Tortella humilis (Hedw.) Jenn. In woods; partly to fully shaded on moist to dry sandy soil, sandstone rock, and on base trunk of trees. C.fr. Occasional. Southeast Kansas. Tortula Hedw. Pottiaceae 1. Leaves with a short + folded mucro tip; lacking both propagula and a long serrulated ‘hyaline awit q. ae-. aout: ane oes ey ee Ce ee be TE caini iv eaves withva hyaline short:or long awn =. (70) os.2 ane 2h ae 2: 2. Leaves with a short, smooth to rough awn; margins of leaf plane; propagulay present axalsof leaves. |i an ee 2. T. pagorum 2. Leaves with a long, serrated awn; distinctly revolute margins; propagula POs) | tae ar haa Mia DS NO AM Ak de gy ts 3. DT ruralis 1. Tortula caintt Crum @ Anderson. In plains grassland; on calcareous soil and rock. Rare. Known thus far from one locality in Kansas, Gove County. This Kansas record was recently reported as new to the United States by Churchill (1980). Previously this 7ortula species had only been recorded from a few sites in Ontario, Canada (Zander & Eckel, 1980). This interesting Tortula should be sought elsewhere in the Plains region. 2. Tortula pagorum (Milde) De Not. In usually open, often dry woods; on bark of trees (including Juglans nigra, Juniperus virginiana, Quercus marilandica, and Ulmus sp.). Propagula present. Frequent. Eastern two-thirds of Kansas. 3. Tortula ruralis (Hedw.) Gaertn., Meyer @ Schreb. In plains grassland; on gypsum, sandy or calcareous soils, and thin soil over gypsum and sandstone rocks. Occasional. Western half of Kansas. Weissia Hedw. Pottiaceae 1. Costa at leaf base ca. 60 um wide; peristome present, slender or often poorly 40 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN GENClOM EGY a5, Haters hoa tee ee Ne Ne Us ee phe Neo 1. W. controversa 1. Costa at leaf base 70 um or more wide; peristome represented by a mere fracilemnemib rane: * isa.) 1 nets ade: - ae ele te eae ge cle hae 2. W. tortilis 1. Weissia controversa Hedw. (Wersza viridula) In open woods, prairies and plains, often in disturbed sites; exposed to shaded soil, and soil over limestone and sandstone. C.fr. Very common. Throughout Kansas. See comments under Astomum. 2. Werssia tortilis (Schwaegr.) C. Muell. In plains grassland; exposed on soil. C.fr. Occasional. Western Kansas. The report by Smith (1966) of W. glauca Bartr is this: taxon. SUMMARY OF CORRECTIONS FOR THE Kansas Moss FLora Acaulon triquetrum (Spruce) C. Muell. reported by Cridland (1960) has not been examined—until verification, this taxon is deleted from the flora. Aloina brevirostris (Hook. G Grev.) Kindb. reported by Cridland (1960) is most likely A. mngeda (Hedw.) Limpr.; specimen not examined, but see Delgadillo M. (1975) for major distribution of the two taxa, and Churchill (19795 & 1980) for A. rigida records for Kansas. Aphanorrhegma_ serratum (J. Hook. © Wils. ex Drumm.) Sull. reported by McGregor (1950) could not be lo- cated at NY and is deleted until verified. Brachythecium calcareum Kindb. (B. flex- wcaule) reported by McGregor (1950) is B. oxycladon (Brid.) Jaeg. ©& Sauerb. Brachythecium salebrosum (Web. G Mohr) B.S.G. (Chamberlainia) reported by McGregor (1950) and Smith (1966) is B. oxycladon (Brid.) Jaeg. @ Sauerb. except in one case (Smith 1944, NY!) which is B. collinum (Schleich. ex C. Muell.) B.S.G. Brachythecium velutinum (Hedw.) B.S.G. (Chamberlainia) reported by Smith (1966) is Rhynchostegiella compacta (C. Muell.) Loeske. Brotherella recurvans (Michx.) Fleisch. re- ported by McGregor & Hartman (1956) is Hypnum curvifolium Hedw. Cratoneuron filicinum (Hedw.) Spruce re- ported by McGregor (1950) is Am- blystegium ripartum (Hedw.) B.S.G. Dichodontium pellucidum (Hedw. ) Schimp. reported by Smith (1966) is Aulacomnium palustre (Hedw.) Schwaegr. Fabronia imperfecta Sharp reported by Smith (1966) as /. cliars-(Bric.) Brid. Fabronia wrighttt Sull. reported by McGregor (1950) is F. celzarts (Brid.) Brid. Fissidens bushit (Card. G Ther.) Card. © Ther. reported by McGregor (1950) is F. subbasilaris Grout. Fissidens osmundioides Hedw. reported by Smith (1966) is F. obtuszfolius Wils. Grimmia olneyi Sull. reported by McGregor and Hartman (1956) is G. pulvinata (Hedw.) Sm. Gymnostomum recurvirostrum Hedw. re- ported by McGregor (1950) could not be located at NY and is deleted until verified. Homomallium mexicanum Card. reported by Smith (1966) is Hypnum vauchert Lesq. Hypnum cupressiforme Hedw. reported by Smith (1966) is H. vauchert Lesq. Leucobryum albidum (Brid. ex P.-Beauv.) Lindb. reported by McGregor A SYNOPSIS OF THE Kansas MosseEs 4{ (1950) is not represented by speci- men at NY, and is excluded. Smith (1961) likewise did not include this species for the Chautauqua Hills where McGregor’s collection was made. Leucodon brachypus Brid. is excluded based on questionable label data on a specimen deposited with NY. Mnium affine Bland ex Funck reported by McGregor (1950), and by Magill and Kirkpatrick (1976) is Plagzo- mnium ciliare (C. Muell.) T. Kop. Orthotrichum sordidum Sull. G Lesq. ex Aust. reported by McGregor (1950) is O. pusillum Mitt., see Vitt (1973), specimen at NY!; a duplicate speci- men (Holland 95, KSC!) contains O. pumilum Sw., as well as O. puszllum. Philonotis glaucescens (Hornsch.) Broth. reported by Flowers in’ Grout (1928-1940) is excluded. No speci- men could be located for this report from Kansas but the distribution is mainly southeastern United States and tropical regions further south. Zales (pers. comm.) has not found any record for Kansas in his revi- sionary work on this genus. Philonotis gracillima Angstr. reported by Flowers in Grout (1928-1940) is P. marchica (Hedw.) Brid. Physcomitrium acuminatum (Schleich.) B.S.G. reported by McGregor & Hartman (1956) is P._ pyriforme (Hedw.) Hampe. Platydicta confervoides (Brid.) Crum (Am- blystegiella) reported by McGregor (1950) is deleted as no specimen could be located at NY. Platydicta subtile (Hedw.) Crum (Am- blystegiella) reported by Smith (1966) is Pesudoleskeella tectorum (Funck ex Brid.) Kindb. ex Broth. Pleuridium sullivantit Aust. reported by McGregor (1950) is deleted from the flora. Likewise, Smith (1961) ex- cluded this taxon from his study on the Chautauqua Hills. No specimen could be located at NY. Pottia arizonica Wareham reported by Smith (1966) is excluded from the flora as too little material is present in Smith’s collection (NY!) to pro- vide a positive identification. Pogonatum pensilvanicum (Hedw.) P.- Beauv. reported by McGregor & Hartman (1956) is P. brachyphyllum (Michx.) P.-Beauv. Sphagnum trinitense C. Muell. reported by Churchill (19796) is S. cuspidatum Hoffm.; see discussion under 5S. cuspidatum. Weissia glauca Bartr. reported by Smith (1966) is Weissia tortelis. Determina- tion made by Ann Stoneburner. 42 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN Fc a a a a eg ee ee | re CHEYENNE | Rawuins Toecatun | norton Tewuues | switm Vyeweue T repuacic [WASHINGTON Tuan Trewana i = ' | | | | \ | ! \ i i i | DONIPHAN | | H | | | | ! ! i i | ! | eee i eas bee RT bie acai a 8 Ale i | ee ene re et thet feces I SHERMAN Tihouas Vsheriban CSS Sexi OeROANE Tuareneue 1 Reiay a RILEY TpoTTAWATOMIE | JACKSON ietanre NY | | i [aeFFERsOn] Gee | ee a as I Fie f H j j . i ! , Slee - SL he =| A, ' | | | | — gryane i rR; Rear TenaWncc i furan E ee Siero es stipe eee at pal Reis Hotels: <5 pe Sa __ | LINCOLN j er te ) ee | (ees Heer’ : LOGAN + GOVE | TREGO eats Taliqusseuu i i j DICKINSON | es I J WABAUNSEE Aimao| er , | | | | i 71 eh 3 "7 | | ' poucvas T yoneson ‘ | ! H | Va yi a ete ste tana | ' | | | | ; j ELLSWORTH | i 7 MORRIS Wee z Pe 7 aa | ae -R te eae eer [En ch oe | ae teeta i | | Woven ' FRANKLIN i GREELEY = | wicniTA | scorr LANE Thess Trusn BARTON | L ! : | i | ‘ ' NinUSHE GS RBARTON. © 4) 0000 ae — | | | | | i ‘ _j MePHERSON — | MARION ie oe oe 2 i | | | | aaa © ; Vonase I 4 | | | | | | j Rice i 1 1 CHAS i eae ee ' i | i | ! j i i t i COFFEY ANOERSON | LINN | H i H } . . . i % { ee ee ade ‘ es Do es i ! ! { t ! ‘ [amicron Tieanne Trinwey 4 HOOGEMAN 7 Tamer! ! C= H | ‘ | } | meses New ees pat 7 H | | | + RENO HARVEY L.-.-i--—-p-— aes S ee OE -1 | | | eres | | ia | | j euTcer i GREENWOOD TL CoDsoN j ALLEN BOURBON | ' | i | oray ne att tn jeowaros | ; ees: | i ! i ‘ L 4 } 4 rORD 1 eee = _+ seocwick | { a { 1 | Seheiientin itettentereeeoe ; H 1 | PRATT Rte re | i == Nie eagne diac eae V stanton | grant Trasneue al | leanne = RGN nie ! i | WILSON ie CRAWFORD | | i | | | \ i | j = = | j . " i , . EuK i ' ! | ' | i | | \ i | le ear aN Ne ae i \ H b menses Failiaiagercs 0s = eer eg al | lcowcey je \i---—-— | es oS eS MEADE CLARK i BARBER ae ee ee eee SUMNER | ETTE — [sonrox ean T sewaro : 1 lcseanciead! Uaarrer | \ Se [ion cone oe | CHEROKEE | ‘ | | | | | | | i | i CHAUTAUQUA i i i | ! : H H \ \ i | | | | | | | ' | | | j | \ i ; i i Seeras ae = nh ee ee | a yes = ae HSS ee at ¥ L - oes oie o os Fic. 5. County locations in Kansas. A SYNOPSIS OF THE KANSAS MOSSES 43 ee “5 HEH AMBLYSTEGIUM TRICHOPODIUM AMBLYSTEGIUM VARIUM 44 Tue UnIversity OF Kansas SCIENCE BULLETIN ANOMODON ATTENUATUS Caine Petts ASTOMUM MUHLENBERGIANUM ASTOMUM PHASCOIDES A SYNOPSIS OF THE KANSAS MOSSES 45 BARBULA INDICA PoP es ores ree SEL leteletet teat te BARBULA UNGUICULATA 46 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Bo ae BRUCHIA TEXANA BRYHNIA GRAMINICOLOR A SYNOPSIS OF THE Kansas MosseEs 47 THe Bueoc BRYUM GEMMIPARUM BRYUM PSEUDOTRIOUETRUM DESMATODON PORTERI THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Bae DICRANELLA HETEROMALLA A SYNOPSIS OF THE KANSAS MOSSES 49 CT ee Pah TT Be DITRICHUM PALLIDUM DREPANOCLADUS ADUNCUS 50 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN mais et 5 ae aa take eee EPHEMERUM SPINULOSUM EUCLADIUM VERTICILLATUM A Synopsis OF THE KANSAS MOSSES 51 poo HEP Ee J pee ee FISSIDENS GARBERI FISSIDENS OBTUSIFOLIUS oy THE UNIVERSITY OF KANSAS SCIENCE BULLETIN GRIMMIA LAEVIGATA GRIMMIA PLAGIOPODIA A SYNOPSIS OF THE Kansas MosseEs 53 HYOPHILA INVOLUTA HYPNUM CURVIFOLIUM 54 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN soe eeea Cee LEPTOBRYUM PYRIFORME LEUCOBRYUM GLAUCUM a LEUCODON JULACEUS A SYNOPSIS OF THE KANSAS MOSSES 5S Spr PHILONOTIS FONTANA PHILONOTIS LONGISETA 56 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN PLATYGYRIUM REPENS PLEURIDIUM SUBULATUM A SYNOPSIS OF THE KANSAS MOSSES Sy Pees POGONATUM BRACHYPHYLLUM POLYTRICHUM JUNIPERINUM POLYTRICHUM PILIFERUM 58 Tue UNIvERSITY OF KANSAS SCIENCE BULLETIN RHODOBRYUM ONTARIENSE RHYNCHOSTEGIELLA COMPACTA A SYNOPSIS OF THE KANSAS MOSSES 59 SPHAGNUM TRINITENSE TAXIPEYLLUM DEPLANATUM 60 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN a TIMMIA MEGAPOLITANA TORTELLA HUMILIS A SYNOPSIS OF THE KANSAS MOosSsEs 61 aan aie grene PoE EE ee TORTULA CAINITI TORTULA RURALIS WEISSIA CONTROVERSA AUTHORS COLLECTION SITES ADDENDUM BRYOANDERSONIA ILLECEBRA WEISSIA TORTILIS D nN ACKNOWLEDGMENTS I am most grateful to those who identifed or verified certain specimens during the course of this study; they include Drs. R. Andrus, H. Crum, C. Delgadillo, R. Ireland, T. Koponen, Re Pursell, P>Redfearn Jr; J. Snider, A. sStoneburner, D). Vitt and R. Zander. I am indebted to Dr. M. Crosby and the Missouri Botanical Garden for providing funds for field work, and to Drs. W. Buck and W. C. Steere and the New York Botanical Garden for travel funds to study the Garden collections. Information and photograph of Minnie Reed were pro- vided by Dr. T. Barkley and the late Dr. R. Thompson, respectively. For permission to use published and un- published vegetation maps, I thank Drs. A. W. Kuchler and P. V. Wells. Drs. H. Crum and P. Redfearn Jr. reviewed the manuscript and gave many critically good comments, and Dr. W. Bloom provided helpful edi- torial suggestions. Sharon Hagen’s as- sistance and = advice concerning graphics is greatly appreciated. I am most appreciative of the Division of Biological Sciences secretarial staff, Jan Elder and Coletta Spencer, for typing the manuscript. LITERATURE CITED Anperson, L. E. 1971. Geographical relationships of the mosses of the Southern Appalachian Moun- tains, p. 101-115. Jn P. C. Holt and R. A. Paterson (eds.), The Distributional History of the Biota of the Southern Appalachians, Part II: Flora. Virginia Polytech. Inst. St. Univ., Res. Div. Monogr. 2: Blacksburg. ——. 1980. Cytology and reproductive biology of mosses, p. 37-76. InR. J. Taylor and A. E. Leviton (eds.), The Mosses of North America. Pacific Div. AAAS. AnpERSON, L. E. and B. E. Lemmon. 1972. Cytologi- cal studies of natural hybrids between species of the moss genera Astomum and Weissta. Ann. Missouri Bot. Gard. 59:382-416. Anpo, H. 1976 (1977). Studies on the genus Hypnum Hedw. (III). J. Sc. Hiroshima Univ., Ser. B, Div. 2. 16:1-46. THE UNIVERSITY OF KANSAS SCIENCE BULLETIN AnonyMous. 1966. Flora of the Maurice L. Brei- denthal Biological Reserve. University of Kansas, Lawrence. (Unpub.). Bare, J. E. and R. L. McGrecor. 1970. An intro- duction to the phytogeography of Kansas. Univ. Kansas Sci. Bull. 48:869-949. Bremer, B. 1980a. A taxonomic revision of Schistidium (Grimmiaceae, Bryophyta) 1. Lindbergia 6:1-16. . 19806. A taxonomic revision of Schistidium (Grimmiaceae, Bryophyta) 2. Lindbergia 6:89-117. Britton, E. G. 1890. New mosses of North America. IlI-IV. Renauld and Cardot. Bot. Gaz. 15:151. 1913. Sphagnales-Bryales. Bruchiaceae. North Amer. Flora 5:47-54. Bryan V. S. and L. E. Anperson. 1957. The Ephemeraceae in North America. The Bryologist 60:67-102. CuurcHIiLL, S. P. 1976. Mosses of the Great Plains: introduction and catalogue. Prairie Natur. 8:44-57. . 1979a. Mosses of the Great Plains III. Addi- tions to Nebraska and the Black Hills of South Dakota and Wyoming. The Bryologist 82:72-75. 19795. New records and distributions for Kansas mosses. Tech. Publ. St. Biol. Surv. Kansas 8:72-86. 1980. New records and distributions for Kansas mosses II. Tech. Publ. St. Biol. Surv. Kansas 9:78-89. . 1981. A phylogenetic analysis, classification and synopsis of the genera of the Grimmiaceae (Musci), p. 127-144. In V. A. Funk and D. R. Brooks (eds.), Advances in Cladistics. New York Bot. Gard., N.Y. 1982. Mosses of the Great Plains VIII. Additions. The Bryologist 86:218-221. . 1983. Mosses of the Great Plains. [X. East- ern Montana. Proc. Montana Acad. Sci. 42:17-23. Cuurcnitt, S. P., P. L. REDFEARN, JR., and G. J. IKENBERRY. 1981. Contributions to the moss flora of Oklahoma. The Bryologist 85:498-504. Conarp, H. S. 1951. Desmatodon obtusifolius in Iowa. The Bryologist 54:128-130. Criptanp, A. A. 1959. The habitat of Aschisma kansanum. The Bryologist 62:132-135. . 1960. Notes on Kansas mosses. The Bryolo- gist 63:51-53. CroizaT, L. 1964(1962). Space, Time, Form: the Biological Synthesis. Publ. by author, Caracas. Crum, H. 1966. A taxonomic account of the genus Thelia. Natl. Mus. Can. Bull. No. 46. Contrib. Bot. 4:123-127. . 1972. The geographic origins of the mosses of North America’s eastern deciduous forest. J. Hat- tori Bot. Lab. No. 35:269-298. ————. 1976. Mosses of the Great Lakes Forest, 2nd ed. Univ. Mich. Herb. 404 pp. Crum, H. and L. E. Anperson. 1964. Notes on Physcomitrium collenchymatum. The Bryologist 67: 350-355. Crum, H., W. C. Steere and L. E. ANpeErson. 1973. A new list of mosses of North America, north of Mexico. The Bryologist 65:85-130. Detcapitto, M. C. 1975. Taxonomic revision of A Synopsis OF THE Kansas Mosses 63 Aloina, Aloinella and Crossidium (Musci). The Bryologist 78:245-303. Duncan, E. S. 1959. Ecology of Kansas prairie bryophytes. MS thesis. Univ. Kansas, Lawrence. Girr, L. J. 1940. Flora of Crawford County (Kansas) State Park: mosses. Trans. Kansas Acad. Sci. 43119-1120. . 1949. Bryophytes of Kansas. Trans. Kansas Acad. Sci. 52:58-65. Grier, L. J. and E. Kennepy. 1955. Thelia lescurri—an environmental form. Trans. Kansas Acad. Sci. 58:334-336. Grout, A. J. 1928-1940. Moss Flora of North Amer- ica, North of Mexico. 3 vol., Newfane. Grucer, J. 1973. Studies on the Late Quaternary vegetation history of northeastern Kansas. Geol. Soc. Amer. Bull. 84:234-250. Hartman, E. L. 1956. The bryophyte flora of the University of Kansas Natural History Reservation. Trans. Kansas Acad. Sci. 59:57-70. Hipparp, C. W. 1955. The Jinglebob interglacial (Sangamon?) fauna from Kansas and its climatic significance. Contrib. Mus. Paleontol., Univ. Michigan 12:179-228. IRELAND, R. R. 1969a. Taxonomic studies on the genus Atrichum in North America. Can. J. Bot. 47:353-368. . 19696. A taxonomic revision of the genus Plagiothecium for North America, north of Mexico. Natl. Mus. Nat. Sci. (Canada) Publ. in Bot. No. 1. 118 pp. . 1970. Validation of the moss name Atrichum altecristatum (Ren. & Card.) Irel. Can. J. Bot. 48:1897. Kapp, R. C. 1965. Illinoian and Sangamon vegeta- tion in southwestern Kansas and adjacent Okla- homa. Contrib. Mus. Paleontol., Univ. Michigan 19:167-255. Kina, J. E. 1973. Late Pleistocene palynology and biogeography of the western Missouri Ozarks. Ecol. Monogr. 43:539-565. Koponen, T. 1968. Generic revision of Mniaceae Mitt. (Bryophyta). Ann. Bot. Fennici 5:117-151. Kucuter, A. W. 1972. The oscillations of the mixed prairie in Kansas. Erdkunde 26:120-126. . 1974. A new vegetation map of Kansas. Ecology 55:586-604. Lane, D. M. 1981. Variation in certain taxa of Sphagnum from the Atlantic Coastal plain. J. Hat- tori Bot. Lab. No. 49:169-245. Lowry, R. J. and W. C. Steere. 1946. A pro- paguliferous form of Aulacomnium heterostichum. The Bryologist 49:30-32. MacArtuur, R. H. 1972. Geographical Ecology. Harper and Row. Maciit, R. E. and R. Kirkpatrick. 1976. Notes on Kansas mosses. The Bryologist 79:101-102. McGrecor, R. L. 1950. Studies on Kansas mosses. I. Univ. Kansas Sci. Bull. 33:291-311. . 1976. Mosses of the Woodson County State Fishing Lake and Game Management Area, p. 4-5. In J. Caldwell (ed.), Preliminary Inventory of the Biota of Woodson County State Fishing Lake and Game Management Area. Report St. Biol. Surv. Kansas No. 5:1-76. ——. 1977. Notes on the genus Sphagnum in Kan- sas. Report St. Biol. Surv. Kansas, No. 18:1-3. McGrecor, R. L. and E. L. Hartman. 1956. Studies on Kansas mosses II. Univ. Kansas Sci. Bull. 38:331-333. Miter, N. G. and J. H. Amsrose. 1976. Growth in culture of wind-blown bryophyte gametophyte frag- ments from arctic Canada. The _ Bryologist 79:55-63. Rau, E. A. 1884. First contribution to the knowledge of Kansas mosses. Bull. Washburn Lab. Nat. Sci. 18: . 1885a. Second contribution to the knowledge of Kansas mosses. Bull. Washburn Lab. Nat. Sci. 1:60-62. . 18856. Third contribution to the knowledge of Kansas mosses. Bull. Washburn Lab. Nat. Sci. 1:114. . 1886. Fourth contribution to the knowledge of Kansas mosses. Bull. Washburn Lab. Nat. Sci. Nei lelys. REDFEARN, P. L., Jr. 1972. Mosses of the Interior Highlands of North America. Ann. Missouri Bot. Gard. 59:1-104. Reep, M. 1893-1894. Kansas mosses. Trans. Kansas Acad. Sci. 14: 152-198. RENAULD, F. and J. Carpor. 1888. New mosses of North America I. Bot. Gaz. 13:197-203. . 1889. New mosses of North America II. Bot. Gaz. 14:91-100. . 1890a. New mosses of North America III. Bot. Gaz. 15:40-41. . 18906. New mosses of North America IV. Bot. Gaz. 15:57-62. . 1892. Enumeration of the Kansas mosses. Bot. Gaz. 17:81-85. Situ, D. K. 1980. Funaria americana Lindb. in North America. The Bryologist 83:335-339. SmitH, H. L. 1961. Mosses of the Chatauqua Hills. MS thesis. Univ. Kansas, Lawrence. . 1966. Mosses of the Great Plains and Arkan- sas River Lowlands of Kansas. Univ. Kansas Sci. Bull. 46:433-474. Smytu, B. B. 1890. Additions to the flora of Kansas. Trans. Kansas Acad. Sci. 12:115-119. . 1891-1892. Additions to the flora of Kansas. Trans. Kansas Acad. Sci. 13:102. . 1893-1894. Additions to the plants of Kansas. Trans. Kansas Acad. Sci. 14:133-134. Smytu, B. B. and L. C. Smytn. 1911. Catalogue of the flora of Kansas. Part 1. Trans. Kansas Acad. Sci. 23-24:280-287. Stewart, J. D. 1979. Paleontology and paleoecology of the Trapshoot Local Fauna, Rooks County, Kansas. MS thesis. Univ. Kansas, Lawrence. Upvarpy, M. D. F. 1969. Dynamic Zoogeography. Van Nostrand-Reinhold, Princeton. Vitt, D. H. 1973. A _ revision of the genus Orthotrichum, in North America, north of Mexico. Bryophyt. Bibliotheca 1:1-208. Wa.ter, W. T. and J. C. Bass. 1967. Sphagnum 64 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN cuspidatum new to Kansas and Missouri. The Bryologist 70:440. Weser, W. A. 1973. Guide to the mosses of Colorado: keys and ecological notes based on field and herbar- ium studies. Institute of Arctic and Alpine Re- search. Occas. Pap. No. 6, Univ. Colorado. 48 pp. Wetcu. W. H. 1960. A Monograph of the Fon- tinalaceae. The Hague. 357 pp. WeLLs, P. V. 1970. Postglacial vegetational history of the Great Plains. Science 167:1574-1582. Wixtarp, J. T. 1909 (1908-1909). William Ashbrook Kellerman. Trans. Kansas Acad. Sci. 22: 46-47. 1 plate. Zates, W. M. 1976. Key to the North American species of Philonotis. (Unpub.). ZANDER, R. H. and P. M. Eckev. 1980. Tortula cainit: additional Ontario stations and behavior in a com- mon garden. The Bryologist 83:209-211. APPENDIX A list of Kansas moss collectors is given below and is based on collections that I have personally examined over several years at various herbaria. To denote where each collectors specimens are deposited I have indicated only the main institutional collections. Her- barium abbreviations are: CANM, National Herbarium of Canada, Ot- tawa; DUKE, Duke University, Dur- ham; KANU, University of Kansas, Lawrence; KSC, Kansas State Univer- sity, Manhattan; MO, Missouri Bo- tanical Garden, St. Louis; NEB, University of Nebraska, Lincoln; NY, New York Botanical Garden, Bronx; PC, Herbier du Laboratoire de Cryp- togamic du Museum National d’ His- toire Naturelle, Paris. Kansas Moss Collectors Agrelus, FU. G..(NY) Barber, M. A. (NY) Barker, William (NY) Bartholomew, E. (NY) Bass, J. C. (MO) Beede, J. W. (NY) Bennett, Rev. John (MO, NY) Brooks, Ralph (KANU) Carleton, M. A. (NY) Churchill} -S: BP, (MO, NY, KANU) Cridland, Arthur A. (DUKE) Demetrio, C. H. (NY) Ellison, Marlon (NY) Fraser, S. V. (NY) Gates, Frank (KSC) Gier, "Es CNY) Hall, Elihu (NY) Hancin, John (NY) Hartman, Emily L. (NY) Herrick, C. Judson (NY) Holland, W. W. (NY) Horn Wel gNyY) Henry, Joseph (NY, PC) Ikenberry, Gilford J. (MO, NY) Ireland, Robert R. (CANM, NEB) Jones, Mary (NY) Kellerman, Williams (NY) Kirkpatrick, Randy (MO) McClung, Frank (NY) McGregor, Ronald L. (KANU, NY) Meeker, Grace (NY) Norton, J. B. S. (NY) Reed, Minnie (KSC, NY) Richards, B. (NY) Riley, ? (NY) Roth, Stan (KANU) Showalter, W. (NY) Smith, Harold L. (NY) Smyth, Bernard B. (NY) Swingle, W. (NY) Thomas; ’O2 E-vGNY) Thompson, Rufus (NY) Ungar, I. At (NY) Vitt, Dale (NY) Volle,. Es Ea( NY) Waller, W. T. (MO) Waters; L7 (NY) Wells, P. V. (MO; KANU) Le wo © wow ww 2 © © oo = © ~~ == .~5-9-9-0'6 0 0 0 0 8 6 8 208 00000902990 F 02S SS 2S 02S eee ees ee ees eo sWe%e%e"e"eTavaTe"e"e7 e707 e070 767070760-9_0,0,0,0,09,0,0,9,0,0,0,0,0,0,0,0,0,0,9,0,0,0,0,0,0,0,0 0 .rsmarerarar arene er erererenerene ero eo ete ete's| "ono 0 0-010. 8. 0.0.0.0. 0.0.0.0. 0,0. 0,0,9 "a" ane ooo 8 on" 0a "a"e'0' 0's" 00's 818 00 60880000 08000606 0 60°60 8' 00's" see's" SCIENCE BULLETIN MCZ LIBRARY APR 17 1986 HARVARD UNIVERSITY Se = wisn A Revision of the Native New World Species of the Ant Genus Monomorium (minimum Group) (Hymenoptera: Formicidae) Se SS ntefatatecatetatotetetenatecetatetatetacccotoceeeseeceswetetetetsteeseeeegenseseseeeeecs By Mark B. DuBois ESTED PRR a St" a aK OR RK RI RRS Sa OR OR OR Ny Vol. 53,.No. 2; pp.°65-119 March 24, 1986 SR RR HH I OR RR ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas University Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its inception, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now exhausted. However, most volumes of the University of Kansas Science Bulletin are still available and are offered, in exchange for similar publications, to learned societies, colleges and universities and other institutions, or may be purchased at $30.00 per volume. Where some of these volumes were issued in parts, individual parts are priced at the rate of 3 cents per page. Current policy, initiated with volume 46, is to issue individual papers as published. Such separata may be purchased individually at the rate of 4.5 cents per page, with a minimum charge of $1.50 per separate. Subscriptions for forthcoming volumes may be entered at the rate of $30.00 per volume. All communications regarding exchanges, sales and subscriptions should be addressed to the EXCHANGE LIBRARIAN, UNIVERSITY OF KANSAS LIBRARIES, LAWRENCE, KANSAS 66045. Reprints of individual papers for personal use by investigators are available gratis for most recent and many older issues of the Bulletin. Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor Charles -D. Michener Editorial Board William L. Bloom Philip W. Hedrick Rudolf Jander Delbert Shankel Norman A. Slade George W. Byers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. 53, No. 2, pp. 65-119 March 24, 1986 A REVISION OF THE NATIVE NEw WorLD SPECIES OF THE ANT GENUS MonomoriuMm (MINIMUM GROUP) (HYMENOPTERA: FORMICIDAE)! Mark B. DuBots2 CONTENTS ZANE SGU A Gish ee ed oer Aa ae EE Oana AE ska aed as S OM e Ak 66 NER @ 1) CANON Mee hg es Se ee oe roe eek oR cee Ens AV Pi ols olen ane 66 GENERA ViEMH@ 1 Swtien seer ike Otic: hak Ween hese Daa: te A eee at neg eneee ik 69 GES Ie ere en aeons Sana CaM eM LBerme Cit cet cite ere Stak eae han ee ADE ty AE eek aN Co 74 SVS MeEIVIAGIg Cees AM NIE Nulek Paka eet eweees gS ciety Sepa WO STEM 6 kot og phe Ud MoONOMORIUM (MONOMORIUM) MINIMUM SPECIES GROUP ............... The) VA TON ADINTENA Cag (ISLE KATE) penn ea esas A ck Ae ci esate Oe SL Sec oe U5) VISCCANEUNMGNIER MER 2 axe so. 19 ead een he. Spo 2, | eR ee eS ae 82 WMeERCATOGYNA.\NIFRRERR pie, ions. old 2. 1.8 oe een 2 eee 86 MEMVHEEDERORUM NEW SPECIES igs «s.2 “ses knee eee GAs Se ee 91 WVIMCOMPRESSUM VWITREIUBR: © 2s ctidh-24 « oS: See eee 93 MER UERED UATE ROWING Searles ae els 4a 5-2 a. ae RE Le eee 96 MISEMARGCINATUMENEW ‘SPECIES# is = 0 = 6.6.0 ed Hae eee eo ee ee 101 IVI RSP BENIN CIM ALOR G¥ ha. Poy 2es acs 5

-Santschi, 1927; 1936: Ettershank, 1966). Three Old World species are widely distributed ‘‘tramps’’ in the New World (predominantly in tropical and subtropical regions): Monomorium (Monomorium) floricola (Jerdon), M. (M.) pharaonis (Linnaeus), and M. (Parholcomyrmex) destructor (Jerdon) (Creighton, 1950; Brown, 1964; Kempf, 19723 Smith, 1979): In addition to these species, Kempf (1972) lists the tropical species occurring in the New World. Two of them, MZ. minutum brastliense Forel (1908) and M. subcoecum Emery (1894), may belong to the group herein revised. Lack of queens for com- parisons prevented me from including them. All remaining New World species belong to different species groups. Emery (1921) considered the minimum group closely related to M. minutum Mayr (which occurs in southern Europe). Based on published descriptions of M. minutum and comparison of 2 queens and 12 work- ers of this species (from France, Italy and Yugoslavia) with members of the minimum group, it appears unlikely that this species is Closely related to any of the native New World Monomorium. Other evidence which indirectly sup- ports the above contention lies with plate tectonics of North America and Europe. Since Pangea probably fragmented before the Formicidae evolved, it is unlikely that Monomorium could have dispersed directly to North America from Europe (Wilson et al., 1967a, 1967b; Dietz and Holden, 1970; Burnham, 1978). It seems more likely that Monomorium arrived in North America via the Bering land bridge since a number of other species are thought to have arrived via this route (Buren, 1958). Furthermore, comparisons between work- ers of North American Monomorium and Asiatic Monomortum (M. chinensis Santschi, 1925) reveal many similarities (they are difficult to differentiate). However, no queens of Asiatic species were available for study. Most studies on native New World members of the mznizmum group have been isolated descriptions of taxa with little effort made to synthesize this information. Confusion has resulted from descriptions which compare taxa being described with that author’s concept of M. minimum. Thus, Wheeler (1904) described M. mini- mum ergatogyna and compared it with ‘‘M. minimum’ from New England (now known as M. emarginatum). Later, Wheeler (1914b) described M. minimum compressum and M. minimum cyaneum and compared them with M. minimum from Texas (which may have been M. minimum or M. cyaneum). GENERAL BIOLOGY Little biological information has been recorded for most species of Monomorium Ant GENUS MONOMORIUM 67 FIGS. 1-5. Monomorium ergatogyna. Scanning electron microscope photographs. Some photographs enlarged more than others; for actual dimensions, refer to text. 1—Head of male, frontal view. 2— Head of worker, frontal view. 3—Head of queen, frontal view. 4— Male, lateral view (legs and wings removed). 5—Queen, lateral view (legs removed). (Ettershank, 1966). One notable exception is the detailed study of M. pharaonis (Pea- cock and Baxter, 1949, 1950; Peacock, 1950a, 1950b, 1951; Hall and Smith 1951, 1952,.1953; Smith and Peacock, 1957). Most studies of biology of the mznimum group emphasized information useful in constructing higher classifications of ants. Recent studies dealt with larvae (Wheeler and Wheeler, 1955, 1960a, 1960b, 1973, 1976), chromosomes (Crozier, 1970, 1975), sting morphology (Kugler, 1978), and venom components (Jones et al., 1980, 1982). INQUILINISM Inquilinous relationships between two species are those in which a socially para- sitic species spends its entire life cycle in nests of its host species (Wilson, 1971). Since the inquilines rely on their hosts for food and care of their brood, the worker caste is often lacking (no workers have ever been discovered in the North American inquilinous Monomorium). This relationship is sometimes called ‘“‘permanent para- sitism.”’ Three species of inquilinous Monomorium have been discovered in North America: M. inquilinum (known only from one lo- cality in central Mexico), M. pergandei (known only from Washington, D.C.), and M. talbotae (known only from one locality in central Michigan). These species follow the distributional pattern of most inquilinous ants, being known only from isolated, lim- ited areas. These inquilines are remarkably similar in general morphology, presumably be- cause of convergence. In M. pergandei and 68 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN FIGS. 6-11. Monomorium queens, heads, frontal views. Scanning electron microscope photographs. Some photographs enlarged more than others to aid in comparison; for actual dimensions, refer to text. 6—M. minimum. 7—M. trageri. 8—M. ebenium. 9—M. viridum. 10—M. cyaneum. 11—M. wheelerorum. M. talbotae, males and females are quite similar to one another in overall mor- phology. In fact, the only method of deter- mining gender is to examine their genitalia. This phenomenon has yet to be explained since non-inquilinous Mono- moritum have marked morphological differ- ences between sexes. DuBois (1981a) recognized the following common traits of inquilinous Monomorium (numbers of traits correspond to those of Wilson, 1971: 374-375): (1) the worker caste is absent, (4) both male and female are reduced in size, (8) venation in both forewing and hindwing is diluted — most ‘veins’ in forewing are represented as ab- sence of microtrichia; hindwing venation is equally diminished, (9) mouthparts are re- duced, (13) petiole and postpetiole are thickened, (14) ventral surface of postpetiole becomes armed with a spine, and (16) cuticular sculpturing is reduced. One additional character, probably associ- ated with diminished size, is reduction in size and complexity of male genitalia (Du- Bois, 1981a). Fossit History Although genus Monomorium is repre- sented in the fossil record by two species, M. mayrianum Wheeler and M. pilipes Mayr, both species are known only as workers in Baltic Amber (Wheeler, 1914a; Burnham, 1978). No fossil Monomorium have been discovered in North America (see, for example, Carpenter, 1930; Brown, 1973; Burnham, 1978). Neither fossil species appears to be closely related to native North American Monomorium. Monomorium mayrianum has 5 AntT GENUS MONOMORIUM 69 FIGS. 12-15. Monomorium queens, lateral views (gasters and legs excluded). Scanning electron microscope photographs. Some photographs enlarged more than others to aid in comparison; for actual dimensions, refer to text. 12—M. minimum. 13—M. cyaneum. 14—M. ebenium. 15—M. viridum. mandibular teeth, 12-jointed (segmented?) antenna, and a deep metanotal sulcus (Wheeler, 1914). Monomorium pilipes has 11- jointed (segmented?) antenna, and (proba- bly) 5 mandibular teeth based on Wheeler’s (1914a) description and accom- panying figure. GENERAL METHODS As specimens were received, they were sorted into morphologically distinct groups; each group was compared with specimens previously received. These ‘‘morphospecies’’ were segregated with no effort to name each taxon due to prevailing confusion in the systematics of this group. When the majority of specimens was as- sembled, collection localities were plotted on a map to ascertain which geographic areas lacked representatitives. Efforts were then made (by contacting individuals or institutions in those regions) to cover defi- cient areas. Finally, names were assigned to taxa after comparison with type material or with descriptions when types were not readily available. When possible, an effort was made to assign previously recorded information to one species or another. A reference is listed in synonymy if either one of two criteria is met: (1) a specimen from that study was examined and identified by me, or (2) the literature reference in question referred to an area where only one species of Mono- mortum probably occurs naturally. Ob- viously, this latter criterion is the weaker of the two. ‘TERMINOLOGY Most terms used are conventional to hymenopteran systematics and were de- fined by Snodgrass (1935, 1941). Some special terms, although known to myr- mecologists, may cause difficulty; these have been listed below to clarify their use within this publication. Most of these terms have been used previously in revisions of other ant taxa (see, for example, Snelling, 70 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN FIGS. 16-21. Monomorium workers, lateral views (gasters and legs excluded). Scanning electron microscope photographs. Some photographs enlarged more than others to aid in comparison; for actual dimensions, refer to text. 16—M. ebenium. 17—M. wheelerorum. 18—M. ergatogyna. 19—M. minimum. 20—M. tragert. 21—M. viridum. 1976; Ettershank, 1966). Terms are listed in the order they appear within descrip- tions. 1. Head length (HL). Maximum length of head (in full face view), measured from midpoint of occipital margin to midpoint of anterior clypeal margin (Fig. 22, no. 1). 2. Head width (HW). Maximum width of head (excluding compound eyes) (Fig. 22, no. 2). 3. Scape length (SL). Maximum length of scape (excluding basal radicle) (Fig. 22, no:: 3): 4. Interocellar distance (LOD). Queen and male only; minimum distance between in- ner margins of posterior ocelli, measured from dorsal view (Fig. 22, no. 4). 5. Ocellar diameter (OD). Queen and male only; maximum diameter of anterior ocellus (Fig. 22, no. 5). 6. Eye length (EL). Maximum length of compound eye, measured from lateral view (Fig. 22, no.,6). 7. Maximum ocular diameter (MOD). Max- imum width of compound eye, measured from lateral view. 8. Cephalic index (CI). Head width X 100 / head length. 9. Scape index length comparison (SIL). Scape length X 100 / head length. 10. Scape index width comparison (SIW). Scape length X 100 / head width. 11. Pilosity. Setal terminology used to indi- cate degrees of inclination following Wilson (1955) including following terms: (1) erect, (2) suberect, (3) subdecumbent, (4) decum- bent, and (5) appressed (Fig. 23). Since these terms are discontinuous expressions of a continuous variable, the two extremes are used to delimit variation (for example, Ant GENUS MoNnoMORIUM FIGS. 22-25. Terminology. Fig. 22. Frontal view of head, generalized to show measurements. Numbers refer to following measurements: 1—head length, 2—head width, 3—scape length, 4— interocellar distance, 5 - ocellar diameter, 6—eye length. Fig. 23. Schematic representation of terminology of angle of declination of vestiture (after Wilson 1955). Numbers represent following declinations: 1—erect, 2—suberect, 3—subdecumbent, 4—decumbent, 5—appressed. Fig. 24. Dorsal view of scutum. 1—mesonotal furrow, 2—measurement of pronotal width (pronotum extends laterally along alitrunk and is usually the widest directly underneath this part of the scutum; refer also to Figure 25, number 4). Fig. 25. Lateral view of queen alitrunk. 1—propodeal length, 2—Weber’s length (actual measurements are taken at a slight diagonal from front to rear instead of in a horizontal line as figure indicates), 3—scutum, 4—pronotum, 5—scutellum, 6—metanotum, 7—anepisternum, 8— katepisternum, 9—propodeum. Further explanations are provided in text. setae erect to subdecumbent). First term used denotes most common inclination of setae on a given structure. 12. Sculpture. Usually reflected in terms of punctation or rugosity (see below). When surfaces are described as smooth and shin- ing, this is their appearance at magnifica- tions of 80X or lower; all cuticular surfaces appear rough at higher magnifications. 13. Punctation. Punctures can be piliferous (with setae) or non-piliferous (without setae). Sizes are given in relation to aver- age diameter of one eye ommatidium. Se- lect 5 ommatidia from the central portion of the compound eye and average their diameters. Small punctures are smaller than the average diameter of an om- matidium, moderate punctures are ap- proximately the same diameter, and large punctures are larger than the average di- ameter. 14. Rugosity. Size as for punctation except size is measured as distance between two consecutive ridge summits (i.e., width of one trough). 15. Pronotal width (PW). Maximam width of pronotum, measured dorsally, perpen- dicular to longitudinal axis of thorax. 16. Propodeal length (PL). Maximum length of propodeum measured along longitudinal 72 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN axis of body from metanotum to apex of inferior propodeal plates (Fig. 25, no. 1). 17. Weber’s length (WL). Maximum length of alitrunk, measured diagonally from dor- sal margin of pronotal collar to apex of inferior propodeal plate (Fig. 25, no. 2). 18. Propodeal index (P1). Propodeal length X 100/Weber’s length. 19. Color. Although all taxa examined ap- pear piceous black at low magnifications, color refers to coloration observed at mag- nifications of 30X% and above under flores- cent lighting. Many of the specimens examined have faded with age; hence, the colors in fresh specimens may be darker than indicated in the descriptions. MEASUREMENTS All measurements were made from dry, mounted specimens; most measurements were made at 80X% magnification. Meas- urements were made with an ocular mi- crometer (100 units), converted to millimeters, and rounded to the nearest 0.01 mm. Where possible, these measure- ments were made from individuals (se- lected at random) from different localities (one individual per locality). All measure- ments are expressed as range and mean (mean in parentheses) with a maximum sample size of 10 per caste. Initial measure- ments (including mean, standard devia- tion, median, minimum value, maximum value, and correlation coefficients) are listed in DuBois (1981c). Although both indexes involving scape length are listed for each species, it appears that the scape length to head length index is slightly more reliable; these indexes are closely correlated (most species r= 0.900 or greater). Since previous authors have em- ployed either index in various revisions (cf. Francoeur, 1973; Snelling, 1976), both in- dices are included for comparative pur- poses. Several steps were taken to avoid as many errors as possible. First, no measure- ments were recorded until I was able to obtain like measurements of the same structure on the same specimen on differ- ent days. Second, after measurements were taken, a number were selected (at random) and repeated. If the values differed for a given specimen, additional measurements were taken until an agreement was reached. Third, all measurements used in statistical analyses were checked against the original data sheets in search of ty- pographical errors. Similarly, the final manuscript was checked against original data sheets. SpEcIES DESCRIPTION FORMAT Descriptive features pertaining to holo- type only are surrounded by brackets {} (with the exception of M. inquilinum, which is only known from the holotype). Features which rarely occur in a species are sur- rounded by the following symbols )¢. If specimens are probably typical, overlook information within these symbols. ORIENTATION Specimens should be oriented in the same manner I examined them when de- scribing a species. The head (excluding gular region and compound eye measure- ments) should always be examined in full frontal view. This is particularly true when examining setae, since their degree of in- clination sometimes changes when view- point is radically changed. The gular region, compound eye measurements, al- itrunk, petiole, postpetiole (excluding their dorsa), and gaster should be examined in lateral view. Both petiolar and postpetiolar dorsa should be examined by looking over the posterior end of the gaster, up the gastral dorsum. DISSECTIONS The following structures must be dis- sected before features can be observed: wings, male mouthparts, and male geni- talia. Wings — Most wings curl around their longitudinal axes in pinned specimens. Furthermore, wing venation is difficult to observe in specimens preserved in alcohol. Therefore, wings were removed, placed on a microscope slide, smoothed out, covered with a cover slip, and affixed to the slide with tiny droplets of mounting medium on corners of the cover slip. Since the mount- ing medium has nearly the same refractive Ant GENUS MONOMORIUM index as the wing itself, venation is nearly invisible if the entire wing is immersed in the medium. Venation in all species exam- ined is fairly uniform; therefore, one exam- ple is depicted (M. ergatogyna, Fig. 71). Male mouthparts — Mouthparts of males were selected for detailed study, since they are used less than queen and worker mouthparts. Additionally, female mouthparts are less variable among spe- cies. Mouthparts (labial palp, maxillary palp, and mandible) were examined in glycerine at 400X; illustrations were made using an ocular grid. Male genitalia — The male genital capsule was removed entirely (after being drawn in situ) and dissected so that individual parts could be examined (examinations and illus- trations same as for mouthparts). The eighth and ninth sterna were examined (in glycerine) after being flattened by a cover slip. PRESERVATION OF SPECIMENS Due to their small size, most specimens should be preserved in alcohol. If necessary (for detailed examination) a subset can be pinned on points. Although the legs of the right side should be pushed into the glue, the entire body should never contact glue as the glue may spread by capillary action between setae and obscure details on the body. ComPuTER TECHNIQUES Preliminary data analysis was conducted using MINITAB statistics programs which were run on the University of Kansas Academic Computer Center’s Honeywell 66/60 computer. REFERENCE COLLECTIONS Specimens examined during this revi- sion were borrowed from the following individuals and institutions (abbreviations are those used in listing localities). I am extremely grateful to those who cooperated by sending specimens. AFPC—Andre Francoeur, personal col- lection AMNH—American Museum of Natu- ral History, New York (Marjorie Favreau) ~J] SX) BPPC—B. Poldi, personal collection (Mantova, Italy) CAS—California Academy of Sciences, San Francisco (Paul Arnaud, Jr.) CGPC—Chris George, personal collec- tion GCWPC—George C. and Jeanette Wheeler, personal collection INHS—lIllinois Natural History Sur- vey, Urbana (Wallace LaBerge) JMPC—James Moody, personal collec- tion JTPC—James Trager, personal collec- tion KSU—Kansas State University, Man- hattan (Jim Johnson) KU—Snow Entomological Museum, University of Kansas, Lawrence (George Byers) LACM—Natural History Museum of Los Angeles County (Roy Snelling) MBDPC—Mark DuBois, personal col- lection MCZ—Museum of Comparative Zool- ogy, Cambridge, Massachusetts (Margaret Thayer and Ron McGinley) MHNG—Museum d’Histoire Natu- relle, Geneva, Switzerland (Claude Besuchet) MTPC—Mary Talbot, personal collec- tion NCSU—North Carolina State Univer- sity, Raleigh (C. Parron) NMNH—National Museum of Natural History, Washington, D.C. (David Smith) OHSU—Ohio State University, Co- lumbus (R. Triplehorn) ORSU—Oregon State University, Cor- valis (Gary Stonedahl) OSU—Oklahoma State University, Stillwater (W. Drew) PWPC—Petr Werner, personal collec- tion (Prague, Czechoslovakia) RHPC—Robert Hamton, personal col- lection SBMNH-—Santa Barbara Museum of Natural History, California (Scott Miller) SBSK—State Biological Survey of Kan- sas, Lawrence (Mark DuBois) 74 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN UA—University of Arkansas, Fayette- ville (Sheila Hoelscher) UM—University of Missouri, Colum- bia (Edward Riley) UN—University of Nebraska, Lincoln (Brett Ratcliffe) UO— University of Oklahoma, Nor- man (Bill Shepard) KEY TO QUEENS . Head broadest at occiput; surface sculpturing greatly reduced; mas- ticatory margin of mandible with fewer Gloama teeth 2a 145. 8a vg es te S 11 Head broadest near compound eyes; at least some surface sculpturing near mandibular insertion; masticatory margin of mandible with 4 teeth . . 2 . (1). Lateral profile of scutum and scu- tellum convex((Fisure 43)... 2.5. Q Lateral profile of scutum and scutellum flator concave (Fig. 87). ..2.52.. 5 . (2). Scape with lateral fringe of erect setae (Fig. 82); occurs in California, Nevada, and Utah... M. wheelerorum Scape lacking lateral fringe of erect setae . (3). Petiole emarginate; postpetiole not emarginate; head sculpturing reduced (Fig. 120); occurs in Caribbean region Sh na Pee or eee es, esas, oh M. ebenium Petiole not emarginate; postpetiole emarginate; head sculpturing as in Fig. 88: occurs in central Mexico. ..... PN set See Ne GS TL, 2 M. compressum . (2). Mesonotum with notal furrow (Piss 24; rio.) Mesonotum lacking notal furrow . . 7 . (5). Metanotum not projecting to level of propodeum and scutellum; petiole and postpetiole not emarginate; occurs in central and north-central Atlantic COAStadl States 00s M. emarginatum Metanotum projecting to level of pro- podeum and scutellum; petiole and postpetiole emarginate; occurs along coastal regions from New Jersey to POLI as caries metus cece eee M. viridum . (5). Scape with lateral fringe of erect 8. 10. ie 12. 1 setae (Fig. 59); occurs in coastal and southern California. . . . M. ergatogyna Scape lacking lateral fringe of erect setae (7). Basal face of propodeum 2X length of declivity; known only from type locality (Nayarit, Mexico) M. marjoriae Basal face of propodeum never 2X lenathrot declwity 5 Ace: t ent @ . (8). Head sculpturing greatly reduced, rugae only on lateral margin of clypeus; occurs in northern Florida. . eee avi ae AS ees tee eens mane M. trageri Head with rugae in addition to those on lateral margin of clypeus.... . 10 (9). Gyne winged (sclerites near wing base not fused); postpetiole emargi- nate: common, im central U..Si..0%, a: ee ee eer M. minimum Gyne wingless (sclerites fused); postpetiole not emarginate; occurs throughout central Mexico, Arizona, New Mexico, and parts of Texas... 5 yd cis, rade Seber Dae, Seber M. cyaneum (1). Metanotum reaching level of pro- podeum and scutellum; postpetiole CMIPALGINALO 5. < 05 ay ais 5 thks pate 12 Metanotum not reaching level of pro- podeum and scutellum; postpetiole not emarginate; known only from type lo- cality (Livingston Co., Michigan) . . Seen us ue God ea ee, M. talbotae (11). Mesonotum with notal furrow; petiole not emarginate; masticatory margin of mandible with 3 teeth; known only from type locality (Wash- ington, HOCA is e883 2 M. pergandet Mesonotum lacking notal furrow; pe- tiole emarginate; masticatory margin of mandible with 2 teeth; known only from type locality (Estado de Mexico, IMeXICODN pander, ce thiat =, es M. inquilinum Key TO KNown MALES The following species are not included in this key: M. compressum, M. emarginatum, M. inquilinum, M. marjoriae, M. pergandet, and M. wheelerorum. . Eighth sternite emargination lacking Ant GENus MoONOMORIUM Hie PEOJECUME SETAC a a ee: Z Eighth sternite emargination with pro- jecting setae; common along Atlantic coast from New Jersey to Florida... . a ne Aer RATER heme a a M. viridum 2. (1). Aedeagus with toothed margin Sthalchiw ewes Pes ete Balee ese 2 3 Aedeagus with toothed margin curved 3. (2). Cuspis of volsella present; known only from type locality (Livingston Co. Michigan) <5) se M. talbotae Cuspis of volsella absent; occurs throughout Caribbean region .... . Feet eee bar oe ee ne ae M. ebeninum 4. (2). Cuspis of volsella with 3 setae . 5 Cuspis of volsella with 5 setae; occurs throughout central Mexico, Arizona, New Mexico, and parts of Texas... . Sao ee eee Tee M. cyaneum I(t) Aedeacus. quadrate; “occurs throughout central U.S.. M. minimum Aedeagus longer than broad..... 6 6. (5). Eighth sternite emargination 1/2 as deep as wide (Fig. 74); occurs along Califormia coast’... M. ergatogyna Eighth sternite emargination as deep as wide (Fig. 146); occurs in northern loridata-n bans sae aa epee M. tragert Key TO KNown WORKERS The following species are not included in this key and apparently lack a worker caste: M. inquilinum, M. pergandei, and M. talbotae. 1. Propodeum rounded (Fig. 63); occurs along California coast . . M. ergatogyna Propodeum angular (Fig. 31)... .. 2 2. (1). Basal and declivitous faces of pro- podeum of equal length ........ 3 Basal and declivitous faces of pro- podeum of unequal length (either face mays be longer): = la 5 cen n ee. 2% 5 3. (2). Mesopleuron punctate; fewer than 8 setae projecting above dorsum of alitrunk; occurs throughout central Mexico, Arizona, New Mexico, and Pacisvot Dexas'.26 7 ae M. cyaneum Mesopleuron not punctate; more than 10 (rarely 8 or 9) setae projecting above dorsumvof aliirunks 225532. 24 4 4. (3). Petiole and postpetiole convex to flat:voccurs im. central Miexico - 2. =. oe CET eS M. compressum Petiole and postpetiole flat to slightly emarginate; occurs throughout Carib- bean: region. yee er oa M. ebeninum 5. (3). Basal face of propodeum length less: than dechivity:-—-maes es ieee ee 6 Basal face of propodeum length greater than dechivityss ae ce 5 ce en eee 7 6. (5). Fewer than 6 setae projecting above dorsum of alitrunk; occurs in northeastern U.S. . . . M. emarginatum More than 8 setae projecting above dorsum of alitrunk; occurs in northern BlORGas 34h eee Bn eae M. trageri 7. (5). Mesopleuron punctate; occurs along Atlantic coastline from New Jer- sey, tostlomday a4 4. M. viridum Mesopleuron not punctate (for further separation of these species refer to their distributions) % 2)... M. minimum, M. wheelerorum, M. marjoriae SYSTEMATIC TREATMENT Genus Monomorium Mayr A complete synonymy is provided by Ettershank (1966). The following syn- onymy contains only names pertaining to this revision. Similarly, complete syn- onymies are not provided for each species; only references of taxonomic or biological importance are listed. Monomorium Mayr, 1855: 452. Type species: Monomorium minutum Mayr (Monobasic). Epoecus Emery, 1892: cclxxvi-cclxxvul. Type species: Epoecus pergandei Emery (Monobasic). DIAGNOSIS: Queen: Antenna 12- or 11- segmented, rarely 10-segmented, with more or less distinct club of greater than 2 segments; median clypeal setae always present; clypeus usually distinctly bicarinate, carinae sharp or rounded; propodeum never with distinct teeth, rarely angulate, usually rounded; petiole dis- tinctly pedunculate; wing venation similar to that of Solenopsis but radial cell always open; metapleural gland well developed; four Mal- pighian tubules with tips attached to rectum 76 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN (cryptonephritic); diploid chromosome number usually 22. Worker: Usually smaller than queen, remaining diagnosis same as queen, except wings lacking. Mate: Usually smaller than queen but larger than worker; Mayrian furrows lacking; mandible with teeth; antenna with 13 segments. Subgenus Monomorium Mayr Monomorium (Monomorium); Emery, 1921: 170. DIAGNOSIS: Queen: Antenna 12-segmented with distinct 3-segmented club (lengths of club segments increasing distally); propodeum un- armed (small bump may be present); clypeus with carinae usually extended anteriorly into teeth; masticatory border of mandible with 2-5 teeth, usually 4. WorKER: As in queen; prom- esonotal suture reduced or absent; metanotal sulcus present as pronounced constriction. Mate: Antennal scape cylindrical, never sur- passing occiput. Monomorium (Monomorium) minimum species group Monomorium species related to M. minutum Mayr, Emery, 1921: 171 (an part). Monomorium minutum group, Ettershank, 1966: 83 (in part). DIAGNOSIS: Aut Castes: Anepisternum and katepisternum always smooth and shining; pro- podeum with distinct parallel longitudinal rugae on side. This species group as here understood includes only those species which are native to North and Central America. Further revisionary work within this genus may reveal this to be a paraphyletic group and may show that certain South American and Asiatic species are members of this group. However, it is a group of species (some- what isolated geographically) well suited for revisionary study. Monomorium minimum (Buckley) Ficures) 6, 12), 19,.and 26-42 Myrmica (Monomarium) [sic] minima Buckley, 1867: 338. Monomorium minutum var. minimum: Emery, 18952 274-275 (in. part); Wheeler, 19028 27; Wheeler, 1908: 423 (in part). Monomorium carbonarium: Mitchell and Pierce, 1912: 70 (misidentification). Monomorium minimum: Wheeler, 1917a: 464-465, 501 (in part); Talbot, 1934: 420; Smith, 1935: 237; Smith, 1936a: 163; Cole, 1937: 99 (in; part);; Smith, 1943: 300;, Buren, 1944: 289; Gregg, 1944: 456, 466; Smith, 1947: 565; Creighton, 1950: 219-222 (in part); Smith, 1952: 810 (in part); Cole, 1953: 299 (in part); Talbot, 1953: 4; Wheeler and Wheeler, 1955: 122; Kannowski, 1956: 177; Hess, 1958: 26-27; Smith, 1958: 128 (in part); Wheeler and Wheeler, 1960a: 15; Gregg, 1963: 367-368; Wheeler and Wheeler, 1963: 133-135 (in part); Smith, 1965: 33-34 (in part); Ettershank, 1966: 90 (in part); Smith, 1967: 356 (in part); Kempf, 1972: 144: ‘Talbot, 19752 245;. Kugler, 1978: 451-452; Smith, 1979: 1382-1383 (in part); Talbot, 1979: 88; DuBois, 1981la: 33-34; DuBois, 1981b: 35; Jones et al., 1982: 287. Monomorium minutum subsp. minima: Emery, 1921: 172 (in part). Monomorium viridum peninsulatum: Creighton, 1950: 223-224 (in part), Gregg, 1963: 368-370 (misidentification); Smith, 1967: 356 (in part); Smith, 1979: 1384 (in part). Monomorium metoecus Brown and Wilson, 1957: 239-244 (in part); Wilson and Brown, 1958: 33-38 (in part); Jones et al., 1982: 287 (syn. by Brown in Ettershank, 1966: 90). DIAGNOSIS. QurEeN: Winged; petiole as in Figures 26 and 28; scutum and scutellum not depressed; metanotum (in side view) projecting to level of propodeum and _ scutellum; pro- podeum angular. MALE: Genitalia and sterna as in Figures 32 and 36-39. Worker: Propodeum angular; mesopleuron not punctate; petiole as in Figure 31; PI 33-40 (37). DESCRIPTION. Queen: HeEaAb: Measurements (representing different localities; N= 10) {Neo- type measurements in brackets} HL 0.69-0.86 (0.77) {0.79}, HW 0.65-0.81 (0.74) {0.66}, SL 0.40-0.60 (0.50) {0.58}, IOD 0.12-0.20 (0.17) {0.17}, OD 0.04-0.06 (0.05) {0.06}, EL 0.15-0.20 (0.17) {0.20}, MOD 0.11-0.18 (0.13) {0.15}. Structure—Head slightly longer than broad or slightly broader than long, CI 84-106 (95) {84}, distinctly longer than scape, SIL 56-79 (65) {73}; SIW 53-88 (68) {88}. In full frontal view, head broadest slightly above eyes; side convex; occiput rounded laterally, summit flat. Eye moderate in size. Scape reaching but never surpassing occiput. Mandible with 4 teeth; maxillary palp 2-segmented; labial palp 2-segmented. Clypeal teeth sharp, moderate in length. Frontal carinae diverging slightly poste- riorly. Pilosity—Setae erect to suberect on clypeus, frons, gular region, mandible, and occiput, decumbent to appressed elsewhere. ANT GENUS MONOMORIUM 77 39 34 ; 1 FIGS. 26-39. Monomorium minimum. 26—Queen, lateral view. 27—Head of queen, frontal view. 28— Petiole of queen, posterior view. 29—Postpetiole of queen, posterior view. 30—Head of worker, frontal view. 31—Worker, lateral view. 32—Male, lateral view. 33—Labial palp of male. 34—Maxilla and maxillary palp of male. 35—Mandible of male. 36—Volsella of male. 37—Aedeagus of male. 38— Eighth sternite of male. 39—Ninth sternite of male. Scales: Top scale (0.1 mm) for Figure 35. Second scale (1 mm) for Figures 26-32. Third scale (0.5 mm) for Figures 36 and 37. Bottom scale (0.1 mm) for Figures 33 and 34. 78 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Setae of scape suberect to decumbent, of pedicel and flagellum decumbent to appressed with percentage of latter increasing distally. Sculp- ture—Head smooth and shining with small to moderate piliferous punctures evenly distrib- uted except as follows: small, parallel, longitu- dinal rugosities beginning all along lateral margin of clypeus, extending past antennal insertion, converging with rugosities from fron- tal carinae and extending to level of anterior ocellus. Other small, parallel, longitudinal rugosities beginning between frontal carinae, sometimes extending to level of anterior ocellus (Fig 27). ALITRUNK: Measurements PW 0.50-0.72 (0.61) {0.62}, PL 0.31-0.42 (0.38) {0.40}, WL 1.34-1.81 (1.61) {1.37}. Structure—Mesonotum usually lacks notal furrow )rarely, with small emargination¢. Pronotal-scutal suture on dorsal 1/3 of alitrunk. Scutum and _ scutellum not depressed. Mesopleural suture straight or de- flected ventrally on posterior end and deflected dorsally on anterior end (resulting in S-shaped suture). Small pit usually present on posterior end of suture. Metanotum (in lateral view) reaching or exceeding level of propodeum and scutellum. Propodeum angular, basal face ap- proximately 2/3 length of declivitous face. PI 21-29 (24) {29}. Wings present; venation typ- ical (Fig. 71); 6-8 hamuli on hindwing; stigma located directly above vannal emargination {neotype lacks wings but condition of basal sclerites indicate queen once possessed wings}. Pilosity—Dorsal outline of alitrunk with more than 30 erect to suberect setae. Setae erect to suberect on coxae and trochanters, erect to suberect on flexor surfaces of femora (decum- bent to appressed elsewhere), decumbent to appressed on tibiae and tarsi. Scu/pture—Smooth and shining except as follows: moderate to large piliferous punctures on dorsum of alitrunk; lower 1/2 of side of propodeum (below spiracle) with several moderate, parallel, longitudinal rugae )rarely, some rugae occur above spira- cle¢; sculpturing of sutures as in Fig. 26. PeTiove. As in Figures 26 and 28. Dorsum of node convex to flat. Subpetiolar process moder- ate in size, anterior to node. Setae decumbent to appressed on anterior surface, erect to suberect on dorsum, posterior surface, and side, absent from venter. Dorsum of node smooth and shin- ing with moderate piliferous punctures; side with moderate, parallel, longitudinal rugae be- coming transverse near posterior surface of node; posterior surface of node with moderate, semicircular, concentric rugae. POSTPETIOLE: As in Figures 26 and 29. Dorsum of node flat to shghtly emarginate. Anterior subpostpetiolar process reduced, located medially on venter. Setae appressed to decumbent on anterior sur- face of node, suberect to erect on remaining surfaces. Dorsum of node smooth and shining with moderate to small piliferous punctures; remainder of node covered with dense, moder- ate to small, non-piliferous punctures. GasTeER: Setae erect to decumbent on all surfaces with percentage of erect to suberect setae increasing towards posterior end of gaster. Setae of first gastral tergite exceeding level of dorsum of postpetiolar node. All surfaces smooth and shin- ing with small to moderate piliferous punctures. Cotor: Head brown to dark brown with mandi- ble and lateral margin of clypeus yellow brown to brown; scape brown, pedicel and flagellum yellow brown to yellow. Alitrunk brown to dark brown with legs brown; tibiae and tarsi yellow brown to yellow. Petiole, postpetiole, and gaster brown to dark brown. All setae white. Mate: Heap: Measurements (representing dif- ferent localities; N=10) HL 0.65-0.74 (0.71), HiW 0:69-0.78' (0.75); “SE* 0:22-0:30° (0.26); IOD 0:22-0.26 (0.24), OD 0.05-0712 (0:09), EL 0.28-0.34 (0.32), MOD 0.15-0.24 (0.19). Struc- ture—Head slightly broader than long, CI 105-109 (107), distinctly longer than scape, SIL 31-42 (36); SIW 29-38 (34). In full frontal view, head broadest at or slightly above eyes; side straight to slightly convex (usually obscured by compound eye); occiput rounded laterally, su >- mit flat (usually obscured by posterior ocelli). Eye large. Scape not reaching occiput. Mandi- ble with 3 teeth; maxillary palp 2-segmented; labial palp 2-segmented. Clypeal teeth reduced or absent (blunt if present). Frontal carinae diverging slightly posteriorly. Prlosity—Setae erect to decumbent over entire surface of head with erect to suberect setae predominating near clypeus, gular region, mandible, and occiput. Antennal pilosity as in queen. Sculpture—Entire head covered with dense, moderate to large, non-piliferous punctures. Small to moderate, parallel, longitudinal rugae beginning all along lateral margin of clypeus, continuing to level of antennal insertion and fusing with large, con- centric, semicircular rugae surrounding anten- nal insertion. ALITRUNK: Measurements PW 0.:70-0:81 (0:76); PL. 0:44-0:62" (0.52), WE 1.66-1.90 (1.75). Structure—Mesonotum lacking notal furrow. Pronotal-scutal suture on dorsal 1/3 of alitrunk. Scutum and _ scutellum not depressed. Mesopleural suture deflected ven- trally at posterior end and deflected dorsally at anterior end (resulting in an S-shaped suture) with small pit on posterior end. Metanotum (in lateral view) reaching or exceeding level of propodeum and _ scutellum. Propodeum an- gular, basal face 2% as long as declivitous face. Ant GENUS MonomMorIUM Wings present; venation typical; 5-7 hamul on hindwing; stigma located directly above vannal emargination. Pilosity—Dorsal outline of al- itrunk with more than 30 erect to suberect setae; setae absent from side (except near sutures). Leg pilosity as in queen. Sculpture—Entire sur- face smooth and shining except as follows: small to moderate piliferous punctures on dorsum of alitrunk; small to moderate non-piliferous punc- tures loosely distributed on side; lower 1/2 of side of propodeum (below spiracle) with small, parallel, longitudinal rugae; entire surface of propodeum often covered with small, non-pil- iferous punctures (Fig. 32). Perrote: As in Figure 32. Dorsum of node flat to emarginate. Subpetiolar process reduced, anterior to node. Setae erect to decumbent on anterior surface of node, erect to suberect on side and posterior surface of node, absent elsewhere (including dorsum of node). All surfaces smooth and shin- ing except as follows: small to moderate pil- iferous punctures on anterior and_ posterior surfaces of node; side with small to moderate non-piliferous punctures. PosTPETIOLE: As in Figure 32. Dorsum of node flat to slightly emarginate. Anterior subpostpetiolar process absent. Setae erect to decumbent on anterior surface, side, and posterior surface, absent else- where. Dorsum of node smooth and shining; remainder of node covered with moderately aense, non-piliferous punctures. GASTER: Pilosity as in queen. All surfaces smooth and shining with small to moderate piliferous punc- tures. GeniTALiA: As in Figures 32 and 36-39. Eighth sternite with emergination lacking setae, approximately as deep as wide (Fig. 38). Ninth sternite with 6-8 erect setae (Fig. 39). Aedeagus with 12-15 teeth; toothed margin rounded (Fig. 37). Volsella with curved digitus and reduced cuspis; cuspis with 3 setae (Fig. 36). Cozor: Head brown to dark brown, mandible and lateral margin of clypeus yellow brown to yel- low. Alitrunk brown to yellow brown; leg color- ation as in queen. Petiole, postpetiole, and gaster brown to dark brown. Genitalia brown to yellow brown. All setae white. Worker. Heap: Measurements (representing different localities; N= 10) HL 0.46-0.56 (0.52), HW 0.38-0.48 (0.42), SL 0.28-0.40 (0.33), EL 0.06-0.10 (0.09), MOD 0.05-0.08 (0.06). Struc- ture—Head a little longer than broad, CI 76-86 (81), distinctly longer than scape SIL 56-71 (64); SIW 71-90 (79). In full frontal view, head broadest at or slightly above eyes; side convex; occiput rounded laterally, summit flat. Eye small in size. Scape never reaching occiput. Mandible with 4 teeth; maxillary palp 2-seg- ~I Oo mented; labial palp 2-segmented. Clypeal teeth sharp, of moderate length. Frontal carinae di- verging slightly posteriorly )rarely, parallel<. Pilosity—Setae suberect to subdecumbent on clypeus, frons, gular region, mandible, and occiput, decumbent to appressed elsewhere. Setae of scape subdecumbent to decumbent, of pedicel and flagellum decumbent to appressed. Sculpture—All surfaces smooth and shining with small to moderate piliferous punctures. ALITRUNK: Measurements PW 0.22-0.32 (0.28), PE, 0:18-0:25 (0.21), WiLL) 0250-05655 (0257). Structure—Mesonotum lacking notal furrow. An- terior propodeal suture of moderate depth. Pro- podeum angular, basal face 1.5-2X as long as declivitous face. PI 33-40 (37). Pilosity—Dorsal outline of alitrunk with more than 10 erect to suberect setae. Leg pilosity as in queen. Sculpture—Entire surface smooth and _ shining except as follows: small to moderate piliferous punctures on dorsum; lower 1/2 of side of propodeum (below spiracle) with 1-3 small, parallel, longitudinal rugae. Pe7vroze: As in Figure 31. Dorsum of node convex. Subpetiolar process of moderate to reduced size, anterior to node. Setae erect to suberect on dorsum, absent elsewhere. All surfaces smooth and shining with small piliferous punctures on dorsum. PosTreTIoLe: As in Figure 31. Dorsum of node convex. Anterior subpostpetiolar process re- duced, located medially on venter. Setae sub- erect to erect on dorsum, absent elsewhere. All surfaces smooth and shining with small pil- iferous punctures on dorsum. Gas7TER: Setae erect to suberect with percentage of erect setae increasing towards posterior end of gaster. Setae of first gastral tergite not exceeding level of dorsum of postpetiolar node. All surfaces smooth and shining with small to moderate piliferous punctures evenly distributed. Cozor: Head brown to dark brown with trace of bluish reflection, mandible yellow brown to yellow. Alitrunk brown to dark brown with trace of bluish reflection, legs yellow brown with tibiae and tarsi yellow. Petiole, postpetiole, and gaster brown. All setae white. TYPE MATERIAL. None known to exist (Creighton, 1950). Wheeler (1902) stated the probable type locality as Austin, Texas (or vicinity) since Buckley (1867) failed to mention a specific locality. Although a few of Buckley’s specimens remain (Academy of Natural Sciences, Philadelphia and col- lection of Gustav Mayr, Vienna Museum of Natural History, Vienna, Austria), none could be located that were collected in the 80 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN 0.10 0.15 0.20 FIGS. 40-41. Monomorium minimum. 40—Dia- grammatic lateral view of nest. Darkened circles represent tunnels leading away from viewer. Cross-hatched circles represent tunnels leading toward viewer. Stippled chambers represent chambers containing queens and brood. Scale=2 mm. 41—Histogram of larval head widths in one nest. Interval is 0.10 mm. 1860’s or earlier which might have pro- vided the basis for his description. Since two species of Monomorium, M. cyaneum and M. minimum, could occur in the Austin, Texas, vicinity, I designate the following queen [MCZ] as neotype of Monomorium minimum to reduce confusion: TEXAS: Bastrop Co., Bastrop State Park, June 9, 1954, W. Clayd T-119. Since Buckley failed to mention a specific locality in his original description, I followed Wheeler’s suggestion in designating a neo- type from the Austin, Texas, vicinity. This specimen bears a red, handwritten label: Monomorium/ minimum/ (Buckley)/ Neo- type/ M. DuBois 1983/. DISTRIBUTION. This species ranges from Pennsylvania and District of Colum- bia, south to Georgia, west to Texas and New Mexico, and north to Idaho, Mon- tana, and North Dakota (Fig. 42). Al- though this species has been collected in northern Wisconsin and North Dakota, no collections have been made in southern Canada; this species undoubtedly occurs there. In the eastern United States, this species occurs sporadically and is appar- ently replaced by M. emarginatum in the northeastern states and by M. viridum along the Atlantic coast in New Jersey, Georgia, and Florida. In the southern United States, M. minimum is known from collections in northern Georgia (Athens), Alabama (northern Alabama south to Mobile), Mis- sissippi (only two collections, Amory and State College), and numerous collections in Louisiana and Texas. This species overlaps the range of M. cyaneum in Texas and New Mexico. Monomorium cyaneum apparently prefers drier habitats and usually nests under rocks while M. minimum prefers moisture habitats usually near the edges of woods. Most of my records of M. minimum from Colorado are from the eastern plains. I have only examined one worker which was collected from western Colorado (Mesa Verde National Park). However, Gregg (1963) includes many localities from the mountainous regions of the state (his M. viridum peninsulatum records should be included with his records for M. minimum). The furthest western records for MM. mini- mum are three collections from southern Idaho (Rupert, 6.4 km W of St. Anthony, and Holbrook). These three collections consist of workers only. It is possible these are M. wheelerorum although they appear to be M. minimum workers. A detailed list of all localities for this and other Monomorium species 1s provided in DuBois (1981c). DISCUSSION. BioLtocy. Recorded as nesting in soil, under rocks, in logs, man- made structures, and living trees (mes- quite, Prosopis glandulosa) (Dennis, 1938; Buren, 1944; Kannowski, 1956; Hess, 1958; Wheeler and Wheeler, 1963). Talbot (1934) and Gregg (1944) found M. mini- mum most frequently in sandy soil, and Ant GENUS MONOMORIUM 81 FIG. 42 Distribution of Monomorium minimum (closed circles). Gregg (1963) discovered it in a number of microhabitats representing both Upper Sonoran and Transition life zones: foothills meadow, deciduous canyon forest, oak woodland, ponderosa pine-oak woodland, pinyon-cedar woodland, pinyon-cedar-oak woodland, mixed grass prairie, short grass prairie, sagebrush desert, sagebrush- greasewood desert, sandhills grassland, roadside, and exposed bedrock. Nests in soil appear to have a character- istic structure; most are shallow (less than 10 cm deep, Kannowski, 1956) with most brood chambers located from just under the soil surface to depths of 5 cm (Wheeler and Wheeler, 1963). Although this descrip- tion referred to nests in North Dakota, nests in Kansas and Illinois are of a similar internal construction (Fig. 40). However, tumuli vary considerably depending upon environmental conditions. In Kansas, most nests are located near edges of woodlands and have tumuli with a typical crater struc- ture such as that depicted in Figure 40. Nests in areas of scant vegetation usually lack conspicuous tumuli, since soil particles are presumably blown away as soon as they are deposited on the surface. Nests in sheltered areas, such as at the base of grass clumps, possess mounds with many cre- nulations causing the structure to resemble a lump of brain coral. In addition to the wide environmental tolerance of this species, there 1s a wide altitudinal tolerance. In Kansas, collection elevations range from 244 m to over 1220 m (pers. observ.). In Colorado, Gregg (1963) found collection elevations to range from 1067 m through 2591 m. Similar elevational extremes were found in Ten- nessee by Dennis (1938). This species also occurs down to sea level. Monomorium minimum queens live ap- proximately 1 year in laboratory colonies while workers live approximately 4 months. There appear to be three larval instars (Fig. 41) (pers. observ.). Most nests contain multiple queens. For 82 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN example, Gregg (1944) found that nests near Chicago, Illinois usually contained 12-14 queens and Dennis (1938) found a similar number of queens per nest in Ten- nessee. Presumably these are functioning queens since most are associated with small, separate piles of eggs and first instar larvae (pers. observ.). Recent preliminary allozyme studies have demonstrated that several queens produce eggs at the same time in nests near Lawrence, Kansas (pers. observ.). Although it is not known how many queens establish a nest, most nests are probably established by one or two queens since a number of small nests with a small number of offspring and one or two queens are found each autumn in the Law- rence, Kansas, vicinity. Since nuptial flights occur in July in this vicinity, most newly mated queens should have new nests established by autumn. Production of sexuals occurs in late spring and summer; most nests in Kansas contain larvae which will develop into sex- uals by late May. Most sexual adults eclose by mid-July. Nuptual flights have not been observed, although they probably occur, since both males and gynes are winged. Monomorium minimum is one of two spe- cies reported to have inquilines (M. ergatogyna is the other species). Van Pelt and Van Pelt (1972) reported larvae of Microdon baliopterus Loew (Diptera: Syrphidae) from nests of M. minimum in Big Bend National Park, Texas. Monomorium minimum nests have been discovered in close association with nests of other ant species. Wheeler and Wheeler (1963) found M. minimum associated with nests of the following species: Acanthomyops claviger (Roger), Camponotus vicinus Mayr, Formica fusca Linnaeus, Lasius crypticus Wilson, and Pogonomyrmex occidentalis (Cresson). I have found nests of M. mini- mum associated with nests of an additional species, Solenopsis (Diplorhoptrum) molesta (Say), in Kansas. Comparisons. Since Monomorium mint- mum queens are winged, they can easily be separated from other species occurring in the same area, most of which have wingless queens. Only M. viridum queens are winged. Nests of M. viridum appear to be restricted to pure sand while nests of M. minimum only occur in soils containing some clay (W. L. Brown pers. observ.). Workers of this species can usually be separated by the combination of characters listed in the diagnosis, particularly the smooth and shining mesopleuron. Monomorium cyaneum Wheeler Figures 10, 13, and 43-57 Monomorium minimum: Wheeler, 1906: 332, 336 (misidentification); Wheeler, 1908: 423 (in part); Smith, 1936a: 163 (in part); Cole, 1937: 99 (in part); Creighton, 1950: 219 (in part); Cole, 1953: 299 (in part); Smith, 1958: 128 (in part); Ettershank, 1966: 90 (in part); Smith, 1967: 356 (in part); Hunt and Snell- ing, 1975: 21; Smith, 1979: 1382-1383 (in part). Monomorium minimum subsp. cyaneum Wheeler, 1914b: 43; Wheeler, 1917a: 465, 501; Kempf, 1972: 144. Monomorium minutum subsp. minima var. cyanea: Emery, 1921: 173. Monomorium minimum subsp. emersoni Gregg, 1945: 66-67; Smith, 1952: 810 (in part); NEW SYNONYMY. Monomorium cyaneum: Ettershank, 1966: 88; Du- Bois, 1981a: 35-36; Jones et al., 1982: 287. Monomorium near emersoni: Jones et al., 1982: 287. DIAGNOSIS. Queen: Wingless; scutum and scutellum not depressed; metanotum (in lateral view) projecting to level of propodeum and scutellum; propodeum angular; petiole as in Figures 43 and 45; head sculpturing as in Figure 44. Mare: Genitalia and sterna as in Figures 49, 52-55. Worker: Propodeum angular; meso- pleuron punctate; petiole as in Figure 48; PI 29-38: (34): DESCRIPTION. As described for M. mini- mum except as indicated. QuEEN: Heap: Measurements (representing different localities; N=10) HL 0.68-0.88 (0.76), HW 0.62-0.78 (0.68), SE. 0.50-0:55' (0.52), TOD“ 0813-0419 (0.18), OD 0.04-0.05 (0.05), EL 0.16-0.20 (0.18), MOD 0.12-0.16 (0.13). Structure—Cl 83-96 (90), SIL 62-76 (69), SIW 70-84 (77). Scape reaching or barely surpassing occiput. Pilosity—Setae suberect to subdecumbent near clypeus, frons, gular region, mandible, and occiput, decumbent to appressed elsewhere. ALITRUNK: Measurements PW 0.41-0.48 (0.44), Pi0:28-0:32 (0.30), Wi 1228-1538" G32). AnT GENUS MONOMORIUM 83 Se 0.5 ce | ; 55 0.5 EO ET ORNS, 34 0.1 FIGS. 43-56. Monomorium cyaneum. 43—Queen, lateral view. 44—Head of queen, frontal view. 45— Petiole of queen, posterior view. 46—Postpetiole of queen, posterior view. 47— Head of worker, frontal view. 48—Worker, lateral view. 49—Male, lateral view. 50—Labial palp of male. 51—Maxilla and maxillary palp of male. 52—Eighth sternite of male. 53—Ninth sternite of male. 54—Volsella of male. 55—Aedeagus of male. 56—Mandible of male. Scales: Top scale (1 mm) for Figures 43-49. Second scale (0.1 mm) for Figure 56. Third scale (0.5 mm) for Figures 52 and 53. Fourth scale (0.5 mm) for Figures 54 and 55. Bottom scale (0.1 mm) for Figures 50 and 51. 84 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Structure—Mesonotum lacking notal furrow. Mesopleural suture straight with no pits at either end. Propodeum angular, basal face half as long as declivitous face. PI 22-25 (23). Wings absent (fusion of sclerites and presence of callow queens in some nests indicate queens are wing- less). Prlosity—Setae suberect to decumbent on side of alitrunk. Sculpture—Small to moderate piliferous punctures evenly distributed on al- itrunk (moderate punctures predominating dor- sally). Pe7vrore: As in Figures 43 and 45: dorsum of node flat. Subpetiolar process re- duced. Peduncle with dense, small, non- piliferous punctures; remainder of petiole smooth and shining with small to moderate piliferous punctures evenly distributed on ante- rior surface, dorsum, posterior surface, and sides. PosTpeTIOLE: As in Figures 43 and 46: dorsum of node flat. Entire surface covered with dense, small to moderate, nonpiliferous punc- tures. GaSTER: Setae of first gastral tergite not exceeding level of dorsum of postpetiolar node. Cotor: Mandible yellow to yellow brown. Al- itrunk brown, legs yellow brown. Mate: Heap: Measurements (representing 1 locality, N=1) HL 0.68, HW 0.72, SL 0.26, lOD=0.24- OD: 0208; EL 0:30, MOD 0.17. Structure—CI 106, SIL 38, SIW 36. Sculpture— Occiput with several moderate to large, con- centric, semicircular rugae. ALITRUNK: Measurements PW 0.75, PL 0.49, WL 1.72. Structure—Mesopleural suture straight (pits lack- ing on both ends). Metanotum (in lateral view) not reaching level of propodeum and scutellum. Pilosity—Dorsal outline of alitrunk with fewer than 30 erect to suberect setae projecting above outline. Sculpture—Scutum with large, con- centric, semicircular rugae. PeTIoLE: As in Figure 49. Dorsum of node flat. Setae erect to subdecumbent on dorsum of node. Entire sur- face of node covered with small, non-piliferous punctures. PosTPETIOLE: As in Figure 49. Ante- rior subpostpetiolar process reduced. Entire surface of node covered with small, non-pil- iferous punctures. GasTeR: As in Figures 49, and 52-55. Ninth sternite with 11 setae (Fig. 53). Aedeagus with 18 teeth (Fig. 55). Cuspis of volsella with 5 setae (Fig. 54). Cozor: As in queen. Genitalia yellow brown. Worker. Heap: Measurements (representing different localities; N= 10) HL 0.48-0.60 (0.51), HW 0:35-0:42 (0.39), SL 0.30-0.45 (0.35), EL 0.08-0.10 (0.09), MOD 0.05-0.06 (0.05). Struc- ture—CI 70-83 (78), SIL 60-80 (70), SIW 77-107 (90). Pilostty—Setae erect to suberect on clypeus, frons, gular region, mandible, and occiput, decumbent to appressed on remainder. ALITRUNK: Measurements PW 0.20-0.25 (0.22), PL 0: 14-0522 (0.18), WE 0-45-0:62) (0:54). Structure—Propodeum angular, basal and de- clivitous faces of approximately equal length. Pilositty—Dorsal outline of alitrunk with 4-8 erect to suberect setae. Sculpture—Mesopleuron covered with small, dense, non-piliferous punc- tures (leading to a granular appearance); lower 1/3 of side of propodeum (below spiracle) with moderate, parallel, longitudinal rugae. PI 29-38 (34). Peviove: As in Figure 48. Dorsum of node flat to slightly convex. PosTPETIOLE: As in Figure 48. Dorsum of node flat to slightly convex. Anterior subpetiolar process absent. TYPE MATERIAL. Syntypic series con- sisting of ‘‘numerous workers and fe- malés’* (Wheeler, 1914b):- Mexico: HipaLco: Guerrero Mill [spring or sum- mer, 1913], W. M. Mann (4 queens, 40 workers; NMNH). Lectotype queen here designated from NMNH series bears red, handwritten label: Monomorium/cyaneum/ Wheeler/ Lectotype/ M. DuBois 1983/. Synonymous M. emersoni type series con- sists of “1 queen and 79 workers’’ (Gregg, 1945): Unirep STATES: ARIZONA: Gila Co. ; Globe, April, 1937, A. E.. Emerson. Para- type queen and workers: TEXAS. Travis Co. ; Austin, A. E. Emerson [paratype queen and 51 workers; NMNH, examined]. Hol- otype presumably in collection of R. Gregg (Gregg, 1945). DISTRIBUTION. This species ranges from central Mexico (Mexican Plateau), particularly Hidalgo, Mexico, and Queretaro, east to western Vera Cruz, and west to Nayarit. These localities appear to represent the southern extent of this spe- cies. It occurs sporadically northward to southern Texas, New Mexico, and Arizona (Fig. 57). In eastern Texas this species is apparently replaced by M. minimum which seems to prefer moister habitats than A. cyaneum. The furthest eastern collections are from the San Antonio vicinity. The northernmost extensions of M. cyaneum oc- cur in northern Arizona (numerous collec- tions from Grand Canyon vicinity). This species has been collected as far west as the Hualapai Mountains near Kingman, Ari- zona. Further west, M. cyaneum is replaced by M. ergatogyna, while further north, M. wheelerorum predominates. It is possible that Ant GENUS MONOMORIUM 85 FIG. 57. Distribution of Monomorium cyaneum (closed triangles). M. cyaneum may someday be discovered in southern Utah and Nevada since the hab- itat is not significantly different from that of northern Arizona. It appears that while part of the sporadic distribution of this species is due to lack of collections, habitat restriction also plays a part. Most of the localities collected in Arizona and New Mexico represent upland areas (usually canyons within mountain ranges). Thus many populations may be rather isolated. At the southern extents of the range of M. cyaneum, most collections are from the Mex- ican Plateau. Although there are two col- lections from the state of Vera Cruz, both are from upland areas along the Mexican Plateau. Most collections from the plateau were near Mexico City (from such nearby states as Hidalgo and Queretaro). Collec- tions from the western coast of Mexico are spotty. Only one collection has been made on an island (Maria Cleofas Islas, Nayarit). Three collections have been made in the region of Guaymas, Sonora and one collec- tion in the Baja California peninsula (21 km N of La Paz). DISCUSSION. BioLtocy. Wheeler (1914b) indicated this species nests under ‘ stones in ‘‘rather damp places on the sides of canyons.’’ I have observed this to be the case with colonies of this species I have collected in New Mexico and Arizona. For example, one nest of this species (ARIZONA: Gila Co.) was located in such a habitat on the south facing slope of a small canyon. This nest was concealed under a stone measuring approximately 0.1m?. Most of the nest occupied an area about 10 cm in diameter, about 5 cm from the east side of the stone (a populous colony of Iridomyrmex pruinosus (Roger) occupied the western por- tion under this stone). The portion of the M. cyaneum nest which was visible after the stone was removed consisted of a series of small chambers (1 cm in diameter) con- nected by tunnels (3 mm in diameter and 0.5-2 cm in length). The topmost layer of chambers was broken open when the stone was lifted. Most brood was near the sur- face, with different life stages clustered in separate chambers. Although this nest could only be excavated to a depth of about 7.5-10 cm due to large buried stones, I believe I collected the majority of individu- als. Colonies of M. cyaneum have multiple queens, although it is not known whether they are all functional. However, in nests I excavated, each queen was in a separate chamber and surrounded by eggs and first or second instar larvae. Dates on which sexual forms are released are unknown, although the Gila County (Arizona) colony contained many callow queens and developing male pupae on June ZITO? Monomorium cyaneum has a moderately wide altitudinal range (from 1387 m through 2286 m elevation in Arizona and from 1341 m through 2743 m {type lo- cality} elevation in Mexico). Systematics. Although Gregg (1945) believed M. emersoni to be a separate taxon (subspecies), he failed to compare it with M. cyaneum. The most prominent feature used to distinguish MM. emersont was the prominent tubercles at the point where basal and declivitous faces of the pro- podeum meet. Although this seems to be a good character upon superficial examina- 86 Tue UNIVERSITY OF KANSAS SCIENCE BULLETIN tion, one colony (Santa Fe, New Mexico) contained numerous queens which ranged in morphology from the typical M. cyaneum propodeum to one with tubercles nearly developed into teeth. Since all other char- acters indicated this was a queen of M. cyaneum, M. emersoni was synonymized; none of the other characters used by Gregg differed from characters found in M. cyaneum. ComPaRISON. Since Monomorium cyaneum has wingless queens, the following species which occur in or near its range might be confused with it: M. compressum, M. ergatogyna, and M. wheelerorum. Monomorium cyaneum queens can easily be distinguished from M. compressum since the latter species has a depressed scutum and scutellum, an emarginate postpetiole, and fewer than 30 erect to suberect setae projecting above the dorsal profile of the alitrunk. Monomorium cyaneum queens differ from M. ergatogyna queens since the latter species has a fringe of erect to suberect setae on the anterior edge of the scape and less facial sculpturing than M. cyaneum (Fig. 34). Monomorium cyaneum queens are easily distinguished from M. wheelerorum queens since the latter species has an antennal fringe similar to that of M. ergatogyna queens, a flat to slightly depressed scutum and scutellum, and a notal furrow on the mesonotum. Since the ranges of many species overlap with M. cyaneum, it is best to use the keys to separate workers which are not associated with queens. The following species are most likely to be collected within the range of M. cyaneum: M. compressum, M. ebeninum, M. ergatogyna, M. marjoriae, M. minimum, and M. wheelerorum. Monomorium cyaneum workers can easily be separated from M. compressum and M. ebeninum workers since the latter two species have a nonpunctate mesopleuron and more than 10 erect to suberect setae projecting from the dorsum of the alitrunk. Monomorium cyaneum work- ers can be separated from M. ergatogyna workers since the latter species has a more rounded propodeum and occurs mainly along the California coast. Monomorium cyaneum workers can be separated from workers of M. marjoriae, M. minimum, and M. wheelerorum since the latter three species have a nonpunctate mesopleuron and the basal face of the propodeum is longer than the declivitous face. Monomorium ergatogyna Wheeler Figures 1-5, and 58-80 Monomorium minutum: Brues, 1903: 148 (misi- dentification); Wheeler, 1910: 386 (misiden- tification). Monomorium minimum ergatogyna Wheeler, 1904: 269: Wheeler, 1905a: 88; Wheeler, 1917b: 464, 501; Tulloch, 1930: 202; Mallis, 1941: 71 (in part); Smith, 1943: 300; Smith, 1947: 565; Smith, 1952: 810; Cook, 1953: 160-163 (in part). Monomorium minutum subsp. minima var. ergatogyna: Emery, 1921: 173. Monomorium minimum: Essig, 1926: 857 (in part); Mallis, 1941: 71 (in part); Creighton, 1950: 219 (in part); Cook, 1953: 160-163 (in part); Smith, 1958: 128 (in part); Ettershank, 1966: 90 (in part); Smith, 1967: 356 (in part); Smith, 1979: 1382-1383 (in part); Snelling and George, (unpubl.). Monomorium ergatogyna: Essig, 1926: 857. DIAGNOSIS. Queen: Wingless; head (ex- cluding lateral margin of clypeus) lacking non- piliferous punctures and rugae; alitrunk with more than 30 erect to suberect setae projecting above dorsal outline; petiole and postpetiole as in Figures 58, 60 and 61; antennal pilosity as in Figure 59. Mate: Genitalia and sterna as in Figures 65, 69, 70, and 72-78. Worker: Pro- podeum, petiole and postpetiole as in Figure 63; mesopleuron punctate; PI 35-44 (38). DESCRIPTION. As described for M. cyaneum except as indicated. QUEEN: Heap: Measurements (representing different localities; N= 10). HL 0.59-0.71 (0.67), HW 0.55-0.65 (0.61), SL 0.41-0.52 (0.47), IOD 0.12-0.18 (0.17), OD 0.04-0.05 (0.05), EL 0.12-0.15 (0.14), MOD 0.09-0.11 (0.10). Structure—CI 79-95 (90), SIL 58-77 (70), SIW 63-91 (77). Side straight to slightly convex. Frontal carinae diverging strongly posteriorly. Prlosity—Setae erect near clypeus, frons, gula, occiput, and ocelli, sub- erect to decumbent on malar area, appressed on remainder. Setae of scape erect on anterior edge (forming a fringe of setae, Figure 59), decum- bent to appressed on remainder, of pedicel and flagellum (except club) suberect to decumbent, of antennal club appressed. Scu/pture—Smooth and shining except as follows: moderate, paral- lel, longitudinal rugae beginning all along lat- eral margin of clypeus and extending to level of Ant GENUS MONOMORIUM 87 0.1 0.1 FIGS. 58-70. Monomorium ergatogyna. 58—Queen, lateral view. 59—Head of queen, frontal view. 60— Petiole of queen, posterior view. 61—Postpetiole of queen, posterior view. 62—Head of worker, frontal view. 63—Worker, lateral view. 64—Head of male, frontal view. 65—Male, lateral view. 66—Labial palp of male. 67—Maxilla and maxillary palp of male. 68—Mandible of male. 69—Male genitalia (7m situ), dorsal view. 70—Male genitalia (2m sztw), ventral view. Scales: Top left (1 mm) for Figures 58-65, and 69-70. Lower left (0.1 mm) for Figure 68. Right (0.1 mm) for Figures 66 and 67. 88 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN antennal insertion. Small piliferous punctures evenly distributed on head. AL/TRUNK: Measurements PW 0.40-0.48 (0.43), PL 0.21-0.31 (0.28), WL 1.00-1.15 (1.06). Struc- ture—Propodeum evenly rounded (basal and declivitous faces do not meet at an angle). PI 19-28 (26). Pilosity—As in M. minimum. Sculp- ture—Large piliferous punctures on dorsum of scutum, smaller piliferous punctures on dorsum of scutellum, propodeum and side of alitrunk. Several moderate to large semicircular rugae anterior to propodeal spiracle. Pe7roLe: As in Figures 58 and 60. Dorsum of node convex. Setae decumbent to appressed on anterior sur- face of node, erect on dorsum and side of node, and absent from posterior surface of node. Posterior surface of node with large, transverse, concentric, semicircular rugae which extend over half of side of node. PosTpeTioLe: As in Figures 58 and 61. Dorsum of node convex. Dorsum and sides of node smooth with pil- iferous punctures. Posterior 1/5 of node with some non-piliferous punctures. )Sometimes, lower 2/3 of side of node covered with dense, non-piliferous punctures¢. Cozor: Head brown except as follows: antenna, mandible, and lat- eral margin of clypeus yellow to yellow brown. Mate: Heap: Measurements (representing 1 nests. eV = 5) Hb 0.55-0258' (0.56), AW 0.66-0:71 (0:69), SL 0.29-0.35 (0.31), IOD 0.18-0.20 (0.19), OD 0.06 (0.06), EL 0.24-0.26 (0.25), MOD 0.12-0.16 (0.15). Structure—CI W14-127.(123)5 SIL 53-60'(55), SIW. 41-53 (45). Sculpture—Large, concentric, transverse rugae occurring on posterior 1/3 of head. Large rugae beginning all along lateral margin of clypeus, continuing behind antennal insertion and con- verging with frontal carina (Fig. 64). )Occa- sionally, several moderate rugae may converge on anterior ocellus (Fig. 64)¢. ALITRUNK: Measurements PW 0.49-0.56 (0.51), PL 0.38-0.42 (0.40), WL 1.28-1.29 (1.29). Structure—Pro- notal-scutal suture located medially on alitrunk. Propodeum angular (Fig. 65), basal and de- clivitous faces of approximately equal length. PI 29-33 (31). Wings present (Fig. 71); 5-7 hamuli on hindwing. Pilosity—As in M. minimum. Sculp- ture—Anterior edge of mesopleuron, near pro- coxa, with series of small transverse rugae. Sculpturing of sutures as in Figure 65. PETIOLE: As in Figure 65. Dorsum of node convex. Setae subdecumbent to appressed on anterior surface of node, suberect to erect on dorsum, absent from posterior surface of node. Side of petiole with small non-piliferous punctures; dorsum smooth and shining with a few piliferous punc- tures. PosTPETIOLE: As in Figure 65. Dorsum of node convex. Anterior subpostpetiolar process of moderate size, located medially on venter. Setae erect to suberect on dorsum and/or poste- rior surface of node; absent elsewhere. Dorsum of node smooth, remaining surfaces with dense non-piliferous punctures. Geni74L14: As in Fig- ures 65, 69, 70, and 72-78. Ninth sternite with 7-10 erect setae (Fig. 75). Aedeagus with 13 teeth (Fig. 72). Cuspis of volsella with 3 setae (Fig...73); Worker: Heap. Measurements (representing separate localities; N= 10) HL 0.40-0.60 (0.49), HW 0.33-0.42 (0:39), SL 0.31-0:40) (0.35), EL 0.06-0.10 (0.07), MOD 0.04-0.06 (0.05). Struc- ture—CI 70-91 (79), SIL 62-85 (71), SIW 79-98 (89). In full frontal view, side straight. Scape reaching or surpassing occiput by less than length of pedicel. AL7TRUNK: Measurements PW 0.20-0.28 (0.25), PL 0.18-0.24 (0.20), WL 0.48-0.59 (0.52). Structure—Propodeum rounded; declivitous face half length of basal face. PI 35-44 (38). Prlostty—Dorsal surface of alitrunk with 18-25 erect to suberect setae. Sculpture—Anterior propodeal suture with mod- erate, longitudinal rugae. PETIOLE: As in Figure 63. Dorsum of node convex. PosTPETIOLE: As in Figure 63. Anterior subpostpetiolar process of moderate size, located medially on venter. Setae erect on dorsum of node and on venter (beneath node), erect to suberect on sides, remainder bare. Posterior edge of postpetiole with dense, non-piliferous punctures. Cozor: Base of man- dible, lateral margin of clypeus, and antenna yellow brown. TYPE MATERIAL. Syntypic series con- sists of ‘‘numerous workers and eight fe- males taken from three different nests’’ (Wheeler, 1904). Cazirornia: Los Angeles Co., Catalina Island, Baker; (16 queens, 21 workers; AMNH), (17 queens, 12 workers; MCZ). Lectotype queen selected from AMNH series bears red, handwritten la- bel: Monomorium/ ergatogyna/ Wheeler/ Lec- totype/ M. DuBois 1980/. DISTRIBUTION. This species ranges along the Pacific coast of California. It has been collected most frequently in the vicin- ity of San Francisco and in the vicinity of Los Angeles (including several offshore is- lands). The range extends inland in the southern portions of California (Fig. 80). The most northern records for this species are Marin Co. (Fairfax, Mill Valley, Mt. Tamalpias, San Anselmo, and Woodacre). Ant GENUS MONOMORIUM 89 FIGS. 71-79. Monomorium ergatogyna. 71—Wings (right side) of male. 72—Aedeagus of male. 73— Volsella of male. 74—Eighth sternite of male. 75—Ninth sternite of male. 76—Male genital capsule, dorsal view. 77—Male genital capsule, lateral view. 78—Male genital capsule, ventral view. 79— Histogram of larval head capsule widths (interval is 0.11 mm). Scales: Top scale (1 mm) for Figure 71. Middle scale (0.5 mm) for Figures 72-73. Bottom scale (1 mm) for Figures 74-78. 90 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN FIG. 80. Distribution of Monomorium ergatogyna (closed circles) and Monomorium wheelerorum (closed triangles). The southern extent of the San Francisco population is represented by collections from Monterey Co. (Fort Ord and Pacific Grove). Records from the Los Angeles vicinity include collections from many of the Channel Islands (Anacapa, East Ana- capa, San Clemente, San Miguel, San Nicholas, Santa Barbara, Santa Catalina, Santa Cruz, and Santa Rosa islands). Rec- ords in the Los Angeles vicinity range from the San Gabriel Mountains in the north to Camp Pendleton and Mount Palomar in the south. The easternmost record for this species is Palm Springs. Although future collecting may show the northern and southern populations of this species are connected, there are currently no records from Fort Ord south to the Los Angeles Basin, a distance of approximately 320 Km (Fig. 80). This distribution is similar to that of M. viridum on the Atlantic coast which is also represented by two disjunct populations (see further discussion under that species). DISCUSSION. Biotocy. Recorded as nesting under stones (Brues, 1903), in soil or in rotton wood (Essig, 1926). Mallis (1941) records this species (misidentififed as M. minimum) nesting in a dry, unshaded area (in the Berkeley Hills) with tumuli forming tiny craters (2-4 cm in diameter). AntT GENUS MONOMORIUM 91 Further details of nest architecture are unknown. In addition to the variety of nesting sites occupied by M. ergatogyna, it has a wide altitudinal tolerance, from 9 m (Long Beach) through 1500 m (H. James Reserve). Monomorium ergatogyna queens have the longest recorded life span in this genus. Colonies (collected in the field) have been maintained in the laboratory for over two years with the original queens (pers. ob- serv.). This compares with an average queen longevity of 39 weeks for M. phar- aonis (Peacock and Baxter, 1950) and 8 months to 1 year for M. minimum (pers. observ. ). Monomorium ergatogyna worker lon- gevity varies from 4-8 months (4 months appears to be the usual life span for workers of this genus). There appear to be three larval instars (Fig. 79). Colonies of M. ergatogyna have multiple, functioning queens (2-6) (pers. observ.). Sexual forms emerge during July and August {collections of reproductives: Santa Barbara Island, 12 June 1978 (gyne lar- vae); Santa Cruz Island, 23 July 1963 (1 male); East Anacapa Island, 23 August 1978 (10 males and 20 unmated queens)}. Only 2 males have been collected from the mainland {Pasadena, 17 April 1929}. If this date is correct, MM. ergatogyna may pro- duce sexuals twice a year. Another pos- sibility is that an undescribed species occurs sympatrically with M. ergatogyna; no morphological differences have been found to support this theory. Brues (1903) reported a species of Micro- don (Diptera: Syrphidae) from a nest at Pacific Grove. No other symbionts have been reported. It appears that the Argentine Ant, Iridomyrmex humilis (Mayr), occupies much of the habitat once utilized by M. ergatogyna (pers. observ. ). Systematics. A great deal of confusion in previous literature has resulted from the erroneous assumption that M/. minimum and M. viridum peninsulatum occurred in California (see further discussion under M. viridum). Furthermore, M. ergatogyna was thought to inhabit only the coastal islands (Essig, 1926; Mallis, 1941; Cook, 1953). Although the San Francisco and Los An- geles populations of this species currently appear allopatric, much more collecting must be done before this point can be decided. Additionally, there appear to be some minor size differences between queens from island localities versus those from the mainland. It seems best to regard all this material as M. ergatogyna. Comparisons. Although all known pop- ulations of M. ergatogyna are geographically removed from any other native Mono- morium, it is possible that queens of this species might be confused with queens of M. cyaneum or M. wheelerorum. Monomorium ergatogyna queens can be separated from those of M. cyaneum by the fringe of setae on the antennal scape (Figure 61) and the reduced head sculpturing in M. ergatogyna. Although M. ergatogyna and M. wheelerorum are the only species included in this revi- sion which have a fringe of setae on the antennal scape, they can easily be sepa- rated, since M. wheelerorum has a slightly depressed scutum and scutellum, a meso- notum with a notal furrow, and more extensive sculpturing on the head. Workers of M. ergatogyna can be separated by the combination of characters listed in the di- agnosis and in the keys, especially: M. ergatogyna workers have a rounded _ pro- podeum while the prododeum of workers of the other species are more angular. Addi- tionally, workers of M. wheelerorum do not have a punctate mesopleuron. Monomorium wheelerorum new species Figures 11, 17, 80, and 81-86 Monomorium minimum: Wheeler, 1917a: 464-465, 501 (in part). DIAGNOSIS. Queen: Wingless; head sculp- ture as in Figure 82; petiole as in Figures 81 and 83; scutum and scutellum flat or slightly de- pressed dorsally; metanotum projecting to level of propodeum and scutellum; propodeum an- gular. Worker: Propodeum angular; meso- pleuron not punctate; petiole as in Figure 85; PI 26-40 (34). DESCRIPTION. As described for M. cyaneum except as follows: QUEEN (DATA FOR HOLOTYPE IN BRACKETS {}): He4aD.: Measurements (represent- ing different localities; N=10) HL 0.46-0.79 92 THe UNIveRsSITY OF KANSAS SCIENCE BULLETIN (0.73){0.71}, HW 0.54-0.68 (0.62){0.59}, SL 0.37-0.58 (0.52){0.48}, TOD 0.15-0.21 (0.17) {0.15}, OD 0.04-0.05 (0.04){0.04}, EL 0.14-0.17 (0.15){0.17}, MOD 0.10-0.13 (0.11){0.13}. Structure—CI 80-92 (86) {83}, SIL 54-78 (72){68}, SIW 64-95 (84){81}. In full frontal view, side straight. Prlosity—As in M. ergatogyna. Sculpture—Several small, parallel, longitudinal rugae beginning between frontal carinae, extending towards anterior ocellus (but never exceeding 2/3 the distance between poste- rior end of frontal carina and anterior ocellus). ALITRUNK: Measurements PW 0.31-0.46 (0.38){0.46}, PL 0.25-0.34 (0.30){0.25}, WL 1.06-1.29 (1.20){1.08}. Structure—Mesonotum with notal furrow. Scutum and scutellum flat to slightly depressed dorsally. Mesopleural suture straight with pit on anterior end. Metanotum (in lateral view) reaching or exceeding level of propodeum and scutellum. Propodeum angular )rarely rounded<, basal and declivitous faces of approximately equal length. PI 22-27 (25){22}. PeTioce: As in Figures 81 and 83. Subpetiolar process moderate to enlarged. Setae subdecum- bent to decumbent on anterior surface of node, suberect to erect on dorsum, side, and posterior surface of node, absent elsewhere. Posterior surface with small to moderate, semicircular, concentric, transverse rugae. Side with small, parallel, longitudinal rugae extending to and fusing with posterior transverse rugae. PosTPETIOLE: As in Figures 81 and 84. Dorsum of node smooth and shining with few small, piliferous punctures, remainder covered with dense, moderate to large, non-piliferous punc- tures. Cozor: Head brown to dark brown, mandible and lateral margin of clypeus yellow brown. Alitrunk dark brown to brown, legs brown to yellow brown, tibiae and tarsi yellow brown to yellow. Petiole, postpetiole, and gaster dark brown to brown. All setae white )rarely, some setae on gaster light yellow<. Mate: Unknown in M. wheelerorum. Worker. Heap. Measurements (representing different localities; V= 10) HL 0.46-0.54 (0.50), ELW. 0.35-0.42 (0:39), SL 0:30-0.38.(0.36), EL 0.06-0.09 (0.07), MOD 0.05-0.06 (0.06). Struc- ture—CI 71-87 (79), SIL 61-76 (72), SIW 85-95 (91). AxiTRUNK: Measurements PW 0.21-0.26 (0:24), PL: 0:12-0,21) (0:17), WL: 0:46-0.56 (0.50). Structure—Propodeum angular, basal face 1.5X as long as declivitous face. PI 26-40 (34). Pilostty—Dorsal outline of alitrunk with more than 10 erect to suberect setae. Setae absent from side, although leg pilosity as in queen. Sculpture—As in M. minimum. PETIOLE: As in Figure 85. Dorsum of node convex. PosTPETIOLE: As in Figure 85. Dorsum of node convex. TYPE MATERIAL. Nevaba: Clark Co.; Wheeler Spring, Spring Mountains, 2012 m, G. C. & J. Wheeler 1561. Holotype queen [LACM] bears red, handwritten label: Monomorium/ wheelerorum/ Holotype/ M. DuBois 1983/. Two paratype workers [LACM, MBDPC] from same locality bear blue, handwritten label: Monomorium/ wheelerorum/ Paratype/ M. DuBois 1983/. Additional paratypes distributed as follows (all bearing paratype labels similar to one described above). Localities refer to those listed in detail below which are surrounded by brackets []. Davis Creek Park - 1 queen, 2 workers [MBDPC], Mullen Gap - 1 queen, 2 workers [KU], Sand Canyon - 1 queen, 2 workers [MCZ], Ophir Grade - 1 queen, 2 workers [NMNH]. ETYMOLOGY. This species is named in honor of Drs. George and Jeanette Wheeler who provided many specimens of re. DISTRIBUTION. This species is known from scattered localities in California, Ne- vada, and Utah. The majority of collec- tions have come from the vicinity of Lake Tahoe (Fig. 80). Cazirornia: Tulare Co., Kennedy Meadows, 1859 m, R. J. Ham- ton & B. S. Ikeda (under rock). NEvaDa: Clark Co., 4.8 km ENE Charleston Peak, 2134.m, G. GC. &J. Wheeler.839; Douglas Co... 6.4 km WSW Wellington, 1676 m, G. C. & J. Wheeler 1093; [Lyon Co., Sand Canyon, 1981 m, R. Bechtel]; 12.9 km NW Smith, 1432 m, G. C. &. J. Wheeler 1945; [Storey Co:, Ophir Grade; 3.2 km WSW Virginia City, 1981 m, GG: &.) J. Wheeler 2280]; 32) km Es,Reno;, 1195S R20E"s::8. 1463°m-Gw Cs ses). Wheeler 2767; | Washoe Co., Davis Creek Park, 1554 m, G. C. & J. Wheeler 2366]; [Mullen Gap, W of S end of Pyramid Lake, I. LaRivers 1484]; 27.4 km N Sparks, 1341 m, G. C. & J. Wheeler 2751. UTaH: Cache Co., Green Canyon, G. Knowlton; Logan Canyon, G. Knowlton; Kane Co., 32 km N Kanab, A. C. Cole; Salt Lake Co., Mill Creek, Chamberlin; Salt Lake City, P: Miles; Utah Co., Provo, Brigham Young Ant GENUS MONOMORIUM 93 University Campus, M. Tanner; Spring- ville, G. Knowlton; Washington Co., La Verkin, A. Sturtevant. DISCUSSION. Although the known range of M. wheelerorum is allopatric from any other species included in this revision, future collecting may reveal some overlap between this species and M. cyaneum and possibly M. ergatogyna. Although both queens of M. wheelerorum and M. ergatogyna have a similar fringe of erect to suberect setae on the scape, M. wheelerorum can easily be separated since it also has a flat to slightly depressed scutum and scutellum and a notal furrow on the mesonotum. Monomorium wheelerorum queens differ from those of M. cyaneum in that queens of the former have the fringe of setae on their scape as described above. Additionally, queens of M. cyaneum never have a flat to slightly depressed scutum and _ scutellum nor a notal furrow on the mesonotum. Workers of M. wheelerorum may be sepa- rated from workers of M. ergatogyna since the latter have a rounded propodeum. Workers of M. wheelerorum may be distin- guished from workers of MZ. cyaneum since the former possess a propodeum with basal and declivitous faces of equal length while the latter possess a propodeum with the basal face longer than the declivitous face. Monomorium compressum Wheeler Figures 87-93 Monomorium minimum subsp. compressum Wheeler, 1914b: 43; 1917a: 464, 501; Kempf, 1972: 144. Monomorium minutum subsp. minima var. com- pressum: Emery, 1921: 173. Monomorium compressum: Ettershank, 1966: 88. DIAGNOSIS. Queen: Wingless; scutum and scutellum depressed dorsally; metanotum (in lateral view) projecting to level of propodeum and scutellum; propodeum angular; petiole as in Figures 87 and 89. Worker: Propodeum angular; mesopleuron not punctate; petiole as in Figure 91; PI 36-39 (38). DESCRIPTION. As described for M. cyaneum except as follows. QuEEN: Heap. Measurements (representing type locality - probably from same nest; N=3) HL 0.60-0.68 (0.64), HW 0.55-0.60 (0.58), SL 0.45-0.50 (0.48), IOD 0.12-0.14 (0.13), OD 0.02-0.04 (0.03), EL 0.11-0.14 (0.12), MOD 0.10 (0.10). Structure— CI 80-96 (90), SIL 73-75 (74), SIW 77-91 (83). Eye small. Pilosity—Setae erect near clypeus, frontal carinae, mandibles and ocelli, decum- bent to appressed on malar area, appressed on remainder of head (including gular region). Sculpture—Small parallel rugosities extending from distal portion of clypeus (between clypeal teeth) to level of antennal insertions between frontal carinae (Fig. 88). AziTRUNK: Measurements PW 0.38-0.40 (0.39), PL 0.25-0.28 (0.26), WL 0.88-0.90 (0.89). Structure—Scutum and scutellum depressed. Mesopleural suture deflected ventrally at posterior end (with a small pit on anterior end). Propodeum angular (Fig. 87), basal and declivitous faces of approxi- mately equal length. PI 27-32 (29). Pilosity— Dorsal outline of alitrunk with 8-16 erect to suberect setae (projecting over 3/4 their lengths above outline). Pe7roLe: As in Figures 87 and 89. Dorsum of node convex to flat. Anterior surface and side of node with small non-pil- iferous punctures; dorsum smooth with few piliferous punctures; posterior surface with moderate, transverse, concentric, semicircular rugae. PosTreTIOLE. As in Figures 87 and 90. Dorsum of node convex to slightly emarginate. Dorsum of node smooth (with few piliferous punctures), remaining surfaces with dense non- piliferous punctures. GasTeR: As in M. mini- mum. Cotor: Head and antenna brown except as follows: base of mandible and lateral margin of clypeus yellow to yellow brown. Mate: Unknown in M. compressum. Worker. Heap. Measurements (representing different localities; N=9) HL 0.46-0.55 (0.49), HW 0.37-0.44 (0.39), SL 0.27-0.40 (0.33), EL 0.08 (0.08), MOD 0.05-0.06 (0.06). Structure— CI 76-87 (80), SIL 54-78 (68), SIW 71-92 (85). Scape reaching but never surpassing occiput. ALITRUNK. Measurements PW 0.22-0.28 (0.25), PL 0.17-0.24 (0.20), WL 0.44-0.62 (0.54). Structure—PI 36-39 (38). Pilosity—Dorsal outline of alitrunk with 8-14 erect to suberect setae. PeTioLe: As in Figure 91. Dorsum of node convex. Setae erect on dorsum of node, re- mainder bare. Lower 1/3 of side with small, non-piliferous punctures (leading to a granular appearance); remainder smooth and _ shining (dorsum of node with piliferous punctures). PosTPETIOLE: As in Figure 91. Posterior 1/3 of postpetiole with dense, non-piliferous punc- tures; remaining surfaces smooth and shining (dorsum of node with piliferous punctures). GasTER: As in M. minimum. Cotor: Head brown except as follows: base of mandible and antenna yellow brown. 94 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN “OQ 90 FIG. 81-86. Monomorium wheelerorum. 81—Queen, lateral view. 82—Head of queen, frontal view. 83— Petiole of queen, posterior view. 84—Postpetiole of queen, posterior view. 85—Worker, lateral view. 86— Head of worker, frontal view. FIG. 87-92. Monomorium compressum. 87 —Queen, lateral view. 88— Head of queen, frontal view. 89—Petiole of queen, posterior view. 90 —Postpetiole of queen, posterior view. 91—Worker, lateral view. 92—Head of worker, frontal view. Scale: (1 mm) for Figures 81-92. Ant GENUS MONOMORIUM 95 TYPE MATERIAL. Syntypic series con- sisting of ‘*4 females and several workers”’ (Wheeler, 1914b): Mexico: Hidalgo: San Miguel [spring or summer, 1913], W. M. Mann (3 queens, 12 workers; NMNH). Lectotype queen here designated from NMNH series bears red, handwritten la- bel: Monomorium/ minimum/ compressum/ Wheeler/ Lectotype Queen/ M. DuBois 1983/. The lectotype is mounted on same point with 2 workers which could not be removed. DISTRIBUTION. This species is known from scattered localities across the Mexican Plateau and extending west to the Pacific coast (Morelos and Hidalgo to Sinaloa) (Figure 93). Collection data for this species follow: MICHOACAN: San José Purula, R. & A. Hamton, B. Ikeda; More zos: Cuernavaca, N. Krauss; Nayari7: San Blas, R. Hamton; San Blas Playa, E. Schlinger; Tepic; SzvAzoA: Los Mochis; Proterillo 10.5 km E. One collection [ Tzx- AS: Marathon, Garden Springs (2 queens, 8 workers; LACM] may be mislabelled, or this species has a much wider range than is indicated above. DISCUSSION. BioLocy. This species 93 FIG. 93. Distribution of Monomorium compressum (closed triangles). 96 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN nests under stones (Wheeler, 1914b) but nest architecture is unknown. Colonies of M. compressum may have multiple, function- ing queens (Wheeler, 1914b). ComPaRISONS. Queens of M. compressum can be distinguished from queens of the other species that occurs in its range (M. cyaneum), since the former have a depressed scutum and scutellum, an emarginate postpetiole, and fewer than 30 erect to suberect setae projecting above dorsal out- line of alitrunk. Workers of M. compressum are separated from those of M. cyaneum since the former have a smooth meso- pleuron and more than 10 erect to suberect setae projecting above dorsal outline of alitrunk. Monomorium viridum Brown Figures 9, 15, 21, and 94-100 Monomorium minutum var. minimum: Emery, 1895: 274-275 (in part); Wheeler, 1905b: 377 (misidentification). Monomorium minutum subsp. minima: Emery, 1921: 172 (in part). Monomorium minimum: Smith, 1930: 3; Wheeler, 1932: 9; Van Pelt, Jr., 1948: 58-59, 64; Van Pelt.jr:, 19567377, 384; Van Pelt, Jr, 1958: 26 (mididentifications). Monomorium viridum Brown, 1943: 243-248; Smith, 1947: 565; Creighton, 1950: 223; Smith, 1952: 811; Ettershank, 1966: 93; Smith, 1967: 356; Crozier, 1970: 116-117; Crozier, 1975: 50, 55; Smith, 1979: 1384; Jones et alt, 1980:.'789;-791;, Jones et .al., 1982: 287. Monomorium peninsulatum Gregg, 1945: 62-66; Smith, 1952: 811; Ettershank, 1966: 91 (NEW SYNONYMY’). Monomorium viridum peninsulatum: Creighton, 1950: 223-224; Smith, 1958: 128; Smith, 19672. 356;-Smuith, 19792 1384. DIAGNOSIS. Queen: Winged; head with rugae as in Figure 95; altitrunk with greater than 30 erect to suberect setae projecting above dorsal outline; petiole and postpetiole as in Figures 94, 96, and 97. Mate: Mandible with four teeth (Fig. 105); genitalia and sterna as in Figures 102 and 107-110. Worker: Pro- podeum, petiole, and postpetiole as in Figure 99; mesopleuron punctate; PI 31-36 (34). DESCRIPTION. As described for M. mini- mum except as follows: QuEeEN: HeapD: Measurements: (representing different localities, N=8) HL 0.72-0.88 (0.81), HW 0.80-0.95 (0.88), SL 0.58-0.69 (0.63), IOD 0.16-0.22 (O219).OD 0-10-0512, (0:11). EL 016-028 (0.23), MOD 0.14-0.22 (0.18). Structure—Head a little broader than long, CI 102-113 (107), distinctly longer than scape SIL 73-83 (77); SIW 67-76 (72). Scape reaching or surpassing occiput by an amount less than length of ped- icel. Sculpture—Moderate, parallel, longitudinal rugae beginning all along lateral margin of Clypeus, extending behind antennal insertion, and curving towards frontal carina, reaching frontal carina near apex; small, parallel, longi- tudinal rugae beginning along malar area, ex- tending towards compound eye, paralleling ocular suture and vanishing near level of ante- rior ocellus; small, parallel, longitudinal rugae beginning near apex of frontal carina and ex- tending to level of anterior ocellus; moderate, parallel, longitudinal rugae beginning near la- bium and continuing alson gular region ending near occiput. Large piliferous punctures evenly distributed over entire surface of head. ALITRUNK: Measurements PW 0.60-0.85 (0.72), PL. 0,30-0:50. (0.38), “WL, 1295-1290" G70): Structure—Mesonotum with notal furrow. Meso- pleural suture straight (small pits on anterior and posterior ends). Propodeum rounded (Basal and declivitous faces of equal length). PI 18-27 (22). Sculpture—Moderate, piliferous punctures on dorsum of scutum, scutellum and meso- pleural suture. Propodeum covered with mod- erate, parallel, longitudinal rugae throughout. Sculpiuring of sutures as in Figure 94. PETIOLE: As in Figures 94 and 96. Dorsum of node flat to slightly emarginate. Setae appressed on anterior surface of node, erect to decumbent on sides, dorsum, and posterior surface of node. Entire surface of petiole covered with small, non- piliferous punctures (obscuring small piliferous punctures where they occur). Lower half of side of petiole )sometimes including lower parts of node¢ with small, parallel, longitudinal rugae. Posterior surface of node with moderate, trans- verse, concentric, semicircular rugae. Pos7- PETIOLE. As in Figures 94 and 97. Dorsum of node flat to slightly emarginate. Setae decum- bent to appressed on anterior surface of node, suberect to erect on sides, dorsum, and poste- rior surface of node. Entire surface covered with large to small non-piliferous punctures. Cozor: Head dark greenish-brown )green is more pro- nounced in living or freshly preserved spec- imens€ except as follows: mandibles and antenna dark reddish-brown. Altitrunk and legs reddish-brown with green overlay )green tends to fade in older specimens<. Petiole and postpetiole reddish-brown to brown with slight Ant GENUS MONOMORIUM 97 0.1 FIGS. 94-105. Monomorium viridum. 94—Queen, lateral view. 95—Head of queen, frontal view. 96— Petiole of queen, posterior view. 97—Postpetiole of queen, posterior view. 98 —Head of worker, frontal view. 99—Worker, lateral view. 100—Head of ergatogyne, frontal view. 101—Ergatogyne, lateral view. 102—Male, lateral view. 103—Labial palp of male. 104—Maxilla and maxillary palp of male. 105—Mandible of male. Scales: Left scale (1 mm) for Figures 94-102. Middle scale (0.1 mm) for Figure 105. Right scale (0.1 mm) for Figures 103 and 104. 98 THE UNIVERSITY OF Kansas SCIENCE BULLETIN green tint. Gaster black with green or blue hues in some specimens. Mae: Heap: Measurements (representing 2 localities, N=5) HL 0.56-0.69 (0.62), HW 0.90-0.95 (0.93), SL 0.28-0.35 (0.31), IOD 0.21-0.26 (0.24), OD 0.14-0.15 (0.15), EL 0.40-0.44 (0.42), MOD 0.25-0.31 (0.28). Struc- fure—Cl 138-164 (151), SIL 44-55 (60), SIW 30-37 (33). Mandible with 4 teeth (Fig: 105): Pilosity—Setae erect to decumbent over entire head, appressed on gula. Sculpture—Entire head covered with dense, moderate, non-piliferous punctures. Moderate, parallel, rugae beginning along lateral margin of clypeus and converging at antennal insertion; several moderate, paral- lel, rugae beginning at malar area, continuing to level of compound eye then paralleling ocular suture; clypeus with small to moderate, parallel, longitudinal rugae arising between clypeal teeth and extending to level of antennal insertion, moderate, parallel rugae extending from frontal carinae to level of anterior ocellus. ALITRUNK: Measurements PW 0.84-0.97 (0.93), PL 0.44-0.52 (0.48), WL 1.68-1.84 (1.76). Structure—Meso- notum with notal furrow. Scuto-scutellar suture near middle of alitrunk. Mesopleural suture straight. Metanotum not projecting to level of scutellum and propodeum. Propodeum rounded (basal and declivitous faces of equal length, Figure 102). PI 24-29 (28). Sculpture— Moderate piliferous punctures on scutum and scutellum and anterior edge of mesopleuron, small, non-piliferous punctures on propodeum. Sculpturing of sutures as in Figure 102. Periote: As in Figure 102. Dorsum of node emarginate. Subpetiolar process of moderate size with anterior edge forming angle. Setae appressed to decumbent on anterior surface of node, subdecumbent to erect on dorsum, sides, and posterior surface of node. All surfaces except dorsum, with small, non-piliferous punc- tures. Lower 1/3 of side of petiole, below node, gitudinal rugae. PosTPeTIOLE: As in Figure 102. Dorsum of node slightly emarginate. Setae de- cumbent to erect on all surfaces of node. Dor- sum smooth and shining, remaining surfaces covered with small non-piliferous punctures. All surfaces with small piliferous punctures. GeniTaui4: As in Figures 102, and 107-110. Eighth sternite with emargination possessing setae; emargination approximately half as deep as wide (Fig. 107). Ninth sternite with 18-20 erect setae (Fig. 108). Aedeagus with 12 teeth; toothed margin straight (Fig. 109). Cuspis of volsella with 7 setae (Fig. 110). Cozor: As in queen. Genitalia yellow brown to brown. Worker. Heap: Measurements (representing different localities; N= 10) HL 0.45-0.58 (0.51), HW 0.39-0.50 (0.44), SL 0.30-0.40 (0.35), EL 0.08-0.13 (0.09), MOD 0.06-0.08 (0.06). Struc- ture—CI 83-89 (85), SIL 53-78 (69), SIW 65-90 (81). Summit of head concave to slightly emar- ginate. Scape reaching or surpassing occiput by an amount less than length of pedicel. Sculp- ture—Small to moderate, parallel, longitudinal rugae beginning along lateral margin of clypeus, extending to level of antennal inser- tion. AxiTRUNK: Measurements PW 0.24-0.33 (0:27), PE 018-0:22.5(0:20), Wis 0.52-0.66 (0.58). Structure—Anterior propodeal suture deep (almost reaching to level of propodeal suture). Propodeum rounded; declivitous face 1/3 to 1/2 length of basal face. PI 32-36 (34). Pilosity—Dorsal outline of alitrunk with 8-12 erect to suberect setae. Sculpture—Mesopleuron with dense, small, non-piliferous punctures throughout; anterior propodeal suture with short, large, parallel, longitudinal rugae. Petiote: As in Figure 99. Subpetiolar process of moderate size. Setae appressed on anterior surface of node, erect on dorsum of nodes, absent elsewhere. PosTrPeTIOLE: As in Figure 99. Setae apparessed on anterior surface of node, erect to decumbent on sides, dorsum, and venter (beneath node), absent elsewhere. Cotor: Head dark brown to brown except as follows: occiput with green tint (most visible in living or recently killed specimens), antenna, mandible, and lateral margin of clypeus brown to yellow-brown. AziTRUNK: Dark brown with green tint on dorsum; legs brown to yellow- brown. ERGATOGYNE. Heap: Measurements (represent- ing different localities; N=2) HL 0.66-0.69 (0.68), HW 0.64-0.74 (0.69), SL 0.50 (0.50), IOD 0.14-0.15 (0.14), OD 0.05 (0.05), EL 0.15 (0.15), MOD 0.09-0.10 (0.10). Structure—Head as broad as long, CI 97-107 (102), distinctly longer than scape, SIL 72-76 (74); SIW 68-78 (73). In full frontal view, head broadest slightly above eyes; side straight; occiput rounded later- ally, summit flat to slightly convex. Eye moder- ate in size (intermediate between queen and worker). Scape reaching or surpassing occiput by an amount less than length of pedicel. Ocelli present. Mandible, maxillary palp, labial palp, clypeal teeth, and frontal carinae as in queen. Pilosity—Setae erect to decumbent over entire surface of head. Antennal pilosity as in queen. Sculpture—Smooth and shining except as fol- lows: trace of small, longitudinal, parallel rugae extending from lateral margin of clypeus and converging near frontal carinae. Moderate pil- iferous punctures evenly distributed over entire Ant GENUS MONOMORIUM 99 surface (Fig. 100). ALi7TRUNK: Measurements PW 048-0%495(0148); PIE 0230-0231 (0230); Wi 1.11-1.26 (0.18). Structure—Mesonotum with small pit in place of notal furrow. Scuto-scutel- lar suture on dorsal 1/3 of alitrunk. Remaining description of alitrunk as in queen except that wings are absent (fusion of sclerites indicate the ergatogynes are wingless). Pilosity—As in queen with many erect to suberect setae on sides of propodeum and mesopleuron. Sculpture— Smooth and shining with many piliferous punc- tures throughout thorax. Sculpture of pro- podeum as in queen (Fig. 101). Perrove: Dorsum evenly convex to flat. Remaining de- scription as in queen. PosTPETIOLE: Dorsum evenly convex to flat. Remaining description as in queen. Gas7TeER: As in queen. Coxor: As in queen. TYPE MATERIAL. Type series consist- ing of holotype female and numerous para- type females, males and workers (Brown, 1943) New JERSEY: Ocean Co., Lakehurst, August 23, 1940, W. L. Brown, Jr. [Holo- type female - Academy of Natural Sci- ences, Philadelphia, Pennsylvania Number 10561; paratypes in NMNH and MCZ]. Synonymous M. peninsulatum type series consisting of ‘64 females ... and numerous workers’? (Gregg, 1945) Frorrpa: Dade Co., South Miami, E. V. Gregg. | Holotype queen - R. E. Gregg personal collection; paratypes in AMNH, LACM, and NMNH]. DISTRIBUTION. This species occurs in two allopatric populations along the Atlan- tic coast: one in the Pine Barrens of New Jersey and the other in Georgia and Flor- ida (Fig. 106). In spite of intensive collect- ing by W. L. Brown, Jr., E. O. Wilson, and others, only one collection has been made in the coastal plain between New Jersey and Georgia (NORTH CAROLINA: Beaufort Co., Wilmar 1.6 km N, W. L. Brown, Jr., & E. O. Wilson). In many areas, this species is replaced by M. mini- mum. Apparently this difference is due to microhabitat selection for nest sites by queens of both species. Monomorium viridum prefers sand, while M. minimum prefers clay soils (W. L. Brown, Jr. pers. comm.). In the New Jersey population of M. viridum, collections range from Lakehurst (Ocean Co.) and Island Heights (Ocean Co. ) in the north to Cape May (Cape May Co.) in the south. Further north, M. viridum is replaced by M. emarginatum and M. mini- mum replaces both species to the west. In the Florida population, collections are common throughout the eastern and cen- tral portions of the peninsula from Jackson- ville (Duval Co.) in the north to Miami (Dade Co.) in the south. The furthest west- ern records for this species appear to be Sarasota (Sarasota Co.) and Gainesville (Al- achua Co.). This population apparently ranges as far north as Georgia (W. L. Brown, Jr., pers. comm.). Monomorium tragert occurs in portions of northern Flor- ida (Tallahassee and Gainesville) while M. minimum occurs to the north and west of the Florida M. virrdum population. DISCUSSION. BioLtocy. This species nests in open sandy soil. Brown (1943) records it nesting along a roadside with tumuli forming craters 12-20 cm in diame- ter. Gregg (1945) also mentions the crater nests of this species. Although I have at- tempted to excavate several nests (FLOR- IDA, Orlando vicinity and Daytona Beach vicinity), I could not determine additional details of nest architecture due to the shift- ing, loose sand. Colonies of M. viridum may have multi- ple, functioning queens (1-5) (Brown, 1943). It is not known whether nests are established by one or several queens al- though Gregg (1945) encountered over 60 queens (most dealate) in one nest. Nuptial flights apparently occur since males and queens are winged, although no flights have been reported and I have not encoun- tered collections of males or queens found away from nests. Sexual forms are pro- duced during July {collection of males: New JERSEY: Burlington Co., July 31, 1954; Ocean Co., July 29, 1954}. No males have yet been collected outside of New Jersey. SYSTEMATICS. Some confusion in the lit- erature has resulted from Gregg’s re- description of M. viridum as M. peninsulatum (Gregg, 1945). Gregg mentions that the species are quite similar and reported char- acters to separate them (Gregg, 1945: 66). The color characters mentioned are unus- 100 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN 106 FIG. 106. Distribution of Monomorium viridum (closed triangles). able for a number of reasons. First, Brown stated that his specimens were examined in direct sunlight, while Gregg makes no mention of the light source he used. Sec- ond, these colors change and fade over time (the exact sequence is not known). Furthermore, color differences mentioned by Gregg are variable within and among nests. Gregg’s characters dealing with the sculpture of the ocellar triangle and the sculpture of the anterior surface of the petiole and postpetiole are equally variable. Size measurements given by Gregg are not useful since they were based upon total length, including gaster (composed of many membranes which shrink as the spec- imen dries). Length of the gaster can also vary greatly with development of the ov- aries (this is true for workers also). This leaves one character, the shape of the clypeal teeth. Gregg described them as blunt but gave no figure for comparison. In fact, they are of the same general shape in specimens from New Jersey and Florida. This confusion led Creighton (1950) to place M. peninsulatum as a subspecies of M. viridum. While M. viridum continued to be known only from type material, Creighton (using Gregg’s variable characters) ex- tended the range of M. peninsulatum across the southern United States to Arizona and California, confusing it with M. minimum, M. cyaneum, and M. ergatogyna. Creighton’s error has been perpetuated to the present (Gregg, 1963; Smith, 1979). Comparisons. Ranges of three other species (M. emarginatum, M. minimum, and M. trageri) overlap the range of M. viridum. These species can be separated by charac- ters found in the keys and diagnoses. Queens of M. virrdum possess a notal fur- row on the mesonotum while queens of M. minimum and M. trageri do not. Monomorium viridum queens can be separated from M. emarginatum queens since the metanotum of the latter does not project to the level of the propodeum and scutellum. Additionally, head sculpturing is reduced in M. emar- ginatum queens. Furthermore, both petiole and postpetiole are emarginate in M. vir- idum queens. Workers of M. viridum can be separated from M. tragert and M. emarginatum workers since the latter two species have a pro- podeum with basal face shorter than de- clivitous face. Monomorium viridum workers can be separated from those of M. minimum since the former have a punctate meso- pleuron. Ant GENUS MoNnomMorRIUM 101 Monomorium emarginatum new species Figures 111-118 Monomorium minimum: Wheeler, 1916: 584 (misi- dentification). DIAGNOSIS. Queen: Winged; mesonotum with notal furrow; mandible with 4 teeth, basal tooth reduced (Fig. 113); head sculpturing as in Figure 112; petiole and postpetiole as in Figures 111, 114, and 115. Worker: Propodeum not angular (Fig. 117); mesopleuron not punctate (Fig. 117); petiole and postpetiole as in Figure 117; 4-6 erect setae projecting above dorsal outline of alitrunk; PI 34-42 (37). DESCRIPTION. As described for M. viridum except as follows. QuEEN: Heap: Measurements (representing different localities; N=6) HL 0.71-0.80 (0.74){0.80}, HW 0.69-0.81 (0.75){0.81}, SL 0.52-0.64 (0.58){0.62}, IOD 0.16-0.18 (0.17) {0.18}, OD 0.05-0.06 (0.05) {0.05}, EL 0.15-0.19 (0.18) {0.19}, MOD 0.09-0.12 (0.11) {0.12}. Structure—Head as broad as long, CI 96-106 (101){101}, distinctly longer than scape, SIL 68-86 (78) {78}; SIW 68-88 (77){76}. Scape not surpassing oc- ciput. { Holotype lacks left antenna}. Mandible with 4 teeth, basal tooth reduced (Fig. 113). Sculpture—Moderate, faint, parallel, longitudi- nal rugae beginning along lateral margin of clypeus, extending past antennal insertion, and converging with frontal carinae. Large to mod- erate, piliferous punctures evenly distributed (Fig. 112). AzrTRUNK: Measurements PW 0.58-0.66 (0.62) {0.58}, PL 0.29-0.40 (0.34) {0.40}, WL 1.26-1.65 (1.52) {1.52}. Structure— Mesopleural suture straight with small pit on posterior end. Metanotum (in lateral view) not projecting to level of propodeum and scutellum. PI 20-26 (23) {26}. Wings present {lacking on holotype}. Sculpture—Smooth and shining with moderate piliferous punctures except as follows: lower 1/3 of propodeum (below spiracle) with moderate to large, parallel, longitudinal rugae. PeTiove. As in Figures 111 and 114. All surfaces smooth and shining (moderate piliferous punc- tures where applicable) with small non-piliferous punctures (leading to a granular appearance) along lower 1/2 of side. PosTreriote: As in Figures 111 and 115. Dorsum of node flat. Setae appressed to decumbent on anterior surface of node, decumbent to suberect on dorsum, erect on posterior surface and side of node, absent elsewhere. Postpetiolar surface smooth and shin- ing with small piliferous punctures throughout. Coror: Head dark brown to brown except as follows: antenna, mandible, and lateral margin of clypeus brown to yellow. Alitrunk dark brown to brown, legs brown to yellow-brown. Petiole, postpetiole, and gaster dark brown to brown. All setae yellow. Mate: Unknown in M. emarginatum. Worker. Heap: Measurements (representing separate localities; N=5) HL 0.44-0.51 (0.48), HW 0.38-0.44 (0.41), SL 0.28-0.41 (0.35), EL 0.09-0.10 (0.09), MOD 0.06 (0.06). Structure— CI 80-87 (84), SIL 64-80 (72), SIW 74-93 (86). Scape not surpassing occiput. Sculpture—Head smooth and shining with small piliferous punc- tures evenly distributed. AzimTRUNK: Measurements PW 0.22-0.29 (0.26), PL 0.18-0.21 (0.19), WL 0.49-0.62 (0.53). Structure—Ante- rior propodeal suture of moderate depth (Fig. 117). Propodeum rounded, basal face 2/3 as long as declivitous face (Fig. 117). PI 34-42 (37). Pilostty—Dorsal outline of alitrunk with 4-6 erect setae projecting above dorsal outline. Sculpture— Anterior propodeal suture with short, parallel, longitudinal rugae; lower 1/3 of side of propodeum (below spiracle) with small, some- times faint, parallel, longitudinal rugae. PeTIoe: As in Figure 117. Dorsum of node flat to slightly convex. Setae erect on dorsum of node, remainder bare. Cozor: Head dark brown except as follows: antenna and madible yellow-brown to brown. Alitrunk brown, legs brown to yellow-brown. Petiole, postpetiole, and gaster brown to dark brown. All setae yellow. TYPE “MATTE EOLA ie VIER CNT Rappahannock Co.; Amissville 3.2 km W, June-21, 1957, W. Ll. Brown, race ©: Wilson. Holotype queen {[MCZ] bears red, handwritten label: Monomorium/ emar- ginatum/ Holotype/ M. DuBois 1983/. Par- atypes distributed as follows (localities refer to those listed in detail below which are surrounded by brackets []: Brewster, Cape Cod - 1 queen, 3 workers [MCZ], Woods Hole - 2 queens, 4 workers [NMNH]. All paratypes bear blue, handwritten labels: Monomorium/ emarginatum/ Paratype/ M. DuBois 1983/. ETYMOLOGY. This species is named for the furrow (emargination) on the meso- notum. DISTRIBUTION. This species is known from scattered localities throughout the northeastern United States (Fig. 118). Col- lection data follow: MassaCHUSETTS: [Barnstable Co., Cape Cod, Brewster, Au- 102 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN FIGS. 107-110. Monomorium viridum. 107—Eighth sternite of male. 108—Ninth sternite of male. 109— Aedeagus of male. 110—Volsella of male. Scales: Top scale (0.5 mm) for Figures 107 and 108. Bottom scale (0.5 mm) for Figures 109 and 110. FIG. 111-117. Monomorium emarginatum. 111—Queen, lateral view. 112—Head of queen, frontal view. 113—Mandible of queen (schematic). 114—Petiole of queen, posterior view. 115—Postpetiole of queen, posterior view. 116—Head of worker, frontal view. 117— Worker, lateral view. Scale: (1 mm) for Figures 111, 112, 114-117. Ant GENUS MONOMORIUM 103 a > fe 118 FIG. 118. Distribution of Monomorium emar- ginatum (closed triangles). gust 4, 1955, W. L. Brown, Jr.]; [Plymouth Go, Woods Holey July 15, 1930) A. 1. Sturtevant]. New Yorx: New York City [several collectors]; Montgomery Co., John- stown, A. Sturtevant. DISCUSSION. Biotocy. Previous liter- ature records (Wheeler, 1916) indicate that this species prefers to nest in sandy or gravelly soil with tumuli forming tiny cra- ters. Further details of next architecture and biology are unknown. Systematics. Although the collections of this species are sporadic in the northeastern United States, it appears to be the only species which occurs there. Since workers could be easily mistaken for M. minimum, I assume this was the species that Wheeler (1916) discussed. Unfortunately, he did not collect any queens during his study and workers are difficult to separate. It 1s possi- ble this species occurs sympatrically with M. viridum in coastal areas. Comparisons. This species is most likely to be confused with M/. minimum and M. virrdum. Queens of M. minimum can be separated from those of M. emarginatum since the latter possess a furrow on the mesonotum. Queens of M. viridum can be separated from those of M. emarginatum since the petiole and postpetiole of the former are always emarginate while those of the latter are not. Additionally, M. emarginatum queens have less facial sculp- turing (Fig. 112). Workers of M. emar- ginatum can be separated from both these species since the basal face of the pro- podeum is shorter than the declivitous face. Monomorium ebeninum Forel Figures 8, 14, 16 and 119-133 Monomorium carbonarium: Forel, 1881: 8; Forel, 1899: 78; Kusnezov, 1949: 425 (misiden- tifications). Monomorium ebeninum Forel, 1891: 165; Et- tershank, 1966: 88 (misspelled as ebininum); Jjonescet al-, 19822287. Monomorium carbonarium ebeninum: Forel, 1899: 78; Wheeler, 1905a: 88-89; Wheeler, 1908: 423: Wheeler, 1917b: 458; Tulloch, 1930: 203; Smith, 1936b: 832-833; Wheeler, 1936: 199; Wolcott & Martorell, 1937: 577-579; Wolcott, 1948: 814-815; Creighton, 1950: 218 (misspelled as ebininum); Kempf, 1972: 1435, Lavigne, 1977-224. Monomorium carbonarium ebenina: Emery, 1921: Al DIAGNOSIS. QurEEN: Wingless; petiole and postpetiole as in Figures 119, 121, and 122; scutum and scutellum slightly depressed dor- sally or flat; metanotum projecting to level of propodeum and scutellum; propodeum angular. Mate: Genitalia and sterna as in Figures 125, 128-131. Worker: Propodeum angular; meso- pleuron not punctate; petiole as in Figure 121; PI 40-44 (41). DESCRIPTION. As described for M. mini- mum except as follows. QuEEN: Heap: Measurements (representing different localities; N=10) HL 0.65-0.72 (0.68) {0.72}, HW 0.55-0.65 (0.60) {0.61}, SL 0.42-0.52 (0.49) {0:52%, 1OD 010-017, (0:14), {0.12 Op 0.04-0.07 (0.05) {0.05}, EL 0.15-0.18 (0.16) {0.18}, MOD 0.10-0.13 (0.12) {0.13}. Struc- ture—CI 80-93 (88) {85}, SIL 62-77 (72) {72}, SIW 76-88 (82) {85}. Side of head straight. Scape not reaching occiput. Masticatory margin of mandible with 4 )rarely 5¢ teeth. Clypeal 104 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN eoeee \ “Ngee FIGS. 119-132. Monomorium ebeninum. 119—Queen, lateral view. 120—Head of queen, frontal view. 121—Petiole of queen, posterior view. 122—Postpetiole of queen, posterior view. 123—Head of worker, frontal view. 124—Worker, lateral view. 125—Male, lateral view. 126—Labial palp of male. 127—Maxilla and maxillary palp of male. 128—Eighth sternite of male. 129—Ninth sternite of male. 130—Volsella of male. 131—Aedeagus of male. 132—Mandible of male. Scales: Top scale (1 mm) for Figures 119-125. Second scale (0.1 mm) for Figure 132. Third scale (0.5 mm) for Figures 128 and 129. Fourth scale (0.5 mm) for Figures 130 and 131. Bottom scale (0.1mm) for Figures 126 and 127. Ant GENUS MonomMorIUM 105 teeth moderately sharp to blunt, short. Frontal carinae diverging strongly posteriorly. Sculp- ture—Moderate, parallel, longitudinal rugae be- ginning all along lateral margin of clypeus, continuing to level of middle of compound eye after converging slightly near frontal carinae )rugae above frontal carinae may be absent or greatly reduced¢ (Fig. 120). AziTRUNK: Measurements PW 0.40-0.52 (0.42) {0.43}, PL 0.23-0.32 (0.29) {0.23}, WL 1.05-1.18 (1.11) {1.13}. Structure—Scutum and _ scutellum slightly depressed dorsally. Mesopleural suture deflected ventrally at posterior end (with small pit on anterior end and larger pit on posterior end). Propodeum angular, basal and declivitous faces of approximately equal length. PI 20-30 (26) {20}. Wings absent (fusion of sclerites indicate queens are wingless). Scu/pture—Entire propodeum covered with small, non-piliferous punctures (leading to a granular appearance); declivitous surface of propodeum with moder- ate, parallel, transverse rugae. PETIOLE: As in Figures 119 and 121. Dorsum of node flat to slightly emarginate. Setae decumbent to ap- pressed on anterior surface of node, subdecum- bent to decumbent on dorsum, absent elsewhere. Entire petiole smooth and _ shing (with small, piliferous punctures on anterior surface and dorsum) except as follows: posterior 1/5 of petiole with moderate, concentric, semi- circular, transverse rugae. PosTPETIOLE: As in Figures 119 and 122. Dorsum of node flat. Setae subdecumbent to decumbent on posterior sur- face of node, absent elsewhere. Postpetiolar surface smooth and shining except as follows: posterior 1/2 with dense, moderate, non-pil- iferous punctures. Cozor: Entire body and ap- pendages usually brown to dark brown. All setae white. Mate: Heap: Measurements (representing dif- ferent localities; N=3) HL 0.55-0.61 (0.59), HW 0.61-0.68 (0.64), SL 0.24-0.28 (0.26), IOD 0.18-0.20 (0.19), OD 0.06-0.08 (0.07), EL 0.22-0.25 (0.24), MOD 0.12-0.18 (0.16). Struc- ture—CI 100-113 (109), SIL 39-47 (44), SIW 39-41 (40). Maxillary palp 1 - segmented (Fig. 127); labial palp 1 segmented (Fig. 126). Clypeal teeth absent. Frontal carinae not diver- ging posteriorly. Prlosity—Setae erect near clypeus, mandible, and occiput, decumbent to appressed elswhere. Sculpture—Several moder- ate to large, parallel, longitudinal rugae begin- ning all along lateral margin of clypeus, extending to antennal insertion. Several small to moderate, parallel, longitudinal rugae ex- tending from clypeus to level of frontal carinae. Additionally, several moderate, concentric, semicircular rugae extending from level of com- pound eye to occiput and following contour of occiput. ALITRUNK. Measurements PW 0.50-0.60 (0:56); PE, 0530-0336) (0:32), Wily 112-1227 (1.20). Structure—Pronotal-scutal suture on dor- sal 1/2 of alitrunk. Mesopleural suture as in queen except as follows: pit absent on anterior end. Metanotum (in lateral view) not projecting to level of propodeum and scutellum. Pro- podeum angular, basal face 2 times as long as declivitous face. PI 25-28 (27). Sculpture—Sev- eral moderate, semicircular, concentric rugae with vertex at anterior edge of poronotum, extending posteriorly (as parallel, longitudinal rugae) to scutum; sculpturing of sutures as in Figure 125. Periove: As in Figure 125. Setae appressed on anterior surface of node, erect to suberect on dorsum, absent elsewhere. Entire surface of node, erect to suberect on dorsum, absent elsewhere. Entire surface smooth and shining. PosTeTIOLE: As in Figure 125. Setae decumbent to subdecumbent on anterior sur- face of node, erect to suberect on posterior surface of node and venter, absent elsewhere. All surfaces smooth and shining. GasTer: Setae of first gastral tergite reaching or exceeding level of dorsum of postpetiolar node. Geni7aLia: As in Figures 125 and 128-131. Eighth sternite with emargination approximately 1/2 as deep as wide (Fig. 128). Ninth sternite with 8 erect setae (Fig. 129). Aedeagus with 8 teeth; toothed margin straight (Fig. 131). Volsella with re- duced digitus; lacking cuspis (Fig. 130). Cozor: Head and antenna brown to dark brown except as follows: mandible yellow to yellow-brown. Alitrunk brown to dark brown, legs yellow to yellow-brown. Petiole and postpetiole brown to yellow-brown, gaster brown to dark brown. Genitalia brown to yellow-brown. All setae white to yellow. Worker. Heap. Measurements (representing different localities; N= 10) HL 0.48-0.55 (0.51), HW 0.38-0.42 (0.40), SL 0.35-0.40 (0.39), EL 0.07-0.09 (0.08), MOD 0.05-0.06 (0.05). Struc- ture—CI 76-84 (78), SIL 67-95 (78), SIW 71-105 (95). Side of head straight. Clypeal teeth short, blunt. Sculpture—Several small, short, parallel, longitudinal rugae on clypeus; several small, short, parallel, longitudinal rugae begin- ning all along lateral margin of clypeus, con- tinuing to antennal insertion. ALITRUNK: Measurements PW 0.25-0.28 (0.26), PL 0.21-0.25 (0.23), WL 0.52-0.62 (0.56). Structure—Pro- podeum angular, both faces of approximately equal length. PI 40-44 (41). Perroxe: As in Figure 124. Dorsum of node flat to slightly emarginate. Setae erect to suberect on dorsum 106 Tue University OF KANSAS SCIENCE BULLETIN of node, absent elsewhere. PosTPETIOLE: As in Figure 124. Dorsum of node flat to slightly emarginate. GasTeR: Setae of first gastral tergite reaching or exceeding level of dorsum of postpetiolar node. Cozor: Head dark brown to brown except as follows: mandibles yellow- brown to brown. Alitrunk and legs dark brown to brown. Gaster dark brown. TYPE MATERIAL. Syntypic series con- sisting of 2 queens, 1 male, and 16 workers (MHNG) from the following localities: GRENADA (male), St. VINCENT (queens), St. THomas (ANTILLE) (workers). Six workers bear the date 14 X 78. Lectotype queen here designated from MHNG series bears red, handwritten label: Monomortum/ ebeninum/ Forel/ Lectotype/ M. DuBois 1983/. It is from St. Vincent. Forel (1891: 165) described the distin- guishing characters of this species: “‘La race de l’Amérique tropicale que j’avais cru devoir rapporter au carbonarium (Ameisen der Antille St. Thomas, 1881) et que j’ai recue dés lors du Guatémala, etc., se distingue nettement par les deux élévations du métanotum, par sa large échancrure méso-métanotale, par la face déclive plus haute du métanotum plus éléve qui forme presque un angle avec la face basal, par le premier noeud du pédicule plus mince et plus longuement pétiolé. Je propose de |’appeler ebenznum.”’ Presumably, the material from Guatemala was provided through Pergande (W. L. Brown, Jr., pers. comm.); however, none of this material can be discovered in the remains of Pergande’s collection. The re- mainder of this type series is in Geneva (MHNG). A few of these specimens are clearly labelled ‘“TYPUS’’; these include the male, both queens, and 6 workers. I assume that Forel regarded these spec- imens as part of his type series and merely chose to list their localities as ‘‘etc.’’ when he described M. ebeninum. I chose the lec- totype, designated above, in an effort to restrict the type locality and to reduce future confusion since queens are easier to identify than workers. DISTRIBUTION. This species ranges throughout the Caribbean region. It has been collected from many of the islands and from Atlantic and Pacific coastlines in many parts of Central America (Fig. 133). Many collections were examined from the following islands which range from the Bahamas in the north through Tobago in the south: Andros (Bahama Islands), Anti- gua (Lesser Antilles), Barbados (Lesser Antilles), Cuba (Greater Antilles), Domin- ican Republic (Greater Antilles), Grenada (Lesser Antilles), Haiti (Greater Antilles), Isla de Pinos (Greater Antilles), Jamaica (Greater Antilles), Martinique (Lesser Antilles), Montserrat (Lesser Antilles), Nassau (Bahama Islands), Nevis (Lesser Antilles), Puerto Rico (Greater Antilles), St. Croix (Virgin Islands), St. John (Virgin Islands), St. Vincent (Lesser Antilles), and Tobago (Lesser Antilles). Mainland collec- tion localities range from Tamalupias and Vera Cruz in Mexico to Panama along the Atlantic coast. Only a few collections have been made along the Pacific coast of Cen- tral America. These include localities in Oaxaca in Mexico, El Salvador, and Guanacaste Province in Coasta Rica. I was unable to examine any specimens of this species collected in the northern coastal regions of South America or from penin- sular Florida. It is probable that additional collecting will reveal colonies from both these regions. DISCUSSION. Biotocy. This species presumably nests under stones and in plant cavities. Forel (1899) indicated that M. ebeninum lives in stems and hollow branches. Wheeler (1905a) recorded it from under stones, in cavities of 72llandsia spp-, and in dry twigs of buttonwood bushes. Smith (1936b: 832) elaborated fur- ther: ‘‘This is one of the most common ants of the West Indies... It forms populous colonies which are characterized by having many reproductive queens to a colony. Their greath adaptability is indicated by the fact that this species nests in both soil and wood, back of the leaf sheaths or [szc] corn and bananas, in cabbage heads, Tillandsias, and in the fruits of Hzbescus sabdariffa. The workers are exceedingly fond of honeydew. They have been noted attending such insects as Sazssetia hemi- spherica Targ., Coccus viridus Green, the Ant GENUS MONOMORIUM 107 ebeninum I= 12 trageri & A) 4 i) Xv aa o C) 5: 26 ee, - \ siento MS 4 Sd ‘ ~~ a ‘ Tins - N x < ‘ NOE ce = \ } \ S \ FIG. 133. Distribution of Monomorium ebeninum and Monomorium tragert. aphid Svpha flava Forbes, etc. The ants are also highly predacious.’’ Wolcott and Mar- torell (1937) recorded this ant as a predator on the eggs of the sugarcane moth borer, Diatraea saccharalis Fabricius (Lepidoptera: Pyralidae). Although Lavigne (1977) re- corded this species from lawns in Puerto Rico, he provided no details on the nest’s microhabitat nor structure. Wheeler (1905a) recorded multiple queens per nest (up to 12). Although M. ebeninum has a moderately broad altitudinal tolerance (0-1600 m above sea level), it is most frequently collected in coastal areas (Fig. 133). Sexual forms are produced in spring {collections of males: Jamaica: Montego Bay, March 14, 1911; ANpRos IsLAND (Bahama Islands): Mangrove Cay, May 31, 1904}. Wolcott (1948) mentions this ant is eaten by the lizards Anolis stratulus and A. cristatellus. Lavigne (1977) collected workers from stomachs of the following two frog species: Eleutherodactylus cochranae and E. coqut. COMPARISONS. Since Monomorium ebeninum is the only species with wingless queens that occurs in the Caribbean re- gion, it should not be confused with any other species. Three other species occur in areas which border the range of M. ebeninum: M. cyaneum, M. trager, and M. virrdum. Queens of M. ebeninum are sepa- rated from those of these other species since the profile of the scutum and scutellum of M. ebeninum queens is flat or slightly con- cave while the profile of the remaining species is clearly convex. Additionally, the head sculpturing in M. ebeninum is quite reduced. Workers of M. ebeninum can be separated from the other species since their propodeum has basal and declivitous faces of approximately equal length. Although this is also true of M/. cyaneum workers, the mesopleuron in M. ebeninum workers is smooth and shining, while it is punctuate in M. cyaneum workers. 108 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN 147 148 FIGS. 134-148. Monomorium trageri. 134—Queen, lateral view. 135—Head of queen, frontal view. 136—Petiole of queen, posterior view. 137—Postpetiole of queen, posterior view. 138—Head of worker, frontal view. 139—Worker, lateral view. 140—Mandible of queen (schematic). 141—Male, lateral view. 142—Mandible of male. 143—Labial palp of male. 144—Mazxilla and maxillary palp of male. 145—Aedeagus of male. 146—Eighth sternite of male. 147—Ninth sternite of male. 148— Volsella of male. Scales: Top scale (1 mm) for Figures 134-139, 141. Second scale (0.1 mm) for Figures 140 and 142. Third scale (0.5 mm) for Figures 146 and 147. Fourth scale (0.5 mm) for Figures 145 and 148. Bottom scale (0.1 mm) for Figures 143 and 144. AnT GENUS MONOMORIUM 109 Monomorium trageri new species Figures 7, 20, and 133-148 Monomorium new sp. near minimum: Jones et al., 1982: 287. DIAGNOSIS. Quren: Wingless )usually(¢; mandible with 4 teeth, basal 2 reduced (Fig. 140); head smooth and shining except for rugosities near lateral margin of clypeus (Fig. 135); mesonotum lacking notal furrow. MALE: Genitalia and sterna as in Figures 141, and 145-148. Worker: Propodeum angular (basal face 1/2 as long as declivitous face); meso- pleuron punctate; petiole and postpetiole as in Figure 139; 8-12 erect to suberect setae on dorsum of alitrunk; PI 31-38 (35). DESCRIPTION. As described for M. ebeninum except as follows. QuEEN: Heap: Measurements (representing 2 localities; N=5) HL 0.62-0.70 (0.66) {0.68}, HW 0.53-0.62 (0.59) {0.53}, SL 0.38-0.55 (0.46) {0.55}, IOD 0.12-0.18 (0.15) {0.12}, OD 0.05 (0.05) {0.05}, EL 0.14-0.16 (0.15) {0.14}, MOD 0.10-0.11 (0.10) {0.10}. Structure—CI 80-97 (88) {88}, SIL 59-81 (68) {81}, SIW 63-104 (78) {104}. Eye small to moderate in size. Scape usually not surpassing occiput )or surpassing it by less than length of pedicel<. Mandible with 4 teeth (basal two reduced, Figure 140). Clypeal teeth sharp, short to moderate in length. Frontal carinae diverging slightly posteriorly. Prlostty—Setae erect to sub- erect over entire surface of head except as follows: setae suberect to decumbent on mandi- ble. Sculpture—Moderate, parallel, longitudinal rugae beginning all along lateral margin of clypeus, extending past antennal insertion and converging with frontal carina. ALITRUNK: Measurements PW 0.42-0.48 (0.44) {0.42}, PL 0.25-0.31 (0.29) {0.30}, WL 1.02-1.20 (1.13) {1.15}. Structure—Scutum and scutellum not depressed or flat. Mesopleural suture straight with no pits at either end. PI 24-28 (25) {26}. Sculpture—All sutures surrounding scutum and scutellum covered with small, dense, non-pil- iferous punctures (Fig. 134). Propodeum cov- ered with dense, small, non-piliferous punctures throughout (obscuring any trace of rugae on lower side of propodeum). Pe7rove: As in Fig- ures 134 and 136. Setae appressed on anterior surface of node, erect to suberect on dorsum, side, and posterior surface of node, absent elsewhere. Dorsum of node smooth and shining with small piliferous punctures; side covered with moderate to large longitudinal and trans- verse rugae (Fig. 134). PostrpeTio_e: As in Figures 134 and 137. Dorsum of node slightly emarginate. Setae suberect to erect on dorsum, posterior surface, side, and venter of node, absent elsewhere. Dorsal surface of postpetiole smooth and shining with small piliferous punc- tures; lower 2/3 of side (below node) with small to moderate, dense, non-piliferous punctures. GasTer: Setae of first gastral tergite exceeding level of dorsum of postpetiolar node. Mate: Heap: Measurements (representing 1 locality; N=5) HL 0.55-0.60 (0.58), HW 0:70-0:72. (0.71), SL 0.25-0.30 (0:26); TOD 0.18-0.22 (0.19), OD 0.05-0.08 (0.06), EL 0.25-0.28 (0.27), MOD 0.15-0.18 (0.17). Struc- ture—C@I 117-131 (124), SIL) 42-50 (45), SIW 35-42 (36). Maxillary palp 2-segmented (Fig. 144); labial palp 2-segmented (Fig. 143). Fron- tal carinae diverging slightly posteriorly. Sculp- ture—Moderate to large, parallel, longitudinal rugae beginning all along lateral margin of clypeus, extending to level of middle of com- pound eye. Moderate, parallel, longitudinal rugae beginning all along distal border of clypeus (between clypeal teeth) and continuing up clypeus and frons to level of antennal inser- tion. Several moderate, faint, semicircular rugae occurring on occiput and following con- tours laterally around occiput (these rugae are partially obscured by punctation). AZITRUNK: Measurements PW 0.62-0.65 (0.64), PL 0.30-0.38 (0.34), WL 1.30-1.35 (1.33). Structure—Meso- pleural suture straight with small pit present on both anterior and posterior ends. Metanotum (in lateral view) projecting to level of scutellum and propodeum. Propodeum angular, basal face 2/3 as long as declivitous face. Pilosity— Several setae suberect to erect on lower 1/3 of mesopleuron (katepisternum). Propodeum with 2 - 4 erect to suberect setae. Sculpture—Smooth and shining with large piliferous punctures on dorsum. Entire propodeum smooth and shin- ing. PeTiove: As in Figure 141. Dorsum of node flat to slightly emarginate. Setae erect to sub- erect on dorsum, side and posterior surface, absent elsewhere. Entire petiole smooth and shining; lower 1/3 of side with faint, moderate rugae. PosTPETIOLE:. As in Figure 141. Dorsum of node emarginate. Setae erect to suberect on anterior surface, side, posterior surface, and venter, absent elsewhere (including dorsum of node). Entire surface of postpetiole smooth and shining except for posterior 1/2 of node which is covered with small to moderate, non-piliferous punctures. GasTeER. Setae of first gastral tergite not exceeding level of dorsum of postpetiolar node. Geni7Ta4LiA:. As in Figures 141, and 145-148. Eighth sternite with emargination ap- proximately as deep as wide (Fig. 146). Ninth sternite with 10 erect setae (Fig. 147). Aedeagus with 12 teeth, toothed margin rounded (Fig. 110 Tue UnIversiry OF KANSAS SCIENCE BULLETIN 145). Volsella with curved digitus and reduced cuspis; cuspis with 2 setae (Fig. 148). Cozor: Head dark brown except as follows: antenna and clypeus brown. Alitrunk brown, legs yellow brown, wings hyaline. Petiole, postpetiole, and gaster brown to dark brown. Genitalia brown to yellow brown. All setae white. Worker. Heap: Measurements (representing 1 locality, N=5) HE, 0:45-0:52 (0.49), HW 0.35-0.45 (0.42), SL 0.25-0.38 (0.30), EL 0.08-0.10 (0.09), MOD 0.05-0.08 (0.07). Struc- ture—CI 75-93 (86), SIL 48-76 (61), STW 56-84 (72). Side of head convex. Scape reaching oc- ciput but not surpassing it by length greater than length of pedicel. Mandible with 4 teeth Ysometimes, basal 2 reduced¢. Clypeal teeth sharp, of moderate length. Sculpture—Smooth and shining with moderate piliferous punctures evenly distributed throughout. AZITRUNK: Measurements PW 0.25-0.28 (0.27), PL 0.15-0.20 (0.18), WL 0.45-0.53 (0.50). Structure—Pro- podeum angular, basal face 1/2 as long as declivitous face. PI 31-38 (35). Pilostty—Dorsal outline of alitrunk with 8-12 erect to suberect setae. Leg pilosity as in queen. Sculpture—Meso- pleuron covered with dense, small, non-pil- iferous punctures. Anterior propodeal suture with short, parallel, moderate, longitudinal rugae. Periote: As in Figure 139. Dorsum of node convex. Setae erect on dorsum of node, absent elsewhere. PosTreTIOLE: As in Figure 139. Dorsum of node flat. Setae erect on dor- sum and posterior surface of node, absent else- where. GasTER: Setae of first gastral tergite not projecting to level of dorsum of postpetiolar node. Coror: Head dark brown (with bluish reflections) except as follows: antenna and man- dible brown to light brown. Alitrunk brown to dark brown, legs light brown. Petiole, postpetiole, and gaster dark brown with bluish reflections. All setae white. TYPE MATERIAL. HototyPE QUEEN: Fioripa: Alachua Co., Gainesville, April 5, 1980, J. C. Trager. Holotype, deposited in NMNH, bears red, handwritten label: Monomorium/ trageri/ Holotype/ M. DuBois 1983/. Paratypes are deposited as follows: CAS (1 queen, 1 male, 1 worker), AMNH (1 queen, 1 male; 1 worker), JOCTPC (1 queen, 1 male, 1 worker), KU (1 queen, 2 males, 3 workers), LACM (1 queen, 1 male, 1 worker), MBDPG (queen, 2 males, 3 workers), MCZ (1 queen, 1 male, 1 worker), NMNH (1 queen, 1 male, 1 worker). All paratypes bear blue, hand- written labels: Monomorium/ tragert/ para- type/ M. DuBois 1983/. ETYMOLOGY. This species is named in honor of James C. Trager, who provided many specimens of it. DISTRIBUTION. This species is quite rare; it is known from only four localities in Florida. In addition to the type locality, specimens have been collected in Tallahas- see, LaCrosse, and Punta Gorda (Fig. 133). It is possible that earlier records of M. minimum from Florida (e.g., Van Pelt, 1958) may have represented collections of this species. However, no specimens from these earlier studies could be located. It is more probable earlier records referred to M. viridum which is more common in pe- nisular Florida, but does not range as far west as Tallahassee. Additional collecting in northern Florida and southern Georgia may elucidate the distribution of M. tragert. DISCUSSION. Biotocy. Although de- tails of nesting preferences are unknown (little can be inferred from 4 collections), the colony which yielded the holotype and paratypes was collected from beneath a board in a cow pasture (J. Trager, pers. comm.). The colony collected in LaCrosse (Alachua Co.) was collected in cow dung. The type colony was reared in the labo- ratory from 5 April through 20 June when portions of it were killed. At that time, numerous males and queens were present. One male and queen were preserved in coztu indicating that mating may occur between males and closely related queens. Several specimens examined are probably inter- sexes (female head and male alitrunk and gaster). These may have been caused by some form of mechanical damage to the developing individuals (W. L. Brown, Jr., pers. comm. ). Recently, James Trager provided addi- tional specimens of the Gainesville colony. A few queens are winged, indicating that genes which code for wings are still present within this population; approximately 90 % of the queens are wingless and 10% are winged. Comparisons. Monomorium trageri 1s likely to be confused with the following Ant GENUS MoNnoMoRIUM 111 species: M. ebeninum, M. minimum, and M. virrdum. Queens of M. tragert may be sepa- rated from those of M. ebeninum since the latter have a flat or concave scutum and scutellum. Queens of M. tragert may be separated from those of M. viridum since the latter have a furrow on the mesonotum. Queens of M. tragert may be separated from those of M. minimum since the former have reduced sculpturing on the head; the only rugae are along the lateral margin of the clypeus. Monomorium minimum queens have more facial sculpturing. Workers of M_. tragert may be separated from workers of the other three species since they have the basal face length of the propodeum less than the length of the declivitous face. Monomorium marjoriae new species Figures 149-154 DIAGNOSIS. QurEN: Winged; petiole as in Figures 149 and 151; scutum and scutellum not depressed; metanotum (in lateral view) project- ing to level of propodeum and scutellum; pro- podeum angular (basal face 2X length of declivitous face). WorkKER: Propodeum angular (basal face 2X length of declivitous face); meso- pleuron not punctuate; petiole as in Figure 153; PI 35-38 (36). DESCRIPTION. As described for M. mini- mum except as follows. QUEEN: HeEabD: Measurements (representing type locality; N= 3) HL 0.75-0.81 (0.78) {0.81}, HW 0.64-0.68 (0.67) {0.68}, SL 0.52-0.61 (0.56) {0.61}, IOD 0.16-0.18 (0.17) {0.17}, OD 0.04-0.05 (0.05) {0.05}, EL 0.16-0.22 (0.19) {0.22}, MOD 0.12-0.17 (0.14) {0.17}. Structure—CI 84-87 (85) {84}, SIL 67-75 (72){75}, SIW 76-90 (84){90}. Side of head straight. Scape reaching or surpassing occiput by less than length of pedicel. Prlosity—Setae erect to subdecumbent on clypeus, frons, gular region, mandible, and occiput, decumbent to appressed elsewhere. Sculpture—Small, parallel, longitudinal rugae beginning all along lateral margin of clypeus, extending past antennal insertion, and converg- ing with frontal carina. Several small, parallel, longitudinal rugae beginning between clypeal teeth, extending to level of antennal insertion. Several small, parallel, longitudinal rugae be- ginning between frontal carinae (near posterior edge of carinae) and extending towards (but not reaching) anterior ocellus (Fig. 150), ALITRUNK: Measurements PW 0.40-0.48 (0.43){0.48}, PL 0.26-0.40 (0.34) {0.26}, WL 1.30-1.40 (1.35) {1.30}. Structure—Mesopleural suture deflected dorsally at each end (resulting in U-shaped suture), with pits lacking on both ends. Propodeum angular, basal face 2X length of declivitous face. PI 20-30 (26){20}. Pilosity— Fewer than 30 erect to suberect setae projecting above dorsal outline of alitrunk. Pevroie: As in Figures 149 and 151. Setae appressed on ante- rior surface of node, suberect to erect (a few decumbent) on side, dorsum, and _ posterior surface of node, absent elsewhere. POosSTPETIOLE: As in Figures 149 and 152. Setae appressed on anterior surface of node, suberect to erect else- where (including venter). Anterior surface and dorsum of node smooth and shining with small piliferous punctures; remainder covered with, moderately dense, non-piliferous punctures. GASTER: Setae of first gastral tergite not reach- ing level of dorsum of postpetiolar node. Cozor: Head (including mandible and antenna) brown to dark brown. Alitrunk brown to dark brown, legs brown to yellow brown. Petiole, postpetiole, and gaster brown to dark brown. All setae white. Mate: UNKNown IN M. marjoriae. Worker: Heap: Measurements (representing type locality; N=2) HL 0.55-0.61 (0.58), HW 0.45-0.46 (0:46), SL 0:41-0:45 (0:43), EL 0.08-0.09 (0.08), MOD 0.05-0.06 (0.06). Struc- ture—CI 75-82 (78), SIL 74-75 (74), SIW 89-98 (94). Scape reaching or surpassing occiput by an amount less than length of pedicel. Pzlostty— Setae erect to subdecumbent on clypeus, frons, gular region, mandible, and occiput, decum- bent to appressed elsewhere. ALITRUNK: Measurements PW 0.26-0.28 (0.27), PL 0.24-0.25 (0.24), WL 0.65-0.68 (0.66). Structure—Pro- pedeum angular, basal face 2X length of de- clivitous face. PI 35-38 (36). PosTPeTIOLE: As in Figure 153. Moderately dense, non-piliferous punctures on posterior surface of node. Cotor: Head brown to dark brown, mandible and antenna brown to yellow brown. Alitrunk brown, legs yellow brown to yellow. Petiole, postpetiole, and gaster brown to dark brown. All setae white. TYPE MATERIAL. Hototyre Queen: Mexico: Nayant; Punga, May 26, 1923, H. C. Millender. Holotype (NMNH) bears red, handwritten label: Monomorium/ mar- joriae/ Holotype/ M. DuBois 1983/. Two paratype queens and 9 paratype workers collected with holotype. All paratypes bear ty THE UNIVERSITY OF KANSAS SCIENCE BULLETIN THES 151 = 152 FIGS. 149-154. Monomorium marjoriae. 149—Queen, lateral view. 150—Head of queen, frontal view. 151—Petiole of queen, posterior view. 152—Postpetiole of queen, posterior view. 153—Worker, lateral view. 154—Head of worker, frontal view. Scale: (1 mm) for Figures 149-154. blue, handwritten labels: Monomorium/ mar- joriae/ Paratype/ M. DuBois 1983/. ETYMOLOGY. This species is named in honor of my mother, Marjorie A. (Black) DuBois. DISTRIBUTION. This species is known only from the type locality listed above. DISCUSSION. Monomorium marjoriae is most likely to be confused with M. cyaneum or M. compressum which may occur in sim- ilar habitats. Additionally, M4. marjoriae closely resembles M. ebeninum in habitus; however, these species do not appear to overlap. Queens of M. marjoriae may be separated from those of AZ. compressum and M. ebeninum since in the latter two species the scutum and scutellum are concave or flat. Queens of M. marjoriae may be sepa- rated from those of M. cyaneum since the latter are wingless and the former have a propodeum with the length of the basal face 2X the length of the declivitous face. Work- ers of M. marjoriae may be separated from workers of these other species since they possess a propodeum with the basal face length 2X the declivitous face length. Monomorium talbotae DuBois Figures 155-163 Monomorium sp.: Talbot, 1975: 245; Talbot, 1979: 88; DuBois, 1980: 626. Monomorium talbotae DuBois, 1981a: 31-34. DIAGNOSIS. FEMALE: Winged; petiole con- vex (Fig. 155); scutum and scutellum convex; metanotum (in lateral view) not projecting to level of propodeum and scutellum; propodeum rounded; clypeus scarcely emarginate, lacking teeth; anterior edge of scutum not projecting over pronotum; no depression on first gastral tergite. MALE: Genitalia and sterna as in Fig- ures 156, and 160-163. DESCRIPTION. As described for M. mini- mum except as follows. FEMALE: HeEaD: Measurements (representing type locality; N= 8) HL 0.40-0.48 (0.44), HW 0.39-0.43 (0.40), SL 0:31-0:36" (0534); TOD" 0:09-0:10 -(00);-OD 0.04-0.06 (0.05), EL 0.08-0.09 (0.09), MOD 0.05-0.06 (0.06). Structure—CI 81-98 (92), SIL 74-80 (77), STW 80-92 (84). In full frontal view, head broadest at or a little below occiput. Eye small in size. Scape reaching or surpassing occiput by an amount less than length of ped- icel. Mandible with two teeth; maxillary palp 1- segmented; labial palp 1-segmented. Clypeal teeth absent (emargination greatly reduced Ant Genus MonomoriumM 113 where space between teeth should occur). Pil- osity—Setae erect to suberect over entire surface of head. Sculpture—Entire surface of head smooth and shining with small to moderate piliferous punctures. AZITRUNK: Measurements PW '0:27-0:33-(0:32), PE0.19-0°22,(0:21), WL 0.63-0.72 (0.68). Structure—Mesopleural suture deflected ventrally at posterior end. Metanotum (in lateral view) not projecting to level of pro- podeum and scutellum. Propodeum rounded, basal and declivitous faces of approximately equal length. PI 30-32 (31). Three to 5 hamuli on hindwing. Sculpture—Smooth and _ shining throughout with small piliferous punctures. PeTioce: As in Figure 155. Setae erect to suberect on all surfaces except venter which lacks setae. Entire surface smooth and shining with small piliferous punctures. POSTPETIOLE: As in Figure 155. Dorsum of node convex. Anterior subpostpetiolar process enlarged into spine, located medially on venter. Setae erect to suberect on all surfaces, including venter. En- tire surface (except dorsum of node) covered with small, dense, non-piliferous punctures. Dorsum of node smooth and shining with small piliferous punctures. Gas7TeER: Setae of first gas- tral tergite not reaching level of dorsum of postpetiolar node. Cozor: Head and mandible light brown. Alitrunk and legs light brown, except for light yellow tibiae and tarsi. Petiole, postpetiole, and gaster light brown. All setae white. Mate: Heap: Measurements (representing type locality; N=2) HL 0.39-0.43 (0.41), HW 0542-043: 5(0.42), SLE .0:32-0:35- (0:34); 10D 0.07-0.08 (0.08), OD 0.04-0.05 (0.04), EL 0.09-0.10 (0.10), MOD 0.06-0.08 (0.07). Struc- ture—CI 100-108 (104), SIL 81-82 (82), SIW 76-81 (79). Eye small in size. Mandible with 2 teeth (Fig. 157); maxillary palp 1-segmented (Fig. 159); labial palp 1-segmented (Fig. 158). Clypeal teeth and frontal carinae as in female. Pilosity—As in female. Sculpture—As in female. ALITRUNK: Measurements PW 0.29-0.30 (0.30), PL 0.20-0.22 (0.21), WL 0.63-0.68 (0.66). Structure—As in female. PI 32 (32). Wings as in female. Pilosity—As in female. Sculpture—As in female. PETIOLE: As in female. PosTPETIOLE: As in female. GasTeER: As in female. GeniTaia: As in Figures 156 and 160-163. Eighth sternite with emargination approximately 1/2 as deep as wide (Fig. 162). Ninth sternite with 10-12 erect setae (Fig. 163). Aedeagus with 7 teeth; toothed margin straight (Fig. 161). Cozor: As in fe- male. Worker: Unknown and apparently non-exis- tent in M. talbotae. TYPE MATERIAL. Type series consist- ing of 8 females and 2 males (DuBois, 1981a): MicuicaAn: Livingston Co., E. S. George Reserve, 30 June 1966, M. Talbot. Holotype (MCZ) bears red, handwritten label: Monomorium/ talbotae/ M. DuBois 1981/ Holotype/. Allotype male also in MCZ. Paratypes distributed as follows: 6 females, 1 male (MBDPC), 1 female (KU). Allotype and paratypes bear red, handwrit- ten labels: Monomorium/ talbotae/ M. DuBois 1981/ Paratype (or Allotype)/. DISTRIBUTION. This species is cur- rently known only from the type locality listed above. DISCUSSION. Biotocy. This species was discovered in three nests of M. mini- mum. It is presumed to be a social parasite in view of the morphological features it shares with other species of inquilinous Monomorium (see section on inquilinism). Habitat (from notes of M. Talbot): “‘A high, dry field where vegetation was not dense and small patches of bare sandy soil were numerous. Grasses were mostly Poa compressa and Aristida sp. A variety of scat- tered forbes included Lespedeza capitata, Lia- tris aspersa, Rumex acetosella, and Solidago spp. Patches of Polytrichum piliferum moss were large and frequent and the red-tipped lichen, Cladonia cristatella var. vestita, was characteristic.’’? (DuBois, 1981a). Comparisons. Monomorium talbotae spec- imens are easily recognized by the rela- tively small size of the female and male (compared to the size of host male and female) and by their smooth and shining integument. Monomorium talbotae females and males are most likely to be confused with those of M. inguilinum and M. per- gander. The first differs from both the other species since the metanotum does not reach the level of the propodeum and scutellum. Monomorium pergandei (Emery) Figures 164-166 Epoecus pergander Emery, 1892 (1893): cclxxvi- cclxxvil; Emery, 1895: 272-274; Wheeler, 1910: 498; Emery, 1921: 204-205; Smith, 1943> 301-302; (Smith), “19475 569-570; Creighton, 1950: 239-241; Smith, 1952: 815. 162 163 FIGS. 155-163. Monomorium talbotae. 155—Female, lateral view. 156—Male, lateral view. 157— Mandible of male. 158—Labial palp of male. 159—Maxilla and maxillary palp of male. 160—Volsella of male. 161—Aedeagus of male. 162—Eighth sternite of male. 163—Ninth sternite of male. FIGS. 164-166. Monomorium pergandei. 164—Female, lateral view. 165—Petiole of female, posterior view. 166—Postpetiole of female, posterior view. FIGS. 167-170. Monomorium inquilinum. 167—Female, lateral view. 168—Head of female, frontal view. 169—Petiole of female, posterior view. 170— Postpetiole of female, posterior view. Scales: Left (1 mm) for Figures 167-170. Right top (1 mm) for Figures 164-166. Right second (1 mm) for Figures 155-156. Right third (0.1 mm) for Figure 157. Right fourth scale (0.5 mm) for Figures 160-161. Right fifth scale (0.5 mm) for Figures 162-163. Right bottom scale (0.1 mm) for Figures 158-159. Ant GENUS MoONOMORIUM eS Monomorium pergander: Ettershank, 1966: 82-91; Wilson, 1971: 354; Smith, 1979: 1383; Du- Bois, 1981a: 36. DIAGNOSIS. FEMALE: Winged; petiole as in Figures 164 and 165; scutum and _ scutellum convex; metanotum (in lateral view) projecting above level of propodeum and scutellum; pro- podeum angular; clypeus emarginate, teeth blunt and reduced; anterior edge of scutum projecting over pronotum as in Figure 164. DESCRIPTION. As described for M. talbotae except as follows. FEMALE: Heap: Measurements (representing type locality; N= 2) HL 0.44-0.48 (0.46), HW 0.39-0.40 (0.40), SL 0.40-0.42 (0.41), LOD 0.09-0.10 (0.10), OD 0.04-0.05 (0.04), EL 0.09-0.10 (0.10), MOD 0.08-0.09 (0.08). Structure—CI 83-89 (86), SIL 88-90 (89), SIW 103-105 (104). Scape surpassing occiput by an amount slightly greater than length of pedicel. Mandible with three teeth. Clypeal teeth blunt, short. Az77TRUNK: Measurements PW 0:30-0:33) (0332); PL 1022-028 (0:25), WL 0.72-0.81 (0.76). Structure—Mesonotum with small notal furrow. Scutum and scutellum not depressed. Anterior edge of scutum projecting over pronotum as in Figure 164. Mesopleural suture deflected dorsally at posterior end and ventrally at anterior end; smal pit at each end. Metanotum (in lateral view) projecting above level of propodeum and scutellum. Propodeum angular. PI 30-35 (32). Perroze. As in Figures 164 and 165. Dorsum of node flat, smooth and shining with small piliferous punctures. Re- maining surfaces covered with moderately dense, non-piliferous punctures. PosTPETIOLE: As in Figures 164 and 166. Dorsum of node weakly emarginate. GasTeER: As in M. minimum except as follows: first gastral tergite with basal depression. Cozor: Head brown to dark brown except as follows: mandible and antenna yellow brown to yellow. Alitrunk brown, legs yellow brown to yellow. Petiole, postpetiole, and gaster brown. All setae white. Mate: Although males and females of |. pergande: were collected from the same nest, no male specimens could be located for this study. One badly damaged specimen (gaster missing) is labelled as a male in the NMNH collection. The head and alitrunk resemble those of a female. This may actually be a male if the sexes in M. pergandei are as similar as they are in M. talbotae. Worker: Unknown and apparently non-exis- tent in M. pergandet. TYPE MATERIAL. Type series consist- ing of lectotype and 6 paralectotype queens, designated by DuBois (1981a): DisTRICT OF CoLuMBIA: Washington, 12 July (year unknown but before 1892), T. Pergande. Lectotype and 4 paralectotypes in NMNH, remaining paralectotypes in MCZ. All specimens bear red, handwritten labels: Monomorium/ pergandei/ Lectotype (or Paralectotype)/ M. DuBois 1980/. DISTRIBUTION. This species is known only from the type locality. DISCUSSION. BioLocy. This species was discovered in a nest of M. minimum. It is presumed to be a social parasite in view of the morphological features it shares with other species of inquilinous Monomorium (see section on inquilinism). This species has only been collected once (almost 100 years ago), and is probably now extinct. One detail recorded, and per- petuated through the literature (i.e., Wheeler, 1910: 498; (Creighton, 1950: 239-241), was that when the colony con- taining M. pergande: was taken to the labo- ratory, Pergande noted that reproductives of M. minimum were also present in the nest. As he watched, the M. pergandei fe- males attacked and killed some of the M. minimum males. This seems rather unusual behavior if the reproductives of the two species coexisted previously in the same nest. However, Wheeler (1910: 498) points out that Pergande may have collected two colonies in close proximity to each other, with one colony containing M. pergandei reproductives and M. minimum workers, while the other colongy contained M. mini- mum reproductives. Wheeler then supposed these two colonies were combined in the laboratory and the observed behavior en- sued. Comparisons. Monomorium pergandei is easily recognized by the small size of the female (compared to the size of the host), the smooth and shining integument, the abundance of erect to suberect setae over most of the body, the spine projecting where the postpetiolar process is usually located, and the large depression on the dorsum of the first gastral tergite. Females of this species are most similar 116 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN to M. inquilinum. However, these two spe- cies can easily be separated, since the meta- notum only projects to the level of the propodeum and scutellum and there are only two mandibular teeth in M. in- quilinum. The petiolar node is flat and the anterior edge of the scutum does not pro- ject over the pronotum in M. inquilinum. These same characters distinguish females of M. talbctae, which also lack a dorsal impression on the first gastral tergite. Monomorium inquilinum DuBois Figures 167-170 Monomorium sp.: DuBois, 1980: 626. Monomorium inquilinum DuBois, 1981a: 34-36. DIAGNOSIS. FEMALE: Winged; petiole emar- ginate (Fig. 167); scutum and scutellum not depressed; metanotum (in lateral view) project- ing to level of propodeum and scutellum; pro- podeum angular; clypeus emarginate, lacking teeth; anterior edge of pronotum rounded. DESCRIPTION. As described for M. talbotae except as follows. FEMALE: Heap. Measurements {representing holotype only} HL 0.45, HW 0:39, SLE. 0:40; IOD 0:10,,OD:0.04, EL 0.10, MOD 0.08. Structure—CI 89, SIL 89, SIW 100. Scape surpassing occiput by amount slightly greater than length of pedicel. Labial palp 2- segmented. Clypeal teeth absent (clypeus re- taining emargination where space between teeth should occur). AL/TRUNK: Measurements PW 0.30, PL 0.15, WL 0.66. Structure—Mesopleural suture straight. Metanotum (in lateral view) projecting to level of propodeum and scutellum. Propodeum angular, basal face length half length of declivitous face. PI 22. Wings absent (presence of basal sclerites indicate holotype once possessed wings). Prlosity—As in M. cyaneum. Sculpture—As in M. talbotae. PETIOLE: As in Figures 167 and 169. Dorsum of node emarginate. Entire surface (except dorsum of petiolar node) covered with small, dense, non- piliferous punctures (causing a granular appear- ance); dorsum of node smooth and shining with small piliferous punctures. PosTPETIOLE: As in Figures 167 and 170. Dorsum of node emargi- nate. GASTER: As in M. pergandet. Coron: Head dark brown, antenna yellow. Alitrunk dark brown, legs brown (tibiae and tarsi yellow). Petiole, postpetiole, and gaster dark brown. All setae white. Mate: Unknown in M. inqualinum. Worker: Unknown and apparently non-exis- tent in M. inquilinum. TYPE MATERIAL. Known only from the holotype (DuBois, 1981a): Es74D0 DE Mexico: Highway 57 (between Mexico City and Queretaro), km 127 (measured from Mexico City), high desert, Cornell University Mexico Field Party, 9 August 1965 (1 female, MCZ). DISTRIBUTION. This species is known only from the type locality. DISCUSSION. Brotocy. This species was discovered in a nest of M. cyaneum beneath a stone. It is presumed to be a social parasite because of the mor- phological features shared with other in- quilinous Monomorium (see section on inquilinism). Comparisons. The Monomorium in- quilinum female is most similar to M. per- gander and M. talbotae females. It can be easily be separated from these species since the petiole is emarginate only in M. in- quilinum. Furthermore, the metanotum does not reach the level of propodeum and scutellum in M. talbotae, and the meso- notum possesses a furrow in M. pergandei. ACKNOWLEDGMENTS Throughout this investigation I have re- ceived assistance from numerous col- leagues. I am particularly grateful for the continued assistance of my thesis advisor, Charles D. Michener, who provided many useful comments and suggestions. I also wish to thank William L. Brown, Jr., Mary Talbot, Murray Blum, and William F. Buren for enlightening discussions con- cerning Monomorium. My colleagues at the State Biological Survey of Kansas assisted me in collecting ants throughout Kansas. This revision was supported in part by a grant from Sigma Xi. Finally, I wish to thank my wife, Jeri, and son, Benjamin, for assisting me on numerous collecting trips and for their boundless patience while I wrote this man- uscript. LITERATURE CITED Brown, W. L., Jr. 1943. A new metallic ant from the pine barrens of New Jersey. Entomol. News 942243-243, ANT GENUS MONOMORIUM 17 . 1964. Some tramp ants of Old World origin collected in tropical Brazil. Entomol. News (SNS), 1973. 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The first Mesozoic ants with a de- scription of a new subfamily. Psyche 65:108-114. Wotcott, G. N. 1948. Formicidae: ants. Jn The insects of Puerto Rico. J. Agric. Univ. Puerto Rico 32:810-839. Wo tcott, G. N. and L. F. Marroreti. 1937. The ant, Monomorium carbonarium ebeninum Forel, in a new role: as predator on the egg clusters of Diatraea saccharalis in Puerto Rican cane fields. J. Agric. Univ. Puerto Rico 21 1577-579. : Ke BH & s Reatrcesoceseasceseassestatetatatetatenetatenscotes oR ee RoR AR SR RE RR a SR eeahararaatanareta'e" neetocecececeseaatarerata*anaranetas wetetetetetees: etatatet ecatetetetetenecocer’ SCIENCE BULLETIN ~ Larvae of the Crane Fly Genus Tipula In North America (Diptera: Tipulidae) Mc JUL 02 196 HA RVA By UNIVERSI Fy Jon K. Gelhaus Vol. 53, No. 3, pp. 121-182 June 20, 1986 ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas University Quarterly) is an outlet for scholarly scientific investigations carried out at the University of Kansas or by University faculty and students. Since its inception, volumes of the Bulletin have been variously issued as single bound volumes, as two or three multi-paper parts or as series of individual papers. Issuance is at irregular intervals, with each volume prior to volume 50 approximately 1000 pages in length. Effective with volume 50, page size has been enlarged, reducing the length of each volume to about 750 pages. The supply of all volumes of the Kansas University Quarterly is now exhausted. 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Such requests should be directed to the author. The International Standard Serial Number of this publication is US ISSN 0022-8850. Editor Charles.D. Michener Editorial Board William L. Bloom Philip W. Hedrick Rudolf Jander Delbert Shankel Norman A. Slade George W. Byers, Chairman THE UNIVERSITY OF KANSAS SCIENCE BULLETIN June 20, 1986 Vol. 53, No. 3, pp. 121-182 Larvae of the Crane Fly Genus Tipula in North America (Diptera: Tipulidae)! Jon K. GELHAUuS Department of Entomology, University of Kansas CONTENTS ABSTRACT. . fase INTRODUCTION . = 2. = = ANGCKNOWLEDGEMENTS... 2 sea .oe ee eo oe. MATERIALS AND METHODS MorPHOLOGY : HIstToRICAL REVIEW . PE eee THE Genus T7PULA LINNAEUS. . . Kry TO SUBGENERA OF 77PULA SUBGENERIC ACCOUNTS. . SUBGENUS ANGAROTIPULA SAVCHENKO.. . SUBGENUS ARCTOTIPULA ALEXANDER. . . SUBGENUS BELLARDINA EDWARDS. . . SuBGENUS BERINGOTIPULA SAVCHENKO SUBGENUS HESPEROTIPULA ALEXANDER SUBGENUS LUNATIPULA EDWARDS... . SUBGENUS NIPPOTIPULA MATSUMURA. SUBGENUS NOBILOTIPULA ALEXANDER . SUBGENUS ODONATISCA SAVCHENKO.. . SUBGENUS PLATYTIPULA MATSUMURA SUBGENUS PTERELACHISUS RONDANL.. . SUBGENUS SAVTSHENKIA MANNHEIMS. . SUBGENUS SCHUMMELIA EDWARDS .. . SUBGENUS SERRATIPULA ALEXANDER... SUBGENUS SINOTIPULA ALEXANDER . SUBGENUS 77PULA LINNAEUS S. STR. . SUBGENUS 7TRICHOTIPULA ALEXANDER . SUBGENUS 7RIPLICITIPULA ALEXANDER SUBGENUS VESTIPLEX BEzz1 SUBGENUS YAMATOTIPULA MATSUMURA LITERATURE CITED . 122 22 123 123 125 130 130 130 134 134 136 138 140 142 eae 149 ili oe 15S 157 Ua) 162 164 166 168 170 IS 175 177 180 ' Contribution No. 1948 from the Department of Entomology, The University of Kansas, Lawrence, Kansas, 66045, U.S.A 122 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN ABSTRACT This is a taxonomic study concerning larvae of the crane fly genus Tipula. Descriptions and illustrations are given for larvae of at least one species in 20 of the 27 recognized nearctic subgenera; for four of these groups, larvae are described for the first time. A key is presented to the subgenera of 77zpula and to certain similar genera for final-instar larvae. External morphology, concentrating on the diversity found among the subgenera, and various rearing methods are discussed. Included in each subgeneric treatment is a discussion of the characters of the group, and a list of known habitats. Larvae of 7ipula occupy a wide range of aquatic to terrestrial situations and the morphology correspondingly varies among the subgenera. Overall, larval groups are in agreement with the adult subgeneric classification, although the subgenus T7ichotipula contains two extremely different types of larvae. No unique characters of the larvae were found to unify the entire genus, but certain groupings of subgenera have distinctive sets of characters. Some groups within the genus are more similar morphologically to other tipuline genera, such as Holorusia and Prionocera, than to many other subgenera of 77pula. INTRODUCTION At the time of Alexander’s comprehen- sive 1920 paper, ‘‘The Crane-flies of New York. Part II. Biology and Phylogeny,’’ the larvae of only four percent (16 species) of the nearctic species of Tzpula had been described. From 1920 to 1983, the larvae of only five additional nearctic species were described and illustrated, and the life his- tory of only one species was studied in any detail. My work represents a step toward correcting this lack of knowledge about this diverse genus, including descriptions of larvae and information about their biology. The cosmopolitan genus 7?pula contains approximately 450 nearctic species placed in 27 subgenera (Alexander, 1965). Tzpula larvae are by nature of their habitat incon- spicuous, although the larval stages may last from six months to five years. Larvae can be sufficiently abundant locally to have an appreciable ecological impact. The lar- val habitats vary tremendously within the genus. Larvae of some species are fully aquatic and are found in leaf packs or on the bottoms of streams (e.g., Nippotipula), while others occur in the wet soil of stream and lake margins, seepages, algal mats or wet mosses (e.g., Yamatotipula). Many spe- cies such as those in Lunatipula are found in soil under layers of leaf mold or other de- caying vegetation in forested areas. Some species (e.g., Serratipula) are in the soil of pastures or meadows, where they feed on roots and leaves of young grasses. A few of these may become pests in favorable years by denuding hundreds of acres of range- land, thus earning the common name ‘range crane fly.’’ Some larvae are found in dead wood at various stages of decay (e.g., Pterelachisus). Still others, such as Eremotipula (inferring from pupal collec- tions), occur in semiarid areas in the soil under rabbitbrush (Chrysothamnus) and other shrubs. All species appear to be herbivorous or to feed on decaying plant matter. Existing keys, most recently by Alex- ander and Byers (1981), will at best iden- uy larvae of nearctic 77pula to the generic level, and certain species of 77pula cannot even be distinguished from other genera of Tipulinae by these keys. All 42 subgenera are based exclusively on adult characters, and in the present study, inconsistencies are seen between adult and larval group- ings in certain subgenera (e.g., the two larval types of the subgenus 77chotipula). I have examined reared material for as many species as possible (approximately 70 species, 16 percent of the genus in the Nearctic) to ascertain the characters for each subgenus. Larvae were obtained for 20 of the 27 nearctic subgenera, and for four of these subgenera (Bellardina, Hespero- tipula, Serratipula and Sinotipula) the larvae are here described for the first time. A key to the subgenera is included, as well as information on larval biology. LARVAE OF THE CRANE FLy GENUS TIPULA 123 ACKNOWLEDGEMENTS I thank G. W. Byers, who suggested this project and has provided advice and support on numerous occasions, particularly in editing drafts of this paper. Reared and associated larvae and adults of Tipula are not often encountered in museum collections; therefore, everyone who contributed specimens has added significantly to this work. The collection of J. Speed Rogers at the Univer- sity of Michigan (UM), Ann Arbor, contains many specimens reared by Rogers; it also in- cludes the specimens used by C. P. Alexander in his 1920 paper. I thank T. E. Moore, curator of the Michigan collection, for the loan of this important material. Thanks also go to H. J. Teskey, Canadian National Collection (CNC), Ottawa, for the use of his extensive collection from eastern North America. I also used spec- imens from the following museums: Dept. of Entomology, American Museum of Natural History (AMNH), Entomology Museum, Uni- versity of California at Riverside (UCR), Ento- mological Museum, North Carolina State University (NC) and the James Entomological Collection, Washington State University (WSU). Other specimens are from the Snow Entomology Museum, University of Kansas (KU), State Biological Survey of Kansas collec- tion (SBSK), and from my personal collection (JKG). Other individuals provided certain spe- cies or other help and deserve acknowledge- ment: S. A. Teale (ST), State University of New York, Syracuse, identified many of my reared adult 7ipula from the Rocky Mountains and loaned specimens of 7. platymera; G. Pritch- ard (GP), University of Calgary, Alberta, sent specimens of 7. commuiscibilis and sacra; F. B. Brodo, Carleton University, Ottawa, loaned larvae of the genus Prionocera, which allowed me to separate Prionocera from the subgenus An- garotipula; H. M. Knizeski (HK), Mercy Col- lege, Dobbs Ferry, New York, provided specimens of 7° zgnobilis; and P. Oosterbroek, Institute of Taxonomic Zoology, University of Amsterdam, spent a number of hours helping me translate the 1957 paper of B. Theowald. Financial costs for museum visits and fieldwork were partially defrayed by two awards from the Raymond H. Beamer Memorial Fund, Univer- sity of Kansas. I would like to especially thank the staff of the State Biological Survey of Kansas for the use of certain equipment and supplies throughout the period of this study, to C. D. Michener and two anonymous reviewers for their helpful com- ments, and to C. W. Young, now of the Car- negie Museum, Pittsburgh, who spent many hours helping me locate larvae in the field during the initial phase of this study. Finally, and most importantly, I would like to express my appreciation to my wife, Evalyn, for her patience and constant encouragement. MATERIALS AND METHODS Larvae of 68 species of Tipula and of Holorusia rubiginosa and Prionocera dimidiata were exam- ined. I have reared 37 of these species one or more times. I collected larvae primarily in eastern Kansas, northern New Mexico and central California; most other collections stud- ied were made in eastern North America. The diversity of habitats occupied by larvae of Tipula requires a variety of collection meth- ods. Details are given by Gelhaus (1983). A sturdy garden trowel is indispensable for collect- ing in aquatic and terrestrial situations. Uses include clearing aside soil in forest situations; digging up saturated soils bordering aquatic habitats for sieving; or dislodging gravel and rocks on the bottom of fast-flowing streams and rivers. Both a D-frame aquatic net and forceps have limited use when collecting larvae of 77p- ula. The D-frame net is desirable when larvae inhabit leaf packs (e.g., Nippotipula) or are under rocks in rivers and streams (e.g., S?- notipula). Heavy forceps are needed for hand collecting in aquatic habitats that are too small for D-nets or sieves, such as shallow seepages or springs. Detailed examination of mosses or soft, rotting logs requires forceps. As briefly discussed by Brindle (1960c), the use of a sieve when collecting in saturated soils is highly recommended, as larvae are cryptically colored, slow moving and easy to overlook when only hand collecting with forceps. A 1 mm mesh screen is adequate for retaining all late-second instar and older larvae (Brindle suggests '/i6 of an inch). When sieved, a larva usually curls up for a short time, or if small and very active, it may move through the sieve mesh. A sieve works best in fine, sandy areas; it is not as efficient in heavily organic areas with large amounts of plant debris. In all cases, though, sieving reduces the amount of material to be sorted through and washes debris from the larvae, making them more visible. Terrestrial and semiaquatic larvae almost al- ways live within 50 mm of the substrate surface, regardless of climatic conditions. Larvae of the related genus Dolichopeza remain near the sur- face even during the winter when their habitat is frozen (Byers, 1961), and I have observed sim- ilar behaviors in larvae of the subgenera Luna- 124 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN tipula, Pterelachisus and Vestiplex. Hot, dry summer weather also does not appear to force larvae to move deeper into the substrate. 77pula (Trichotipula) stonei, in fact, develops and ma- tures during the summer, and even during a prolonged drought, larvae were found in the top 30-50 mm of dry forest soil. 77pula (Lunatipula) oxytona moves downward to 200 mm during dry periods (Rogers, 1933), but this appears to be an exceptional instance and may be related to this species’ occurrence in sandy soils which rapidly lose moisture near the surface. For species whose larval habitat is unknown, certain predictions can be made by observing the behavior of the adults or by knowledge of larval habitats for related species. Adults may occur away from the specific larval habitat. Teneral adults or ovipositing females give a clearer idea of where larvae occur, as does the activity of males seeking newly emerged fe- males. Pupal collections are useful, for pupae occur near the larval habitat or even among the larvae. In species with highly synchronous adult emergences, however, once pupae are found, few individuals will remain as larvae, and a return visit is required during the next genera- tion to secure larvae. Snap-cap glass vials (60 ml), one-third filled with soil or other habitat material and each with only one or two larvae, are excellent for field use and allow larvae to be easily recovered once in the laboratory. Self-sealing plastic bags are ex- cellent for transporting larvae over a long time period, when many larvae are found in one specific site, or if associated soil or debris from the larval habitat is needed for rearing or sieving in the laboratory. The bags should be one-third to one-half filled with substrate from the larval habitat. The collections are then kept cool until brought into the laboratory, as even short periods of heat (as in a vehicle on a warm day) will kill the larvae. Larvae from aquatic habitats should be transported in damp debris, mosses or leaf litter; although from an aquatic habitat, they are susceptible to drowning. A single microhabitat may yield more than one species of larvae; therefore, larvae should be carefully sorted before placement in rearing chambers. Larvae often expose the spiracular discs at the surface when placed in_ water, permitting identification in life with a micro- scope. Chilling the larvae for a few minutes helps if they are very active. A proper rearing container must admit fresh air and maintain an even moisture level. Both Chiswell (1956) and Byers (1961) recommended Petri dishes, but I found such dishes of limited use. Petri dishes have a small volume, are easily kept too wet, but also quickly dry out. The small volume also allows overcrowding even when there are only a few larvae per dish. Petri dishes are advantageous for observations of larval behavior, but for rearing, I prefer glass containers 40-80 mm high and 100 mm diame- ter. Such jars hold enough food (1.e., leaf litter, rotting wood, etc.) from the habitat to allow late-third or fourth-instar larvae to develop to the pupal stage. Moreover, they do not require daily checking for food depletion and moisture control. Aquatic species need only wet litter or, at most, a small amount of free water; terrestrial species should be kept just damp. Ideally, one larva should be kept per container; practically, I have kept up to six larvae per jar. All Tipula reared have been herbivorous or saprophagous. Wet, decaying leaves for aquatic species and damp leaves or debris for terrestrial larvae suffices for most; some species of Yamato- tipula, Platytipula and Triplicitipula will accept and feed readily on young green shoots of common weeds (Young, 1981; Gelhaus, per- sonal observation). When conditions are ade- quate, larvae remain buried within the debris or soil during the day. Although in most cases it is best to try to duplicate the natural conditions of the larval habitat in the laboratory, in some situations this is not desirable. It is not necessary to maintain species from rapidly flowing streams or rivers, such as some Sinotipula or Nippotipula, in con- stantly aerated water. Gordon Pritchard (an litt. ) reared larvae of 7. (Sinotipula) commiscibilis in containers of wet leaves at 15 degrees C., although they were originally collected from a fast-flowing, cold, mountain stream. Many spe- cies are influenced by photoperiod, particularly when about to pupate; they should be exposed to a source of natural light or to artificial light of similar periodicity (e.g., Platytipula). Since a female 77pula may lay 200-300 eggs, rearing larvae from eggs would appear to be ideal for obtaining many larvae of a single species. Unfortunately, the life histories of many species cause problems in rearing and in main- taining laboratory cultures. Some mated fe- males are reluctant to lay. Although pinching the cervical region of the female will often induce egg laying, it is not reliable. Eggs of many species undergo an obligate diapause that is broken only when they are exposed to certain photoperiods, temperatures or moisture condi- tions. For example, Hartman and Hynes (1977) found that the eggs of 7. (Triplicitipula) simplex would only hatch after exposure to a certain photoperiod and alternating wet and dry condi- tions. I found that eggs of an undescribed LARVAE OF THE CRANE FLy GENUS TIPULA 1255 species of 7. (Platytipula) only hatched after exposure to cold and typical winter photoperi- ods. Other groups have eggs which hatch soon after being laid (approximately one week). Sub- genera with brief egg stages include Nippotipula, Bellardina and most species of Yamatotipula. Eggs of T. (Nobilotipula) nobilis had no apparent di- apause but took approximately three weeks to hatch. Eggs of Lunatipula, Pterelachisus and Sav- tshenkia appeared to have a developmental di- apause but the cues breaking the diapause are unknown. First-instar larvae have a high initial mor- tality, and many eggs never hatch. After larvae start feeding, though, provided food depletion and overcrowding are prevented, mortality 1s minimal and they usually develop rapidly. For example, 7. (Yamatotipula) tricolor completed a generation in 69 days in the laboratory. As larvae reach maturity, it 1s important to watch for inactive larvae or teneral pupae. As noted by Byers (1961) for the tipuline genus Dolichopeza, larvae prior to pupation are con- tracted and inactive. As the pupal molt becomes imminent, the larval head is deflected ventrally and pupal structures become visible through the larval cuticle. Most aquatic larvae leave the water to pupate in drier soil; most pupae im- mersed in water quickly die. Some species have an obligate, fourth-instar, larval diapause in which they remain contracted and inactive dur- ing the summer. I have observed this prepupal diapause in certain species of Yamatotipula (e.g., T. sayt), all reared species of Platytipula and T. (Nippotipula) abdominals. The best association of immature and adult stages of a species is from the last larval skin, pupal skin and emerged adult of a single indi- vidual. All the morphological characters that are needed for identification can be located on the shed skins and there is little likelihood of an erroneous association. The last larval skin is usually found near the posterior end of the pupa and may be crumpled into a compact mass. If found within one or two days after pupation (pupa will still be white or light brown), the skin can be easily stretched and cleaned in water. If the skin does not tear easily when stretched or is known to have been recently shed, I place it in water in an ultrasonic cleaning appliance and vibrate it for 10-15 seconds. The cleaned skin is then quickly restretched and stored in 70-80 percent alcohol to which a small amount of glycerin has been added. Soon after the first pupae are found, I pre- serve some of the remaining larvae. Chiswell (1956) and Brindle (1960c) recommend preserv- ing larvae in Pampel’s fluid, but I have found that it is satisfactory to kill larvae by pouring water near the boiling point on them, then transferring them to storage in alcohol. This heat treatment is necessary for the expansion of the spiracular disc. Killing larvae in alcohol is unsatisfactory and results in shrunken, con- tracted specimens. I rarely preserve larvae in the field but, when necessary, Kahle’s solution gives reasonably good specimens. Before stor- age in alcohol, killed larvae should be placed in the ultrasonic cleaning appliance for 10-15 sec- onds to remove fine dirt particles. Pupae should usually be isolated from larvae, particularly if the rearing container is crowded. Pupae are delicate and should be handled as little as is possible. They can be transferred to a layer of fine damp sand or soil in a large vial with a loose fitting cap. The emerging adult needs a vertical space at least twice the length of the pupa to develop and expand properly. Once the adult has emerged, the pupal skin can be soaked for 5-10 minutes in water, then washed in a stream of water or placed in the ultrasonic vibrator. Unlike the debris on a larva or larval skin, dirt on a pupal skin is difficult to remove completely. MorPHOLOGY Larvae of 7ipula are elongate and cylindrical, tapering slightly at the cephalic end (Fig. 3). Mature larvae range from 12 to 60 mm in length. The black head is only partially visible, as the larval skin is attached near the anterior end of the head. The posterior end of the larva has short to long lobes surrounding two conspic- uous spiracles. Beneath this spiracular area are often whitish lobes, varying in number and length, surrounding the anus. The body varies from white to brown, occasionally having a pinkish tint. Tufts of hairs and dark setae may also be seen, or the larva may appear completely hairy. Sometimes there are patterns of longitu- dinal lines or spots on the abdomen. The head varies slightly from group to group, and the characters that vary are not well-suited for identification purposes. It is not described for any of the taxa treated here. A treatment of the head and its parts is found in Chiswell (1955) for Tzpula and in Byers (1961) for a related genus, Dolichopeza. Three thoracic and eight abdominal segments are clearly visible, although there are actually ten abdominal segments, according to Byers (1961). The obvious segments can be recog- nized by the transverse row of setae found on each segment, dorsally and ventrally. The pro- thoracic segment has two rows of dorsal setae, but this does not indicate two segments (Byers, 126 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN // VA S J Sai 1 =a & (\ y x : hn, ‘ Se —— =e a ‘ vi ovine ovate we = < Figs. 1-3. General morphology of Tipula larvae. 1, spiracular area, aquatic larva. 2, spiracular area, terrestrial larva; AN-anus, AP-anal papilla, DL-dorsal lobe, LL-lateral lobe, MB-marginal band, SP- spiracle, VL-ventral lobe. 3, 7. stone, lateral aspect, abdominal segments I-VIII labelled; length 18 mm. 1961:734). Abdominal segment LX contains the spiracles and the immediately surrounding re- gion. The anus and the perianal region con- stitute abdominal segment X, as in the adult. Abdominal segments I-VIII each appears sub- divided, with a large, anterior portion (annulus) and a narrow, posterior section containing the dorsal and ventral macrosetae (Figs. 3, 16). The cuticle is tough, hence the common name, “‘leather-jackets.’’ Strong sclerotization is almost always restricted to the lobes surround- ing the spiracles; the one exception is in the 7° (Lunatipula) disjuncta group, in which dark scle- rites extend from the lobes onto the dorsum of abdominal segment VIII (Fig. 56). The cuticle itself is light brown; any darker regions are caused by concentrations of hairs. Chiswell (1956) was the first to treat the macrosetae in detail. He illustrated setal ar- rangements for various genera of Tipulinae and consistently applied a numbering system for the setae which I have adopted here; the homolo- gues can be traced from genus to genus. When specimens are roughly handled, the macrosetae can be broken off. Although the positions of broken setae can be determined from the basal sockets, information concerning setal length, thickness, and color is lost. Break- age 1S a primary reason for concentrating on abdominal setal arrangements; thoracic ar- rangements are found on only three segments, with the same arrangement repeated on the mesothorax and metathorax. In contrast, ab- dominal arrangements are repeated on the first seven abdominal segments, so that breakage 1s less of a problem. Macrosetae of aquatic larvae are long, thin, light to dark brown (Fig. 9), and may be ob- scured by dense hairs. Certain aquatic groups possess setae that are branched (setae D6, L3, V1), with the distal halves of these setae divided into two to four branches (Figs. 9, 66). Branched setae have not been noted or illustrated by other authors, although groups in which the setae occur have been described. Macrosetae of terrestrial larvae are almost always short, thick and dark brown to black (Fig. 81). Larvae of the 7. (Lunatipula) disjuncta group and of subgenus QOdonatisca are notable exceptions as the setae are elongate and yellow (Fig. 53). Because of the lack of long hairs on terrestrial larvae, the macrosetae are not usually obscured by hairs. The arrangement of setae often differs be- LARVAE OF THE CRANE FLy GENUS TIPULA 12. tween groups, but seems to be constant within a group (or, at least, within a particular species). Three sets of setae are present on each abdomi- nal segment: dorsal, ventral and pleural. The dorsal and ventral setal arrangements are bilat- erally symmetrical. Generally, there are six pairs of dorsal setae in two rows (Fig. 4a), five pairs of ventral setae in two rows (Fig. 4b), and four pleural setae (Fig. 4c). Seta L3 is below seta L2 and seta L1 is most often behind seta L2, but in Nodilotipula and the palearctic 7. (Platytipula) melanoceros these three setae are in vertical alignment (Fig. 67). Most aquatic or semiaquatic larvae possess the full complement of setae, but reduction in setal size, or complete absence of certain setae, is frequently seen in terrestrial larvae (e.g., Figs. 37, 54). Setae D4, D5, L1, L3, V1 and V5 are often considerably shortened and setae D5 and V1 are occasionally absent. In addition to macrosetae, larvae of T7pula exhibit various kinds, sizes and patterns of finer hairs. Hairs are found on every segment except part of abdominal segment IX and most of segment X. The thoracic segments are usually covered by rather uniform, appressed silky hairs which provide no useful characters. On the other hand, hairs on abdominal segments I-VII exhibit much variation between groups and have been used in this study. On segment VIII and on the spiracular lobes they are usually long and uniform; hairs on the pleura are scattered and usually very short. The hairs can be separated into three groups: macroscopic hairs, long microscopic hairs and short microscopic hairs. An idea of their sizes in ~I relationship to each other and to macrosetae can be obtained from Figs. 105 and 147. In almost all larvae, the hairs are cylindrical and narrow, without longitudinal ridges, and usually dull brown (Fig. 8). In Nobilotipula, however, they are flattened, relatively broad, with longitudinal ridges and are reflective (Figs. 70, 71). Macroscopic hairs are easily seen under low magnification and often equal or exceed the length of the macrosetae. They can be distin- guished from setae because hairs lack the dis- tinctive basal sockets. Although usually single, macroscopic hairs are sometimes organized into clusters or tufts. Hair clusters are made up of many hairs and surround setae D4, D5 and V2 (Fig. 147). They also may be found on abdomi- nal dorsum VIII of Yamatotepula and other groups (Fig. 151) or on the apices of the spi- racular and accessory lobes of Nippotipula (Fig. 60). They also form distinctive abdominal pat- terns in certain species of Yamatotipula (e.g., pruinosa, see Chiswell, 1956). Hair tufts are found only in the subgenus 77pula and are made up of a few, closely set dark hairs located near setae D5 and V2 but not surrounding them (Figs. 116, 117). The predominance of mac- roscopic hairs varies with the group but they are usually abundant only in aquatic larvae. For example, larvae of the subgenus Angarotipula and the 7. (Trchotipula) oropezoides group are covered almost exclusively with these hairs (Figs. 8, 124). In Schummelia, they form a semicircular band on the dorsum of abdominal segment VIII (Fig. 98). Short microscopic hairs are always organized into groups, usually short to long, transverse Fig. 4. Abdominal macrosetal arrangements. a, dorsal setae. b, ventral setae. c, pleural (= lateral) setae. 128 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN rows (Figs. 40, 47), and individual hairs may be difficult to discern at lower magnifications. In most terrestrial subgenera, such as Lunatipula or Pterelachisus, the body is almost exclusively cov- ered with these hairs; a similar condition is found in the aquatic subgenera Nippotipula, Sinotipula and Arctotipula. Short microscopic hairs are entirely absent in T7pula (Trichotipula) stoner. and the subgenus Schummelia. Light and dark short hairs sometimes form contrasting patterns, e.g., a spotted pattern in 7° ( Yamatoti- pula) caloptera (Fig. 150), dark longitudinal lines in 7. (Y.) strepens (Fig. 151) and irregular designs in Sinotipula (Fig. 111) and certain spe- cies of Savtshenkia. Long microscopic hairs vary in length; usu- ally they are much shorter than macroscopic hairs but sometimes are intermediate between macroscopic and short microscopic hairs. These long hairs are always single, unlike the short hairs, and are not found in most subgenera. All three classes of hairs occur together in 7. ( Yama- totipula) strepens (Fig. 147), but long microscopic hairs predominate in Schummelia (Fig. 96) and T. (Trichotipula) stoner (Fig. 126). The abdomen often has cuticular outgrowths that are useful for differentiating certain groups. In Arctotipula, abdominal segments II-VII each possess two pairs of lateral tubercles and one ventral tubercle, all of a similar conical shape (Figs. 15, 16). Nippotipula has one or two pairs of elongate tubercles on abdominal segment VIII and two pairs of prominent outgrowths, which I call accessory lobes, anterior to the normal six spiracular lobes (Figs. 60, 62). One species of Savtshenkia (unreared) has many small tubercles on the dorsum and pleura, each with a small group of hairs at its apex (Fig. 91); possibly these function to anchor the larva among aquat- ic mosses in the fast-flowing streams where it is found. There is a pair of small, lateral swellings in Schummelia on segment VIII under the band of hairs (Fig. 99). In the subgenera Nippotipula and Stinotipula and 7. (Yamatotipula) caloptera, transverse cu- ticular swellings are found on abdominal seg- ments II-VII (Figs. 62, 108). These might be termed ‘‘creeping welts,’ although the swell- ings in T?pula lack recurved hooks or spinules such as those found on creeping welts of other genera (e.g., Limonia). They do appear to have some type of anchoring function in 77pula, as all three groups live in flowing water where a cuticular swelling acting as a wedge between rocks, leaf debris or mosses might maintain the larva’s position against the current. Taxonomically, the most important part is the spiracular region. This consists of the spiracular disc (segment IX, including the two prominent spiracles and the surrounding area) and the six spiracular lobes arising from the posterior bor- der of segment VIII. Some controversy has arisen concerning which segment bears the spi- racles. For example, Teskey (1981) claimed the spiracles are on segment VIII, as did Alexander (1920) and others. I agree with the view of Byers (1961) that the spiracles are on a small segment IX, part of which is later invaginated inside (forming the ‘“‘spiracular yoke’’) during the pupal stage. The spiracular lobes show great variation in size, structure and sclerotization but are con- stant in number and general position. The lobes are paired: two dorsal lobes, two lateral lobes (although they are often slightly dorsolateral) and two ventral lobes (Figs. 1, 2). In aquatic and some semiaquatic larvae, the spiracular lobes are all similar in shape and generally in size. They close against one another and around but not against the spiracles. In terrestrial lar- vae, the dorsal and lateral lobes are usually similar in shape, and sometimes size, but the ventral lobes are often smaller and different in shape than the other lobes. When closed, the ventral lobes, at most, are appressed against the spiracles, not against the other lobes. In aquatic species the ventral lobes are directed ventrally or posteriorly (1.e., sclerotized region facing posteriorly or dorsally, Fig. 1), whereas the ventral lobes in terrestrial species are directed dorsally (i.e., inner sclerotized region facing nearly anteriorly, Fig. 2). Aquatic species have a glabrous region on the posterior surface of the spiracular disc that extends to the apices of the spiracular lobes. This area may be lightly to heavily sclerotized, with virtually no darkened pattern in Sinotepula (Fig. 110), to a distinct pattern in Angarotipula (Fig. 5). Many aquatic species have a dark median line on each ventral lobe and often dark, marginal bands on every lobe (Fig. 145). Ter- restrial species always have the inner surface of the ventral lobes glabrous and often patterned but without dark, median lines. The dorsal and lateral lobes vary from nearly unsclerotized in the 7. (Trichotipula) stoner group or Pterelachisus (Fig. 84), to having large sclerites present but some areas unsclerotized in Serratipula (Fig. 100), and to sclerotization of the entire posterior surfaces, forming acute points at the apices in Lunatipula, Vestiplex and Odonatisca (Figs. 41, 72, 140). Two taxa, Nippotipula and Arctotipula, ap- pear anomalous in that they are aquatic but have the posterior surfaces of all the lobes and the spiracular disc generally unsclerotized and covered with short hairs (Fig. 14). The aquatic LARVAE OF THE CRANE FLy GENUS TIPULA 129 and semiaquatic species nearly always possess a border of apparently hydrophobic setae around the outer margins of all spiracular lobes. In Angarotipula, these hairs are long and dense (Fig. 5); in Yamatotipula, they are shorter (Fig. 145), and in some palearctic species of Savtshenkia the hairs are very short. In Beringotipula, the short hairs encircle only the spiracular disc but do not border the lobes (Fig. 27). The border of setae is absent in fully terrestrial larvae. A number of functions have been attributed to the spiracular lobes, although none has been investigated in detail. The similarly-shaped lobes of aquatic species, with the well-developed border of hydrophobic setae, appear suited for closing around the spiracles and trapping an air bubble. The air bubble may serve to prevent the entrance of water into the spiracles (Theowald, 1957), or may act as a physical gill (Teskey, 1981;Pritchard, 1983). Glabrous — spiracular lobes with well-developed hydrophobic setae, when opened at the water surface, can keep a larva afloat, particularly if it is a small larva in the second or third instar. Brindle (1960c) dis- agrees with the view that the enclosed air bubble might prevent the entrance of water into the spiracles, but suggests that in semiaquatic hab- itats soil would be excluded. In terrestrial larvae, only the ventral lobes are closed against the spiracles, apparently prevent- ing the entrance of water or soil particles into the spiracles. The heavily sclerotized dorsal and lateral lobes of Lunatipula, Odonatisca and others probably function in locomotion; the sclero- tized, pointed apices might be thrust against soil and debris (i.e., when the lobes are slightly contracted posteriorly) and serve as a brace, allowing the larva to extend its forward region. The unsclerotized lobes of Arctotspula and Nippotipula appear to have a quite different use according to Pritchard and Stewart (1982). They found the spiracles of these aquatic taxa to be non-functional with the spiracular lobes serv- ing as a site for cutaneous respiration. This may also be a function for the various abdominal swellings and accessory lobes of both these taxa. The spiracles vary in size, shape and color. Nearly all groups possess circular or slightly elliptical spiracles (Figs. 125, 136), but those in Tipula (Trichotipula) oropezoides are distinctly tri- lobed (Fig. 119). The spiracles are small and widely separated in Arctotipula and some others (Fig. 14), and quite large and close together in a number of groups, including Platytepula and Bellardina (Figs. 20, 76). The spiracles have a central circular area, usually dark brown or black, and an outer brown ring with many slit- like openings (aeropyles). The spiracles of Arcto- tipula and Nippotipula are thickened and raised slightly above the surrounding surface (Fig. 14); most other groups have spiracles which are nearly flush with the surface. The anal segment, containing the anus, is a pale, membranous region below the spiracular lobes and disc. It is always bordered basally by a thin, brown or black line called the marginal band (Fig. 2). In one subgenus, Vestiplex, there is an additional band extending between the anus and the more ventral papillae (Fig. 140). The anal opening is transverse. The various lobes on this segment, called anal papillae, are taxonomically important. They vary in num- ber, placement and size and are completely absent in a few taxa. The most usual arrange- ment is a pair of lateral papillae and one or two pairs of median papillae (Figs. 99, 149); some- times the median papillae are somewhat ante- rior to the lateral one, not strictly medial in position (Fig. 75). The size, shape and number of the papillae are distinctly correlated with habitat. They are large, elongate and/or numer- ous in aquatic species (Figs. 6, 90) and reduced in size and number in semiaquatic and ter- restrial species (Figs. 57; 1/13). Brndle7 (1957; 1960c) discussed the specialization of the pa- pillae and postulated an osmoregulatory func- tion; Pritchard (1983) cited the possibility of their use in gas exchange (hence the earlier use of the term ‘‘anal gills’’). The stouter medial papillae in terrestrial species are of use in locomotion, according to Chiswell (1956); they act as a pushing structure in the forward exten- sion of the body, similar to the function postu- lated for hook-like dorsal and lateral lobes. The overall morphology of 7Tzpula larvae does change considerably from the first through the fourth (final) instars, and this work is based only on the morphology of the fourth-instar larva. The first-instar larva usually lacks distinct spi- racular lobes, and the sclerotization of the spi- racular region is rudimentary (see Hennig 1950, Fig. 205). The first instar also has a sim- plified macrosetal arrangement and lacks most hairs. The anal papillae may also be reduced in number from that of later instars. Second-, third- and fourth-instar larvae have spiracular regions of similar shapes, but the sclerotized patterns may change among these instars; in addition, the abdominal hairs increase in number and complexity as the larva gets older. Therefore, second and third instars may possi- bly be identified successfully using characters found in this work, particularly if the distinctive taxonomic features are not hair or sclerite pat- terns. First-instar larvae cannot be identified using the characters employed in this work. 130 Tue UNIvERSITY OF KANSAS SCIENCE BULLETIN HistoricAL REVIEW Much of the early taxonomic work on larvae of Tipula concerned only one or two species, with inadequate descriptions or illustrations. Alexander’s 1920 paper remains the most rele- vant of these early works, for he comparatively treated the larvae of twelve species of nearctic Tipula and recognized the value of using a variety of characters. J. R. Chiswell (1956) published the first major study of larval Tipula. Dealing with a small, local fauna (Great Britain), he was able to rear over forty percent of the species. He pub- lished complete descriptions with clear drawings and presented a key to the specific level for the fourth (final), larval instars. This study was quickly followed by a similar paper by The- owald (1957), which overlapped with Chiswell’s work in some of the species treated, but addi- tionally considered the larvae within a sub- generic Classification. Theowald was the first to place the larvae of European Tipulinae in sub- genera and species groupings and to discuss relationships among subgenera, based on larval characters. In the same time period, A. Brindle pub- lished a series of smaller papers (1957-1960) describing the larvae of British 7zpula not de- scribed previously. Brindle (1960c) gave a key to the larvae of almost 50 British species and discussed the transition of larvae from terrestrial to aquatic habitats. Savchenko (1954, 1961, 1964) also illustrated various palearctic Tzpula larvae; in 1966 (translated into German by Theowald and Theischinger, 1979) he con- structed a phylogeny for the European tipuline groups, based in part on characters of the larvae. THE Genus T7PULA LINNAEUS A useful diagnosis for larvae of the entire genus Tipula cannot be constructed, as every character varies enormously within the genus, and may be found in a similar state in other tipuline genera. Only a few genera of this subfamily can be separated from all 7zpula on the basis of characters which do not occur somewhere within 77%p- ula: Nephrotoma has distinct, thickened pro- thoracic welts; Leptotarsus (Longurio) has pinnately branched anal papillae; and Brachypremna shows a distinct contrast be- tween short, thoracic hairs and long, dense abdominal hairs. Yet these genera are very similar in other characters to certain larvae of Tipula. For example, the mostly un- sclerotized spiracular area of Nephrotoma is similar to that of 7. (T7ichotipula) stoner; the same area of Leptotarsus is similar to that of T. (Arctotipula) and the spiracular area of Brachypremna suggests that of 7. (Schum- melia) and T. (Trichotipula) oropezoides. The remaining tipuline genera are not easily separable from Tzpula. The similarities be- tween larvae of the genus Prionocera and those of 7. (Angarotipula) are remarkable, and the two groups can only be separated on the basis of a minor difference in hairs. A similar case exists with the genus Holo- rusta and the subgenera Platytipula and Bel- lardina. Larvae remain unknown for approxi- mately one-fourth of the nearctic subgen- era of Tipula; these subgenera are Evremotip- ula, Eumicrotipula, Labiotipula, Lindnerina, Nesotipula, Setitipula and Tipulodinodes. Nearly half of the nearctic subgenera have had the larvae of only one or two of their included species reared and associated. Therefore, it seems premature to offer subgeneric descriptions now, and instead I describe one or more species within each group, and discuss the characters of the subgenus based on all known larvae. The following key to the subgenera of Tipula should be used in conjunction with the generic key by Alexander and Byers (1981). Larvae of Tipula will key to cou- plets 9 and 14 in the generic key; they can then be further separated to the subgeneric level using the key presented here. Larvae of the genera Prionocera and Holorusia can- not be reliably separated from those of Tipula in Alexander and Byers (1981) and are therefore included in the subgeneric key for Tipula. Distributions given in the key refer to the entire subgenus, not indi- vidual species or species groups. The key is designed for identifying fourth-instar lar- vae only. KEY TO THE SUBGENERA OF T7PULA AND SIMILAR GENERA 3 Usually six to eight anal papillae (Figs. 90, 149), occasionally four (Figs. 99, 123), all papillae with distinctly narrowing apices; spi- 3(2) LARVAE OF THE CRANE FLY GENUS TIPULA Sul racular lobes with developed border of setae (exceptions: Nip- potipula, Arctotipula, Figs. 14, 60); all spiracular lobes usually similar in size and shape (Figs. 89, 145); lateral lobes equal to or smaller than ventral lobes (Figs. 5, 68, 119); when closed, all lobes are against one another; in aquatic or semiaquatic situations. ...... 2 Four or fewer distinct papillae (Figs. 27, 57, 140), median and often lateral papillae reduced to broad protuberances; spiracular lobes without border of setae (ex- ceptions: Tzpula s. str., Savtshenkia, in part, Fig. 113); dorsal and lat- eral lobes usually not similar in size and shape to ventral lobes (Figs. 84, 136); ventral lobes close against spiracles, but not against dorsal and lateral lobes; in semiaquatic to terrestrial situations! 7. . > 3% 15 Border of setae around spiracular lobes reduced or absent (Figs. 14, 60); posterior surfaces of spiracular lobes with little sclerotization and covered with short microscopic hanns<(1e7 Olt) aes eee ote ee: 3) Border of setae around spiracular lobes strongly developed (Figs. 5, 145); posterior surfaces of spi- racular lobes sclerotized and glab- GOMS 3 oe ethene rane ae te eee eses ae Transverse swellings on dorsum and venter of each abdominal seg- ment I-VII (Fig. 62); two pairs of accessory lobes in addition to usual six spiracular lobes (Fig. 60); all spiracular and accessory lobes (ex- cept dorsal) with at least bifurcate apices; spiracular lobes with re- duced border of setae; eastern and central North America =. 2 22 2: Seach aCe te Nippotipula Conical swellings, one ventral and two lateral (Figs. 15, 16) on each abdominal segment II-VII (re- duced or absent in some species); accessory lobes and border of setae absent from spiracular area (Fig. 14); spiracular lobes with simple 4(2) 5(4) apices; northern North America . ES See Arctotipula Abdominal segment VIII with dor- sal semicircular row of macroscopic hairs and pair of broad lateral swellings (Figs. 98, 99); posterior surface of each dorsal lobe with extensive dark sclerite (Fig. 93); four, short anal papillae; wide- Spread 4 an, sce eee Schummelia Abdominal segment VIII without row of hairs; dorsal lobes with usu- ally only light brown sclerites; usu- ally six to eight anal papillae . . 5 Abdominal segment II-VII each with transverse ventral swelling, width often twice height (Fig. 108); abdominal dorsum with light and dark patches of short microscopic hairs, the lighter hairs forming a pair of pale, longitudinal, marginal lines or an ‘‘H’’-shaped pattern (Fig. 111); macroscopic hairs ab- sent; spiracles separated by more than 1.5X diameter of a spiracle (usually twice or more); spiracular lobes mostly unmarked (Fig. 110); western North) Americas te AE is ce ee eee Stnotipula No conspicuous transverse abdom- inal swellings (exception: T. (Yamatotipula) caloptera, with swell- ings, length 3X height, Fig. 150); abdominal dorsum usually unpat- terned or, if pattern present (Figs. 150, 151), then not as above and macroscopic hairs present; spira- cles separated by less than 1.5 X diameter of a spiracle; spiracular lobes usually distinctly marked (Fige76,"1:19) ae eee ee 6 Spiracular lobes each 2-3 X as long as basal widths (Fig. 5)...... 7 Spiracular lobes each less than twice as long as width at base (Figs. Abdomen with abundant mac- roscopic hairs (Fig. 8) and few mi- croscopic hairs; northern North ASNCLICae aes here Angarotipula Abdomen with rows of short micro- scopic hairs (Fig. 7), macroscopic 10(8) 11(10) 12(11) THE UNIVERSITY OF KANSAS SCIENCE BULLETIN hairs only around bases of mac- rosetae; northern North America eee Ae ee genus Prionocera Four anal papillae (Fig: 123). - 9 Six or more anal papillae... . 10 Spiracles trilobed (Fig. 119); ab- domen covered with macroscopic hairs (Fig. 124); microscopic hairs absent; widespread _. Trichotipula, in part (oropezordes) Spiracles oval or circular (Fig. 145); macroscopic hairs only in isolated clusters (Fig. 147); microscopic hairs abundant and generally dis- tributed> widéspreadvas .)s.... 22 . Yamatotipula, in part (ludoviciana) Eight, subequal anal papillae (Fig. 90); widespread Letras. a tome Savtshenkia, in part Six-anali:papillae. ts. sc 2nd. =< 11 Microscopic and macroscopic hairs distinctly flattened (Fig. 70, 71), broad and reflective of light; setae L1-L3 vertically aligned (Fig. 67); éastern-North America... .... nee eRe bce nayl hd 95 SS fs Nobilotipula All hairs cylindrical and narrow (Fig. 147); pleural seta L1 poste- rior to seta L2 (Figs. 19, 146), setae L1-L3 not vertically aligned (ex- ception: some species of Platytipula) Se IO ter ee ee ae ey en ae ae 1 Dorsum of abdomen primarily with long and short microscopic hairs, short hairs in rows (Fig. 147); macroscopic hairs in distinct clusters around setae D4, D5, V2 (Figs. 147, 148) and on dorsum of segment VIII (anterior to separa- tion between dorsal and _ lateral lobes); often a conspicuous pattern on dorsum of abdomen formed by microscopic and (rarely) mac- roscopic hairs (Fig. 150, 151); pos- terior surface of each lateral lobe with at most a faint median line (Fig. 145), setae of spiracular lobe hair border short, setal length '/2 or less of ventral lobe width; wide- spread Yamatotipula, in part Dorsum of abdomen primarily 13(12) 14(13) 15(1) 16(15) with macroscopic hairs, with at most indistinct clusters around setae D4 and V2 (Figs. 17, 18, 74, 79); microscopic hairs not in com- pact rows; no conspicuous hair pat- terns on abdomen; each lateral lobe usually with a dark median line, although lhne may be short (Figs. 20, 76); setae of spiracular lobe hair border long, setal length ‘/s or more of ventral lobe basal WiGUELa gS £5 &, 02 Was 8 Mh mee, ge 13 Lateral and one medial pair of anal papillae curled dorsally around ab- dominal segment VIII (Fig. 23); abdominal setae D5 and V2 ob- scured by macroscopic hairs (Figs. 17, 18); each dorsal spiracular lobe sometimes with median dark line CBG DOM Caines crite os A ee ge 14 All anal papillae directed laterad or ventrad, not dorsad (Fig. 75); setae D5 and V2 not obscured by mac- roscopic hair, all setae clearly visi- ble (Figs. 74, 79); each dorsal lobe without dark median line (Fig. 76); widespread. tes = © Cadeguala Mydrosomella oO _ WY W Mydrosoma © @ (a ) a a —_ Zz Ptiloglossidia Figure 1. Cladistic relationships of the genera of Diphaglossinae. The letters represent apomorphic characters listed in the section on cladistics; numbers following certain letters represent steps in variables having a series of apomorphic conditions. eroup have been prepared by McGinley (1981:285 and in Otis et al., 1982). Larvae of the genera Caupolicana, Crawfordapis, Ptiloglossa, and Cadeguala (with Policana) were included. The 1982 cladogram 1s in complete agreement with Figure 1 in the relative positions of branching points. For the 1981 paper the same is true of the preferred cladogram (Crawfordapis larvae were not included) although in an alternate cladogram Caupolicana is the sister group to all the others. Thus the cladogram shown in Figure 1 based on adult characters is strengthened by independently prepared cladograms based on larval characters. Un- fortunately no larvae of Dissoglottini are known. A key to the genera of known mature larvae was presented by McGinley (1981) and the larva of Crawfordapis was described in Otis et al. (1982). Key TO THE TRIBES OF DIPHAGLOSSINAE (Modified from Michener, 1966) 1.. Pre-episternal complete; flagellomere 1 nearly as long as to longer than scape, much longer than . Caupolicanini groove others, petiolate — Pre-episternal groove absent below scrobal groove; flagellomere 1 much shorter than scape, less than twice as long as middle flagellar segments, not or only moderately petiolate. 2. Notaulus represented by deep groove in anterior part of meso- scutum; malar space nearly one third as long as eye or longer... .. ee ae ee ee ee Diphaglossini — Notaulus weak or absent; malar space short or absent . . Dissoglottini There is no important new material to add to the account of Caupolicanini pub- lished in 1966, except that Crawfordapis luctuosa (Smith) has now been found as far south as western Panama and as far north as the state of Jalisco, México, and its larva and nesting biology have been studied by various authors (Otis et al., 1982; Roubik and Michener, 1985). The other two tribes are treated below. TriBE DIPHAGLOSSINI Diphaglossinae Vachal, 1909:33. 1. Malar space at least 1.5 times as long as basal mandibular width and well over one fourth as long as eye. 2. Mandible of female with two small preapical teeth, thus tridentate, but often worn so that only one preapical tooth is evident. 3. Labrum produced to acute, hairy, medial apical point. 4. Clypeus usually strongly REVIEW OF DIPHAGLOSSINI AND DISSOGLOTTINI 1! convex, often protuberant in front of eye by distance as much as eye width (lateral view); apex produced downward well below lower ends of eyes. 5. Vertex seen from front depressed at least laterally, lateral part lower than upper end of eye. 6. Pre-episternal groove absent below scrobal groove. 7. Notaulus deeply impressed in anterior part of scutum. 8. Basitibial plate of female entirely absent or represented by weak carina along posterior margin; hind basitarsus of male half as long as tibia or less. 9. Vein M of forewing meeting or basal to cu-v; second recur- rent vein near middle of second submarginal cell. 10. Jugal lobe of hind wing less than half as long as vannal lobe. 11. Pygidial plate of female broadened at base (under T V). 12. Gonostylus not recognizable or if recognized, much shorter than gonocoxite, not expanded distally. 13. Penis not reaching apices of penis valves, with- out sclerotized rods on under side. This tribe appeared to be restricted to the Chilean or Araucanian faunal region in 1966, but a genus described below is now known from the Andean area as far north as Venezuela. The deep notauli are unique among bees and constitute a strong synapomorphy. The long malar areas are also a synapo- morphy, although they occur elsewhere in scattered groups of Apoidea. Key TO THE GENERA OF DIPHAGLOSSINI 1. Malar space about two thirds as long as eye in female, three fourths in male: S) lil of male with, broad: median, apical projection bearing a directed stiff setae . OD che = Wala « space Hla one aire as long aSteye;so LIL @tmale simple. Miales:,. srnt).).8 a Se eee eee 24 20. Tibial and sternal scopal hairs with numerous short, fine side branches projecting at right angles to rachis or curled basad |. 5, 2Ms0--sro) anaes Sarocolletes —. Tibial scopal hairs with long branches directed distad; sternum with hairs simple or with branches directed, distad ):. 845.0n ee ae 21 21. Inner hind tibial spur coarsely pec- tinate with 10 teeth or less.......-. Pa —. Inner hind tibial spur ciliate or finely pectinate with over a dozen teeth cura AR ass Se 23 22. Scopa of hind tibia formed around CLASSIFICATION OF AMERICAN COLLETINAE tibia without long, loose hairs ex- tending above and below; basitibial plate with carinate margins not hid- denvbyjhair, 22.70) Perditomorpha (part) —. Hind tibia with a few long, loose hairs fully half as long as tibia on upper and lower margins; basitibial plate hidden by hair except some- iiMeS atibase a meeeis) cee aa: Kylopasiphae 23. Inner hind tibial spur finely pecti- nate; claws simple or with inner ramus reduced to small tooth, shorter than basal diameter of outer ramus. Body metallic blue or green- ISIMe rene Oto Acar seca Spinolapis —. Inner hind tibial spur ciliate; inner ramus of claw strong, longer than basal diameter of outer ramus, so that claws are bifid. Body without metallic coloration: 7. @ ae ere Perditomorpha (part) 24. T7 with shiny, hairless, irregularly rough pygidial area, not narrowed posteriorly, defined across posterior border by weak carina, this area occupying much of dorsum of tergum; S7 with apical lobes much reduced, in a single plane (Fig. 5e) Bet AG bE Bee ee ee eee oe Kylopasiphae —. T7 with dull or shiny, usually ill- defined pygidial area, sometimes a longitudinal strip, sometimes a broader area narrowed posteriorly; S7 with well developed apical lobes, two to four, usually at two levels, on CACMSIGeit vw o.s9e ha Seen er ee 25 25. Body metallic bluish or greenish; margin of S6 produced midapically as rounded hairy lobe about one third as wide as sternum, notched medially ieee hehe Tek. ee ee Spinolapis —. Body almost always nonmetallic; margin of S6 broadly rounded with median notch, often broad and shal- a) al cares Mer tregt Oe ren Med ea cig Mera 26 26. Metasoma rather broad and flat- tened, resembling that of female in Shapes cryerein eaten tek, Se Me Sarocolletes —. Metasoma commonly _ rather slender, not flattened, usually dis- tinctly different in shape from that of feriales #.ai82 5) beret poe ee Perditomorpha 635 Subgenus Perditomorpha Ashmead Perditomorpha Ashmead, 1899:86; Moure, 1954:165. Type species: Perditomorpha brunertt Ashmead, 1899 (monobasic). Bicolletes Friese, 1908:11; Moure, 1954:170. Type spe- cies: Bicolletes neotropica Friese, 1908, by designation of Cockerell, 1915:342. Edwynia Moure, 1951:195 (not Aldrich, 1930) (new synonym). Type species: Pasiphae flavicornis Spinola, 1851, by original designation. Edwyniana Moure, 1954:165 (new name for Edwynia) (new synonym). Type species: Pasiphae flavicornis Spinola, 1851 (autobasic). Belopria Moure, 1956:305 (new synonym). Type spe- cies: Belopria zonata Moure, 1956. Lewoproctus (Bicolletes, Belopria, Edwyniana and Perditomor- pha); Michener, 1965:41. This is the largest South American subgenus of Lewoproctus. Because it is common, its characters are described in detail and other subgenera are mostly characterized by comparison to the usual (rather than exceptional) features of Perditomor- pha. (Figs. 2; 3, 10amib, ce) 19a, ibs eBigs.—o. Moure, 1954; Figs. 1-3, Moure, 1956; Figs. 1- 6, Toro, 1968; Figs. 1-9, Toro and Rojas, 1970; Figs. 121-126, Toro, 1973b). Description: a. Integument black, nonmetallic or sometimes with barely perceptible bluish tint, metasoma sometimes red (blue in a rather large, undescribed Argentine species). Length 5-13 mm. Pubescence all pale to all black, pale haired species usually with weak apical bands of pale hairs on terga. b. Inner orbits converging below. c. Clypeus usually rather uniformly punctured and hairy but sometimes largely hairless medially (as in male L. ruber) or nearly all smooth and shining (as in L. stzlborhinus) or sometimes with lower and lateral parts sparsely punctate and shining, upper central part flat and closely punc- tate, suggesting Lonchopria (Biglossa) and its rela- tives (as in L. mourec and females of L. herrerae, nitidior, flavicornis, and weakly in L. trstis). d. Labrum of both sexes over three times as wide as long (often five times), apical margin straight, weakly convex, or weakly concave, apical bristles usually longer than labrum; nearly whole surface or at least basal two fifths forming elevated, convex bare zone so that distal depressed zone is usually narrow but sometimes wider than ele- vated basal zone; apicolateral lobe absent (illus- trated by Toro, 1973b, for L. flavitarsis). e. Max- illary palpus usually long, about last three segments beyond apex of galea; in L. theringi and ruber only one or two segments beyond apex of galea, and in L. arnauellus, brunerit, chrysostomus, inconspicuus, and leucostomus palpus scarcely attain- ing apex of galea. Mandible with preapical tooth on upper margin in both sexes (illustrated by Toro, 1973b, for L. flavitarsis). Glossa with lobes usually broader than long, sometimes about as broad as long. f. Ocelli near summit of vertex, 636 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Fic. 2. a-f. Letoproctus (Perditomorpha) eulonchopriodes n. sp., male. a, b. Dorsal, ventral and lateral views of genitalia. c, d. Dorsal, ventral, and lateral views of S8. e. Labrum. f. Dorsal and ventral views of S7. g. L. (P.) leucostomus (Cockerell), dorsal and ventral views of S7. anterior margin of median ocellus usually well behind midpoint between antennal bases and posterior edge of vertex but sometimes near that midpoint, at midpoint in L. arnauellus and female of L. brunertt. g. Male flagellum usually reaching tegula, sometimes as in L. ruber not attaining tegula; middle segments usually as broad as long or slightly longer than broad, sometimes broader than long as in L. ruber and eulonchopriodes, much broader than long in L. arnauellus and brunerit. h. Propodeum with subhorizontal surface usually shorter than but often longer than metanotum, steeply slanting and unusually short in mourei, nitidior, and zonatus; propodeal triangle smooth, less commonly weakly roughened or transversely wrinkled, marginal groove usually pitted but sometimes not. i. Anterior basitarsus of female without comb or row of hairs along outer margin. Basitibial plate defined by carinae, apex rounded (as in both sexes of L. theringi and zonatus) to almost pointed. }. Femoral scopa sparse; hairs on anterior side of femoral corbicula with several long branches and no well marked rachis (except in L. arnauellus, brunerit, inconspicuus, and leuco- stomus, see discussion below), similar to hairs of outer side of tibia but smaller; on posterior side of corbicula hairs usually sparse and simple, some- times a few branched as in L. neotropicus and saltensis, or less sparse and simple (L. wagen- knechtt), with plumose hairs near base (L. herrerae), or with branched hairs (L. flavicornis, theringi, nitidior, stilborhinus, and zonatus), these hairs un- usually long and dense in L. nitidior and zonatus. Tibial scopa sparse to well developed, white to black; long scopal hairs of lower margin of tibia curved upward, these and hairs of outer surface with long apical and preapical branches, several branches of each hair of more or less equal length, no axis extending far beyond branches (Fig. 19a) (except in L. arnauellus, bruneri, inconspicuus, and CLASSIFICATION OF AMERICAN COLLETINAE 637 ao Fic. 3. a-f. Leioproctus (Perditomorpha) inconspicuus n. sp., male genitalia, S8 and S7; female inner hind tibial spur. g-j. L. (P.) brunerii (Ashmead), male genitalia and S8 (lateral views), S7; female inner hind tibial spur. 638 leucostomus, see discussion below). k. Tibial spurs straight, inner margin of inner spur of female pectinate (Fig. 3f), ciliate in L. arnauellus and brunerii (Fig. 3j), other margins ciliate (see illus- tration of L. flavitarsis, Toro, 1973b). 1. Wings with minute hairs throughout or sometimes basal half or more hairless (as in L. neotropicus and stilborhinus); two submarginal cells, cell 2+3 sube- qual to 1; basal vein of forewing meeting or beyond cu-v; jugal lobe of hind wing usually reaching but sometimes not reaching or exceed- ing level of cu-v, in L. neotropicus and stilborhinus attaining midpoint between level of cu-v and apex of vannal lobe (as in Kylopasiphae). m. Sterna of female without well developed scopa but with apical bands of long hair (sometimes with pollen and functionally a scopa), hair suberect or sloping posteriorly, shorter than exposed parts of sterna, usually dense in band across posterior part of each sternum but sometimes sparse as in L. flavitarsis and mourei, usually simple or with only a few plumose hairs on median basal or on lateral part of each band, but sometimes coarsely branched as in L. theringi, saltensis, and neotropicus; in L. arnauellus, brunerit, and inconspicuum sternal scopa moderately developed, hairs with numer- ous rather short side branches, mostly directed apicad. S6 of male unmodified, ordinarily with small apical emargination. Pygidial plate of male absent but T7 with median, often flattened, sparsely hairy or bare area (convex and bare, sharply different from adjacent areas, in L. zona- tus). n. S7, S8, and genitalia of male as illustrated by Moure (1954), Toro and Rojas (1970) and for L. zonatus by Moure (1956) and L. flavitarsis by Toro (1973b), and in Figures 2 and 3 herein. Apicolateral lobes of S7 of the male two on each side, usually both broad and more or less rounded (Moure, 1954; Toro and Rojas, 1970; Toro, 1973b; Fig. 2f); lobes sometimes narrower and elongate, e.g. in L. mourei (Toro, 1968) and zonatus (Moure, 1956) or rather small and one somewhat elongate, e.g. L. herrerae (Toro, 1968), or broadened and extended basally, each bilobed so that there are four apicolateral lobes on each side in arnauellus and brunerii (Fig. 3). Process of S8 little downcurved; distal part of gonoforceps broad so that there is no stylus-like structure when viewed from above or below although slender distally in lateral view, under surface near apex with ill-defined membranous area. Distribution: This large subgenus is abundant in temperate South America on both sides of the Andes and ranges north to Peru and the state of Ceara, Brazil. Included species: (*marks species not seen by me): Levoproctus (Perditomorpha) abdominis Michener, 1965 (= Pasiphae abdominalis Jorgensen, 1912, preoc- cupied in Levoproctus); *alismatis (Ducke, 1908) | Panurginus|; *andinus (Herbst, 1923) [Paszphae]; arnauellus. Michener, named in the Appendix ( = Perditomorpha arnaui Moure, 1954, preoccupied in Lewoproctus); atacama Toro & Rojas, 1970; *basz- THE UNIVERSITY OF KANSAS SCIENCE BULLETIN rufus (Schrottky, 1920) [Paszphae]; bicellularis (Ducke, 1910) [Lonchopria|; bruner (Ashmead, 1899) [= Pasiphae leaena (Vachal, 1909)]; chryso- stomus (Cockerell, 1917) [Pastphae]; *coloratipes (Cockerell, 1933) [Pastphae]; *decoloratus (Ducke, 1908) | Panurginus|; erithrogaster Toro & Rojas, 1970; eulonchopriodes Michener n. sp.; *fasciatus (Schrot- tky, 1920) [Pasiphae]; *ferrugineus (Moure, 1954) [ Bicolletes|; flavicornis (Spinola, 1851) [| Pasephae]; flav- itarsus Toro, 1973; *franki (Friese, 1908) | Brcolletes]; *frieset (Jorgensen, 1912) [Pasiphae]; herrerae Toro, 1968; thering’ (Schrottky, 1910) [Paszphae] (= Pa- stphae cestri Ducke, 1912); inconspicuus Michener n. sp.; leucostomus (Cockerell, 1917) [Pasiphae]; *longi- pes (Jorgensen, 1912) [Pasiphae]; lucidulus (Cock- erell, 1933) [Pasiphae]; *moerens (Vachal, 1909) [Pasiphae|; mouret Toro, 1968; neotropicus (Friese, 1908) [Bicolletes|; nitidior (Moure, 1956) [Belopria]; *paraguayensis (Schrottky, 1907) [Perditomorpha); penai Toro & Rojas, 1970; perezi Toro & Rojas, 1970; peruvianus (Cockerell, 1926) [| Pasephae]; phar- cidodes (Moure, 1954) [Bicolletes|; pseudozonatus (Moure, 1954) [Bicolletes]; ruber Toro & Rojas, 1970; rufiventris (Spinola, 1851) [Pasiphae]; saltensis (Friese, 1908) [Bicolletes]; *seydi (Strand, 1910) [Bicolletes]; *spegazzin’ (Jorgensen, 1912) [Cal- liopsis|; *stictus (Moure, 1954) [Bicolletes]; stilbo- rhinus (Moure, 1954) [Bicolletes]; tristes (Spinola, 1851) [Pasiphae|; wagenknechti ‘Toro & Rojas, 1970; zonatus (Moure, 1956) [Belopria}. Comments: In spite of the name of this sub- genus, there is no resemblance to the panurgine genus Perdita. The following characters that are often of subgeneric or generic importance in bees vary within the subgenus: Facial foveae of females are usually completely absent or represented only by areas that are less densely punctate or slightly more shiny than surrounding regions. However, the foveae are distinct, well defined, and depressed in females of L. herrerae and tristis. Certain species are very coarsely punctate, unlike most species which are only moderately to finely punctate. One of the most coarsely punc- tate is L. theringi. It also has the basal, sub- horizontal part of the propodeum as seen in profile, slanting and unusually short, only about half as long as the metanotum, and the margin of the propodeal triangle marked by a row of strong pits. L. eulonchopriodes is similarly coarsely punc- tate, with yellow apical integumental bands on T1, 3, and 4, and with the apical tergal margins upturned and carinate as in Eulonchopria psaenythi- oides. L. luctdulus is intermediate in coarseness of punctation between L. thering: and ordinary spe- cles. Unusual features of certain other species are as follows: In L. stilborhinus the front tarsus of the CLASSIFICATION OF AMERICAN COLLETINAE female is only about as long as the tibia, the basitarsus is particularly shortened, bristled api- cally, with a flat, hairless translucent apical proc- ess extending distad about to the apex of segment 3. In L. neotropicus (or a close relative) the lines margining the propodeal triangle are evanescent, so that the triangle, which is so clearly defined in most colletids, is often difficult to recognize. In L. chrysostomus the claws of each leg are asymmetri- cal, the principal ramus of the inner claw being enlarged and blunt. The species that have been placed in Edwyniana are diverse. The type species, L. flavicornis, and L. herrerae are similar in being rather large, with strong apical bands of often erect white hair on T2-4, and with red legs. In these respects they differ from the other species placed in Edwyniana, L. mouret and tristis, which look like ordinary Perditomorpha. | can recognize no characters that justify separation of Edwyniana from the Bicolletes sort of Perditomorpha. Toro and Rojas (1970) already noted problems in the recognition of Edwyniana, some characters of L. erithrogaster, a rather ordinary looking Perditomorpha, being those attributed to Edwyniana. The species that have been placed in Belopria (L. zonatus, nitidior) are slightly more distinctive and obviously closely related to one another. Both have the subhorizontal basal part of the propod- eum steeply sloping and very short, so that the profile of the propodeum is almost wholly de- clivous. The same is true of ZL. moure: and 1s approached by L. theringi. More noteworthy is the scopa which is better developed on the hind femora and trochanters than in any other Perdito- morpha. Furthermore, the pygidial area of the male is represented by a convex bare area, nar- rowly rounded at the apex. Although not mar- gined it is better developed than in other Perditomorpha. On the basis of these characters, Belopria could be recognized as a subgenus but it would be weakly differentiated compared to other subgenera and [ have therefore chosen to place its species in Perditomorpha. The subgenus Perditomorpha in the old sense contains only two previously described species (L. arnauellus, brunerit) that have long been separated from Bicolletes at the generic or subgeneric level (Moure, 1954; Michener, 1965). It no longer seems practical to maintain the distinction be- tween Bicolletes and Perditomorpha, even at the subgeneric level. The following are the distinctive features of the two species listed above, lettered as in the subgeneric descriptions: g. Flagellum of both sexes very short, not over twice as long as scape, not attaining tegula, middle segments much broader than long. j. Scopal hairs on anterior side of femoral corbicula and on outer 639 side of hind tibia plumose with branches along rachis (Fig. 19b). k. Margins of tibial spurs ciliate (Fig. 3}). m. Sternal hairs long, forming moder- ately distinct metasomal scopa, with numerous rather short, fine branches. n. S7 of male with lateroapical lobes large, each bilobed so that there are four lobes on each side, two of them extending far basally and two apically, apodemes extending laterally in opposite directions (Fig. 31). In the features listed above, L. inconspicuus agrees with Perditomorpha in the old sense except for the hind tibial spur (k) which is coarsely pectinate in the female, as in typical Bicolletes (Fig. 3f). Also the emargination between two of the lobes of S7 of the male is less deep, although the sternum is of the same type as that of Perditomor- pha in the old sense (Fig. 3e). L. leucostomus occupies a more clearly interme- diate position between the old Perditomorpha and Bicolletes, and indeed has been placed at different times in each group. The scopal hairs of the hind leg (j) are like those of Perditomorpha in the old sense or those of the tibia with somewhat longer branches. The sternal scopa (m), however, is scarcely developed and the hairs are simple or with long branches, 1.e., as in Bicolletes. The inner hind tibial spur of the female is pectinate but finely so, with about ten teeth, thus deviating from the ordinary coarsely pectinate type (with five or six teeth) in the direction of the old Perditomorpha. The flagellum of the male (g) 1s more elongate than in the old Perditomorpha. S7 of the male (n) has the two apicolateral lobes on each side large and extended basad but not divided into more lobes (Fig. 2g), thus intermediate between Bicolletes and the old Perditomorpha, but as in the latter, the apodemes extend laterally in opposite directions. L. leucostomus also differs from the old Perditomorpha in some less decisive charac- ters (because they vary within Bicolletes) as fol- lows: anterior ocellus well behind midpoint between antennal bases and posterior margin of vertex; basitibial plate rounded at apex, espe- cially in female. J. L. Neff (in litt.) has noted that Perditomorpha in the old sense (L. arnauellus, brunerii, and incon- spicuus) seems to specialize in collecting pollen from Malvaceae, and the coarse pollen on spec- imens in collections supports this view. Specimens of L. leucostomus, however, have much finer pol- len. The correct position of L. bicellularis is in doubt although it is listed above as a Perditomorpha. It has strong apical enamel-like pale yellow bands on T1-4 of the female and T1-5 of the male, suggest- ing the subgenus Nomuocolletes, but does not re- semble that subgenus in other features. Similar bands but on fewer terga are found on L. (Perdito- 640 morpha) eulonchopriodes, but the bands of L. bicellu- lars are flat and smooth, not upturned and carinate at the margins, and the body is not especially coarsely punctate as in L. eu- lonchopriodes. Features that do not agree with other Perditomorpha include the middle flagellar seg- ments of the male which are fully 1.25 times as long as wide; the scopal hairs are as noted for L. (P.) leucostomus and Perditomorpha s. str. The claws of the female have the inner ramus reduced to a minute tooth so that they are almost simple; in the male the inner ramus is much smaller than usual. Unfortunately the only male of L. bicellu- laris seen [AMNH] lacks the apex of the meta- soma so that the genitalia and hidden sterna remain unknown. Chilicolletes new subgenus Type species: Lezoproctus delahozi Toro, 1973. This subgenus closely resembles Perditomorpha but has three submarginal cells. It differs from Sarocolletes by the very different scopal hairs of the latter, and from American Leioproctus s. str. by the coarsely pectinate inner hind tibial spur of the female. (Figs. 4, 10c; Figs. 127-135, Toro, 1973b). Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha except as follows: a. Length 7-11 mm. Pubes- cence largely dusky, if with considerable pale hair, then usually with tergal hair bands. d. Labrum of male three times, of female over three times, as wide as long; apical margin gently concave (fig- ures in Toro, 1973b). e. Maxillary palpus with THE UNIVERSITY OF KANSAS SCIENCE BULLETIN about 2.5 segments beyond apex of galea. g. Male flagellum scarcely reaching tegula, middle seg- ments slightly broader than long. h. Propodeum with subhorizontal zone about as long as meta- notum; groove margining triangle weakly pitted or not pitted. i. Anterior basitarsus of female with weakly evident row of hairs along outer margin. }. Hairs along posterior margin of femoral corbicula with several short barbs. Hairs of tibial scopa with several long branches as in most Perditomor- pha but with one (the rachis) distinctly longer than others. 1. Wings with parts of basal cells hairless. Three submarginal cells, 2 much shorter than 3 and receiving first recurrent vein near middle or at distal end. m. Sternal scopa sparse, longer hairs almost as long as exposed part of sternum, simple or nearly so. T7 of male with large, bare, shining somewhat convex area, narrowed posteriorly, sharply defined by absence of hairs. n. S7, S8, and genitalia illustrated by Toro (1973b) and Fig. 4. S7 with apicolateral lobes elongate, unlike most Perditomorpha. Distribution: Central Chile. Included species: Leioproctus (Chilicolletes) delahozu Toro (1973) and at least one related undescribed species. Comments: This subgenus is related to Perdito- morpha and my first inclination was to include it in that subgenus in spite of the three submarginal cells. However, presence of a comb of hairs on the anterior basitarsus of the female is also unlike any Perditomorpha. This weakly developed comb, how- ever, is not like that of Nomuocolletes, Reedapis, and Cephalocolletes. Instead it seems to be merely the abrupt line ending the vestiture of long hairs on the upper surface of the basitarsus, the lower Fic. 4. a-c. Leioproctus (Chilicolletes) delahozii Toro, male, lateral views of genitalia and $8, dorsal and ventral views of S7. CLASSIFICATION OF AMERICAN COLLETINAE surface being nearly bare, unlike other species. The other characters listed above are within the range of variation found in Perditomorpha. The trochanteral and femoral scopas are better devel- oped than in most Perditomorpha. The presence of three submarginal cells suggests that this might be a surviving representative of the group from which Perditomorpha was derived. It cannot be a specialized derivative of Perditomorpha like the little groups here included in that subgenus, such as Edwyniana, Belopria, and Perditomorpha s. str. Etymology: The subgeneric name is based on Chile, the country where these bees occur, and the generic name Colletes. Kylopasiphae new subgenus Type species: Lezoproctus (Kylopasiphae) pruinosus new species. This subgenus is similar to Perditomorpha. The greatly reduced lobes of S7 of the male are the most distinctive feature; others include the hid- den basitibial plate and the features of the tibial scopa of the female (see below). The long anten- nae of the male also differentiate this subgenus from Perditomorpha; the short antenna of Perdito- morpha is likely to be a derived feature uniting species of that subgenus. (Figs. 5a-f, 10f, 23e) Description: Agreeing with usual (rather than exceptional) characters of Perditomorpha except as follows: a. Length 7-8.5 mm. Pubescence all pale, covering metasomal terga which lack apical hair bands or have weak bands in male. c. Clypeus shiny and impunctate except for lateral extremities and longitudinal median punctate depression; male with band of punctures across upper margin. d. Labrum about three times as wide as long; apical margin gently convex; basal two thirds shining, convex; female with transverse carina before depressed apical zone which lacks short hairs before apical fringe; longest apical bristles about as long as labrum in male, little longer in female; apicolateral lobe absent. g. Male flagellum reaching beyond tegula, middle segments slightly over 1.5 times as long as broad. h. Propodeum with subhorizontal surface 1.5 times as long as metanotum, not much sloping; propodeal triangle smooth, shin- ing; marginal groove not conspicuously pitted, evanescent in female [as in L. (Perditomorpha) neotropicus|. 1. Basitibial plate of female small, hidden by hair (even margin invisible without removing hair) except at base, that of male nearly hairless. j. Scopa rather dense. Hairs of trochanter and in front of femoral corbicula with several long branches directed apicad, most hairs with rachis exceeding branches (suggesting tibial scopal hairs of Lewoproctus s. str., Fig. 19c). Hairs behind femoral corbicula abundant, mostly simple, a few branched ones near base of femur. Tibia on both lower and upper margins with a few extremely long (half as long as tibia or more), loose, hairs 641 (Fig. 23e) with short branches directed apicad, these hairs extending as far below as above tibia; hairs on outer side of tibia like those of most Perditomorpha (Fig. 19a) but near margins grading through types like those illustrated for Lezoproctus s. str. and Perditomorpha s. str. (Fig. 19b, c) to very long hairs; hairs of lower margin not strongly curved upward. |. Forewing almost hairless ex- cept for short hairs beyond venation and in distal part of marginal cell. Jugal lobe of hind wing extending to point half way between vein cu-v and apex of vannal lobe. n. S7 of male with apical lobes greatly reduced, not layered one above another but all on a single plane (Fig. 5e); S8 and genitalia as in Fig. 5a-d. Distribution: Argentina, desert areas in the provinces of Tucuman, Catamarca, La Rioja, and Neuquén. Included species: Levoproctus (Kylopasiphae) pruino- sus Michener, n. sp. Comments: This subgenus may be a derivative of some of the group of small species of Perditomorpha found in the same region. The relatively long Jugal lobe of the hind wing suggests L. (P_) neotropicus and stilborhinus; the largely smooth female clypeus and supraclypeal area suggest the latter, and the near absence of the lines delimiting the propodeal triangle of the female suggest the former. Etymology: The subgeneric name is based on the Greek kylon, face below the eyes, with reference to the shining clypeus, and Pasiphae, an unused generic name for Paracolletinae with two submar- ginal cells (see Spznolapis). Subgenus Leioproctus Smith, s. str. Letoproctus Smith, 1853:8; Michener, 1965:39. Type species: Levoproctus imitatus Smith, 1853, by designa- tion of Cockerell, 1905a:348. Lamprocolletes Smith, 1853:10. Type species: Andrena chalybeata Erickson, 1851, by designation of Cock- erell, 1905a:344. Dasycolletes Smith, 1853:14. Type species: Dasycolletes metallicus Smith, 1853, by designation of Cockerell, 1905a:347. The comments and description below are based entirely on the single American species provision- ally assigned to this subgenus and are not intended to encompass the variation exhibited by the many species in the Australian region. The three submarginal cells, the apical lobes of S7 of the male and the short, robust process of the male S8 are suggestive of L. (Chilicolletes) delahozit. The ciliate inner hind tibial spur, the longer male antennae, and other characters italicized below distinguish our species of Lezoproctus s. str. from Chiltcolletes and Perditomorpha. (Figs. 5g-l, 19c) Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha 642 except as follows: a. Length 10-11 mm. Pubes- cence largely dark but fulvous on dorsum of thorax; tergal hair bands absent. c. Clypeus with irregularly defined, narrow preapical zone shin- ing and largely impunctate, especially in female. d. Labrum of both sexes four or more times as THE UNIVERSITY OF KANSAS SCIENCE BULLETIN wide as long, apex gently convex, basal half elevated, convex, bare, shining, in male medially lower; depressed distal zone and distal slope of elevated zone with small hairs basal to fringe. g. Male flagellum reaching beyond tegula. h. Pro- podeum with subhorizontal surface a little shorter Fic. 5. a-f. Leioproctus (Kylopasiphae) pruinosus n. sp. Genitalia, S8 and S7 of male, inner hind tibial spur of female. g-l. L. (Leioproctus) fulvoniger n. sp. Inner hind tibial spur of female and genitalia, $8 and S7 of male. CLASSIFICATION OF AMERICAN COLLETINAE than metanotum, sloping rather steeply; triangle dull, transversely striate; marginal line pitted especially anteriorly. i. Basitibial plate of female pointed, of male narrowly rounded. j. Hairs of femoral scopa black, rather short, straight, those on anterior side of corbicula with a few rather long branches diverging from rachis, thus similar to but smaller than tibial scopal hairs; on poste- rior side of corbicula hairs simple. Tibial scopa sparse, black, long hairs of lower margin not much curved, thus scopa spreads as far below tibia as above it, branches of these hairs of moderate length, rather few in number, rachis of hair extending beyond branches (Fig. 19c). k. Margins of tibial spurs ciliate. 1. Three submarginal cells, 2 much shorter than 3 and recelving first recurrent vein beyond middle; basal vein distal to cu-v. m. Sterna of female with abundant hair sloping posteriorly, shorter than usual in Perditomorpha, not much over half length of exposed part of sternum. n. S7, S8, and genitalia as in Fig. 5h-l. Apical lobes of S7 rather slender, more or less as in L. (Chilicolletes) dela- hozu, not broad as in most Perditomorpha; S8 with apical process short and very robust; gonobase enormous; gonocoxite with hairs nearly to base and with strong dorsal inner angle. Distribution: Santa Catarina, Brazil (and the Australian region). Included species: Levoproctus (Lewoproctus) fulvoniger Michener, n. sp., and about 120 species in the Australian region. Glossopasiphae new subgenus Type species: Lezoproctus (Glossopasiphae) plaumanni new species. The most distinctive character of this subgenus is the two long glossal lobes which resemble those of the subgenus Tetraglossula. However, the struc- ture must have evolved independently, for Tetra- glossula belongs with Protodiscelis in a group having simple mandibles in the male and an extremely short labrum with lateroapical lobes at least in females. Glossopasiphae, however, like many Leio- proctus, has bidentate male mandibles and a rela- tively long labrum without lateroapical lobes. (Figs. 6, 19d) Description: Agreeing with usual (rather than exceptional) characters of Perditomorpha except as follows: a: Length 12-14 mm. Pubescence largely black or dusky except for thoracic dorsum; tergal hair bands absent. b. Inner orbits very slightly diverging below (both sexes), almost without concave region so that upper parts of orbits are not greatly different in orientation from remainder. d. La- brum of male about twice as broad as long, of female over 2.5 times as broad as long; apex produced to rounded angle; male with basal two fifths gently convex, convexity slightly depressed medi- ally; female with basal three fifths strongly ele- vated and convex; apical bristles scarcely longer than labrum; apical four fifths of labrum punc- 643 tate, with hairs. e. Glossa deeply bifid, each lobe attenuate, hairy, about five times as long as basal width (Fig. 6h). Galea about as long as stipes, postpalpal part about six times as long as wide, not or slightly exceeded by maxillary palpus. f. Ocelli in front of summit of vertex; vertex convex and rounded behind ocelli; anterior margin of median ocellus in front of midpoint between antennal bases and posterior margin of vertex. g. Male flagellum not attaining tegula, middle flagellar segments slightly broader than long. h. Propodeum with basal zone steeply sloping in profile, shorter than metanotum; propodeal tri- angle transversely striate granulate, marginal groove not pitted. i. Basitibial plate rounded at apex, narrowly so in male. j. Femoral corbicula with a few short hairs on surface basally; scopa on anterior side of corbicula long, dense, hairs with many fine, short lateral branches as on tibial scopa; trochanteral hairs similar; hairs on poste- rior side of corbicula simple or toward base of femur with fine, short side branches. Tibial scopa dense, long, extending farther below tibia than above, hairs of lower margin gently curved apicad and slightly upward; hazrs with many fine, short branches at more or less right angles to rachis and often curved basad (Fig. 19d), rachis extending well beyond branches. k. Tibial spurs gently curved near api- ces, inner hind spur of female coarsely pectinate, other margins coarsely ciliate. |. Basal vein of forewing meeting cu-v. m. S2-5 of female with dense scopa of hairs as long as exposed part of sternum, similar to those of tibial scopa. n. S7, S8, and genitalia of male as in Fig. 6. S8 with apical process not bent down, without ventral tubercle. Gonostylus fused to gonocoxite, with distal ventral membranous area. Distribution: Santa Catarina, Brazil. Included species: Leioproctus (Glossopasiphae) plau- manni Michener, n. sp. Etymology: The subgeneric name is based on the Greek glossa, tongue, with reference to the elon- gate glossa, plus Pasiphae, a generic name used for various paracolletines with two submarginal cells (see Spinolapis). Sarocolletes new subgenus Type species: Lonchopria rufipennis Cockerell, 1917. The appearance of the scopal hairs, with nu- merous short fine side branches, is a striking feature of this subgenus shared only with Glosso- pasiphae and Protodiscelis. Most other characters agree with those of Chilicolletes, or in the case of two-celled species, with Perditomorpha. Sarocolletes differs further from those subgenera, however, in the well developed ventral scopa of the female and in the relatively broad metasoma of the male which therefore resembles a female. (Figs. 7a-h, 10g) Description: Agreeing with usual (rather than 644 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Fic. 6. Letoproctus (Glossopasiphae) plaumanni n. sp. a-f. Genitalia, S8 and S7, and mandible of male. g, h. Mandible and glossa of female. i, j. Labrum of male and female. k. Inner hind tibial spur of female. exceptional) characters listed for Perditomorpha except as follows: a. Length 7-11 mm. Pubes- cence mostly pale, forming apical bands of pale hair on terga. Metasoma of male broad so that body form is like that of female. b. Inner orbits only slightly converging below. d. Labrum two to three times as broad as long, apex gently convex or with small median concavity, apical bristles about as long as labrum; basal three fifths to four fifths a convex bare zone, more elevated in female than male, this convexity weakly depressed medi- ally. Maxillary palpus reaching little if any be- yond apex of rather long galea, postpalpal part of which is as long as prepalpal part. f. Ocellar position variable as described for Perditomorpha. Vertex behind ocelli well developed, convex. g. Male flagellum sometimes, as in L. (S.) rufipennis, not reaching tegula, middle segments slightly broader than long to longer than broad. h. Pro- podeum variable as in Perditomorpha. 1. Basitibial CLASSIFICATION OF AMERICAN COLLETINAE 645 Fic. 7. a-h. Letoproctus (Sarocolletes) duplex n. sp. a-e. Genitalia, S8 and S7 of male. f, g. Labrum of male and female. h. Inner hind tibial spur of female. i-o. L. (Holmbergeria) rubriventris (Friese), male. i. S7. j-m. Genitalia and S8. n, o. Mandible and labrum. p, q. L. (Hoplocolletes) ventralis (Friese), female, inner hind tibial spur and labrum. 646 plate variable as in Perditomorpha, rounded in L. (S.) rufipennis. j. Femoral corbicula with a few short hairs on surface; hairs anterior to corbicula and on trochanter with numerous rather short, fine branches along rachis, often shorter than those on tibial scopal hairs; hairs posterior to corbicula simple. Tibial scopa with hairs of lower margin not curbed upward, scopa extending far- ther below tibia than above, hairs of entire tibial scopa with numerous short, fine side branches along rachis similar to those illustrated for L. (Protodiscelis) palpalis (Fig. 19e). 1. Wings usually with three submarginal cells, two only in L. (S.) duplex. m. S2-5 of female with strong scopa of long (as long as exposed part of sternum), dense, sloping hairs with numerous short, fine side branches that tend to be at right angles to rachis and curled basad, as on tibial hairs of L. (Glossopasiphae) plaumanni (Fig. 19d). S7 of male with median bare area convex, tapering posteriorly, as in L. (Perditomor- pha) zonatus. n. S7, S8, and male genitalia as shown in Fig. 7a-e. Lateroapical lobes of S7 rather slender, somewhat as in L. (Chilicolletes) delahozu, all in same plane; ventral projection of S8 extremely weak (Fig. 7d). Distribution: Argentina, the provinces of Buenos Aires and Entre Rios to Tucuman, including Santiago del Estero, Santa Fe, Catamarca, and La Rioja. Included species: Leioproctus (Sarocolletes) rufipennis (Cockerell, 1917) [Lonchopria] and its probable synonym, parvus (Cockerell, 1917), proposed as a variety of rufipennis. In addition, it contains at least five apparently undescribed species. All of the above have three submarginal cells. It also includes L. (S.) duplex Michener, n. sp., described in the Appendix, which is exceptional among species of Sarocolletes only in having two instead of three submarginal cells. Comments: Padre J. S. Moure recognized the species with three submarginal cells as a group and named it with specimen labels before 1956. I have not accepted his name since it indicated a relationship to Lonchopria. The possibility that this subgenus is a synonym of Holmbergeria is discussed under that subgenus. Etymology: The name Sarocolletes is based on the Greek saron, brush or broom, plus the generic name Colletes, with reference to the large ventral scopa in females of this subgenus. Subgenus Holmbergeria Jorgensen Holmbergeria Jorgensen, 1912:100. Type species: Holm- bergeria cristariae Jorgensen, 1912 (monobasic). This subgenus is known only from the male. When female characters are known, its rela- tionships may be clarified. If the female scopa turns out to be like that of Sarocolletes, that subgenus might well be regarded as a synonym of THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Holmbergeria in spite of the peculiar facial features (italicized below) of the males of the latter. (Figs. 7i-o, 10d) Description: Agreeing with usual (rather than exceptional) characters of Perditomorpha except as follows: a. Metasoma sometimes partly red (T1-2 or 1-3). Length 9-11 mm. Pubescence pale, dusky on vertex and distal metasomal segments, with strong apical bands of pale hair on terga. b. Face broad, inner orbits parallel. c. Supraclypeal area small, convex, hairless, shining and impunctate; subantennal area shorter than diameter of antennal socket, flat, hairless, shining, impunctate, in contrast to rest of face which is punctate and hairy. d. Labrum about three times as wide as long; apical bristles mostly not longer than labrum; basal half ele- vated, smooth, shining; distal half with small hairs basal to apical bristles. e. Maxillary palpus with little more than last segment extending beyond apex of galea. f. Anterior margin of median ocellus at midpoint between antennal bases and posterior margin of vertex or nearer to the former; vertex strongly developed, convex. g. Flagellum reaching tegula or not, middle seg- ments as long as broad or slightly longer than broad. h. Propodeum with subhorizontal surface (in profile) shorter than metanotum, rounding onto sloping (slanting) posterior surface; triangle smooth or feebly wrinkled, marginal line not pitted. i. Basitibial plate with apex rounded, surface hairy except at margins. k. Tibial spurs (male) with margins ciliate. 1. Three submarginal cells, 2+3 about as long as 1, 2 shorter than 3, receiving first recurrent vein in apical half; jugal lobe of hind wing extending beyond level of cu-v. m. S6 (male) unmodified. Pygidial plate absent but with hairless longitudinal median bare area, narrowed toward posterior margin of tergum. n. S7, S8 and genitalia as in Fig. 7j-m. S7 with two slender apicolateral lobes; S8 with apical process not bent downward, without ventral protuberance but with longitudinal ridge in its place; membranous area at apex of gonostylus well defined. Distribution: Argentina, provinces of Mendoza to Santiago del Estero; Paraguay. Included species: Leioproctus (Holmbergerta) cristar- tae (Jorgensen, 1912) and L. (H.) rubriventris (Friese, 1909) [Bzglossa]. These forms are mor- phologically very similar; the former has the legs and metasoma black, while the latter has the legs and first two or three metasomal terga red. (See Appendix for details on L. cristariae.). Subgenus Hoplocolletes Michener Leioproctus (Hoplocolletes) Michener, 1965:42. Type spe- cies: Dasycolletes ventralis Friese, 1924, by original designation. This subgenus is recognized only on the basis of a combination of female characters most of which are duplicated in other Lezoproctus. The combination of the characters found in L. (Hoplo- colletes) ventralis, however, suggests a form quite CLASSIFICATION OF AMERICAN COLLETINAE dissimilar to other subgenera; discovery of the male will, of course, greatly assist in placement of Hoplocolletes. (Fig. 7, p-q) Description: Agreeing with usual (not excep- tional) female characters listed for Perditomorpha except as follows: a. Length 12 mm. Pubescence short, sparse, blackish except on hind legs and metasomal sterna where there are long, pale hairs; metasomal hair bands absent. b. Jnner orbits subparallel. d. Labrum with apical margin concave medially; elevated zone highest medially, occupy- ing about basal half of labrum. f. Ocelli well forward so that anterior margin of median ocellus is nearer to antennal bases than to posterior margin of vertex. h. Propodeum with sub- horizontal surface about as long as metanotum, triangle smooth, marginal line pitted. 1. Basitibial plate rather slender, pointed. j. Femoral scopa sparse, hairs in front of corbicula mostly simple or bifurcate, those behind corbicula and on trochanter long but simple. Tibial scopa sparse but hairs long, those of lower margin curved apicad, branched as in Perditomorpha, but many hairs of outer and upper surfaces with main axes extending beyond the one or few long branches; scopa extending farther above tibia than below; hairs of inner surface of tibia as long as scopal hairs and thus not like keirotrichia. 1. Three submarginal cells, second much shorter than third and receiving recurrent vein beyond middle; basal vein of fore- wing slightly beyond cu-v. m. $3-5 with dense, long (shorter than exposed part of sternum), pale yellow, simple hairs (some hooked at tips) forming band occupying apical half of each sternum, S2 with similar but sparser hair band. Distribution: Rio de Janeiro, Brazil. Included species: Leioproctus (Hoplocolletes) ventralis (Friese, 1924). Comments: The very coarsely punctate head and thorax of Hoplocolletes suggest L. (Perditomorpha) theringi and eulonchopriodes, although the dorsum of the thorax is even more coarsely and less closely punctate. The metasoma, however, is nearly im- punctate (T1-2) or finely punctate (T3, etc.) in L. ventralis, and the subparallel inner orbits, the three submarginal cells, the very deeply im- pressed median line of the anterior half of the scutum, the deeply impressed parapsidal lines, the simple hind trochanteral hairs, and the strong ventral scopa further differentiate Hoplocolletes. (A superficially similar black species known in both sexes, L. (Tetraglossula) anthracinus Michener n. sp., from the same area as L. ventralis, has been identified as that species in some collections.) The only known species of this subgenus, L. (H. ) ventralis (Friese), was described from Sydney, Australia. The specimens must have been mis- labeled, for the species is now known from the vicinity of Rio de Janeiro, Brazil (Michener, 1965:42); one specimen [Lawrence]. 647 Pygopasiphae new subgenus Type species: Levoproctus mourellus new species. The most distinctive characters of Pygopasiphae are the short, erect, unbranched, capitate hairs of the metasomal sterna of the female, and the distinct pygidial plate and long, slender hind tarsi of the male. (Figs. 8, 10h, 19f) Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha except as follows: a. Metasoma and legs red in some females of L. mourellus, black in other species. Length 7-11 mm. Pubescence largely pale (sometimes partly dusky on vertex, tibiae, and metasomal terga), forming apical pale bands on T1 or 2 to 4. b. Inner orbits slightly converging below to subparallel in females, converging in males; upper parts converging above. e. Maxillary pal- pus with about one segment extending beyond end of galea. g. Male flagellum reaching beyond tegula, middle segments about 1.5 to 1.8 times as long as broad. h. Propodeum with subhorizontal basal zone almost as long as to slightly longer than metanotum; triangle smooth, marginal line pitted or not. i. Anterior basitarsus of female with weakly differentiated row of hairs on outer mar- gin. Basitibial plate pointed at apex, largely covered by hair in female. j. Femoral scopa rather short and sparse (hairs on trochanter simple), hairs simple or (in L. wagner?) those on distal half of anterior side of femoral corbicula with several preapical branches surpassed by rachis. Tibial scopa whitish or upper outer zone along tibia dusky; long scopal hairs along lower margin of tibia not curved upward, spreading as far below tibia as scopa spreads above tibia, these hairs in L. mourellus plumose with several long branches exceeded by rachis, like hairs on outer face of tibia, in L. wagner hairs of lower margin simple or with few short branches (Fig. 19f) although quickly grading to hairs with long branches (as in Perditomorpha) on outer sur- face of tibia. k. Hind trochanter and tibia of male unmodified in L. mourellus, in L. wagnert trochan- ter with strongly produced angle or tooth on under side and hind tibia enlarged and truncate apically, tibial spurs arising far from one another on truncate apex. Hind tarsus of male elongate, segment 2 several times as long as broad. Tibial spurs of female nearly straight or slightly curved, apices blunt, margins almost entire or with a few small teeth except inner margin of inner spur which is pectinate with two to five large teeth. Tibial spurs of male nearly straight, pointed, finely and briefly ciliate. 1. Wings with distal two fifths with minute hairs, basal parts much more sparsely hairy but with a few relatively long hairs. m. Sterna of female without scopa, hairs erect, of uniform length, uniformly dispersed (not forming api- cal bands or fringes), tips bent slightly backward and enlarged like ketrotrichia. S6 of male un- modified, short hairs directed laterad from broad longitudinal median elevation. Pygidial plate of female unusually large, apex rounded, not at all truncate. T7 of male with well defined (by carina 648 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Fic. 8. a-f. Levoproctus (Pygopasiphae) mourellus n. sp. a-d and f. Genitalia, S8 and S7 of male. e. Inner hind tibial spur of female. g-n. L. (P.) wagneri (Vachal). g-k. Genitalia, S8 and S7 of male. 1. T7 of male. m. Hind leg of male. n. Inner hind tibial spur of female. CLASSIFICATION OF AMERICAN COLLETINAE posteriorly) pygidial plate, rounded posteriorly (in L. wagnert slightly narrowed basally). S7, S8, and genitalia of male as in Fig. 8. S7 with apical lobes variable in shape (Fig. 8f, k), Aazrs all or nearly all on ventral surfaces; S8 with process nearly straight or in L. wagneri strongly down-curved; gonostylus not separate from gonocoxite or separation visible only on ventral side. Distribution: Xeric parts of Argentina (prov- inces of Santiago del Estero, Catamarca, La Rioja). Included species: Levoproctus (Pygopasiphae) wagnert (Vachal, 1909) [Pastphae], L. (P.) mourellus Michener, n. sp., and several undescribed spe- cles. Comments: L. mourellus is less extreme in various ways than the larger species with modified male legs such as L. wagnert. | have made L. mourellus the type species because I have not seen type specimens of L. wagneri. The subgenus was first recognized (as a genus) by Moure over 30 years ago but, although he labeled specimens, he never published a description. Michener (1965) in- cluded L. wagnert in the subgenus Bicolletes (= 649 Perditomorpha). Etymology: The subgeneric name is based on the Greek pyge, rump, with reference to the pygidial plate of the male, and pasiphae, an invalid generic name for paracolletines with only two submargi- nal cells (see subgenus Spinolapis). Subgenus Halictanthrena Ducke Halictanthrena Ducke, 1907:364. Type species: Halictanth- rena malpighiacearum Ducke, 1907 (monobasic). Lewoproctus (Halictanthrena), Michener, 1965:42. Distinctive features of the subgenus are the sharp tooth on the dorsolateral angle of the pronotum and the presence of erect hairs and lack of distinctive short hairs on the basitibial plate of the female. The subgeneric name suggests a bee with the aspect of Halictus or Andrena, probably with pale pubescence and a banded metasoma. Halictanthrena, however, is a rather slender bee with largely dark hair, no metasomal hair bands, and with the partially red metasoma suggesting Sphecodes more than Halictus. (Figs. 9, 101) Description: Agreeing with usual (rather than Fic. 9. a-f. Leioproctus (Halictanthrena) malpighiacearum (Ducke). a-e. Genitalia, S8 and S7 of male. f. Inner hind tibial spur of female. 650 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN exceptional) characters listed for Perditomorpha except as follows: a. First two metasomal terga dark red in female, metasoma largely red-brown in male. Length 6.5-9 mm. Pubescence of female blackish on dorsum, tibiae and basitarsi, pale yellow on face below antennae, sides and venter of body, femora, and lower margin of hind tibia; f male pale yellowish, dusky on dorsum, some D black hairs intermixed on scutellum. c. Clypeus with scattered coarse punctures, only sparsely haired. d. Labrum over four times as wide as long, apical margin weakly convex. e. Mandible of female (not male) with convexity on upper margin basal to preapical tooth, suggesting a tridentate condition. g. Male flagellum reaching scutellum, middle segments about 1.5 times as long as wide. h. Fic. 10. Forewings. a. Letoproctus (Perditomorpha) brunerti (Ashmead). b. L. (P.) flavicornis (Spinola). c. L. (Chilicolletes) delahozii Toro. d. L. (Holmbergeria) rubriventris (Friese). e. L. (Perditomorpha) neotropicus (Friese). f. L. (Kylopasiphae) pruinosus n. sp. g. L. (Sarocolletes) rufipennis (Cockerell). h. L. (Pygopasiphae) mourellus n. sp. 1. L. (Halictanthrena) malpighiacearum (Ducke). CLASSIFICATION OF AMERICAN COLLETINAE 651 Dorsolateral angle of pronotum with small, sharp tooth, projecting dorsolaterally, shorter than nearby hairs, in male so small as to be difficult to see except in profile. Propodeum with subhorizontal surface strongly sloping, about as long as meta- notum, gradually curving onto declivous surface. 1. Front basitarsus of female with abundant coarse bristles, strongly curved at apices, on outer sur- face. j. Scopal hairs on outer side of tibia often with rachis extending slightly beyond other branches. Scopal hairs near upper margin of tibia particularly coarse near bases, shorter near base of tibia, similar coarse, nearly erect hairs arising from and nearly hiding basitibial plate which is pointed apically; anterior carina of basitibial plate of female weak for short distance preapically. 1. Three submargi- nal cells, 2 much shorter than 3; basal vein of forewing approximately meeting cu-v; jugal lobe of hind wing reaching distinctly beyond level of cu-v. m. S6 of male broadly convexly subtrun- cate. Pygidial plate of male represented by broad, smooth hairless area delimited posteriorly by distinct, convex carina. n. S7, S8, and genitalia of male as in Fig. 9a-e; S7 with a single hairy apicolateral lobe on each side, $8 with apical process short and shoulders sloping. Distribution: State of Minas Gerais, Brazil. Included species: Leioproctus (Halictanthrena) mal- pighiacearum (Ducke, 1907). Comments: This distinctive subgenus is known to me only from a single pair of specimens [Berlin]. Characters not listed above include the unusually large stigma (Fig. 101), larger than in any other American colletine, and the opposed teeth on the male volsella (not exposed in ventral view). Torocolletes new subgenus Grupo Fazii Toro, 1973a:156. Type species: Lonchopria fazu Herbst, 1923. The largely dull integument without recogniz- able punctures except on the clypeus is character- istic of this subgenus, as are the protuberant clypeus, distinct malar area, reduced jugal lobe of the hind wing, and other characters italicized below. (Fig. 16a; Figs. 115-132, Toro, 1973a) Description: a. Integument dark blue to black, except for clypeus with only minute and widely separated punctures on finely roughened surface, that of head and thorax dull. Length 7-11 mm. Pubescence mostly pale, not forming bands on terga. b. Inner orbits converging below. c. Clypeus moderately coarsely punctate, lower lateral areas more sparsely so and shiny than flat median area. Clypeus produced well below lower tangent of eyes and protuberant in front of eye by about eye width. Malar area longer than flagellar diameter. d. Labrum of both sexes convex, shining, twice as broad as long or slightly more; apex strongly convex; fringe little if any longer than labrum; apicolateral lobe absent (illustrated by Toro, 1973). e. Maxillary palpus with segment 3 reaching beyond apex of galea, segments 4-6 together slightly longer than 1-3 together. Mandible with preapical tooth on upper margin in both sexes (illustrated by Toro, 1973). Glossa with lobes broader than long. f. Ocelli as usual in Perditomorpha, vertex not at all elevated, flat or concave between ocelli and sum- mit of eye. g. Male flagellum reaching front of scutellum, middle segments about 1.5 times as long as broad. h. Propodeum with subhorizontal surface sloping, about as long as metanotum, propodeal triangle dull, minutely and closely areolate, marginal line not pitted. i. Anterior basitarsus of female without comb or row of hairs. Basitibial plate defined by carina, narrowly rounded at apex in both sexes. j. Femoral and tibial scopa sparse, hairs not very long; femoral corbicula with scattered short hairs, mostly with one or two branches, on surface; hairs anterior to corbicula with a few long branches surpassed by rachis; hairs posterior to corbicula simple in contrast to long, loosely plumose hairs on trochanter; tibial scopa with hairs of lower margin as described for Perditomorpha but usually with one branch (rachis) surpassing the others, hairs of outer surface and especially upper margin of tibia with two or three long branches but rachis strong and bare beyond the branches. k. Tibial spurs straight, inner margin of inner hind spur finely pectinate to coarsely ciliate (illustrated by Toro, 1973). 1. Wings with minute hairs distally, hairs longer and sparser basally; three submarginal cells, cell 2 much shorter than 3 and receiving first recurrent vein near middle; basal vein meeting or beyond cu-v; jugal lobe of hind wing terminating well before cu- v and scarcely over half as long as vannal lobe. m. S2-5 of female with distinct preapical bands of mostly simple, suberect hair similar to that of Perditomor- pha; hairs markedly shorter than length of ster- num. S6 of male unmodified. Pygidial plate of male absent, not represented by area of sparser hair. n. S7, S8, and genitalia of male as illustrated by Toro (1973), not differing from those of Perdito- morpha; S7 with lobes slender as in L. (Chilicolletes) delahozit. Distribution Central Chile. Included species: Lewoproctus (Torocolletes) fazit (Herbst, 1923) [Lonchopria] and ibanez (Ruiz, 1944) [Lonchopria|]. (Some specimens at Lawrence are probably erroneously labeled as from a lo- cality in Cordoba Proy., Argentina; see Daly et al., 1987, for comments on similarly labeled Manuelia.) Comments: Both species of Torocolletes were er- roneously included in Lonchopria by Michener (1965). Although the sternal hairs of the female resem- ble those of most Perditomorpha and Chilicolletes, specimens with tibial pollen loads do not have pollen on the sternum as is usual in Perditomorpha. In its distinct malar space, Torocolletes resembles Nesocolletes from New Zealand. The latter sub- genus, however, has only one pair of small later- 652 oapical lobes on S7 of the male, a subvertical propodeal profile (1.e., without the long sub- horizontal surface found in Torocolletes), and is punctate and shining (1.e., without the distinctive dull cuticle of Torocolletes). The somewhat elongate malar space is presumably of independent origin in the two groups. Some undescribed Australian species of Lezoproctus also have elongate malar areas but in other characters are unrelated to Nesocolletes and Torocolletes. Etymology: ‘The subgeneric name is a_pat- ronymic for Haroldo Toro of Valparaiso, Chile, who first characterized the group, plus the generic name Colletes. Subgenus Hexantheda Ogloblin Hexantheda Ogloblin, 1948:172. Type species: Hexantheda missionica Ogloblin, 1948, by original designation. The long, six- or seven-segmented labial palpi are unique. The metasoma of the male is robust like that of a female. (Figs. 11, 16b; Figs. 16-23, Ogloblin, 1948) Description: Agreeing with usual (not excep- tional) characters of Perditomorpha except as fol- lows: a. Length 8-12 mm. Male metasoma broad like that of female. Pubescence mostly whitish, forming at most weak pale tergal bands. b. Inner orbits slightly converging below (except for upper ex- THE UNIVERSITY OF KANSAS SCIENCE BULLETIN tremities) in female, almost parallel in male. c. Clypeus protuberant in front of eye by about eye width. d. Labrum over four times as wide as long; apical margin feebly convex in female, weakly concave in male; bare convex surface depressed medially in male. e. Mandible of male with upper margin basal to preapical tooth convex (Fig. lle). Max- illary palpus not attaining apex of galea. Labial palpus two-thirds as long as prementum, much longer than maxillary palpus, six- or seven-segmented (illus- trated as 6-segmented by Ogloblin, 1948). f. Vertex considerably extended and convex behind the unusually small ocelli. g. Male flagellum scarcely reaching mesoscutum, middle segments much broader than long (illustrated by Ogloblin, 1948). h. Propodeum with subhorizontal surface a little shorter than metanotum; triangle shining, transversely wrinkled, marginal line not pitted. 1. Basitibial plate narrowly rounded at apex in female, pointed in male. Male with strong carina extending from apex of basitibial plate to apex of tibia. }. Tibial scopa rather sparse, some hairs on lower margin simple and nearly straight so that scopa extends farther below tibia than above, but most hairs near lower margin curved upward and with long branches as in Perditomorpha; hairs of most of outer surface simple or with one or two branches directed apicad, rachis surpassing branches. k. Tibial spurs gently curved, those of male and of mid tibia of female ciliate, hind spurs of female serrate except inner margin of inner spur which is i/- “Ns (\ Fic. 11. Levoproctus (Hexantheda) missionica (Ogloblin). a-c. Genitalia and lateral view of S8 of male. d. Inner hind tibial spur of female. e, f. Mandibles of male and female. CLASSIFICATION OF AMERICAN COLLETINAE pectinate with slender teeth (Fig. 11d). 1. Basal vein of forewing meeting cu-v. m. S5 of male with long apical fringe. T7 of male with convex, bare pygidial area, tapering posteriorly, as in L. (Per- ditomorpha) zonatus. n. S7, S8, and genitalia of male as illustrated by Ogloblin (1948); see also Fig. 1la-c. S7 with apical lobes not broad and rounded as is common in Perditomorpha, basal one deeply divided so that there are three lobes on each side instead of two. Process of S8 downcurved. Distribution: Southern Brazil (state of Parana) and northern Argentina (provinces of Misiones and Formosa). Included species: Leioproctus (Hexantheda) missionica (Ogloblin, 1948), new combination. Comments: Michener (1965) considered Hexanth- eda to be a genus distinct from Lezoproctus but the long labial palpi with a variable number of segments are the single character that might justify such placement. Hexantheda is probably a derivative of Perditomorpha-like ancestors with a few striking autapomorphies such as the labial palpi. Subgenus Protodiscelis Brethes Protodiscelis Brethes, 1909:245. Type species: Protodiscelis fiebrigi Brethes, 1909 (monobasic). Leioproctus (Protodiscelis); Michener, 1965:42. Protodiscelis resembles Tetraglossula in the simple mandibles of the male (a character found other- wise in American Lezoproctus only in the unrelated subgenus Cephalocolletes) and in the presence of apicolateral lobes on the very short labrum of the female. These are both apomorphies that unite the two subgenera. Protodiscelis differs from Tetra- glossula, however, in numerous features such as the scarcely elongate glossa and the plumose scopal hairs of both metasomal sterna and hind legss(Figs. 12; 16e, 196) Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha except as follows: a. Length 6-9 mm. Pubescence all pale or that of dorsum brown to dusky, sometimes scopa dusky; hairs forming at most weak apical bands on metasomal terga which usually lack hair bands. b. Inner orbits converg- ing below to nearly parallel; paraocular area convex in female except upper parts where there 1s a shiny, depressed, usually ill-defined fovea extending onto vertex and mesad toward ocelli. d. Labrum a transverse band six times as wide as long or nearly so, surface entirely occupied by convexity which in males and some females is lower medially, some- times with small weak median convexity between larger lateral ones; apical margin rather straight, with bristles longer than labrum, female with apicolateral lobe {usually smaller than in L. (P.) spathigerus, Fig. 12f, 1]. e. Maxillary palpus long, apex of third segment beyond apex of galea. Mandible of male not toothed, of female unusually slender with 653 preapical tooth small. Glossa with lobes unusually prominent, as broad as long or longer. h. Propod- eum with subhorizontal surface nearly as long as to longer than metanotum; triangle shining and nearly smooth or transversely striate, marginal line usually not pitted. Basitibial plate narrowly rounded to pointed at apex. j. Femoral corbicula usually with rather long, mostly simple hairs arising from surface. Hairs on anterior and posterior sides of corbicula, also hairs of trochanter, long, rather dense, with numer- ous short branches (similar to those of tibial scopa. Tibial scopa strong but not obscuring surface, hairs of lower margin bent apicad but not up- ward, hairs extending farther below than above tibia, strongly plumose, rachis surpassing the numerous short branches (Fig. 19e); hairs of inner surface of tibia mostly plumose but on distal half an area of long, unbranched but apically capitate kei- rotrichia; hairs of hind basitarsus on outer surface and upper margin plumose. k. Tibial spurs straight or gently curved, inner hind tibial spur finely pectinate (Fig. 121, k), others ciliate. 1. Jugal lobe of hind wing slightly to distinctly exceeding level of cu-v. m. 82-6 of female with apical bands of sloping to erect hair; hairs considerably shorter than exposed parts of sterna, plumose with nu- merous short branches like scopal hairs of hind leg. n. S7, S8, and genitalia as in Fig. 12a-d. Gonostylus much narrower than body of gono- coxite, Distribution: Northeastern Brazil (Paraiba) to the state of Parana and to Paraguay. Included species: Lewoproctus (Protodiscelis) fiebrigi (Brethes, 1909), spathigerus Michener, n. sp., and probably fpalpalis (Ducke, 1908) [Panurginus]. There are also several undescribed species. In the American Museum of Natural History there are ‘‘typus’’ specimens of an apparently unpublished Friese species named for its piceous coloration. An undescribed species from the State of Rio de Janeiro, Brazil, is unusually large (9 mm long) with the head longer than broad. The male is unknown; two females are at [Lawrence]. I have not seen authentic specimens of L. palpalis; its tentative placement here is based on specimens identified largely because of Ducke’s remarks on its large maxillary palpi. Comments: Recognition of the subgeneric name Protodiscelis has been difficult. Ducke (1912) and Cockerell (1919) placed it with Oedtscelis (= Chili- cola) in the Xeromelissinae, perhaps largely be- cause of the name, while Schrottky (1913) included it in the ‘‘Prosopidae’’ (= Hylaeinae). For one familiar with ordinary, rather hairy paracolletines, the subgeneric name and place- ments in Xeromelissinae or Hylaeinae seem to indicate a form not in the Paracolletinae. How- ever, some species of Protodiscelis such as L. (P.) jiebrigi (Brethes) have such short and sparse hair that the males superficially resemble Chilicola or other Xeromelissinae. As explained in the Appen- 654 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN “ 7, CteZ Zz, O Fic. 12. a-d. Letoproctus (Protodiscelis) fiebrigi (Brethes), male holotype, genitalia, S8 and S7. e-i. L. (P.) palpalis (Ducke)? e, f. Labrum, male and female. g, h. Mandible, male and female. i. Inner hind tibial spur of female. j-o. L. (P.) spathigerus n. sp., female. j. Mandible. k. Inner hind tibial spur. 1. Labrum. m, n. Ventral and anterior views of middle tibial spur, apex of tibia, and basitarsus. o. Ventral view of hind tibial spurs, inner spur above. dix, Brethes described a male, thinking that it was a female; of course it did not have a scopa and other features of female Paracolletini. Moure (1951) discussed Protodiscelis briefly among groups now included in the Paracolletini, but gave no characters or indications of the reasons for that placement. It is here properly placed, near Tétraglossula, for the first time. Subgenus Tetraglossula Ogloblin Tetraglossula Ogloblin, 1948:165. Type species: Tetra- glossula deltivaga Ogloblin, 1948, by original designa- tion. This subgenus was not included in Lezoproctus by Michener (1965) but it is as much part of that genus as are the other subgenera. The simple mandible of the male shows its relationship to Protodiscelis, which has a slightly more bifid glossa than most subgenera, thus approaching Tetragloss- ula. The deeply bifid glossa suggests Glossopasi- phae, which, however, has a subapical mandibular tooth in the male and plumose scopal hairs. (Figs. 13, 16f; Figs. 1-15, Ogloblin, 1948) Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha except as follows: a. Metasoma sometimes partly or largely yellowish red. Length 6-12 mm. Meta- somal hair bands absent or weak. b. Jnner orbits not or very slightly converging below (both sexes), almost CLASSIFICATION OF AMERICAN COLLETINAE 655 without concave region so that upper parts of orbit are not greatly different in orientation from re- mainder. d. Labrum about six times as wide as long; apical margin weakly concave to weakly convex; male with basal three fifths or more shining, convex, weakly binodulose; female with most of surface strongly elevated, binodulose, be- tween nodules with transverse depression surrounded by carinae; lateral lobes weak in male, conspicuous and serrate or pectinate in female; apical bristles much longer than labrum. Apzcal margin of clypeus of male with small median convex lobe overhanging labrum. e. J Maxillary palpus long, last or last two segments extending beyond apex of galea. Mandible simple in male (Ogloblin, 1948, Fig. 8), slender in female. Glossa deeply bifid, each lobe attenuate, hairy, seven to ten times as long as basal width (Ogloblin, 1948, Fig. 9) annulate part extending well out into base of each lobe. Galea much longer than stipes, postpalpal part pointed apically, four or five times as long as basal width (Ogloblin, 1948, Fig. 9). f. Ocelli in front of summit of vertex; vertex convex and rounded behind ocelli; ante- rior margin of median ocellus in front of to Fic. 13. Letoproctus (Tetraglossula) fucosus n. sp. a-e. Genitalia, S8 and S7 of male. f. T7 of male. g, h. Labrum of male and female (g at twice the magnification of h). i. S6 of male. j, k. Mandibles of female and male. 656 behind midpoint between antennal bases and posterior margin of vertex. g. Male flagellum not attaining tegula, middle flagellar segments dis- tinctly broader than long. h. Propodeum with basal zone in profile sloping, about as long as metanotum; triangle smooth with marginal groove strongly pitted. i. Apex of basitibial plate narrowly rounded. j. Corbicular surface with some simple hairs on basal half. Femoral and tibial scopa, and that of distal part of trochanter, of sparse simple hairs (in L. fucosus many hairs of outer surface of tibia, except lower margin, with one or two branches, often three on hairs of upper margin); hairs of lower margin of tibia curved apicad but not extending farther below tibia than above. k. Tibial spurs straight or gently curved, inner margin of inner hind tibial spur pectinate, other margins ciliate (Ogloblin, 1948, Fig. 4). 1. Basal vein of forewing meeting or distal to cu-v (Ogloblin, 1948, Fig. 3); jugal lobe of hind wing exceeding level of cu-v. m. 77 of male with tr- angular, bare pygidial plate, tapering to narrow rounded apex on tergal margin and margined laterally by weak carina, beyond which surface is densely hairy. S2-5 of female with scopa of simple hairs longer than length of sternum. n. S7, S8, and genitalia of male as in Fig. 13a-e (also Ogloblin, 1948, Figs. 5-7, 13-15). Distribution: Argentina (provinces of Buenos Aires, Tucuman, and Misiones) and Brazil (states of Parana, Sao Paulo, and Para). Included species (*marks species not seen by me): Letoproctus (Tetraglossula) anthracinus Michener, n. sp.; bigamicus (Strand, 1910) (= Tetraglossula bira- beni Ogloblin, 1948, according to Moure, 1951), new combination; deltivagus (Ogloblin, 1948), new combination; fucosus Michener, n. sp.; and *stigmaticus (Strand, 1910), new combination. Comments: The species of Tétraglossula are quite diverse in appearance, although morphologically similar. L. (T_) deltivagus, bigamicus, and stigmaticus are black with pale pubescence and thus rather ordinary looking gray bees. L. (7!) anthracinus is black haired. L. (T.) fucosus is a smaller species with a largely reddish metasoma; superficially it exactly resembles females of L. (Perditomorpha) brunerit. Reedapis new subgenus Grupo L. semicyaneus Toro 1973a:162. Type species: Leioproctus bathycyaneus ‘Toro, 1973: Superficially the rather large, robust species of Reedapis closely resemble those of Spinolapis that are found in the same area. Reedapis differs from Spinolapis in having three submarginal cells, bifid claws in the female, rather uniformly punctured and hairy clypeus, rather short male flagellum, etc. The two groups are not closely related in spite of superficial appearance. Reedapis is actually more THE UNIVERSITY OF KANSAS SCIENCE BULLETIN similar to Perditomorpha and Chilicolletes. It differs from those subgenera not only in appearance (and from Perditomorpha in having three submarginal cells) but also in the distinct row or comb of hairs on the outer margin of the front basitarsus of the female and in the strongly pectinate condition in females and some males of both inner and outer hind tibial spurs and of the mid tibial spur. The closest relative of Reedapis is Cephalocolletes; these subgenera differ conspicuously in the characters listed in the key. (Figs. 14h, i, 16c, 19g; Figs. 110-114, 133-152, Toro, 1973a) Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha except as follows: a. Integument black with the metasoma at least weakly metallic blue. Length 9 to 15 mm. Head, venter of thorax, and metasoma with much black hair. T1-3 with apical bands of white hair, sometimes weak on T1 and weakly indicated on T4. b. Inner orbits slightly to strongly converging below. d. Labrum over three times as wide as long, apex straight or broadly emarginate with bristles longer than labrum; basal elevated bare zone strong and well defined in female, longer than distal depressed zone, in male not well defined but occupying much of labral surface (for figures, see Toro, 1973a). f. Anterior margin of median ocellus at or behind midpoint between antennal bases and posterior margin of vertex. g. Middle flagellar segments of male a little longer than broad. h. Propodeum with subhorizontal surface about as long as meta- notum, which is convex and elevated medially in L. semicyaneus; propodeal triangle smooth to transversely wrinkled. i. Anterior basitarsus of female with comb-like row of hairs on outer margin. }. Femoral and tibial scopa well developed, blackish. Hairs in front of femoral corbicula with several long branches but main rachis usually surpassing api- ces of branches; hairs behind femoral corbicula with only few short branches. Tibial scopa with lower hairs not much curved upward, not extend- ing farther below tibia than upper hairs extend above it, lower hairs with several long branches usually surpassed by rachis (Fig. 19g), hairs of upper part with lateral branches conspicuously surpassed by rachis. k. Middle and hind tibial spurs straight to curved, all coarsely pectinate in both sexes of L. (R.) semicyaneus, all pectinate in female and finely pectinate in male of L. bathy- cyaneus (illustrated by Toro, 1973a, whose illustra- tions represent inner and outer find tibial spurs, contrary to the captions). 1. Wings hairy through- out but basal halves with hairs sparse and rela- tively long; three submarginal cells, second usually subequal to or longer than third on posterior margin except in occasional individuals in which second is much shorter than third; second receiving first recurrent vein near middle. m. $2-5 of female with scopa of dusky or blackish somewhat appressed plumose hairs nearly as long as exposed parts of sterna, branches mostly di- CLASSIFICATION OF AMERICAN COLLETINAE 657 Fic. 14. a-f. Letoproctus (Cephalocolletes) laticeps (Friese). a-e. Genitalia, S8 and S7 of male. f, g. Mandibles of female and male. h, i. L. (Reedapis) bathycyaneus Toro, male, genitalia and S8 (lateral view). rected apicad (Fig. 19h). S5 of male with long, in- curved marginal hairs. n. S7, S8, and genitalia of male as illustrated by Toro (1973a). Gonostylus and gonocoxite fused on dorsal surface but sepa- rable on ventral surface, gonostylus about as long as but much more slender than gonocoxite. S7 with lateroapical lobe deeply bifid, or in L. bathycyaneus according to Toro (1973a), basal divi- sion much reduced. (In a specimen determined by Toro as L. bathcyaneus, however, the lobe is divided more or less as in semicyaneus; perhaps there are more species.) Distribution: Central Chile. Included species: Levoproctus (Reedapis) semicyaneus (Spinola, 1851) [Colletes]; bathycyaneus Toro, 1973; and *melanocyaneus Toro, 1973. Biglossa caerulescens Friese, 1906, is a synonym of L. (Reedapis) semi- cyaneus (Spinola) and a homonym of L. (Spinol- apis) caerulescens (Spinola, 1851). This synonymy was omitted by Toro (1973a). Comments: An unusual feature of L. semicyaneus is the presence of two subantennal ‘‘sutures’’ or lines below each antennal base. They converge toward the clypeus and meet just above the upper margin of the clypeus, so that the subantennal area is triangular. It is smooth, impunctate, unlike adjacent parts of the face. The inner line probably represents merely a change in surface sculpture but it is fully as conspicuous as suban- tennal sutures in many Andrenidae. L. bathy- cyaneus has no such subantennal areas. The subgeneric name was used (at the genus level) on labels over 30 years ago by Padre J. S. Moure but has not been published. I use Reedapis as a patronymic in recognition of Edwyn C. Reed, well known for a study of Chilean bees, plus the generic name Afzs. Since I have not seen type or authentic material of L. semicyaneus, I have designated L. bathycyaneus as the type species, although I have no reason to doubt the customary identification of the common species, L. semicy- aneus. Cephalocolletes new subgenus Type species: Biglossa laticeps Friese, 1906. 658 Cephalocolletes differs from Reedapis in having the head very broad so that the interocular distance is much greater than the eye length, the eyes scarcely converging below in males and parallel in females, the vertex much enlarged (see char- acter f below), the labrum slightly less than three times as wide as long (with the apex convex in the female), the mandible of the male edentate (Fig. 14¢), the female sternal scopal hairs with branches somewhat longer and less numerous than illustrated for L. bathycyaneus, and T7 of the male with a longitudinal median bare line. The relationship of Cephalocolletes to Reedapis is shown by the male genitalia and sterna, the scopa, the large second submarginal cell, etc. (Figs. 14a-g, 16d) Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha except as follows: a. Integument and pubescence black. Length 10-16 mm. b. Inner orbits parallel in female, feebly converging below in male; head unusually broad, interocular distance much greater than eye length. c. Clypeal hairs short, especially in female. d. Labrum nearly three times as wide as long, apex convex in female, weakly emarginate in male, bristles as long as labrum; basal elevated zone strong and well defined in both sexes, longer than distal depressed zone. e. Mandible simple in male. f. Ocelli far from posterior margin of much developed, convex vertex; all ocelli closer to antennal bases than to posterior margin of vertex. g. Male flagellum with middle segments a little longer than broad. h. Propodeum with horizontal surface a little longer than metanotum. 1. Anterior basitarsus of female with comb-like row of hairs on outer margin. }. Femoral and tibial scopa as in Reedapis. k. Inner hind tibial spur of female with inner margin coarsely pecti- nate; otherwise spurs finely pectinate (female) to ciliate (male). 1. Wings as described for Reedapis. m. Exposed sterna and terga as described for Reedapis but scopal hairs mostly with longer and less numerous branches than illustrated for Reed- apis and T7 of male with longitudinal median bare line. n. S7, S8, and genitalia of male as in Fig. 14; S7 with only one large lateroapical lobe on each side, basal lobe greatly reduced. Distribution: Argentina (provinces of Tucuman, La Rioja, Catamarca and Mendoza). The coarse pollen on the females suggests that this bee is a cactus visitor. Included species: Leioproctus (Cephalocolletes ) laticeps (Friese, 1906) [Biglossa| Comments: Cephalocolletes superficially resembles black species of Brachyglossula or even the large black-haired species of Lonchopria (Biglossa); it differs from those groups by the generic charac- ters, Cephalocolletes and Reedapis are the only subgen- era of Levoproctus with the second submarginal cell rather long compared to the third (except in a few THE UNIVERSITY OF KANSAS SCIENCE BULLETIN individuals of Reedapis). See couplet 3 of the key to subgenera for details. This is a Lonchopria-like feature but in other respects these are clearly groups of Lezoproctus. Subgenus Spinolapis Moure Pasiphae Spinola, 1851:226 (not Latreille, 1819). Type species: Pasiphae caerulescens Spinola, 1851, designated by Sandhouse, 1943:585. Spinolapis Moure, 1951:193. Type species: Pasiphae cae- rulescens Spinola, 1851, by original designation. Lewoproctus (Spinolapis); Michener, 1965:41. From the superficially similar Reedapis, Spinol- apis differs by having only two submarginal cells, by the reduced (or absent) inner tooth on the claws of the female, by the finely pectinate or ciliate tibial spurs, etc. (Figs. 15, 16g; Figs. 136-139, Toro, 1973b) Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha except as follows: a. Integument largely metallic blue or head and thorax black. Length 9-12 mm. Head, venter of thorax, and metasoma with much of hair black; metasoma without hair bands. c. Clypeus somewhat protuberant with broad, transverse, shining, sparsely punctate or impunctate, preapical band; basal part flat and closely punctate. d. Labrum as described for Reedapis (Fig. 15i, j, and Toro, 1973b for L. cyaneus = cerdai). e. Maxillary palpus with seg- ments 5 and 6 beyond apex of galea. Mandible with preapical tooth often worn completely away in females. f. Ocelli as described for Reedapis. g. Male flagellum reaching at least to scutellum, middle segments about twice as long as broad, or in L. cyaneus only reaching tegula and middle segments scarcely 1.5 times as long as broad. h. Propodeum with subhorizontal zone slanting, about as long as or shorter than metanotum, triangle shining or dull and granular in L. cyaneus; marginal line not or scarcely pitted. 1. Anterior basitarsus of female without or (in cyaneus) with comb-like row of hairs on outer margin. Claws of female simple or (in LL. cyaneus) with small tooth. j. Femoral and tibial scopa well developed, pale (in L. cyaneus) to black. Femoral scopal hairs rather short, not much curved, those in front of cor- bicula with two or three long branches but main rachis usually surpassing other branches; hairs behind femoral corbicula almost simple. Tibial scopa with lower hairs not much curved upward, not extending farther below tibia than upper hairs extend above it, lower and outer hairs with a few long branches, some with and others without a rachis surpassing other branches. k. Middle and hind tibial spurs of both sexes straight, ciliate (more coarsely so than in Perditomorpha s. str.) or in female of L. cyaneus, inner hind tibial spur finely pectinate (see Toro, 1973, as cerdai Toro). 1. Wing hairs as described for Reedapis; two submar- ginal cells; basal vein meeting or distal to cu-v; CLASSIFICATION OF AMERICAN COLLETINAE 659 Fic. 15. Letoproctus (Spinolapis) caerulescens (Spinola). a-e. Genitalia, S8 and S7 of male. f, g. Mandibles of male and female. h. Inner hind tibial spur of female. i, j. Labrum of female and male. Jugal lobe reaching or surpassing level of cu-v. m. S2-5 of female with rather short pollen carrying hairs, shorter than exposed parts of sterna, often somewhat erect, some simple, others with rather short, apically directed branches. S6 of male with narrower midapical lobe than in other subgenera, otherwise unmodified except for apicolateral densely hairy shoulders in L. caerulescens. Pygidial plate of male not defined but T7 of male with broad, longitudinal, median hairless ridge which in L. cyaneus is broad basally and tapers toward apex like a pygidial plate; apex of T7 densely hairy. Pygidial plate of female narrow and paral- lel-sided apically. n. S7, S8, and genitalia of male as in Figure 15a-e; in L. (S.) cyaneus the two apical lobes on each side of S7 are both broad and rounded, the apical process of S8 is shorter than in caerulescens, and the apical half of the gonofor- ceps is slender. Distribution: Central Chile and the Lake Dis- trict of Argentina to Tierra del Fuego. Included species: Levoproctus (Spinolapis) caerulescens (Spinola, 1851) [Paszphae| and a closely related species. L. (Spinolapis) cyaneus (Cockerell, 1915) [| Pasiphae| (= L. cerdai Toro, 1973) and a related species, perhaps L. (Spinolapis) melanurus (Cock- erell, 1917) [Pasiphae], probably belong here. They are smaller and more brightly metallic than the caerulescens group; other differences are indi- cated in the description above. An indication of their relationship to L. (S.) caerulescens is the reduced inner tooth of the claw of the female; the tooth is absent in caerulescens. Comments: The sternal scopa of the L. caerules- cens group is reasonably convincing as a scopa, although the hairs are shorter and more erect than in Reedapis. In the L. cyaneus group the hairs are still shorter and sparser, although in one of the two females available they are largely covered by a dense mass of pollen, showing that they func- tion as a scopa. 660 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Fic. 16. Forewings. a. Leioproctus (Torocolletes) fazi (Herbst). b. L. (Hexantheda) missionica (Ogloblin). c. L. (Reedapis) bathycyaneus Toro. d. L. (Cephalocolletes) laticeps (Friese). e. L. (Protodiscelis) palpalis (Ducke)? f. L. (Tetraglossula) fucosus n. sp. g. L. (Spinolapis) caerulescens (Spinola). h. L. (Nomiocolletes) Joergensent (Friese). Subgenus Nomiocolletes Brethes Nomuocolletes Brethes, 1909:455. Type species: Nomia Jjorgenseni Friese, 1908, by original designation. Letoproctus (Nomuiocolletes); Michener, 1965:41. In spite of its distinctive appearance due to the enamel-like metasomal bands, Nomziocolletes is similar to Perditomorpha although with three sub- marginal cells. The modified hind legs of most males suggest Pygopasiphae but are quite different and no doubt independently evolved features. As in Reedapis, Cephalocolletes, and some Spinolapis, there is a distinct row of hairs on the front basitarsus of the female. (Figs. 16h, 17, 18) Description: Agreeing with usual (rather than exceptional) characters listed for Perditomorpha except as follows: a. Metasoma black, in male broad like that of female (except for L. szmplici- crus), posterior margins of T1-4 of female, T1-5 or 6 of male with smooth, hairless, shiny apical bands of enamel- like green, yellowish, or whitish, punctate and hairy basally in L. simplicicrus. Length 10-12.5 mm; 8 mm in L. simplicicrus. Pubescence largely pale, CLASSIFICATION OF AMERICAN COLLETINAE 661 with or without black on vertex and dorsum of thorax; with dusky or black on posterior part of metasoma; not forming pale metasomal hair bands. f. Ocelli in front of summit of convex vertex, anterior margin of median ocellus in front of or near midpoint between antennal bases and posterior edge of vertex. g. Male flagellum reach- ing middle of tegula, middle segments longer than broad, or in L. simplicicrus reaching scutel- lum with middle segments 1.5 times as long as broad. h. Propodeum with base sloping, much shorter than metanotum, so that profile of pro- podeum is mostly steeply declivous; propodeal triangle smooth except in L. stmplicicrus, marginal line pitted or not. i. Anterior basitarsus of female with well-formed row of hairs along outer margin. Basitibial plate pointed (both sexes); male with narrow, smooth, shining, hairless ridge from behind basi- tibial plate to apex of tibia, or in L. simplicicrus with strong but hairy carina originating near apex of basitibial plate. j. Femoral and tibial scopa well developed, white or yellow, sometimes Fic. 17. Letoproctus (Nomuocolletes) joergenseni (Friese). a-e. Genitalia, S8 and S7 of male. f. Inner hind tibial spur of female. g. Hind leg of male. 662 blackish along upper margin of tibia; femoral corbicula with a few hairs arising from its surface; hairs on anterior side of corbicula abundant, long but not curled, form as in Perditomorpha; hairs on posterior side of corbicula abundant, mostly sim- ple except for few branched ones adjacent to basal half of corbicula; trochanteral hairs loosely plumose. Tibial scopa well developed, hairs of lower part as in most Perditomorpha, grading to- ward upper margin to hairs with the rachis much exceeding moderately long lateral branches. k. Hind femur of male swollen and hind tibia broad- ened apically (Fig. 17g) except in L. simplicicrus which has slender, unmodified hind legs. 1. Wing hairs on basal half of wing longer than on distal part of wing; three submarginal cells, basal vein of forewing meeting or basal to cu-v except distal to cu-v in L. simplicicrus; jugal lobe of hind wing reaching level of cu-v or not attaining that level in simplicicrus. m. S2-4 of female with apical bands of suberect, mostly simple hairs less than half as long as sterna; S5 with apical band of plumose hairs. S6 of male unmodified, posterior margin : it Z NGG i Palit, we WZ e SW SR wEs Ne SWS By THE UNIVERSITY OF KANSAS SCIENCE BULLETIN with shallow median concavity. T7 of male with- out pygidial plate, with sparsely hairy, shiny longitudinal median zone which is widest anteri- orly, through which runs longitudinal median ridge except in L. simplicicrus. n. S7, S8, and genitalia as in Figs. 17, 18. S7 with three or four lateral apical lobes, (two in simplicicrus), distal one with coarse apical teeth (except in simplicicrus); proc- ess of S8 directed somewhat downward, spiculum rather long, acute; apex of gonoforceps (except in simplicicrus) directed downward and mesad, perhaps a gonostylus separated from gonocoxite except on lateral surface, largely membranous. Distribution: From Bolivia and central Argen- tina (Rio Negro) to the northeast of Brazil (Ceara), mostly in xeric areas, with the odd species, L. semplicicrus, in Peru. Included species: Leioproctus (Nomuocolletes) arnaut (Moure, 1949); *cearensis (Ducke, 1908) [Noma]; *ensent (Friese, 1906); joergenseni (Friese, 1908) [Nomia]; and simplicicrus Michener, n. sp. Comments: As is obvious from the above de- Fic. 18. Leioproctus (Nomiocolletes) simplicicrus n. sp., male holotype. a-e. Genitalia, S8 and S7. f. Mandible. CLASSIFICATION OF AMERICAN COLLETINAE 663 scription, L. s¢mplicicrus is quite different in nu- merous characters from the other species of the subgenus. The female might further emphasize the difference from other Nomuocolletes; it must be noted that in the above description characters recorded for females are based only on the other species for the female of L. szmplicicrus is un- known. There are enough similarities between that species and the others to suggest that they are sister groups. If one ignores the tergal banding, elaborate S7, and hind tibial carina of the male, characters that indicate its association with Nomi- ocolletes, L. simplicicrus agrees best with such rela- tively plesiomorphic subgenera as Chilicolletes. Genus Brachyglossula Hedicke Brachyglossa Friese, 1922:577 (not Boisduval, 1829, and others). Type species: Brachyglossa rufocaerulea Friese, 1922 (monobasic). Brachyglossula| Hedicke, 1922:427 (replacement for Brachyglossa Friese). Type species: Brachyglossa rufo- caerulea Friese, 1922 (autobasic). Lewproctus (Brachyglossula); Michener, 1965:41. This genus of large, dark haired, unbanded bees is distinctive in appearance [except for the superficially similar Lezoproctus (Cephalocolletes) lat- iceps|. In some major features such as the ves- titure and form of the hind basitarsus of the female and the shape of the process of S8 of the male, Brachyglossula resembles Lezoproctus. The italicized characters in the description are sufh- ciently marked and unique, however, to support recognition at the genus level even though the result at least for the time being is a paraphyletic H Fic. 19. a-g. Hairs from the lower part of the tibial scopa. a. Lewoproctus (Perditomorpha) erithrogaster Toro and Rojas. b. L. (Perditomorpha) brunerit (Ashmead). c. L. (Letoproctus) fulvoniger n. sp. d. L. (Glossopasiphae) plaumanni n. sp. e. L. (Protodiscelis) palpalis (Ducke)? f. L. (Pygopasiphae) wagneri (Vachal). g. L. (Reedapis) bathycyaneus Toro. h. Hair from sternal scopa of L. (Reedapis) bathycyaneus Toro. 664 genus Leoproctus (see section on Relationships Among Paracolletine Bees). (Figs. 20, 31a) Description: 1. Integument black or metallic blue, metasoma sometimes red. Length 12-16 mm. Vestiture largely black, metasomal hair bands completely absent. 2. Face rather flat, supraclypeal area not elevated above frons or other adjacent areas. Inner orbits diverging below. Malar space short. Clypeus somewhat protuberant, punctured throughout. 3. Facial fovea as de- scribed for Lonchopria. 4. Head often markedly convex both in front of and behind ocelli; vertex convex seen from front; development of vertex behind ocelli variable, so that anterior margin of median ocellus is behind to in front of midpoint between antennal bases and posterior margin of vertex. Preoccipital carina absent. 5. Mandible of female with strong preapical tooth on upper margin, of male with that tooth broad and weakly to THE UNIVERSITY OF KANSAS SCIENCE BULLETIN strongly emarginate so that mandible is at least weakly tridentate. 6. Labrum of female at least four times as wide as long, almost entirely a broad, shiny convexity, wth a strong transverse carina before de- clivity to depressed margin; apical depressed margin narrow, apical fringe much longer than labrum. Labrum of male two to three times as wide as long, smooth, shining, flat or with lateral basal convex- ities, without usual transverse convexity; apical margin weakly concave to straight, apical fringe slightly to much longer than labrum. 7. Proboscis without unusual features except long palpi; labial palpus over half as long as prementum; maxillary palpus with last four segments beyond apex of galea. 8. Male flagellum short, reaching pronotal lobe, middle segments much broader than long. 9. Propodeum almost entirely steeply declivous in profile, only upper margin sometimes slanting; posterior part of tho- racic dorsum is the convex metanotum, posterior Fic. 20. Brachyglossula bouvieri (Vachal). a-e. Genitalia, S8 and S7 of male. f, g. Mandibles of male and female. h, i. Inner hind tibial spur of female, inner and outer margins. j. Inner margin of inner spur of male. k, 1. Labrum of male and female. CLASSIFICATION OF AMERICAN COLLETINAE margin of which is vertical; propodeal triangle shining, finely roughened, marginal line weakly pitted anteriorly, not pitted posteriorly. 10. Front basitarsus of female hairy, without well developed comb. 11. Middle and hind tibial spurs strongly curved, coarsely pectinate with long teeth in female (inner margin of inner hind spur, outer margin of outer hind spur, outer margin of middle spur; other margins coarsely serrate); teeth shorter in male, inner margin of inner hind tibial spur nearly edentate in some males. 12. Femoral and tro- chanteral scopa not well developed, hairs little longer than femoral diameter; tibial scopa rather short, not hiding surface, longest hairs on lower margin of tibia, these not curled upward or scarcely so; all scopal hairs with strong axes and fine side branches at more or less right angles to axes except that posterior to femoral corbicula branches are short, inconspicuous and directed apicad and on upper part of tibial scopa hairs are dense with apically directed branches. Kei- rotrichia of female of moderate length, well differ- entiated from other tibial hairs, of male ordinary. 13. Hind basitarsus of female slender, scarcely tapering apically, apex more than half as wide as width near base, outer surface rather flat; hairs of outer surface longer than those of inner surface, espe- cially basally. Hind basitarsus of male scarcely wider than apex of second tarsal segment. 14, 15. Basitibial plate about one fifth as long as tibia or in males even shorter, defined by carinae, rounded at apex and covered by hair except marginally in female, apex narrowly rounded in male and surface hairy but not hidden; no carina extending apicad. 16. Wings hairy throughout or basal cells partly bare, all hairs minute; two submarginal cells; basal vein meeting or basal to cu-v. Stigma small, costal margin of marginal cell 2.5 to 3 times as long as stigma which is not broader than prestigma (measured to wing margin), vem r arising well beyond middle, margin within marginal cell slightly convex. Apex of marginal cell as described for Lonchopria. 17. Jugal lobe of hind wing reaching level of cu-v. 18. T1 markedly narrower than T2, in female about three fourths and in male two thirds width of T2. S2-5 of female with well developed, dense, plumose scopa, hairs with numerous fine side branches more or less at right angles to axes. 19. S6 of male unmodified, apical margin rounded or with small median emargination. Pygidial plate of male absent but T7 with longitu- dinal, median, shiny line. Pygidial plate of female with apex narrowly rounded, lateral margins diverging strongly anteriorly, nearly straight. 20-22. S7, S8, and genitalia of male as in Fig. 20. S7 with a single apicolateral lobe; S8 without usual ventral bump, apical process as in Leioproctus Group. Gonocoxite attenuate apically, this por- tion possibly a gonostylus; volsella vertically ex- panded, reaching dorsum of genital capsule, deeply bifid in dorsal or ventral view; penis valve strongly curved downward and deeply bifid apically, distal part free from penis which is broadly sclerotized laterally. 665 Distribution: Bolivia and Argentina (provinces of Misiones, Jujuy, La Rioja, Catamarca, etc.). Some and perhaps all visit flowers of Cactaceae. Included species: Brachyglossula boliviensis (Vachal, 1901) [Pasiphae], new combination; bouviert (Vachal, 1901) [Pasitphae]; personata (Cockerell, 1939) |Pasiphae|]; and rufocoerulea (Friese, 1922) [Brachyglossa]. There seem to be a few other species, evidently undescribed. Genus Niltonia Moure Niltonia Moure, 1964:52. Type species: Neltonia virgilu Moure, 1964, by original designation. The extremely long labial palpi are the out- standing feature of this genus but this is by no means a Leioproctus with long palpi, as shown by a whole series of other unique features italicized in the description below. (Figs. 21, 31b; Fig. 1, Moure, 1964; Figs. 1-3, Laroca and Almeida, 1985). Description: 1. Integument black, nonmetallic. Length 10-12.5 mm. Vestiture of female largely black, of male mostly gray, metasomal hair bands completely absent. 2. Supraclypeal area not much elevated above frons or clypeus. Inner orbits converging below. Malar space short. Clypeus convex, protuberant (less than eye width), punc- tured throughout. 3. Facial fovea as described for the Lonchopria group. 4. Head strongly biconvex in front of ocelli; vertex convex behind ocelli and seen from front; anterior margin of median ocellus behind midpoint between antennal bases and posterior margin of vertex. Preoccipital car- ina absent. 5. Mandible with preapical tooth on upper margin. 6. Labrum about four times as wide as long with strong median elevation over one third as wide as labrum in female, less than one third as wide in male; apical margin feebly convex, apical fringe much longer than labrum. 7. Proboscis short, galea rounded at apex, glossal lobe much shorter than basal width; labzal palpus enormous, 8-9 mm long, in repose reaching S3 or S4, fourth segment much longer than first three together, tapering, often extended beyond apex of metasoma, irreg- ularly toothed (Fig. 1 of Moure, 1964; Fig. 3 of Laroca and Almeida, 1985); maxillary palpus with last three segments beyond apex of galea. 8. Male flagellum short, reaching pronotal lobe, middle segments longer than broad. 9. Propod- eum mostly steeply declivous in profile but with slanting basal zone shorter than metanotum, propodeal triangle finely roughened, marginal line not pitted. 10. Front basitarsus of female hairy, except under side abruptly only sparsely so, no well developed comb. 11. Middle and hind tibial spurs robust, curved, with numerous fine teeth, finer in male than female, longest on inner margin of inner hind spur and outer margins of middle and outer hind spurs; inner hind tebial spur much longer than outer. 12. Scopa on leg rather 666 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Fic. 21. Niltonia virgilit Moure. a-e. Genitalia, S8 and S7 of male. f. Claw of male. g, h. Inner and outer hind tibial spurs of female. i, j. Same, of male. k, 1. Mandibles of male and female. m, n. Labrum of male and female. short, hairs with major branches such that there is no distinct rachis distally except on upper part of tibia; hairs behind femoral corbicula with few branches, many merely bifid; lower hairs of tibial scopa curved distad but not upward. Keirotrichia of moderate length, well differentiated from other hairs. 13. Hind basitarsus of female slender, slightly tapering apically, apex more than half as wide as width near base, outer surface flat; hairs of outer surface longer than those of inner sur- face. Hind basitarsus of male only slightly wider than apex of second tarsal segment. Claws (both sexes) with inner and outer rami similar in shape, slender, of almost equal length, so that claws are deeply bifid. 14, 15. Basitibial plate a little less than one third as long as tibia in female, one fourth in male, defined by carinae, almost pointed, with short hairs not obscuring surface; male with strong ridge, almost a carina but hairy, from apex of basitibial plate to apex of tibia. 16. Wings minutely hairy through- out; two submarginal cells; basal vein meeting or slightly distal to cu-v. Stigma rather small, costal margin of marginal cell over twice length of stigma which is not broader than prestigma (measured to wing margin), vein r arising well beyond middle, margin within marginal cell convex. Apex of marginal cell as described for the Lon- chopria group. 17. Jugal lobe of hind wing exceeding level of cu-v. 18. S2-5 of female with well developed scopa of hairs that have several long branches so that there is no clear rachis apically. 19. S6 of male with median, broad, shallow, shining, impunctate concavity; apical margin shallowly emarginate. T7 of male with triangular pygidial plate, sharply pointed apically, margined by carinae laterally. Pygidial plate of female rounded apically. 20-22. $7, S8, and genitalia of male as in Fig. 20a-e and Fig. 1 in Laroca and Almeida (1985). S7 with a single apicolateral lobe plus small basal membranous lobe, hairs on these lobes marginal, i.e., neither dorsal nor ventral; S8 without usual ventral bump; apical process extremely short, terminating as in Leio- proctus group. Apex of gonoforceps bilobed, volsella with apex expanded dorsally, reaching dorsum of genital CLASSIFICATION OF AMERICAN COLLETINAE capsule; penis valve strongly curved downward at apex, apodemes of penis valves parallel, not curved upward proximally; penis with long sclerotic strips on lower surface. Distribution: Brazil, states of Santa Catarina to Rio de Janeiro. Included species: Niltonia virgilii Moure, 1964. Comments: In certain features Niltonia resembles Brachyglossula. ‘The dorsally expanded volsellae and the distal origin of vein r on the stigma are the most apparent such characters. Perhaps both genera arose from a common ancestor within the Leioproctus group. Lonchorhyncha new genus Type species: Diphaglossa ecuadoria Friese, 1925. The elongate clypeus and malar area (about as long as the eye) led to placement of this bee in the genus Diphaglossa (subfamily Diphaglossinae) where it has remained until now. No other Ameri- can paracolletine has such a head. Other charac- ters quite unlike Leioproctus include those of S7, S8, and the genitalia of the male, as described below. The two ventral prongs on the penis valve suggest Lonchopria s. str. but other characters do not indicate a close relationship to that taxon. (Figs. 22a-1, 23a-d, 31c) Description: 1. Nonmetallic, metasomal hair bands and integumental color bands absent. Punctation mostly fine and weak but surface, especially of head and thorax, minutely rough- ened and dull. 2. Face protuberant and extremely produced; upper margin of clypeus below lower ends of eyes; supraclypeal area (measured to lower mar- gins of antennal sockets) over half as long as clypeus, longer than wide, not elevated above level of clypeus but strongly elevated above frons and paraocular area, frontal tubercle strong. Orbits subparallel in female, converging below in male. Malar space nearly as long as eye in female, slightly longer than eye in male. Clypeus convex with longitudinal median flat or slightly depressed zone but no differentiated depressed area as in most Lonchopria. 3. Facial foveae not recognizable although surface mesal to upper part of eye broadly concave. 4. Vertex extending only an ocellar diameter or less behind ocelli, seen from front not broadly convex, laterally (between ocelli and eye) concave and lower than summit of eye, medially a little elevated, bearing ocelli slightly higher than summit of eye. Distance from anterior margin of median ocellus to posterior margin of vertex about half of distance to antennal sockets. 5. Mandible of female short, slender medially but broadened apically, cap of rutellum large, pre- apical tooth on upper margin; of male slender, almost parallel sided except for broader base, with preapical tooth on upper margin. 6. Labrum of female unknown. Labrum of male over twice as wide as long; apical margin straight medially, 667 with long bristles; rest of labrum thickened, irregularly roughened, hairless. 7. Proboscis long but otherwise not unusual, stipes about three fourths as long as head; galeal comb of 15 bristles; apices of galeae broken off in only specimens so that length is unknown; maxillary palpus about half as long as stipes, of six subequal segments; glossa shallowly emarginate, lateral lobe about as long as broad; labial palpus over two fifths as long as prementum, segments subequal in length. 8. Antenna of male reaching base of scutellum, flagellar segment 1<2+ 3, nearly twice as long as broad, 2 about as long as broad, segments 3-9 ranging from less than 1.5 (seg. 3) to over 1.5 (seg. 9) times as long as wide, segment 10 over twice as long as wide. Antenna of female reaching tegula, flagellar segment 1 equal to 2+3, over twice as long as wide, 2 slightly broader than long, remaining segments about 1.5 times as long as broad except segment 10 which is twice as long as broad. 9. Propodeal triangle minutely rough- ened, nearly dull, marginal grooves not pitted, profile nearly vertical, dropping from posterior mar- gin of metanotum which is convex but largely declivous. 10. Front basitarsus of female, on outer margin with weakly differentiated comb of hairs or bristles longer than other basitarsal hairs, hairs of comb well separated (i.e., not closely placed), tapering, curved. 11. Inner hind tibial spur of female pectinate with about 10 teeth, the middle ones longest, bases well separated; of male ciliate with many fine, rather long teeth, some nearly as long as diameter of spur. 12. Femoral scopa weak; long, rather dense, straight, loosely plumose hairs present on underside of hind trochanter and base of femur; rest of femur with only short sparse hairs on lower surface, simple on distal half; long, branched, downcurved hairs on anterior surface of female partially enclose femoral corbicula. Hind tibia (female) with long branched hairs of lower surface curved outward and upward, along with hairs of outer surface forming a strong scopa which is not so dense as to fully obscure the tibial surface; hairs of inner surface simple, moderately long, not forming zone of short keirotrichia. 13. Hind basitarsus of female slender with outer surface convex, hairs longer, finer, and sparser than those of inner surface, mostly simple but those near basal half of upper margin plumose like those of tibial scopa, not obscuring surface. 14. Basitibial plate of female well defined, strongly elevated, with shining, marginal carina; apex narrowly rounded, almost pointed; surface largely hidden by abundant short hairs. Claws with strong inner tooth. 15. Basitibial plate of male similar to that of female but hairs not hiding surface; no carina. 16. Wings hairy throughout. Basal vein slightly distal to cu-v. Stigma slender, as wide as prestigma (measured to wing margin), vein r arising near middle, margin within mar- ginal cell slightly convex. Three submarginal cells, second and third subequal in length meas- ured either on anterior or posterior margins. Apex of marginal cell narrowly rounded or 668 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Fic. 22. a-i. Lonchorhyncha ecuadoria (Friese). a-d. Genitalia, S8 and S7 of male. e, f. Mandibles of male and female. g. T7 of male. h. Labrum of male. i. Inner hind tibial spur of female. j. Eulonchopria punctatissima, male genitalia. pointed, separated from costal margin by three or four vein widths. 17. Jugal lobe of hind wing half as long as vannal, only two thirds as long as cell Cu, far short of level of cu-v. 18. Metasomal sterna with hair of moderate length, sparser and shorter posteriorly in male. 19. Pygidial plate of male represented by large, slightly concave, hair- less area, not sharply defined except for apical emarginate flange which extends slightly beyond morphological posterior tergal margin. 20. S7 of male moderately narrowed medially, flat, with one pair of apical hairy lobes, each lobe with mesal, posteriorly directed process. 21. S8 of male weakly sclerotized, almost quadrate, without apical process and without spiculum. 22. Male genitalia without distinct gonostylus, gonoforceps robust. Volsella largely membranous, weakly sclerotized only laterally, without denticles. Penis valve compressed, down- curved at apex and with long median process projecting downward. Distribution: Ecuador. Included species: Lonchorhyncha ecuadoria (Friese, 1925). An account of the species is included in the Appendix. CLASSIFICATION OF AMERICAN COLLETINAE 669 Fic. 23. a-d. Lonchorhyncha ecuadoria (Friese). a. Head of female (type). b, c. Head of male. d. Outer side of hind leg of female. e. Outer side of hind leg of female, Lezoproctus (Kylopasiphae) pruinosus n. sp. Genus Eulonchopria Brethes This is a genus of coarsely punctured and sculptured bees with yellow integumental bands on some of the metasomal terga and often plaited forewings with darkened costal margins so that there is a superficial resemblance to eumenid wasps. The many unique features are italicized below. Description: 1. Body nonmetallic, largely coarsely punctate and coarsely sculptured; pubescence short, in Eulonchopna s. str. at anterior and posterior scutal angles and often elsewhere hairs so short that each fits within a puncture and is_ broadly plumose. Metasomal hair bands absent but some terga with apical integumental pale yellow bands in depressed marginal zones. Length 7-11 mm. 2. Face convex, supraclypeal area more protuberant than clypeus. Orbits converging below. Malar space very short. 3. Facial fovea absent or deeply impressed, well defined in both sexes, elongate, low on face (not reaching summit of eye). 4. Vertex convex and elevated far above upper ocular tangent seen from front. Anterior margin of median ocellus near or in front of midpoint between antennal bases and posterior margin of vertex. Preoccipital carina strong, often lamella-like. 5. Mandible (male and female) with preapical tooth on upper margin. 6. Labrum much broader than long with apical margin convex. 7. Proboscis without unusual features; labial palpus about one third as long as prementum; maxillary palpus with last one to two segments extending beyond apex of galea. 8. Antenna of male reaching scutellum, middle flagellar segments usually less than twice as long as broad. 9. Pronotum dor- solaterally with strong transverse carina or lamella extend- ing onto lobe. Metepisternal area deeply impressed and almost hairless. Propodeal triangle with large, deep pits; some ridges margining pits produced, lamella-like or tooth-like. Horizontal base of propodeum as long as or shorter than metanotum, less than one third as long as vertical surface in profile, horizontal and vertical surfaces separated by sharp angle or lamella. 670 10. Front basitarsus of female with outer apical process from which comb extends basad on outer edge of basitarsus. 11. Inner hind tibial spur of female coarsely pectinate (3 to 5 teeth); of male coarsely toothed or ciliate or hind tibial spurs completely absent. 12. Femoral scopa of rather short, curved hairs densely plumose toward apices*. Hind tibia with hairs of outer surface short and not scopa-like, especially on distal half of tibia, hairs of lower margin and inner surface longer, on inner surface (except basally) not plumose but not forming zone of short keirotrichia. 13. Hind basitarsus of female elongate, tapering, apex more than half as wide as maximum width near base, outer surface slightly concave below upper margin; hairs as long as or shorter than those of inner surface, not very different from those of outer surface of tibia, not obscuring surface. 14. Basitibial plate of female well defined, pointed or rounded apically, about one third as long as tibia. 15. Basitibial plate of male well defined, carina extending to apex of tibia. 16. Forewing commonly folded longitudinally (as in Vespidae) (see Danforth and Michener, 1988). Basal vein distal to or meeting cu-v. Stigma nearly parallel sided, vein r arising near apex and part of margin within marginal cell oblique, not convex. Three submarginal cells. Apex of mar- ginal cell obliquely bent away from wing margin or obliquely truncate. 17. Jugal lobe of hind wing variable. 18. Metasomal sternal hairs (both sexes) usually extremely short. Prepygidial fimbria of female strongest medially, fading away laterally. * 19. Pygidial plate of female with margin continued basolaterally nearly to lateral margin of T6, isolating depressed area on which pygidial fimbria arises. Pygidial plate of male ab- sent, surface of T7 extremely coarsely punctate (like preceding terga). 20. S7 of male narrow medially, with two or three pairs of broad lobes distally, these lobes bent so that hairs are on ventral surfaces. 21. S8 of male rather large and broad, with broad basal spiculum and broad apical region, sometimes continued as small apical process which is flat, not beveled like a pygidial plate, at apex. 22. Male genitalia sometimes with distinct slender gonostylus. Volsella small, without distinct denticles. Penis valve not strongly downcurved apically, with preapical lobes or processes on inner margin. Distribution: Arizona to Argentina, mostly in xeric or savanna areas. It occurs principally from central Brazil to Argentina and Bolivia and from Oaxaca through western Mexico to Arizona. It has also been collected in Nicaragua and Vene- zuela. It is the only paracolletine in North Amer- ica. Comments: Characters marked by an asterisk in the above description have not been verified for the subgenus Ethalonchopria. This is a genus of remarkable bees, with nu- merous distinctive characters italicized above. Most of these characters are unusual among bees, not shared by potential outgroups (Lezoproctus and Colletidae other than Paracolletini) and are ap- ‘THE UNIVERSITY OF KANSAS SCIENCE BULLETIN omorphies. The deep, rather slender, bare, well defined facial foveae (one subgenus), however, are shared with the Australian paracolletine Cal- lomelitta and with other colletid subfamilies such as Hylaeinae. This is probably a plesiomorphy. Likewise the distinct, slender, male gonostyli of some species, unique for the Colletidae, are a possible plesiomorphy. Its apparently disjunct distribution (absent in wet tropics) combined with its unusual characters and the diversity of the species suggest that Eulonchopria is an archaic group that possesses many specialized features. Recent studies of the genus are by Michener (1963, 1985). Genitalia and hidden sterna of males are illustrated in the first paper listed. An additional illustration of genitalia is included herein to provide a needed lateral view. KEY TO THE SUBGENERA OF EULONCHOPRIA 1. Facial fovea absent; anterior and lateral surfaces of mesepisternum not separated by carina...... Ethalonchopria —. Facial fovea distinct; anterior and lateral surfaces of mesepisternum sep- arated by carma,.... >, Eulonchopna s. str. Ethalonchopria new subgenus Type species: Apista gaullei Vachal, 1909. This subgenus differs in many features from the other subgenus. In nearly all of the subgeneric characters it is less strange than Eulonchopria s. str., 1.e., more like other paracolletines. Notewor- thy are the punctate and only slightly concave foveal areas on the face, so that distinct foveae are absent, and the simple axillae. (Michener, 1985, Fig. 1) Description: a. Facial fovea absent. b. Preoccipi- tal carina strong but not a lamella. c. Ridge mesal to posterior ocellus absent. d. Base of labrum convex in female, binodulose in male. e. Margin of dorsolateral pronotal carina straight. f. Ante- rior and lateral surfaces of mesepisternum not separated by carina. g. Axilla not produced and angulate posteriorly. h. Margins of basitibial plate largely hidden by hair in female; carina on upper margin of hind tibia of male not toothed. 1. First submarginal cell longer than second plus third (measured on posterior margins), second very small. Basal vein of forewing meeting cu-v. j. Jugal lobe of hind wing extending little more than half way from base to level of cu-v. k. Apex of T7 of male entire. Distribution: Bolivia; Santa Catarina, Brazil. Included species: Eulonchopria (Ethalonchopria) gaulle: (Vachal, 1909) and limbella (Vachal, 1909), CLASSIFICATION OF AMERICAN COLLETINAE which may be the same species (see Michener, 1985). Comments: The above description is based on a single male from Santa Catarina, Brazil, kindly loaned by T. L. Griswold [Logan] and on my earlier observations of the female types (the only known specimens) of the two nominal species from Bolivia (Michener, 1985). No females were available for supplementary observations. The male is clearly a member of this subgenus but its species is uncertain since there are no associated females. Etymology: The subgeneric name is based on the Greek ethas, ordinary, plus the generic name Lonchopria, with reference to the fact that these are more ordinary bees than the other Eulonchopria. Subgenus Eulonchopria Brethes s. str. Eulonchopria Bréthes, 1909:247. Type species: Euloncho- pria psaenythioides Brethes, 1909 (monobasic). This subgenus contains the more ornate and extraordinary members of the genus. The cari- nate mesepisterna and produced and angulate axillae are unique among Paracolletinae. (Fig. 22j; Figs. 1-15, Michener, 1963; Fig. 3, Danforth and Michener, 1988) Description: a. Facial fovea of female distinct. b. Preoccipital carina forming a strong lamella. c. Brow-like ridge mesal to posterior ocellus distinct to absent. d. Base of labrum binodulose. e. Margin of dorsolateral pronotal carina convex. f. Anterior and lateral surfaces of mesepisternum separated by angle that is carinate above. g. Axilla slightly produced and angulate posteriorly. h. Margins of basitibial plate fully exposed in both sexes; carina on upper margin of hind tibia of male toothed. i. First submarginal cell about as long as second plus third (measured on posterior margins). Basal vein of forewing beyond cu-yv. J. Jugal lobe of hind wing nearly attaining level of cu-v. k. Apex of T7 of male bilobed or bidentate. Distribution: Argentina (Tucuman Province) and Paraguay to Brazil (states of Santa Catarina to Minas Gerais); Venezuela; Nicaragua; Mexico (states of Oaxaca to Sonora), U.S.A. (southern Arizona). Floral records on labels and my obser- vations near Chamela, Nayarit, Mexico, indicate that E. punctatissima and oaxacana may be oligolec- tic visitors to flowers of Acacza. Included species: Eulonchopria (Eulonchopria) oa- xacana Michener (1963), psaenythioides Bréthes (1909), and punctatissima Michener (1963). There appear to be additional South American species. Comments: Eulonchopria s. str. contains two spe- cies groups. In the South American E. psaenythi- oides and its undescribed relatives the hind tibial spurs of the male are present, the brow-like ridge mesal to the posterior ocellus is strong, S8 of the 671 male lacks an apical process, and there are no slender male gonostyli. In the two North Ameri- can species, E. punctatissima and oaxacana, the hind tibial spurs of the male are absent, the brow-like ridge mesal to the posterior ocellus is weak to absent, S8 of the male has a small, apically expanded process, and there is a slender, well defined genital process, probably the gonostylus. Genus Lonchopria Vachal The very dense tibial scopa of the female, contrasting with short, sparse, simple or re- strictedly plumose hairs on the outer side of the hind basitarsus, is the most striking distinction from other American paracolletines. Lonchopria (Lonchoprella) is an exception to this character, however. Other characters italicized below also distinguish Lezoproctus and Lonchopria. Males of Lonchopria differ from those of Lezoproctus in the lack of a beveled apex of the process of S8. In Leioproctus the apex of the process of S8 is ex- posed, beveled, and superficially resembles a pygidial plate. The subgenus Ctenosibyne, which clearly is related to the subgenus Lonchopria, has an apex of the process of S8 that seems suggestive of that of Leioproctus. It is, however, broad, thin, and hairy, as in some species of the subgenus Biglossa, and is in reality not like that of Letoproc- tus. Description: 1. Integument black or metallic bluish, metasomal hair bands present or absent, terga without colored integumental bands. Length 7-15 mm. 2. Face flat to moderately convex, supraclypeal area not much elevated above clypeus but elevated above frons. Inner orbits converging below. Malar space very short. Clypeus commonly with depressed, median closely punc- tate area, lateral and distal to which are more shining convex areas (not so in Lonchopma s. str.). 3. Facial fovea not recognizable or indicated only by broad, undefined area of slightly different texture than rest of face. 4. Vertex weakly convex seen from front to strongly convex and elevated well above upper ocular tangent. Anterior margin of median ocellus variable in position. Preoccipital carina absent. 5. Mandible of female and most males slender with preapical tooth on upper margin (two such teeth in male of one species of subgenus Biglossa), commonly with apex in male broadened by expansion of lower margin or with process on lower margin. 6. Labrum variable, usually less than three times as wide as long. 7. Proboscis without unusual features or glossa moderately [L. (Biglossa) robertsi| to greatly (Porterapis) elongate and deeply bifid; labial palpus about one third as long as prementum (longer in Lonchoprella); max- illary palpus with little more than last segment (if any) extending beyond apex of galea, which is short (or most of last two segments exceeding galea in Lonchoprella). 8. Antenna of male reach- 672 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN ing tegula or beyond, middle flagellar segments 1.5 to over 2 times as long as broad. 9. Propodeal triangle smooth or finely transversely wrinkled, dull to shining; base of propodeum sloping, shorter than or equal to metanotum, curving onto declivous surface, about one third as long as declivous surface in profile. 10. Front basitarsus of female with comb of hairs on outer margin. 11. Inner hind tibial spur of female pectinate with four to eight very long teeth, their bases about as close as they can be so that they diverge from a sometimes somewhat thickened part of spur; of male ciliate with many fine and somewhat elongate teeth. 12. Femoral scopa of abundant, long hairs with numerous fine side branches, femoral corbicula closed basally by similar long, curved, plumose hairs arising on trochanter almost like floccus of Andrena. Hind tibia (female) with hairs of lower surface curled outward and up- ward, along with hairs of outer surface forming dense scopa of long, branched hairs usually en- tirely obscuring tibial surface, many of hairs lacking a single strong rachis but with several branches of more or less equal length, as in Leioproctus (Perditomorpha) (Fig. 19a) but often with more branches; hairs of inner surface of hind tibia simple, moderately long to long, not forming zone of short keirotrichia. (Lonchoprella is an exception to most scopal characters; see descrip- tion below.) 13. Hind basitarsus of female tapering toward extreme apex which is only about half as wide as width near base. Hind (also middle) basitarsus of female with outer surface usually slightly longitudinally concave below upper margin, vestiture entirely different from that of tibia, hairs short, not longer than those of inner surface [or slightly longer in L. (Biglossa) nivosa and Lonchoprella|, not obscuring surface. (Hairs of upper margin of hind basitarsus long, not to be confused with those of outer surface). 14. Basitib- ial plate of female well defined but often hidden by hair, pointed or rounded, about one fourth as long as tibia or less. 15. Basitibial plate of male well defined at least on posterior margin and apex; no carina extending apicad. 16. Wing vestiture variable; three submarginal cells; basal vein distal to or occasionally meeting cu-v. Stigma slender, not or little broader than prestigma (measured to wing margin), somewhat broader in the subgenus Porterapis; marginal cell less than 2.0 to 2.5 times as long as stigma; vein r arising near middle of stigma, margin within marginal cell convex, usually somewhat angulate. Apex of marginal cell narrowly rounded or pointed, sepa- rated from wing margin by a few vein widths. 17. Jugal lobe of hind wing not attaining to surpass- ing level of cu-v. 18. Metasomal sterna with abundant short hair and apical fringes of longer hair. 19. Pygidial plate of male absent or repre- sented by ill-defined elevated area that tapers toward apex of T7 which is rounded, truncate, or emarginate. 20. S7 of male narrow medially, with two pairs of apical lobes, one or both of which is rather narrow or small. 21. S8 of male variable, with base rounded or truncate, rarely pointed; apical process simple and pointed to large and ornate, not beveled like a pygidial plate. 22. Male genitalia with or without recognizable gonostylus. Volsella without distinct denticles except in Lon- choprella. Penis valves usually large and elaborate, not strongly downcurved. Distribution: Southern South America, north in the Andean uplift to Colombia. Comments: This genus contains numerous spe- cies. They are diverse in appearance, ranging from rather small forms superficially not easily distinguished from small or middle-sized Levoproc- tus to large and easily recognized forms like Lonchopria s. str. KEY TO THE SUBGENERA OF LONCHOPRIA 1. Mandible of male with large tooth or process on lower margin. Clypeus of female without or with weakly differentiated closely punctate upper median area; lateral clypeal area, if differentiated, merely less closely punctate than upper median area... . 2 —. Mandible of male often with ob- tuse preapical angle on lower mar- gin but without large tooth or process. Clypeus of female with flat or depressed, relatively closely punctate, upper median area con- trasting with impunctate or sparsely punctate, convex, U-shaped lateral anid apical: FEQIOM. 3.2)... ne tee ee oe 3 2. Pollex tooth of male mandible enor- mous, separated from apical part of mandible (rutellum) by curved emargination (Fig. 30e); S8 of male with apical process downcurved, apex expanded and quadrangular; mandible of female unusually slender (Fig. 30g), preapical (pollex) tooth weakly developed... . . Ctenosibyne —. Pollex of male mandible not forming the usual preapical tooth; S8 of male with apical process short, pointed, scarcely downcurved; mandible of fe- male more robust with well developed preapicaliteoth: 47.5 42: Lonchopria s. str. 3. Glossa deeply bifid, each lobe longer than prementum; apex of mandible scoop-shaped (Fig. 291, j), preapical (pollex) tooth reduced in female, absent in male...... Porterapis —.Glossa of the usual bilobed form (sometimes rather deeply so), each lobe less than one third length of CLASSIFICATION OF AMERICAN COLLETINAE prementum; apex of mandible pointed, usually with strong pollex tooth (which is sometimes double) .. . 4 4. Female with tibial scopal hairs sparse (as in Levoproctus), not hiding surface, which is therefore not greatly different from that of basitar- sus; body length 7.5-9 mm; jugal lobe of hind wing reaching or sur- passing cu-v, over two thirds as long as vannal lobe measured from wing DASE