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Seen oo tan nasa ec See DORN ere ee noe ae Seen erat aeRO Se oetnc ie soon ennk—orpencence area an Leeccloneepep epee aerator vere oereenr aremmpemre eros etemereneet esto aeaa Ta ate p ns Opa nS cEDRRSRSERSSRONS BRPREC EOE > NNN eet coat eran tT Ee eI ON ee ee eens a eee a wenanne aoa ret Saas = Eeesken ae ar f ==; oP = Po. ‘ F ff : bt ase t = es = rn a? tes ——_ os cn c= Si =a Se x ne " m, ; aie ipas My ib oh | A i ij : ul te ar] P y Tid I Fe, iN 5 im Hr a i i th ? 1 " ’ 1A 7 ‘i ra a | To ; ; : i - vi a Py, i Al { ) f i F hme j ’ a \ j | i ¥ ‘ } 5 ; - { ’ mn 4 ’ ; i « ‘ ¥ ' r Y 4 * 7 PR Ni 7 i — = Z a nee { : : ee i, eee ee — Po So SMITHSONIAN INSTITUTION UNITED STATES NATIONAL MUSEUM PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM VOLUME 108 NUMBERS 3391-3410 UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON : 1960 ADVERTISEMENT The scientific publications of the National Museum include two series, known, respectively, as Proceedings and Bulletin. The Proceedings, begun in 1878, are intended primarily as a medium for the publication of original papers, based on the collections of the National Museum, that set forth newly acquired facts in biology, anthropology, and geology, with descriptions of new forms and revisions of limited groups. Copies of each paper, in pamphlet form, are distributed as published to libraries and scientific organizations and to specialists and others interested in the different subjects. The dates at which these separate papers are published are recorded in the tables of contents of each of the volumes. The present volume is the hundred and eighth of this series. The Bulletin, the first of which was issued in 1875, consists of a series of separate publications comprising monographs of large zoological groups and other general systematic treatises (occasionally in several volumes), faunal works, reports of expeditions, catalogs ef type specimens, special collections, and other material of similar nature. The majority of the volumes are octavo in size, but a quarto size has been adopted in a few instances in which large plates were regarded as indispensable. In the Bulletin series appear volumes under the heading Contributions from the United States National Herbarium, in octavo form, published by the National Museum since 1902, which contain papers relating to the botanical collections of the Museum. Reminaton KeELioaa, Director, United States National Museum. AUG 29 1960 GBRAR* ~— CONTENTS Buaxe, Doris H. A review of some galerucine beetles with excised middle tibiae in the male.’ Six figures. No. 3395, published July 16, 1958 New genera: Lepionesiotes, Iceloceras. New species: Iceloceras maximum, I. flavipes, I. verbesinae, I. laliplagiatum, I. parviplagiatum, I. vittatum, I. biplagiatum, I. flavocyaneum, Luperosoma nigrum, Ectmesopus leonardorum, Platymorpha albiventris. New subspecies: Iceloceras parviplagiatum teapense. CarrweicHt, O. L. Scarab beetles of the genus Bothynus in the United States (Coleoptera : Scarabaeidae). Six fig- ures. No. 3409, published March 10, 1959 . New species: Bothynus selandert. Deienan, H.G. The systematic position of the bird genus Apalopieron. One plate. No. 3397, published June 16, 1958 .. ee ee eee a ee ee Dunxkiz, Davin H. Three North American Cretaceous fishes. Three plates. No. 3401, published October 21, Medes ee eae ieee erat BE Sn “tale TR To? oti tenn ee, eT ee a 8h. New genus: Paraciichthys. . Fietp, Wizu1am D. A redefinition of the butterfly genera Tatochila, Phulia, Piercolias, and Baltia, with descriptions of related genera and subgenera. Forty figures. No. 3396, published June 9, 1958 . New genus: Theochila. New subgenus: Infraphulia, Pierphulia. Frick, Kmnneru EK. Synopsis of the species of agromyzid leaf miners described from North America.” One hundred seventy figures. No. 3407, published March 5, 1959 . FrRiepMANN, Hersert. Advances in our knowledge of the honey-guides. No. 3404, published October 21, 1958 . Pages 59-101 515-541 133-136 269-277 103-131 047-465 308-320 1 Errata: On page 68, figure 6, h, for ‘“Ecfmesopus malachoides” read “ Ectmesopus malachioides’’; on page 97 (key), for ‘“Ectmesopus malachiodes” read “‘ Ectmesopus malachioides.”’ 2 Erratum: On page 397, line 1, for ‘“‘Phyfobia (Xenophytomuza)} illinoensis (Malloch)”’ read ‘‘Cercdontha (Xenophytomy za) illinoensis (Malloch).” m1 IV CONTENTS Herrera, José, and Fieytp, Witu1am D. A revision of the butterfly genera Theochila and Tatochila (Lepidoptera : Pieridae). Ninety-three figures. No. 3408, published July 24, 1959 FOE TS, Bi al og beg New subspecies: Tatochila theodice staudingeri, T. microdice fue- guensis. Horrman, Ricuarp L. Revision of the milliped genus Pachydesmus (Polydesmida : Xystodesmidae). ‘Twelve figures. No. 3399, published August 20, 1958 . New subspecies: Pachydesmus crassicutis hubrichti, P. c. adsini- colus. Hyman, Lispre H. A further study of Micronesian poly- clad flatworms. Seventeen figures. No. 3410, published March 6, 1959 . New genera: Asolenia, Nymphozoon. New species: Plehnia tropica, Stylochoplana minuta, Notoplana micronesiana, Aquaplana pacifica, Paraplanocera fritillata, Asolenia deilogyna, Pseudoceros habroptilus, P. caeruleocinctus, P. ferrugineus, P. ater, P. fulvogriseus, P. fuscogriseus, P. tri- striatus, Nymphozoon bayeri, Acanthozoon nigropapillosus, A. albopapillosus, Acerotisa rugosa, Prosthiostomum exiguum, P. grvseum., New name: Pseudoceros perviolaceus. JOHNSON, Puytuis T. Type specimens of lice (order Ano- plura) in the United States National Museum. No. 3393, published May 2%, 1958). sat) 2... pc sabe einer meres Kanazawa, Ropert H. A revision of the eels of the genus Conger with descriptions of four new species. Four plates, seven figures. No. 3400, published October 6, 1958 . New species: Conger jordani, C. oligoporus, C. macrocephalus, C. philippinus. New name: Conger triporiceps. Lyncu, JAMES E. Branchinecta cornigera, a new species of anostracan phyllopod from the State of Washington. Sixteen figures. No. 3392, published April 15, 1958 . New species: Branchinecta cornigera. Masuin, Paut T.; BerpLheMANn, Ricoarp G.; and Lowe, Cuarues H., Jr. The status of the lizard Cnemidophorus perplexus Baird and Girard (Telidae). No. 3406, pub- lished December 31, 1958 . HOY Wit HK Pages 467-514 181-218 543-597 39-49 219-267 25-37 331-345 CONTENTS Orvic, Tor. Pycnaspis splendens, new genus, new species, a new ostracoderm from the Upper Ordovician of North America. Three plates, five figures. No. 3391, pub- lished April 10, 1958 . New genus: Pycnaspis. New species: Pycnaspis splendens. Parkes, Kennetao C. Taxonomy and nomenclature of three species of Lonchura (Aves: Hstrildinae). One figure. No. 3402, published October 21, 1958 . New subspecies: Lonchura leucogastra manueli, L. l. smythiesi, L. punctulata yunnanensis, L. malacca deignant. RemnHArpb, Epwarp G. Rhizocephala of the family Pelto- gastridae parasitic on West Indian species of Galatheidae. One plate, four figures. No. 3403, published November 20,1958 . New genera: Cyphosaccus, Boschmaia. New species: Cyphosaccus chacei, C. cornutus, Boschmaia muni- dicola. Scuuttz, Leonarp P. Three new serranid fishes, genus Pikea, from the western Atlantic. Two figures. No. 3405, published November 17, 1958 . New species: Pikea cubensis, P. mexicanus. New subspecies: Pikea mexicanus mexicanus, P. m. atlaniicus. Smit, F.G. A. M. A new North American bird-flea. Ten figures. No. 3394, published May 15, 1958 . New subspecies: Ceratophyllus affinis neglecius. Witson, Miniprep Srratron. A review of the copepod genus Ridgewayia (Calanoida) with descriptions of new species from the Dry Tortugas, Florida. Thirty-seven figures. No. 3398, published August 11, 1958 . New family: Ridgewayiidae. New species: Ridgewayia gracilis, R. shoemakeri. 279-293 295-307 51-57 137-179 PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1958 No. 3391 oo ————————————————————————————ooooooeeoOooeoeyoyoeo®oq0qQqqqqqSSSSSSSS ao PYCNASPIS SPLENDENS, NEW GENUS, NEW SPECIES, A NEW OSTRACODERM FROM THE UPPER ORDOVICIAN OF NORTH AMERICA By Tor @rvic! Introduction In this paper a short description is given of a new ostracoderm, Pycnaspis splendens, new genus, new species, from the Upper Ordo- vician of the eastern slope of the Bighorn Mountains in north-central Wyoming. This new form is related to Astraspis desiderata from the Harding Sandstone, but it differs from the latter in the shape and microstructure of the tubercles of the exoskeleton. At the type locality it is associated with Astraspis? sp. indet. and Eriptychiida gen. and sp. indet. Exoskeletal plates referable to Pycnaspis, new genus, but at present indeterminable as to species are represented in the Winnipeg formation of the Williston Basin in Montana and in the Whitewood formation of the Black Hills in South Dakota, but such plates are missing entirely in the Harding Sandstone of Colorado. Ordovician vertebrates from North America were first recorded by Walcott who, in 1892, described small exoskeletal plates of two forms, Astraspis desiderata Walcott and Eriptychius americanus Walcott, from the Harding Sandstone outcrops near Canyon City, Colo. These two forms Walcott referred tentatively to the Asterolepidae and Holoptychiidae, respectively, but they are now generally recognized as true representatives of the Heterostraci. Walcott also figured a third fossil from the Harding Sandstone, Dictyorhabdus priscus Wal- 1 Curator, Department of Paleozoology, Swedish Museum of Natural History, Stockholm. 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 cott, which he regarded as the ‘calcified chordal sheath” of a fish allied to the bradyodonts, but which, as later researches have clearly shown (Dean, 1906, pp. 132-135; Bryant, 1936, p. 410; Flower, 1952, pp. 516-517), cannot possibly be of vertebrate origin. The Harding Sandstone vertebrate remains have been dealt with by Jaekel (in an appendix to Walcott’s paper of 1892) and subsequently by, among others, Vaillant (1902), Eastman (1907, p. 33; 1917, pp. 236-239, pl. 12, figs. 5, 6), Woodward (1921, p. 179), Stensid (1927, pp. 314-315, 333), Stetson (1931, p. 153), Bryant (1936), Berg (1940, pp. 107-108, 360-361), Kvam (1946, pp. 19-20, fig. 1), Gross (1950, p. 73; 1954, p. 80, pl. 3, fig. 2, pl. 5, figs. 3, 4, 7), Orvig (1951, pp. 381-382, 387, 393, 415, 433, fig. 22 8, pl. 3, figs. 3, 4), Gregory (1951, vol. 1, pp. 102- 104, vol. 2, figs. 6.2 (A), 6.3, 6.4), Denison (1956), J. D. Robertson (1957), and James (1957, p. 9, pl. 1). A description of Astraspis desiderata, based on the only fairly complete carapace of this form known so far,’ will be given in a forthcoming paper by the present writer (Orvig, in MS., b). Various notes on the invertebrate fauna and/or the stratigraphy of the Harding Sandstone have been given by Walcott (1892), Darton (1906b), S. R. Kirk (1929), Miller (1930), Stauffer (1930, p. 83), E. Kirk (1930), Branson and Mehl (1933); Behre and Johnson (1933), Ulrich (1938), Johnson (1944, pp. 320-322), Flower (1952), Frederickson and Pollack (1952), Twenhofel et al. (1954), Sweet (1954; 1955), and Denison (1956, pp. 368-369). This formation is now generally believed to be of Middle Ordovician (Trenton) age. Vertebrates to some extent resembling those of the Harding Sand- stone have long been known to occur in certain sandstone beds beneath the Bighorn Dolomite of north-central Wyoming and in the lower sandstone and siltstone members of the Whitewood formation of the Black Hills region in South Dakota, but none of this material has yet been properly described (see Darton, 1906a, p. 29; 1906b, pp. 550-551, fig. 3; 1909; Cockerell, 1913, p. 247; E. Kirk, 1930; Miller, 1930, p. 206; Romer and Grove, 1935, pp. 810-811; Furnish, Barragy, and Miller, 1936, pp. 1835-1338, pl. 2, figs. 14-16; Amsden and Miller, 1942, p. 304; Miller, Cullison, and Youngquist, 1947, p. 31; McCoy, 1952; Denison, 1956, p. 367). Dermal elements of Thelodontida and Osteostraci (Ateleaspis?) have been recorded from strata of a presumed lower Middle Ordovician age in the Bighorn Mountains of Wyoming (Tieje, 1924), but everything seems to indicate that those fossils are incorrectly determined. A new occurrence of Ordovician vertebrates has recently been detected farther north, in the sandstone and shale of the Winnipeg formation of the Williston Basin. This latter fauna, "a This specimen (USNM 8121), mentioned already by Walcott (1892, footnote on p. 167) and later de- scribed by Eastman (1917, pp. 238-239, pl. 12, figs. 5, 6) and by Bryant (1936, pp. 416-417, pl. 1), is of par- ticular interest in that it shows part of the lateral line canal system on the anterior part of the carapace. NEW OSTRACODERM—@RVIG 3 which is contained in drill cores from various places in eastern Mon- tana, has proved to be of considerable interest, and together with other undescribed vertebrate material from the Ordovician of Wyoming and South Dakota it will be dealt with by the writer on a future occasion. A ete Ny ¥). a SAE aif CLE AA, SNA Granite Carboniferous, Permo- Triassic RSGQQ - 7 te Pe Jurassic ormation ; Cretaceous, BEB Bighorn aad’ younger formation e--- faults Ficure 1.—Map showing the outcrops of Ordovician rocks on the eastern slope of the Bighorn Mountains in the area west of Lake De Smet and Buffalo, Johnson County, Wyo. Redrawn and to some extent simplified from the geological maps of the Bighorn Mountain Region published by Darton (1906a, pl. 47; 1906b, pli: 73): The new astraspid, Pycnaspis splendens, new genus, new species, described in this paper comes from the Upper Ordovician deposits of the Bighorn Mountains of Wyoming. The histology of the exoskele- ton in this form will be dealt with more in detail in another connection (Orvig, in MS., a). 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Fossils presumed to be remains of early vertebrates have been recorded on two occasions from North American deposits that are older than the Harding Sandstone of Colorado and the other verte- brate-bearing beds of Wyoming and South Dakota referred to above, viz.: (a) small, tuberculated plates from the Middle Cambrian St. Albans Shale of Vermont, described by Bryant (1927) and Howell (1937, pp. 1200-1202, p. 2, figs. 7, 8) under the name of Hoichthys howelli Bryant, and (b) Archeognathus primus Cullison, a peculiar V-shaped element with tooth-like cusps along one margin, which comes from the Lower Ordovician Dutchtown formation of Missouri (Culli- son, 1938, p. 227, pl. 29, fig. 16a,b; Miller, Cullison, and Youngquist, 1947, pl.1). It is by no means certain, however, if any of these fossils really are of vertebrate origin (Mrvig, 1951, footnote 1, on p. 381). The Eoichthys plates, at any rate, which I had the opportunity to study in 1953 during my visit to the department of geology of Prince- ton University, Princeton, N. J., cannot possibly have belonged to any true chordate. According to Rhodes and Wingard (1957, p. 453), Archeognathus primus is, in several ways, reminiscent of the Neurodontiformes (the “fibrous conodonts’ of previous writers). Whatever its affinities are, this fossil cannot possibly be a detached cornu of an early representative of the Osteostraci (as tentatively suggested by G. M. Robertson, 1954, p. 733). Outside North America, pre-Silurian vertebrates are only known with certainty from the lower Ordovician Glauconitic Sand of Es- thonia. From these beds Rohon (1889) has described small tooth-like fossiles belonging to two genera, Palaeodus and Archodus, which, by Russian paleoichthyologists, are nowadays referred to the Thelo- dontida (Obrutchev, 1948, p. 285; Bystrow, 1955, p. 473; Berg, 1955, p. 33; according to Denison, 1956, p. 367, they are indeterminable as to group). As will be shown in another connection (@rvig, in MS., a), however, it is fully evident that Palaeodus represents broken-off tubercles from the exoskeleton of a form belonging to the Astraspida. That the Astraspida are met with not only in various places in the Cordilleran Region of the United States (Colorado, Wyoming, South Dakota, and Montana) but also in Northern Europe is a circumstance of some interest as it indicates that this ancient group of the Hetero- straci may have had a worldwide distribution in Ordovician times. MeETHODS AND MATERIALS: All of the specimens figured in this paper except two (figs. 1-3 of pl. 2 and figs. 1-4 of pl. 3) are contained in beds of Upper Ordovician age from the Rock Creek section on the eastern slope of the Bighorn Mountains. As far as can be judged from the limited material at my disposal, these beds are similar to the other vertebrate-bearing rocks from the Ordovician of North America in that they contain a thanatocoenotic assemblage of detached exo- NEW OSTRACODERM—9@RVIG 5 skeletal plates and scales, more or less fragmentary; however, no single specimen shows any of these elements in their natural associa- tion. The average size of the plates and scales is somewhat larger than that in the Harding Sandstone of Colorado and the sandstone and shale of the Winnipeg formation of the Williston Basin in Mon- tana, and very small elements are missing altogether, a circumstance which probably indicates that the material has been sorted mechani- cally (by currents, etc.) prior to fossilization. ‘The plates and scales have been abraded by rolling to some extent, but cannot, nevertheless, have been transported very far from their place of origin. The state of preservation of the material is unusually favorable both for macroscopial and microscopical investigation. The plates and scales are easily freed of adhering sand-grains, e. g., by means of a sharp needle-scalpel, and the thin coat of reddish iron oxides which frequently obscures the fine details of their ornamentation can be removed, either with sodium hexametaphosphate or by boiling, for a very short time only, in a 10 percent solution of potassium hydrox- ide. The histological structure of the hard tissues is excellently dis- played in those specimens which are light yellowish in color, but it has become obscured to some extent in those which are impregnated with iron oxides. For the preparation of exactly orientated thin sec- tions, it is advantageous to embed the specimens in thin rods of acrylatic plastic by means of a bakelite press, according to the method recently described by the writer (Orvig, 1957, p. 370). The exoskeletal plates figured on plate 2, figures 1-3, and plate 3, figures 1-4, are from the gray, fissile shale of the Winnipeg formation of the Williston Basin (see, e. g., Ehlers, 1943, p. 1620; McCabe, 1954, p. 1998, fig. 10) contained in the drill core material from eastern Montana at my disposal. In this shale, which also exhibits phos- phatic nodules, various invertebrate fossils such as lingulid shells, etc., and conodonts, the vertebrate remains are dark brownish or black. Their histological structure is frequently well preserved but is, in places, obscured by post-mortem changes caused by penetrative Algae or Fungi (see, e. g., Peyer, 1945; Bystrow, 1956). It is worth mentioning here, finally, that the plates and scales of the Harding Sandstone—whose ornamentation is, In many cases, difficult to bring out satisfactorily by the ordinary methods of me- chanical preparation—can be studied to great advantage on latex micro-moulds made from the impressions in the rock after the re- moval of the hard tissues with hydrochloric acid. A perfect repro- duction of even very minute details of the ornamentation of those elements can be obtained by the method (described by Baird, 1955, p. 202; see also Gill, Caster, and Boswell, 1956, p. 198) of treating the etched surface of the sandstone with liquid detergent (e. g., Johnson 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Wetting Agent 326) immediately before the first thin latex coat is applied. All figured specimens belong to the U. S. National Museum. The technical work in connection with this paper has been carried out by members of the staff of the Swedish Museum of Natural History in Stockholm. ACKNOWLEDGMENTS: For their generosity in placing Ordovician vertebrate material from a series of localities in North America at my disposal, I wish to express my indebtedness to Drs. P. E. Cloud, Jr. and R. J. Ross, Jr., of the U. S. Geological Survey in Washington, D. C., and Denver, Colo., respectively, and to Drs. A. K. Miller and W. M. Furnish of the University of Iowa. My thanks also are due to Dr. D. H. Dunkle of the U. S. National Museum for his kind assistance in procuring the material and to Dr. Jean M. Berdan of the U. S. Geological Survey, Washington, D. C., for her information concerning the stratigraphy at the type locality of Pycnaspis splen- dens, new genus, new species. Subclass PTERASPIDOMORPHI Superorder HE'TEROSTRACI Order ASTRASPIDA 3 Pycnaspis, new genus Diacnosis: A genus of the Astraspida of large size. Plates of the exoskeleton thick. Ornamentation in early stages of growth con- sisting of a shagreen of small tubercles exhibiting deep, radiating grooves on their crowns; in succeeding stages of growth consisting of stout, mushroom-like tubercles, smooth on top and ranging in outline from round to elongated or somewhat kidney-shaped. All tubercles consisting of an external layer of dentinous tissue and a basal portion of bone (aspidin) surrounding a “pulp” cavity. In tubercles belong- ing to early stages of growth the dentinous tissue is penetrated through- out its height by very fine canals originally housing cell-processes, and the aspidin is devoid of vascular canals issuing from the upper part of the ‘‘pulp” cavity. In tubercles of succeeding stages of growth the dentinous tissue in its basal part contains short, stemlike dentinal tubes which, in a superficial direction, rapidly subdivide into very fine ramifications, and the basal, bony portion is pierced by narrow, straight vascular canals radiating upwards and outwards from the roof of the “pulp” cavity and extending to about the boundary between the aspidin and the dentinous tissue. 3 New rank, from Astraspidae Eastman, 1917. Synonym: Astraspiformes Berg, 1940. NEW OSTRACODERM—9YRVIG 7 Type species: Pycnaspis splendens, new species. GEOLOGICAL OCCURRENCE: Upper Ordovician: Upper part of the Bighorn formation, eastern slope of the Bighorn Mountains, Wyoming. Middle Ordovician: Whitewood formation, “Icebox’”’ shale, Black Hills Region of South Dakota; Winnipeg formation of Williston Basin, Montana. Remarks: The order Astraspida, comprising the genera Palacodus Rohon, Astraspzs Walcott, and Pycnaspis, new genus, has been defined in another connection (Orvig, in MS., b). The diagnosis of Pycnaspis given here lists mainly such characters by which this genus is dis- tinguishable from Asiraspis. The generic name is derived from the Greek zuxvos, thick, closely set, referring to the shape and distribution of the tubercles of the exoskeleton; and domes, shield. Pycnaspis splendens, new species Dracnosis: Same as for genus (single species). Hototrre: USNM 21333; detached exoskeletal plate, showing typical ornamentation (pl. 1, figs. 1, 2). LOcALITY AND GEOLOGICAL HORIZON: The holotype and the other material of Pycnaspis splendens, new genus, new species, dealt with here were collected in 1951 by Dr. Jean M. Berdan, of the U. S. Geological Survey, at the South Fork of Rock Creek, center, N4NE% SWYSWY, sec. 25, T. 52 N., R. 84 W., Johnson County, Wyo. (locality number USGS 1362-CO; see fig. 1 and Darton, 1906a, p. 28). According to personal communication by Dr. Berdan, there follows, at this locality, on top of the massive Bighorn dolomite: (a) red shales (“Red Bighorn’’) containing a rich fauna of Upper Ordovician in- vertebrate fossils which seems to correlate with the Maquoketa of Iowa, including, e. g., Lepidocyclus perlamellosus (Whitfield), ZL. cf. rectangularis Wang, Sceptropora facula Ulrich, and Streptelasma trilo- batum Whiteaves (cf. also the fauna list in Darton, 1906a, p. 28); (b) red and green mottled calcareous siltstones and sandstones; and (c) an upper sandy layer with numerous Ostracoderm remains. The latter is directly overlaid by the Lower Carboniferous (Mis- sissippian) Madison formation. By weathering, the fossiliferous shales above the Bighorn Dolomite disintegrate into soil, and in this soil the vertebrate material was found. Since, however, the plates and scales always are associated with remnants of a coarse-grained sand- stone, there is every reason to believe that they did not originally belong to the red shales but are, in fact, derived from the beds above, in all probability from the topmost sandy layer underneath the Madi- son Limestone or, in other words, from the very youngest part of the Bighorn formation exposed in the Rock Creek section. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 There has been some difference of opinion in recent years concerning the correlation of the Bighorn formation with Ordovician strata else- where in the United States (see, e. g., Twenhofel et al., 1954). Ac- cording to what has been said above, however, there can be little doubt with regard to the dating of the red fossiliferous shale which, in the Rock Creek section, follows on top of the massive dolomite mem- bers of this formation. Despite the fact that their exact position in the section cannot, at the moment, be made out with certainty, it is clear that the plates and scales of Pycnaspis splendens, new genus, new species, are no older than these red shales and that, therefore, they are of an Upper Ordovician age. It is true that there are also outcrops of red vertebrate-bearing sandstone beds underneath the Bighorn Dolomite in the Rock Creek section similar to those further south in the same area (see E. Kirk, 1930, p. 462; Amsden and Miller, 1942, fig. 1) but, according to Dr. Berdan (in litt.), there is no evidence at this locality of tectonic movements of such an order as to bring these sandstone beds in contact with the upper part of the Bighorn formation. Remarks: In the other material at my disposal, I have been able to identify some plates of Pycnaspis sp. indet., to some extent rem- iniscent of those of Pycnaspis splendens, new genus, new species, both in the sandstone and shale of the Winnipeg formation of the Williston Basin in Montana (pl. 2, figs. 1-3) and in a phosphatic conglomerate from the lower portion of the ‘‘Icebox’”’ shale member of the White- wood formation of the Black Hills region in South Dakota (White- wood Creek section, near Deadwood). On the other hand, Pycnaspis plates and scales are missing altogether in all the numerous samples of the Harding Sandstone of Colorado which I have had the oppor- tunity of examining in American and European museums, a circum- stance which may be of some importance from the point of view of stratigraphy. The specific name refers to the shining crown of the tubercles of the exoskeleton. Description: In the material of Pyenaspis splendens, new genus, new species, one may distinguish thick exoskeletal plates and scales of mainly three different categories: (a) polygonal plates from the carapace; (b) ridge-scales, for the most part, at least, from the cara- pace; and (c) other scales probably belonging to the trunk behind the carapace. The polygonal plates and ridge-scales are in several respects rather reminiscent of those in the carapace of Astraspis desiderata (Orvig, in MS., b), and everything goes to prove that they originally occupied much the same position in relation to each other as in the latter form. Thus, the ridge-scales very likely are detached com- ponents of a system of longitudinal crests extending, like in Astras- NEW OSTRACODERM—9ORVIG 9 pis, over the whole post-orbitopineal portion of the carapace (Eastman, 1917, p. 239, pl. 12, figs. 5, 6; Bryant, 1936, pp. 416-417, pl. 1; Orvig, in MS., b), and possibly continuing backwards on to the trunk behind the carapace, and the polygonal plates probably formed a fairly regu- lar pattern in the interspaces between the crests and anteriorly to Ficure 2.—Pycnaspis splendens, new genus, new species. Ornamentation of the exoskeleton: a, shagreen of small tubercles belonging to an early stage of growth, in super- ficial view (mainly after USNM 21334, X30); d, large tubercles belonging to a late stage of growth, in oblique lateral view (USNM 21335, X30). them. On the other hand, there are no plates and scales in the material similar to those which, in Astraspis, exhibit shallow lateral line grooves on their external face (Orvig, in MS., b; see also Walcott, 1892, pl. 4, fig. 1). If not purely coincidental, this circumstance may 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 either mean that the lateral line system was rather incompletely developed as compared with that in Astraspis, or, perhaps still more probable, that it was situated to a large extent in canals in the interior of the exoskeletal elements so that it was invisible in external view (cf. in the Eriptychiida, p. 18). can‘pca peo fi ib. fi Ficure 3.—Pycnaspis splendens, new genus, new species. Vertical section of a tubercle of the exoskeleton (section No. S 1477, from @rvig, in MS., a; 150). Explanation of sym- bols: b.as, aspidin; bzul, inner part of dentinous tissue; can. pea, vascular canals issuing from the upper part of the “pulp” cavity; d?’”’, fine ramifications of dentinal tubes; ezul, outer part of dentinous tissue; fi, fibers of Sharpey; /am, concentric lamellation of aspidin; pea, “pulp” cavity. The polygonal plates of the carapace, which are most commonly met with in the material, are generally elongated rhomboid or more or less distinctly pentagonal in shape. They are often somewhat convex externally, but as far as I can find they are not raised upwards into a central cone-like prominence as frequently is the case in Astraspis. In some places they overlap each other to some extent, but as a rule they show no imbricating arrangement, being situated beside each other in much the same manner as the exoskeletal units, e. g., in the carapace of Tesseraspis (Wills, 1935), in the cephalic shield of certain late Osteostraci (Orvig, in MS., a), and in the armor of various Ostra- ciontidae. The ridge-scales, which are symmetrical, vary to some extent where their shape is concerned. Thus, some of them—which, judgmg by the condition in Astraspis desiderata (Orvig, in MS., b), quite con- ceivably occupied a position in the anterior half, or thereabouts, of the longitudinal crests of the carapace—are longer than broad and NEW OSTRACODERM—9@RVIG tl] have a truncated, or slightly rounded, posterior margin (pl. 1, figs. 3-6). Others, which probably were situated further backwards in the crests, are equally as long as broad, and their posterior margin is more or less obtusely V-shaped. In all the ridge-scales one may, just as in those of Astraspis desiderata, distinguish the following faces: (a) a convex external face, consisting of an anterior portion which slopes forwards and downwards and which is overlapped by the ridge-scale in front (od,, pl. 1, figs. 3, 4), and a posterior, exposed portion orna- mented with tubercles throughout its extent (pl. 1, figs. 3, 4); (b) a concave basal face having the appearance of a broad, longitudinal groove (Of, pl. 1, figs. 5, 6); (c) a concave posterior face overlapping the anterior portion of the external face of the ridge-scale following next behind (0g,, pl. 1, figs. 5, 6); and, finally, (d) a paired lateral face, a little concave and sometimes fairly low, overlapping the adjoining polygonal plates on each side (0q,, pl. 1, figs. 3, 5). As fully evidenced by their structure, the ridge-scales were firmly attached to each other. In the material at my disposal, however, there is no indication what- soever that they tended to fuse with their fellows in the individual crests in the same manner as in Astraspis desiderata (Orvig, in MS., b). The scales presumably belonging to the trunk behind the carapace are far less common in the material than the polygonal plates of the carapace and the ridge-scales. They are more or less rhomboid in shape, frequently broader than long, and deeply imbricating. Their external face is made up of a large anterior overlapped portion and a fairly narrow posterior exposed portion ornamented with tubercles. Scales of this particular kind are, as far as I can find, not met with in the Astraspis material from the Harding Sandstone or from other formations. The ornamentation of the plates and scales is subject to considerable variation. Some of the elements that are clearly preserved in early stages of growth show an immature type of ornamentation, made up of a shagreen of fairly small and low tubercles (fig. 2,a, ef. t,, pl. 1, figs. 1, 2, 4), which in superficial view are rather similar to those on the exoskeleton of various Psammosteida and certain Osteostraci as well (see Obrutchev, 1956, pl. 1, fig. 4). These tubercles are highly reminiscent of those in Astraspis (fig. 4; Walcott, 1892, pl. 3, figs. 13, 14; Bryant, 1936, p. 418, pl. 2, fig. 2), and are, in fact, only distinguish- able from the latter by the circumstance that, in the individual plates and scales, they are remarkably uniform in size and, as a rule, placed close beside each other with about equal interspaces. They consist of a bulbous crown, exhibiting deep grooves radiating in a basal direc- tion from the apex, and a fairly low neck-portion which is somewhat constricted below the crown but otherwise not sharply defined from the latter. LD PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 In the great majority of the plates and scales, on the other hand, the early shagreen of starlike tubercles has, during subsequent stages of growth, been supplanted completely by another, mature type of ornamentation consisting only of thick, mushroom-like tubercles (fig. 2,5; ts, pl. 1, figs. 1, 2, 4), frequently of large size, which are in several respects different from those on the exoskeleton of Astraspis. In top view the crown of these tubercles is rounded, elongated, or even some- what kidney-shaped; it is always smooth and shining superficially, and at its basal circumference it is generally more or less crenulated. The neck-portion of the tubercles, which is always well developed, is a little constricted immediately below the crown, and is separated from the latter by a distinct nick. Contrary to what might be ex- pected in view of the state of preservation of the material, the smooth- ness of the crown is certainly a primary condition. As in Astraspis (Bryant, 1936, p. 418, pl. 2, fig. 1), the tubercles were in certain cases subject to some post-mortem abrasion, but, as far as I can find, this cannot possibly account for the fact that in tubercles of different height, situated close together, and sometimes even partly superim- posed on each other, the crown invariably shows the same smoothly rounded contour in vertical section. In the material of Pycnaspis splendens, new genus, new species, at my disposal, at a rough estimate only 0.4 percent of the specimens (6 out of a total 1,500, or more) display both the immature and mature types of ornamentation simultaneously (e. g., the holotype, pl. 1, figs. 1, 2). If such specimens were lacking altogether—which might easily have happened in a less comprehensive material—one should no doubt refer the plates and scales, where the ornamentation is of the immature type (fig. 2,¢), to another species and (in view of the microstructure of the tubercles, see below) probably even to another genus than those where it is of the mature kind only (fig. 2,b). It is true that, as far as one can tell, a correspondingly marked and abrupt change in the ornamentation of the exoskeleton with advancing age is not met with in any other representative of the Heterostraci and rarely, if ever, in placoderms and early teleostomians either (cf., e. g., in Bothriolepis, Stensid, 1948, pp. 169-171, 376-386). But the condition of Pycnaspis splendens, new genus, new species, referred to above nevertheless shows that in dealing with early Paleo- zoic bone bed material from the taxonomic point of view one has, at least in certain cases, to allow for a considerable margin of error. It is worthy of mention, finally, that the ornamentation of the plates and scales was subject to variation not only in different stages of growth but even to some extent in different parts of the body. Thus in the polygonal plates, which presumably lay mainly on the anterior portion of the carapace, all tubercles, large and small alike, rise straight NEW OSTRACODERM—9@RVIG 13 upwards and are blunt on top (fig. 2,6, pl. 1, fig. 2). On the other hand, in the ridge-scales of the longitudinal crests and in the polygonal plates which probably occupied a position in the interspaces between these crests on the posterior portion of the carapace, the tubercles are inclined backwards (pl. 1, figs. 3, 4) and are frequently tapering Ficure 4.—Astraspis? sp. Large exoskeletal plate from the Pycnaspis-bearing beds of the Rock Creek section, in oblique lateral view. (USNM 21338; X20.) to a point posteriorly. In Astraspis desiderata from the Harding Sandstone there is little or no indication of a smilar variation in the shape of the tubercles from one part of the carapace to another (Orvig, in MS., b). The plates and scales of the exoskeleton are all remarkably stout, with their average thickness being, as in Astraspis and Eriptychius, probably greater in relation to the size of the animals as a whole than in any of the geologically younger members of the Heterostraci. They are built up of three layers—a superficial tubercle-layer; a middle, vascular layer; and a basal layer. In their general propor- tions these layers correspond closely to the similar layers in the exo- skeleton of Astraspis (Bryant, 1936, pp. 418-421, pl. 2, fig. 3). The tubercles are made up of two, sharply defined, histological constituents: (a) a thick upper layer of compact, semitransparent, sometimes faintly bluish white substance (ezul, bzul, fig. 3) which, as I shall have the opportunity of showing in another connection (Orvig, in 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 MS., a), is not enamel, nor in any way enamel-like, but clearly a special variety of dentinous tissue (mesodentine); and (b) a lower part of typical aspidin which forms thick walls around a basal, ‘pulp’ cavity (b.as, pea, fig. 3). In the tubercles of early stages of growth, the upper layer is, just as in Astraspis (Orvig, in MS., a), devoid of real dentinal tubes similar to those in dentine proper (e. g., in the tubercles and ridges of the exoskeleton in Hriptychius and post-Ordo- vician Heterostraci in general), but it is, however, pierced throughout its height by numerous very fine canals which correspond, in a broad sense, to delicate ramifications of such tubes. In the tubercles of consecutive stages of growth, on the other hand, this layer is to some extent suggestive of the dentinous tissues in the exoskeleton of various early Osteostraci (Orvig, in MS., a) in that it consists of a thick outer part with the same fine canals as those just referred to (ezul, fig. 3), and, in addition, of a thin, inner part containing a system of short, stemlike dentinal tubes or, in some cases, elongated cell-spaces, from which these fine canals issue in a superficial direction (bzul, dt’’, fig. 3). A similar inner part of the dentinous tissue, exhibiting den- tinal tubes or cell-spaces, is absent altogether in all tubercles of the exoskeleton in Astraspis. The aspidin in the lower part of the tubercles is perfectly similar histologically to that in the middle and basal layers of the plates and scales. As in Asiraspis, it is penetrated by fine fibers of Sharpey radiating from the “pulp” cavity (Bryant, 1936, p. 419, pl. 3; pl. 4, fig. 2; pl. 7, fig. 2; Gross, 1954, p. 80, pl. 8, fig. 2; pl. 5, figs. 3, 4). In the tubercles of early stages of growth this cavity is always undivided and completely separated from the upper dentinous tissue by the thick layer of aspidin which forms its roof (see Gross, 1954, pl. 5, figs. 3, 4). In the tubercles of subsequent stages of growth, however, a system of fairly thin, straight vascular canals issue in a radiating manner from the upper part of the “pulp” cavity, terminating close beneath and sometimes almost at the boundary between the aspidin and the outer dentinous tissue (can.pea, fig. 3). This system of vascular canals has no connection whatever with the short dentinal tubes or cell- spaces in the inner part of the dentinous tissue, A similar canal system is not developed in the tubercles in Astraspis, but may possibly have existed in the predecessors of that form (@rvig, in MS., a). The nature of the hard tissues of the tubercles will be dealt with elsewhere (Orvig, in MS., a) and will not be commented upon further here. Of interest to us in the present connection is above all the fact that, by their microstructure, the tubercles of early stages of growth invariably are sharply distinguishable from those of subse- quent stages of growth (clearly shown, e. g., by sections prepared of NEW OSTRACODERM—®@RVIG 15 specimens exhibiting simultaneously both the immature and mature types of ornamentation). This is surprising in view of the condition of the exoskeleton in a great many other lower vertebrates where the dentine tubercles belonging to consecutive generations certainly may show varying degrees of complexity and hence may differ to some extent from each other (see, e. g., Orvig, 1957, p. 388), but neverthe- less are always basically similar in their histological structure. The middle, vascular layer is thick, and its vascular canals are irregularly distributed except in some of the polygonal plates of the carapace where they may, occasionally, show a more or less distinct “transsutural” arrangement near the marginal faces (see Delpey, 1942, pp. 52-53; Orvig, in MS., a). Ontogenetically this layer arose as a system of thin, bony trabeculae separated by fairly wide vascular spaces, but later on it became rather compact (more so, in fact, than the corresponding layer of the exoskeleton in most post-Ordovician Heterostraci) by the deposition of thick-walled primary osteons on the margins of the trabecles. The osteons, which are perfectly similar to those in Astraspis (Bryant, 1936, pl. 6, pl. 7, fig. 1; Orvig, 1951, pl. 3, figs. 3, 4) and to those in various Psammosteids as well (Gross, 1954, pl. 5, fig. 11), are penetrated by fine fibers of Sharpey radiating from the vascular canals. The basal layer was somewhat thin in early ontogenetic stages, but subsequently it increased considerably in thickness by the apposition of new aspidin lamellae on its inner side. As far as one can judge, it is not continuous basally with layers of perichondral bone. As in Astraspis and Eriptychius (fig. 5, 6; see Bryant, 1936, pp. 421, 425, pl. 2, fig. 3; pl. 13, fig. 1), but contrary to the case in all Silurian and Devonian Heterostraci, it is penetrated throughout its extent by a great many vascular canals, ascending with about equal interspaces from the subcutaneous (subaponeurotic) vascular plexus beneath the exoskeleton. The presence of this system of ascending vascular canals, whose more or less funnel-like openings are spread all over the basal face of the plates and scales (pl. 1, fig. 5; Bryant, 1936, pl. 5, fig. 2), is no doubt a primitive condition (Mrvig, in MS., a). It is worthy of mention that the Pycnaspis plates from the Winnipeg formation of the Williston Basin in Montana and the Whitewood formation in South Dakota referred to above (p. 8) exhibit stout, mushroom-like tubercles which in shape and microstructure are similar to those belonging to mature stages of growth in Pycnaspis splendens, new genus, new species (pl. 2, figs. 1-3). These plates are undoubtedly referable to Pycnaspis, new genus, but for the time being, at least, they cannot be determined as to species. 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Other vertebrates The great majority (at least 75 percent) of the plates and scales in the material from the Rock Creek locality at my disposal belongs to Pycnaspis splendens, new genus, new species, but there are also several (about 20 percent) referable to the Eriptychiida, probably to a new genus of this group, and some few to Astraspis? sp. In addition, the fauna may also include exoskeletal elements of other forms whose systematic position is at present indeterminable. On the other hand, the Pycnaspis-bearing beds do not seem to contain pieces of bone tissue with cell-spaces or pieces of globular calcified cartilage, like those met with in the Harding Sandstone of Colorado (@rvig, 1951, p. 381, fig. 18a) and the Winnipeg formation of the Williston Basin in Montana. To what extent those beds are devoid of other micro- fossils, such as lingulid shells, conodonts, etc., cannot be decided with any certainty on the basis of the limited material dealt with here. The plates belonging to Astraspis? sp. (fig. 4) are of approximately the same order of size as those of Pycnaspis splendens, new genus, new species, with the mature type of ornamentation, but clearly distin- guishable from the latter by the shape and microstructure of their tubercles (fig. 2,6, pl. 1, figs. 1-4; cf. also pl. 2, figs. 1-3). These tubercles, which vary considerably in size, all consist of a pointed crown with deep, radiating grooves, and a conical neck-portion, which is frequently comparatively high. In their microstructure, they are similar to those in Astrapis desiderata (Bryant, 1936, pp. 418-420, pl..2, fig. 3;_pls..3, 4; pl..5,-figs..1,3;Orvmeg, 1951, fie<22R8: im. MS... a: Gross, 1954, pl. 3, fig. 2; pl. 5, figs. 3, 4) and those belonging to early stages of growth in Pycnaspis splendens, new genus, new species. Most of the material of Eriptychiida gen. and sp. indet. comprises fragments of plates, frequently of large size, whose ornamentation consists of fairly low and broad tubercles. These tubercules (fig. 5,a)— which as a rule are considerably coarser than those of the Eriptychius plates from the Harding Sandstone of Colorado (Walcott, 1892, pl. 4, figs. 8, 9; Bryant, 1936, pl. 8, fig. 1) and the Winnipeg formation of the Williston Basin in Montana (@rvig, in MS., a)—are fairly low, elongated pearshaped or irregular in outline, and smooth on top; they have no well-defined neckportion and they are frequently arranged in more or less distinct rows (pl. 2, figs. 4, 5, 7). As in EHriptychius (fig. 5,b,¢; Bryant, 1936, pl. 11) and in the post-Ordovician Heterostraci in general, these tubercles consist entirely of ordinary dentine. In their basal part, there is no large pulp cavity but a system of vascular canals from which the dentinal tubes issue in a superficial direction (for further details concerning the microstructure, see Mrvig, in MS., a). In only one of the plates is the position in the cuirass of the living NEW OSTRACODERM—9ORVIG Vi animal determinable with any degree of certainty. This specimen, which is particularly large and well preserved, in all probability repre- sents a branchio-cornual plate, unknown so far in any of the pre- Silurian Heterostraci (pl. 2, figs. 4-7). A second specimen, which may o.can.d ~ ~ can.asc dt mae / a b se YS GS | a f by v 7 ae ‘ 7 ‘ can. asc}, bal FIGURE 5.—a, Eriptychiida gen. and sp. indet.: Ornamentation of a plate-fragment from the Pycnaspis-bearing beds of the Rock Creek section, in oblique lateral view (mainly after USNM 21339, 36). b, c, Eriptychius sp., Winnipeg formation, Williston Basin, Mont., sections of scales (from Mrvig, in MS, a): 5, Vertical section showing the dentine ridges of the external face (cut somewhat obliquely to their axes of length), the middle, vascular layer, and the basal layer (section No. S 1438; X36); c, horizontal section of the dentine ridges and the adjoining part of the middle, vascular layer (section No. S 1454; X36). Explanation of symbols: bal, basal layer of the scales; can.ascy, ascending vascular canals piercing the basal layer; can.asc,, vascular canals in the upper part of the middle layer of the scales, ascending towards the dentine ridges; can.d, short vascular canals extending outwards in the basal part of the dentine ridges and opening to the ex- terior at the basal circumference of those ridges; can.J, longitudinal vascular canals in the basal part of the dentine ridges; dt, dentinal tubes; mil, middle, vascular layer of the scales; o.can.d, the openings of vascular canals at the basal circumference of the dentine ridges; sul, superficial layers of the scales. 438551—58——2 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 well be the broken-off anterior part of another branchio-cornual plate, shows a wide canal in its interior that may, perhaps, be a lateral line canal (lle?, pl. 2, fig. 8). If this interpretation is correct, there may be some reason to believe that in the Eriptychiida the lateral line system had no such superficial position as in Astraspis (Orvig, in MS., b), but that, like in several younger Heterostraci, it lay partly, or entirely, in the middle, vascular layer of the exoskeleton. Apart from the plates dealt with here, the material of Eriptychiida gen. and sp. indet. from the Pycnaspis-bearing beds also includes a certain amount of scales of much the same kind as those of Hripty- chius americanus from the Harding Sandstone of Colorado (Walcott, 1892, pl. 4, figs. 5, 6, 11; Bryant, 1936, pp. 423-424, pl. 8, fig. 2; pl. 9, fig. 2) and Eriptychius spp. from the Winnipeg formation of the Williston Basin in Montana and the Whitewood formation of South Dakota. These scales, which vary somewhat in size, consist of an anterior overlapped portion without ornamentation and a pos- terior exposed portion exhibiting a system of roughly parallel dentine ridges. As in the scales of Hriptychius from the Harding Sandstone and other formations, the dentine ridges are highly reminiscent of those of the exoskeleton in the Cyathaspida and Pteraspida (see Lindstrém, 1895, fig. 2; Kier and Heintz, 1935, pl. 37; Gross, 1935, fig. 5c; and others) in that they contain, in their basal part, a longi- tudinal vascular canal from which a series of short vascular canals issue with fairly regular interspaces to both sides (can.l, can.d, fig. 5,b,c; concerning this canal system see also Mrvig in MS., a). In the material from the Pycnaspis-bearing beds there are also a few scales of large size ornamented with fairly coarse tubercles (pl. 3, figs. 5, 6) that are to some extent suggestive of the median dorsal and median ventral ridge-scales of the trunk in several post-Ordovi- cian Heterostraci, e. g. the Cyathaspida, Pteraspida, and Psammos- teida (see Kier and Heintz, 1935, figs. 41-44, md and mv of fig. 50; White, 1935, figs. 1, 2, 63, 65; Obrutchev, 1945, figs. 2-5; and others). As far as one can tell at present, scales of this particular kind are not met with in the Eriptychius material from the Harding Sandstone of Colorado and the Winnipeg formation of the Williston Basin in Montana. Discussion There is surely reason to believe that, as they become better known, the Ordovician vertebrates of North America will prove highly use- ful for correlation purposes, but up to now there has been very little information in the literature as regards their stratigraphical distribu- tion. It may be of some interest in this respect that, according to NEW OSTRACODERM—RVIG 19 what has been said in this paper, one may now distinguish two differ- ent vertebrate faunas in the Ordovician of the Cordilleran Region, one in the Harding Sandstone of Colorado and another in the Pyc- naspis-bearing beds of the Bighorn Mountains of Wyoming. In addition, there is probably also a third vertebrate fauna—intermedi- ary, to some extent at least, between the other two—in the material at my disposal from the Winnipeg formation of the Williston Basin in Montana. The latter material has not yet been investigated in detail, but a preliminary examination has shown that it includes plates and scales of Eriptychius sp. (possibly a new species), Astraspis sp., Pycnaspis sp. indet. (see p. 8 and pl. 2, figs. 1-3) and surely also of other forms at present indeterminable as to genus. A par- ticularly interesting specimen from this material is a large, keel-like plate, ornamented on its external face with a system of broad, inoscu- lating dentine ridges (pl. 3, figs. 1-4); it may, perhaps, belong to a new representative of the Eriptychiida. Bibliography AMSDEN, T. W., and Miuter, A. K. 1942. Ordovician conodonts from the Bighorn Mountains of Wyoming. Journ. Paleont., vol. 16, No. 3, pp. 301-306, 2 figs., 1 pl. Barrp, DonaLp 1955. Latex micro-molding and latex-plaster molding mixture. Science, new ser., vol. 122, p. 202. BEuHRE, C. H., and JoHnson, J. H. 1933. Ordovician and Devonian fish horizons in Colorado. Amer. Journ. Sci., ser. 5, vol. 25 (225), pp. 477-486. Berg, L. S. 1940. Classification of fishes, both recent and fossil. Trudy, Zool. Inst. Acad. Sci. U. R. 8. 8., vol. 5, No. 2, pp. 85-517, 190 figs. (Russian and English.) 1955, Classification of fishes and fish-like animals, living and fossil. Ed. 2. Trudy, Zool. Inst. Acad. Sci. U. R. 8. 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A review of the habitat of the earliest vertebrates. Fieldiana, Geology, vol. 11, No. 8, pp. 359-457. EastMan, C. R. 1907. Devonic fishes of the New York formations. Mem. New York State Mus., No. 10, pp. 1-235, 34 figs., 15 pls. 1917. Fossil fishes in the collection of the U. 8. National Museum. Proc. U. 8. Nat Mus., vol. 52, No. 2177, pp. 235-304, 9 figs., 23 pls. EHLERS, ALLEN 1943. Williston Basin wildcat tests, Oliver County, North Dakota. Bull. Amer. Assoc. Petr. Geol., vol. 27, No. 12, pp. 1618-1622. FLower, R. H. 1952. Cephalopods from the Harding and Manitou formations of Colorado. Journ. Paleont., vol. 26, No. 8, pp. 505-518, 2 pls. FREDERICKSON, E. A., AND Poxuack, J. M. 1952. Two trilobite genera from the Harding formation (Ordovician) of Colorado. Journ. Paleont., vol. 26, No. 4, pp. 641-644, 6 figs. Furnisu, W. M.; Barragy, E. J.; anp MiIuuer, A. K. 1936. Ordovician fossils from upper part of type section of Deadwood formation, South Dakota. Bull. Amer. Assoc. Petr. Geol., vol. 20, No. 10, pp. 13829-1341, 1 fig., 2 pls. Gi1, E. D.; Castsr, K. E.; anp BosweE 1, P. C. R. 1956. Simple apparatus for vacuum injection of moulding latex. Australian Journ. Sci., vol. 18, No. 6, pp. 198-199. Grecory, W. K. 1951. Evolution emerging. A survey of changing patterns from primeval life to man. Vol. 1, 762 pp.; vol. 2, 1013 pp., illus. NEW OSTRACODERM—®@RVIG ai Gross, WALTER 1935. Histologische Studien am Aussenskelett fossiler Agnathen und Fische. Palaeontographica, vol. 83, Abt. a, 60 pp., 30 figs., 7 pls. 1950. Die paléontologische und stratigraphische Bedeutaug der Wirbeltier- faunen des Old Reds und der marinen altpaliozoischen Schichten. Abh. Deutsch. Akad. Wiss., Math.-Naturw. Kl. Jahrg. 1949, No. 1, 130 pp., 1 fig. 1954. Zur Conodonten-Frage. Senckenbergiana Lethaea, vol. 35, pp. 73-85, 2 figs., 5 pls. Howe .t, B. F. 1937. Cambrian Centropleura vermontensis fauna of northwestern Vermont. Bull. Geol. Soc. Amer., vol. 48, No. 8, pp. 1147-1210, 6 pls. JAMES, W. WARWICK 1957. A further study of dentine. Trans. Zool. Soc. London, vol. 29, pt. 1, 66 pp., 4 figs., 10 pls. JoHNsON, J. N. 1944. Paleozoic stratigraphy of the Sawatch Range, Colorado. Bull. Geol. Soc. Amer., vol. 55, No. 3, pp. 303-378, 7 figs., 11 pls. Kiar, JOHAN, AND Heintz, ANATOL 1935. The Downtonian and Devonian vertebrates of Spitsbergen. 5. Sub- order Cyathaspida. Skrift. Svalb. Ish., No. 40, 138 pp., 57 figs., 40 pls. Kirx, EpwIn 1930. The Harding Sandstone of Colorado. Amer. Journ. Sci., ser. 5, vol. 20 (220), No. 120, pp. 456-465. Kirg, S. R. 1929. Conodonts associated with the Ordovician fish fauna of Colorado— a preliminary note. Amer. Journ. Sci., ser. 5, vol. 18 (218), No. 108, pp. 4938-496, 1 pl. Kvam, THORVALD 1946. Comparative study of the ontogenetic and phylogenetic development of dental enamel. Norsk Tannlegeforenings Tidende, vol. 56, suppl., 198 pp., 147 figs. LInpstrROM, GuSTAF 1895. On remains of a Cyathaspis from the Silurian strata of Gotland. Svensk. Vet. Akad. Handl. Bih., vol. 2, afd. 4, No. 3, 15 pp., 2 figs., 2 pls. McCabe, W. S. 1954. Williston Basin Paleozoic unconformities. Bull. Amer. Assoc. Petr. Geol., vol. 38, No. 9, pp. 1997-2010, 10 figs. McCoy, M. R. 1952. Ordovician sediments in the Northern Black Fills. Billings Geol. Soc., Guidebook, 3d Ann. Field Conf., pp. 44-47. (Not seen, cited after Denison, 1956.) Mier, A. K. 1930. The age and correlation of the Bighorn formation of northwestern United States. Amer. Journ. Sci., ser. 5, vol. 20 (220), No. 117, pp. 195-2138. Miter, A. K.; Cunison, J. 8.; AnD YouNGQuIST, WALTER 1947. Lower Ordovician fish remains from Missouri. Amer. Journ. Sci., vol. 245, No. 1, pp. 31-34, 1 pl. 22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 OBRUTCHEV, D. V. 1945. The evolution of Agnatha. Zoologicheskii Zhurnal, vol. 24, pp. 257-272, 10 figs. (In Russian with English summary.) 1948. On the evolution of the exoskeleton in teleostomian fishes. Bull. Acad. Sci. U. R. §.8., Biol. Ser., vol. 3, pp. 281-286. (In Russian.) 1956. Cephalaspids from the lower Devonian of Tova. Comptes Rendus (Doklady) Acad. Sci. U. R. §. 8., vol. 106, No. 5, pp. 917-919, 1 fig., 1 pl. (In Russian.) Orvic, Tor 1951. Histologic studies of placoderms and fossil elasmobranchs. 1. The endoskeleton, with remarks on the hard tissues of lower vertebrates in general. Svensk. Vet. Akad., Ark. Zool., ser. 2, vol. 2, No. 2, pp. 321-456, 22 figs., 8 pls. 1957. Remarks on the vertebrate fauna of the lower Upper Devonian of Escuminac Bay, P. Q., Canada, with special reference to the porolepiform crossopterygians. Svensk. Vet. Akad., Ark. Zool., ser 2, vol. 10, No. 6; pp. 367-426, 18 figs. In Studies on North American ostracoderms. 1: On certain Osteost- MS.,a —_raci from the lower Upper Devonian of Escuminac Bay in Canada, with remarks on the phylogeny of the dentinous tissues in lower vertebrates. In Studies on North American ostracoderms. 2: On Astraspis desiderata MS.,b Walcott. PEYER, BERNHARD 1945. Uber Algen und Pilze in tierischen Hartsubstanzen. Zurich, Arch. Julius Klaus-Stiftung Erg. Bd., vol. 20, pp. 496-546, 48 figs. Ruoves, F. H. T., anp WinearD, PAUL 1957. Chemical composition, microstructure, and affinities of the Neuro- dontiformes. Journ. Paleont., vol. 31, No. 2, pp. 448-454, 2 figs. Rosertson, G. M. 1954. Some attempts at the phylogeny of the early vertebrates. Proc. Iowa Acad. Sci. (1953), vol. 60, pp. 725-737, 2 figs. RoBERTSON, J. D. 1957. The habitat of the early vertebrates. Quart. Rev. Biol., vol. 32, No. 2, pp. 156-183. Ronon, J. V. 1889. Uber unter-silurische Fische. Bull. Acad. Imp. Sci., St. Petersbourg, new ser., vol. 1 (83), No. 2, pp. 269-277, 1 pl. Romer, A. §S. 1955. Fish origins—fresh or salt water? Jn Papers in Marine Biology and Oceanography, dedicated to Henry Bryant Bigelow. . . 1955. Deep-Sea Research, suppl., vol. 3, pp. 261-280. Romer, A. S., anp Grove, B. H. 1935. Environment of the early vertebrates. Amer. Midl. Nat., vol. 16, No. 6, pp. 805-856. STAUFFER, C. R. 1930. The Devonian of California. Sci. Bull. Univ. California, Dept. Geol., vol. 19, No. 4, pp. 81-118, 5 pls. NEW OSTRACODERM—®@RVIG 92 Strensi6, HE. A. 1927. The Downtonian and Devonian vertebrates of Spitsbergen. 1: Family Cephalaspidae. Skrift. Svalb. Nordish., No. 12, 402 pp., 103 figs., 112 pls. 1948. On the Placodermi of the upper Devonian of East Greenland. 2: Antiarchi. Subfamily Bothriolepinae. . . . Palaeozoologica Groen- landica, vol. 2, 622 pp., 308 figs., 75 pls. Stetson, H. C. 1931. Studies on the morphology of the Heterestract. Journ. Geol., vol. 39, No. 2, pp. 141-154, 7 figs. Sweet, W. C. 1954. Harding and Fremont formations, Colorado. Bull. Amer. Assoc. Petr. Geol., vol. 38, pp. 284-305, 4 figs. 1955. Conodonts from the Harding formation (middle Ordovician) of Colorado. Journ. Paleont., vol. 29, No. 2, pp. 226-262, 17 figs., 3 pls. Tinyz, A. J. 1924. New earliest known vertebrates. (Abstract.) Bull. Geol. Soc. Amer., vol. 35, No. 1, p. 1620. TWENHOFEL, W. H., ET AL. 1954. Correlation of the Ordovician formations of North America, by the Ordovician Subcommittee of the Committee on Stratigraphy of the National Research Council. Bull. Geol. Soc. Amer., vol. 65, No. 3, pp. 247-298, 2 figs., 1 pl. Unricn, E. O. 1938. Age of the Harding Sandstone fish remains. (Abstract.) Proc. Geol. Soc. Amer. (1937), pp 289-290. ee L£OoN 1902. Sur la presénce du tissu osseux chez certains poissons des terrains palaeozoiques de Canyon City (Colorado). Comptes Rendus Acad. Sci. Paris, vol. 184, No. 22, pp. 1821-1322. Watcort, C. D. 1892. Notes on the discovery of a vertebrate fauna in Silurian (Ordovician) strata. Bull. Geol. Soc. Amer., vol. 3, pp. 153-172, pls. 3-5. Waite, E. I. 1935. The Ostracoderm Pteraspis Kner and the relationships of the agna- thous vertebrates. Phil. Trans. Roy. Soc. London, ser. B, vol. 225, No. 527, pp. 381-457, 97 figs., pls. 25-27. Wits, L. J. 1935. Rare and new ostracoderm fishes from the Downtonian of Shropshire. Trans. Roy. Soc. Edinburgh, vol. 58, pt. 2, No. 18, pp. 427-447, 4 figs., 7 pls. Woopwarp, A. S. 1921. Visit to the gallery of fossil fishes, British Museum (Natural History). Proc. Geol. Assoc., London, vol. 32, pt. 38, pp. 179-181, 2 figs. U.S, GOVERNMENT PRINTING OFFICE: 1958 mi sf Mies: fk i sane ih aorshatt ee raed Pee at b; panied ae eStore IE i ted Pile Iepseiltonina ta. is a 2 N , : : “_ a (as jay 7 te he oe ic diy ¥ i a ee; a mae, ef: aluniee ih i ne - |) gael at af RE a : pe 9 De Hh uF 7 asain ey: sae, ta eat Nek) i ia tile er 1 ana i Ge Whe) Figlwaceirhet Pit ae a a i ; ; Te - 7 ioe Pirates 1-3: PrateE 1 Pycnaspis splendens, new genus, new species. 1, 2, USNM 21333, holotype, from Upper Ordovician, Rock Creek section, Johnson County, Wyo.: 1, detached exoskeletal plate, somewhat incomplete, showing tubercles of early and subsequent stages of growth, super- ficial view; 2, same, oblique lateral view. 3-6, USNM 21336, from same layer and locality as the holotype, detached ridge-scale oriented (3-5) with its anterior margin upwards: 3, lateral view; 4, external view; 5, basal view; 6, posterior view. Magnification: 1, 2, X 32; 3-6, X 24. Explanation of symbols: bf, basal face; oda, anterior portion of external face, overlapped by the ridge-scale in front; og, lateral faces, overlapping adjoining polygonal plates; og», posterior face, overlapping the ridge-scale following next behind; tg, tubercles belonging to early stages of growth; ¢,, tubercles belonging to subsequent stages of growth. PROC. U. S. NAT. MUS. VOL. 108 ORVIG, PLATE 1 Pycnaspis splendens, new genus, new species. [xplanation on facing page. Piate 2 1-3, Pycnaspis sp. indet. USNM 21337 from Middle Ordovician, Winnipeg formation, Shell Pine Unit No. 1 core, depth 9525-9531 feet, sec. 30, T. 12 N., R. 57 E., Wibaux County, Mont.: 1, detached exoskeletal plate, somewhat incomplete, external view; 2, 3, same, lateral views from opposite sides. 4-8, Eriptychiida gen. and sp. indet. 4-7, USNM 21340, branchio-cornual plate from the same layer and locality as the holotype of Pycnaspis splendens, new genus, new species: 4, lateral view; 5, ventrolateral view; 6, posterolateral view; 7, medial view, showing the basal face of the plate after removal of the matrix. 8, USNM 21341, fragment, possibly of another branchio-cornual plate, from the same layer and locality as the foregoing; a short wide canal in the interior of the plate, which may be a lateral line canal, is shown on the broken-off surface. Magnification: 1-3, X 12; 4-7, * 2.4;8, X 4. Explanation of symbols: br. 0, branchial opening; //c?, possibly a lateral line canal. PROC. U. S. NAT. MUS. VOL. 108 ORVIG, PLATE 2 Pycnaspis sp. indet. and Eriptychiida gen. and sp. indet. [explanation on facing page. Prare.3 Eriptychiida gen. and sp. indet. 1-4, USNM 21342, a detached keel-like plate, presum- ably belonging to a new genus of the Eriptychiida, from the same core and the same depth as the Pycnaspis specimen shown in figures 1-3 of plate 2: 1, external view; 2, 3, lateral views from opposite sides; 4, basal view. 5, 6, USNM 21818, a detached ridge-scale, some- what incomplete, from the same layer and locality as the holotype of Pycnaspis splendens, new genus, new species: 5, external view; 6, anterior view. Magnification: 1+, X 7; 35.6, 2% 16.55 QYRVIG, PLATE 3 PROc. U.S. NAT. MUS. VOL. 108 Mes Be eT TE RS Meee, ak ty Fao aa on facing page. and sp. indet. Explanation gen. Eriptychida PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM PGA INCRE hee Service iw ECD LAN ES (SEEN ey BD oF are 525) by the sy x Id WAT HONS, 42¥5, HSOX9 45, 7ITVT Oo. cron” SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM pa eo} KN, issued Vol. 108 Washington : 1958 No. 3392 BRANCHINECTA CORNIGERA, A NEW SPECIES OF ANOSTRA- CAN PHYLLOPOD FROM THE STATE OF WASHINGTON By James E. Lynca’ The fairy shrimp described below has been repeatedly collected by the writer since 1935 from numerous ephemeral fresh-water ponds in eastern Washington. The data in the specific diagnosis are based on measurements of ten apparently full-grown individuals of each sex. The specific name, cornigera, refers to the dorsolateral protuberances on the head of the female, which remind one of incipient horns. Genus Branchinecta Verrill, 1869 Branchinecta cornigera, new species Speciric pracNosis: Male: Total length, from front to end of shaft of cercopods, 29 (25-36.5) mm. Average ratio of length of head and thorax to genital segments, abdomen, and cercopods 1 :1.3. Nuchal organ transversely oblong with rounded corners on top of head. Compound eye in lateral view slightly flattened dorsoventrally, 0.9 (0.75-1.05) mm. by 1 (0.95-1.2) mm. Antennule 2.6 (2-3) mm. long, terminated by three setae and eight aesthetascs. Antenna biarticulate, 7 (6-8) mm. long, extending back to the level of the fifth or sixth thoracic segment if flexed posteriorly. Proximal article 4.5 1 School of Fisheries, University of Washington, Seattle, Wash. 25 44627458 26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 (4-5) mm. long, without apophysis; near its junction with the head an anteromedian bulging area (pulvillus) bearing minute spinules 7-10 microns long; a padlike area of cuticular verrucae on the medioanterior and anteromedian sides of its lower half, with a prominent, uncorni- fied, anterior protuberance near the proximal end of the verrucose area. The proximal article presents a more or less prominent genicu- late bend as seen in lateral view, with the uncornified bulge at the angle of the bend. Distal article of the antenna 2.5 (2-3) mm. long, somewhat flattened laterally, curving gently inward to a pointed tip with a roughened area. ‘Thoracic appendages typical of the genus; branchial lamina entire with serrated margins; endopodite produced medially, the heavy spines of the distal end and median border thickly beset on all sides with distally directed spinules. Second genital segment with a pair of hollow eminences on ventral side. The penes bear each a chitinized spur 0.26 (0.22-0.31) mm. high near the tip of the rigid portion; the retractile portions when fully protruded present two distal lobes, each with 6 to 10 pyramidal teeth. Cercopods 2.3 (2-2.75) mm. long, distinctly articulated to the telson, with long, plumose hairs on both lateral and median margins. Female: Total length, from front to end of cercopods, 29 (21.5-33.5) mm. Average ratio of length of head and thorax to genital segments, abdomen, and cercopods 1:1.48. Transversely oblong nuchal organ on top of head. A retrorse, papillose protuberance, rising 0.35—0.55 mm. above the surface, is located on each side of the head dorsal to the mandibular articulation. Compound eye in lateral view slightly flattened dorsoventrally, 0.62 (0.6-0.7) mm. by 0.76 (0.6-0.85) mm. Antennule 1.8 (1.5-2.5) mm. long, terminated by three setae and eight aesthetascs. Antenna 3 (2.2-3.5) mm. long, cylindrical, with a me- dian spur 0.18—-0.35 mm. long at the junction of the middle and distal thirds, distal to which the antenna tapers to a sharp, incurved point. Thoracic segments 6 to 11 each with a right and left dorsolateral, conical protuberance, of which the last four, only, are large enough to be conspicuous. The largest protuberance, 0.5 to 0.9 mm. high, usu- ally on segment 10, occasionally on segment 11. Ovisac fusiform, ex- tending backward for the length of 4.75 (4.3-5.8) abdominal segments; may contain up to 416 eggs. Eggs from preserved specimens 0.29- 0.34 mm. in diameter. Cement glands consist of two closely ap- pressed masses of large cells at anterior end of ovisac; from each mass a row of cells extends for from 65 percent to 80 percent of the length of the ovisac on its dorsal side and widens at its posterior end, and similar rows of cells extend posteriorly along the midventral line for about half the length of the ovisac, without the posterior expansion. Shaft of cercopods 2.2 (1.9-2.8) mm. long, with long plumose setae on lateral and median borders. 27 ANOSTRACAN PHYLLOPOD—LYNCH (SZ X YyI0g) *q'2 {xvJOY] Jo apis uO saqo] [eJole[esiop -laqnjoid snosu103 ‘ oe ) suo] ‘wu QZ a[eulay “T *sotoads mau ‘¢ “ZUO] “WU FOZ BRP ‘Z “YLT pros ul poywidap oie spuvjs Juewes S(pesy Jo opis [vio}e] uo aoue rp p4931U409 DIIIULYIUDIG *7 INV [| Saunory 28 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 108 Smallest specimens which obviously were sexually mature were females 11 mm. long with one or two eggs in the ovisac and males 12 mm. long. In small females, 11-12.5 mm. long, the ovisac is relatively shorter than in full-grown ones, extending only the length of 2% to 3% abdominal segments. Typr Locauiry: A pond in Lincoln County, Wash., about 8% miles southeast of Creston, and about 50 yards north of U. S. Highway 2. Elevation about 2,400 ft. above sea level. TYPE SPECIMENS: One male holotype (USNM 100912) and 8 male and 8 female paratypes (USNM 100913) have been deposited in the U.S. National Museum. DIFFERENTIATING CHARACTERS: Male: The clasping antenna of the male has an extensive verrucose area on its anteromedian and medio- anterior sides and a prominent rounded bulge near the upper part of the verrucose area. The proximal article is more or less pronouncedly geniculate in lateral view. The degree of geniculation varies with muscular contractions, but is constant enough to constitute a specific character. Female: The second antenna of the female, near the junction of the middle and terminal thirds, has a medially directed spur, distal to which the antenna tapers to an inwardly curved point. No North American species described to date has this feature, although it occurs in the South American Br. pollicifera Harding, 1940. The female has a pair of conspicuous papillose protuberances on the dorsolateral sides of the head. This hornlike outgrowth is either absent or rudimentary on other North American species, and has not been recorded from exotic species. Dorsolateral lobes, of conical shape, occur from thoracic segment 6 or 7 to segment 11, that on segment 10, or sometimes on segment 11, being the largest. Other species lack such lobes, or have them differing as to number, or arranged in different sequences as to size. CoLor OF LIVING SPECIMENS: Br. cornigera, new species, is more brightly colored than most species of the genus in western North America. The females are more brightly colored than the males, and their coloration is less variable. Indeed, the variation in shade and intensity of color of the males is so great that it is difficult to describe their colors adequately or briefly. Male: The over-all color varies from a burnt orange to a pale greenish or gray-green; occasionally they are almost colorless except for the greenish head and pale green appendages. The antennule is pale yellow, the basal article of the clasping antenna may be dark blue-green, yellow-green, or nearly colorless; the distal article yellow to orange with greenish punctations on the lateral side, and gray on the roughened area near the tip. The front and top of the head are ANOSTRACAN PHYLLOPOD—LYNCH 29 = SS LZ, = — Ss SSS aS IW SS |omm Ficures 3-6: Branchinecta cornigera, new species. 3, Anterior aspect of head, male (pu, pulvillus). 4, Same, female. 5, Spine from median border of endopodite of thoracic appendage of a male. 6, Spines from median border of thoracic appendage of a female. (3, 4, X 10; 5, 6, X 340.) 30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 often yellow, but with scattered punctations of green. There is a large bluish spot on the front on each side of the ocellus. The peduncle of the eye is yellow, the corneal portion black. The dorsal side of the thorax is grayish, grayish green, or sometimes faint orange, or colorless. The anterior third of the intestine, as seen through the body wall, is yellow; the posterior two-thirds is black from accumulated food residues. Ficure 7: Right fifth thoracic appendage of a male, anterior aspect (br.1., branchial lamina; br.s., branchial sac; ent., endites; end., endopodite). (XX 18.) See note, figure 8. The axis or corm of the appendages is pale green, blue-green, or orange-yellow, and contains numerous large cells with colorless, ~ yellow, or orange oil globules. These cells range upward on the sides of the thorax between the appendicular muscles, thus tinting the lower half of the thorax yellowish. The branchial lamina and gill are color- less; the exopodite and endopodite nearly colorless to pale green or dark yellow; the endites pale green bordered with yellow. The dorsal and lateral sides of the genital segments are like adjacent parts of the abdomen and are nearly colorless; the ventral sides are colorless or yellow. The spur, and border of the penis anterior to it, ANOSTRACAN PHYLLOPOD—LYNCH 31 are dark yellow. The abdomen may be colorless, except for the black, food-packed intestine, but often is pale gray-green or grayish yellow. Cercopods are colorless, to yellow, orange-yellow, or pinkish. Female: The antennule is colorless to pale yellow or salmon-pink. The antenna is yellow with a greenish wash, or green with a yellow tip. The top of the head is faint blue-green, but often the entire head appears yellow from the yellow digestive caeca. The “horns” on the side of the head may be yellow, blue, or blue-gray. The corneal part of the eye is black. The labrum is greenish, spotted with blue, and may contain large, brownish, internal cells visible through the integu- ment. The dorsal side of the thorax may be colorless, whitish, or faint green; the dorsolateral lobes colorless to faint yellow. The anterior third of the intestine, as seen through the body wall, is yellow to pink; the posterior two-thirds is black with contained food residues. The lateral sides of the thorax, above the appendages, are blue; the blue extends further dorsally in the last four segments. The blue color is \\ HT [In Ly, ‘YS May Ficure 8: Right fifth thoracic appendage of a female, anterior aspect (ex., exopodite). (< 18.) Note: No attempt has been made to represent the setules of the plumose setae which fringe the thoracic appendages. On the ventral border of the endopodite the plumose setae gradually become thicker and assume the character of spines, with spinules replacing the setules. 32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 caused by large, vividly blue cells which occur among the appendicular muscles, and also as a layer applied to the dorsal side of the intestine throughout its length, and to the ventral side in the thoracic region. The axis, or corm, of the appendages is colorless or pale green, some- times contains blue cells, and in addition has large cells, less numerous than in the male, with yellow, orange, or brown oil globules. Endites, exites, and endopodite and exopodite are colored as in the male. The ovary appears yellowish white to light blue, depending on the stage of development of the large odcytes, and can be plainly discerned through the body wall from the fourth or third abdominal segment to the eighth or seventh thoracic segment. The color pattern of the ovisac is characteristic of the species. The proximal third of its wall is colorless to pale yellow; the middle third yellow, orange, or pinkish yellow; about two-thirds of the posterior third is bright blue, and the tip, about 12% percent of the total length, is white or faintly blue. In the case of living females, the species can be easily recognized by a conspicuous blue area, which can be seen by the unaided eye, on the posterior half of the ovisac. The cement glands appear brownish yellow to pale brown under low magnification, and have a characteristic arrangement. Eggs, within the ovisac, vary from dull yellow to yellow-brown. The walls of the abdomen are nearly colorless except for the black intestine and the conspicuous layer of blue cells along the dorsal side of the intestine. The cercopods are colorless in most, but sometimes are pale yellow, pale orange, or colorless with yellow or pinkish yellow borders. Discussion: The number and arrangement of the dorsolateral lobes on the thorax of the female is probably a good specific character in many species of Branchinecta. In fact, they are conspicuous in size, shape, and arrangement in many species of other genera, but have rarely been mentioned or depicted in taxonomic descriptions. They have been present in every specimen of Br. cornigera, new species, and also in every specimen of Br. packardi Pearse and Br. lindahli Packard (as revised by Shantz)? that I have examined. In Br. coloradensis Packard (as revised by Shantz)? and in Br. paludosa (O. F. Miiller) they are conspicuous in populations from some localities and absent in those from other localities. Other characters of some taxonomic value are the pulvillus and the dorsolateral bosses. The pulvillus, or swollen area on the median side of the basal article of the antenna of the male near its junction with the head, is present and conspicuous in some species of Branchinecta and absent, or barely represented, in others. When present, it is covered with minute spinules, verrucae, or denticles 2 See under heading of ‘‘ Remarks,” page 34. ANOSTRACAN PHYLLOPOD—LYNCH 33 pie 11 Ficures 9-12: Branchinecta cornigera. 9, Right antenna of male: a, anterior view (pu., pulvillus; bu., uncornified protuberance of second antenna); b, lateral view. 10, Right antenna of male (another specimen): a, anterior view; b, lateral view. The right antennae from the two specimens show that although the antennae of the male vary in shape with muscular contractions, nevertheless the uncornified bulge and the angular outline of the anterior border remain evident. 11, Telson and cercopods of a female 27 mm. long. 12, Hornlike protuberance from left side of head of female, lateral aspect. (9, 10, X 11.5; 11, X 18; 12, X 80.) 34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 which differ in shape in each species. Although the pulvillus has been noted and depicted by some specialists on the Anostraca, I can find no description of the cuticular elaborations of its surface. Most species of Branchinecta have, on the dorsolateral sides of the entire trunk, more or less conspicuous rounded elevations, or bosses, covered with cuticular papillae, with one or several sensory hairs near the center (see Linder, 1941, figs. 20 and 22). These papillose bosses are very poorly developed in Br. cornigera, new species, and often appear to be absent on one or both sides of some segments. The shape of the endopodite of the thoracic feet and the structure of the spines on its distal end and median border are both strikingly different in the male and female (figs. 7, 8). Such a sexual dimor- phism in the endopodites appears to be the rule in the genus, although Br. gigas, in which the thoracic feet are practically alike in both sexes, is an exception. The endopodital spines, especially those of the males, constitute a minor specific character (figs. 5, 6). DIstRIBUTION AND ENVIRONMENT: Branchinecta cornigera, new species, has been collected from 30 temporary ponds in Grant, Lincoln, Spokane, and Adams Counties in eastern Washington. The total number of separate collections is 43, all of which were made from late March to the middle of May. The species is typically present in clear or slightly turbid water with a variety of other fresh-water organisms, and often is associated with other species of phyllopods. Br. cornigera was the sole phyllopod in 25 of the 43 collections; it was associated with Hubranchipus serratus 13 times; with Hu. serratus and Lynceus sp. once; with Hu. bundyi once; with Br. mackini twice, and with Br. mackini and Br. gigas once. The temperature of the milieu water ranged from 42° to 59° F. On 15 occasions the pH of the water was ascertained with a Hellige Pocket Comparator. The water was only moderately alkaline, the pH ranging from 7 to 8.8. In the pond where Br. cornigera was associated with Br. gigas and Br. mackini the water undoubtedly was much more alkaline (probably pH 10 or more), but unfortunately the pH was not taken in that case. Remarks: I have been unable to accept the shuffling of specific names proposed by Dr. J. G. Mackin (1952) for the following reasons: Packard’s (1883) description of Br. lindahli is so deficient and brief as to be meaningless, and is not accompanied by illustrations. It is not possible to recognize the species from the description. Packard’s original (1874) description of Br. coloradensis is entirely inadequate, and the drawings are so trivial and devoid of essential detail as to be without taxonomic utility. This description was based upon one female from an altitude of 12,500 ft. in the Colorado Rocky Mountains, and about 100 specimens of both sexes from the indefinite locality of ““Colorado.”” Packard’s second (1883) description ANOSTRACAN PHYLLOPOD—LYNCH 35 ia. >... Ficures 13-16: 13, Genital segments and first abdominal segment of a male 23 mm. long, lateral aspect (sp., spur on copulatory appendage). 14, Genital segments of a male 23 mm. long, ventral aspect. 15, Cuticular verrucae from the verrucose area of the antenna of a male. 16, Cuticular spinules from the pulvillus of the antenna of a male. (13, 14, xX 24; 15, X 460; 16, * 1066.) 36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 of Br. coloradensis, accompanied by the drawings of 1874, contains statements contradicted by the drawings and by the first description, as well as internal contradictions and confusions. The material for the second description came from a third locality on Gray’s Peak at an elevation of 12,000 ft. On the basis of the second description Br. coloradensis remains incognizable, and it is most likely that the two descriptions are based on a careless examination of two species. It seems appropriate, at this point, to call attention to the untrust- worthiness of Packard’s work on phyllopods. His descriptions are not only deficient in differential characters, but they also contain errors, contradictions, vague and indefinite comparisons with other species, invalid distinctions, and are replete with blunders and mis- labeling in the illustrations. He apparently left no type specimens of any of his species of fairy shrimp. In my opinion, neither of Packard’s species of Branchinecta can be recognized from his publications, and his specific names should be regarded as nomina dubia. Dr. Mackin’s distinction between alpine, permanent-water species and species inhabiting the plains and foothills at lower altitudes is not in complete accordance with my own experience. I have collected both Br. coloradensis and Br. paludosa in the Medicine Bow Mountains at elevations above 10,000 ft. and also on the plains in Montana at 4,200 ft. I have in my collection six distinct species of Branchinecta collected from the plains regions of Montana and Wyoming. These circumstances make most uncertain any surmises as to the species Packard had before him, based on their supposed occurrence only in an alpine or a plains locality. Shantz (1905) was the first reviser of Packard’s work on Branchi- necta, and the species he described as being, in his belief, Packard’s coloradensis and lindahli are entitled to stand. Pearse (1912) gave a description adequate for the recognition of a species which up to that time had not been recognizably described, and there is no con- vincing proof that Br. packardi is, in fact, a synonym of one of Packard’s species. As a consequence, Br. shantzi Mackin, 1952, must be regarded as an invalid synonym of Br. coloradensis Packard (as revised by Shantz, 1905); the Br. coloradensis of Mackin’s revision is an invalid synonym of Br. lindahli Packard (as revised by Shantz, 1905), and the Br. lindahli of Mackin’s revision is an invalid synonym of Br. packardi Pearse, 1912. ANOSTRACAN PHYLLOPOD—LYNCH 37 Literature cited LINDER, FOLKE 1941. Contributions to the morphology and taxonomy of the Branchiopoda Anostraca. Zool. Bidrag f. Uppsala, vol. 20, pp. 101-302, 45 text figs., 1 pl. Mackin, J. G. 1952. On the correct specific name of several North American species of the phyllopod genus Branchinecta Verrill. Amer. Mid]. Nat., vol. 47, No. 1, pp. 61-65. Packarp, A. S. 1874. Synopsis of the fresh-water phyllopod Crustacea of North America. Ann. Rep. U. 8S. Geol. Geogr. Survey Territories . . for the year 1873, pp. 613-622, pls. 1-4. 1883. A monograph of the phyllopod Crustacea of North America, with remarks on the order Phyllocarida. 12th Ann. Rep. U. S. Geol. Geogr. Survey Territories . for the year 1878, pp. 295-457, pls. 1-38, 73 text figs. Pearse, A. S. 1912. Notes on phyllopod Crustacea. 194, pls. 1-3. SHantz, H. L. 14th Rep. Mich. Acad. Sci., pp. 191- 1905. Notes on the North American species of Branchinecta and their habitats. Biol. Bull., vol. 9, No. 4, pp. 249-259, pls. 10-12. U. S$. 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oT i en PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1958 No. 3393 TYPE SPECIMENS OF LICE (ORDER ANOPLURA) IN THE UNITED STATES NATIONAL MUSEUM By Paytuts T. Jonnson' It has been recommended in the Copenhagen Decisions on Zoological Nomenclature (1953) that: “All institutions maintaining zoological collections should prepare and publish lists of type material in their possession.” (Recommendation 75, p. 78, par. 150.) The present paper includes the names of all the Anoplura represented by the type in the U. S. National Museum as of Dec. 31, 1956. As arule, reference is made only to holotypes and syntypes, paratypes being mentioned only where it is necessary to clarify the data or otherwise aid in the fixing of holotypes and lectotypes. The specific name is given under the genus to which it is now assigned, with a brief comment on its present status. If host names given in the type data have been changed since the original description of the Anoplura species, the currently accepted name is placed in brackets following the original form of the name. Appended is an alphabetical listing of the specific names of the Anoplura together with their currently accepted generic names. If the generic name differs from that designated by the author of the species, the original genus is given in brackets. 1Entomology Research Branch, Agricultural Research Service, U. 8, Department of Agriculture. 453663—58 39 40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 I am grateful to Dr. Henry W. Setzer of the Division of Mammals, U. S. National Museum, who checked the host names used in this paper. Genus Enderleinellus Fahrenholz Haematopinus suturalis Osborn, U.S. Dep. Agr. Div. Ent. Bull. (old series) 7, p. 27, fig. 15, 1891. Types pata: “Plentiful’” on Spermophilus franklini [Citellus franklini] and S. 13-lineatus [Citellus tridecemlineatus] at Ames, Iowa. There is one slide with eight specimens of suturalis in the collection with an Osborn label bearing the data ‘on Spermophilus 13-lineata, 4/24/84, Ames, Iowa.” There is no determination on this label. On the opposite side, in H. E. Ewing’s handwriting, is given: ‘“Cyclo- phthirus [Enderleinellus (crossed out)] suturalis (Osborn),’”’ and a sketch of the position of the specimens with one marked ‘“o holotype.” Osborn did not designate a holotype, and Ewing never published a lectotype selection; therefore, Ewing’s label means nothing from a nomenclatural standpoint. Although there is no species determination by Osborn, it was Ewing’s habit to rewrite type labels at times with a new determination or a different generic combination. It is there- fore accepted that Ewing recognized these to be the type series of suturalis, and the male designated by Ewing as ‘‘* holotype’’ is here selected as lectotype. Present status: Enderleinellus suturalis (Osborn), 1891. Genus Eulinognathus Cummings Eulinognathus americanus Ewing, Journ. Washington Acad, Sci., vol. 13, No 8, p. 149, 1923. Tyre pata: USNM 23761. Holotype female from Ctenomys brasiliensis (USNM 1939/3252) taken at Salade River, Paraguay. PresEeNT status: As originally described. Genus Haematopinoides Osborn Haematopinoides squamosus Osborn, U.S. Dep. Agr. Div. Ent. Bull. (old series) 7, p. 28, fig. 16, 1891. Type pata: Two females from pocket or pouched gopher, Geomys bursarius, Ames, Iowa. There is in the collection a slide bearing data as above, a deter- mination label in Osborn’s handwriting, and containing two syntype females. Further labels have been placed on the slide by H. E. Ewing, the type No. 24137 is given and an accessory label gives a sketch of the position of the two females with the one on the right (next to the type label) labeled as lectotype. Ewing did not publish TYPE SPECIMENS OF LICE IN USNM—JOHNSON 41 the lectotype selection. The specimen so designated on the slide is the better of the two and is here selected as lectotype. PRESENT status: As originally described. Genus Haemodipsus Enderlein Haemodipsus setoni Ewing, Amer. Journ. Trop. Med., vol. 4, No. 6, p. 548, 1924 Typr pata: USNM 23768. One female from Lepus californicus melanotis (USNM 123846) and two females and two nymphs from another skin (USNM 123847) of the same host subspecies, collected at Wichita, Kans., by Ernest Thompson Seton, and one male and one nymph from Lepus c. californicus (USNM 60907) collected at summit of Coast Range Mountains, San Diego County, Calif., by the Inter- national Boundary Commission. All the syntype specimens listed above are in the collection under USNM 23768. None of the specimens are very well preserved. The single female from USNM 123846, although some of the legs are missing, is best preserved and is designated lectotype. PreEsENT sTaTus: As originally described. Genus Hoplopleura Enderlein Hoplopleura erismata Ferris, Stanford Univ. Publ. Biol. Sci., vol 2, No. 2, p. 113, figs. 72B,u,F, 1921. Typrk pata. USNM 201408. Holotype female, allotype male from Sciurus ferrugineus cinnamomeus [Callosciurus ferrugineus cinnam- omeus], South East Siam. The type slide, containing a male and a female, bears the following data: ‘Ferris Col. 457, Type o& allot. 9, Hoplopleura erismata n. sp., from Sciurus ferrugineus cinnamomeus, 8. E. Siam. USNM 201408.” The holotype is accepted as being the female on this slide, ‘type # allot. 2’’ as written on the slide label being a lapsus. PRESENT status: As originally described. Haematopinus hesperomydis Osborn, U. S. Dep. Agr. Div. Ent. Bull. (old series) 7, p. 26, fig. 14, 1891. Typr pata: Male(s) and female(s) from white-footed mouse or deer mouse, Hesperomys leucopus [Peromyscus leucopus subsp.] at Ames, Iowa. There is in the collection one slide bearing a label on the right in Osborn’s handwriting: ‘“Haematopinus hesperomydis n. sp., from white-footed mouse, H. O. 1885.’’ There are three male and one female syntypes on the slide; a well-preserved male is chosen lectotype and has been circled with a diamond point pencil. PRESENT status: Hoplopleura hesperomydis (Osborn), 1891. 42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Hoplopleura oryzomydis Pratt and Lane, Journ. Parasit., vol. 37, No. 2, p. 141, pl. 1, figs. 1-8. Type pata: USNM 60412. Holotype female and allotype male from Oryzomys palustris palustris (Harlan) No. 78, Oatland Island, Chatham County, Ga., Feb. 26, 1948. Both the holotype and allotype are in the collection. PRESENT status: As originally described. Genus Lemurphthirus Bedford Lemurphthirus verruculosus Ward, Ent. News, vol. 62, No. 6, p. 190, figs. 1, 2, 1951. Type pata: USNM 64237. Holotype female from a mouse lemur (formalin specimen), Bemangidy, Fort-Dauphin district, Tuléar Province, Madagascar, 1948, H. Hoogstraal and R. Alison. PRESENT status: As originally described. Genus Nechaematopinus Mjoberg Haematopinus antennatus Osborn, U.S. Dep. Agr. Div. Ent. Bull. (old series) 7, p. 25, fig. 13, 1891. Type pata: Collected from a fox squirrel, Sciurus cinereus var. ludovicianus |Sciurus niger subsp.], at Ames, Iowa. There is in the collection a slide bearing labels as follows: Left: USNM type label, in Ewing’s handwriting, ‘‘Neohaematopinus an- tennatus (Osborn), remounted on original slide Nov. 22, ’20 by H. E. E., Type No. 24136 U.S.N.M.” Right Label: In Osborn’s handwriting, ‘“H. [sphaerocephalus (crossed out)] antennatus n. sp., on Fox Squirrel, 5/1/82, S. K. Chauri 10/5/[?].””. There are three female syntypes in a row on this slide. I select as lectotype the best preserved female, in the middle of the row, and have circled it with a diamond point pencil. PresEnT status: A junior homonym of Haematopinus antennatus Piaget, 1880. Earliest available name: Acanthopinus sciurinus Mjéberg, 1910, now known as Neohaematopinus sciurinus (Mjéberg), 1910. Haematopinus columbianus Osborn, Canad. Ent. vol. 32, No. 7, p. 215, 1900. Typr DATA: Described “from a number of specimens taken from the Columbian Spermophile, Spermophilus columbianus [Citellus colum- bianus subsp.], at Pullman, Washington, by Prof. C. V. Piper in July, 1896. Type material in USNM.” In the collection are two slides which must be considered in lecto- type selection. The first has on the left, in Ewing’s handwriting, on a USNM type label: ‘=Linognathoides laeviusculus (Grube)” and on the right: “(8526, Linognathoides montanus (Osb.) (in Ewing’s handwriting)], Haematopinus columbianus Osb., type material. TYPE SPECIMENS OF LICE IN USNM—JOHNSON 43 Type No. 5178, U.S. N.M.” There are five poorly preserved females on the slide. The second slide has on the left-hand label in Osborn’s handwriting: ‘8526 (80/13), Haematopinus columbianus Osb., on Spermophilus columbianus, Pullman, Wash., July ’96, C. V. Piper,” and contains eight females and six nymphs. The best preserved female on this second slide has been chosen as lectotype and has been circled with a diamond point pencil. PRESENT status: A junior synonym of Neohaematopinus laevius- culus (Grube), 1851. Neohaematopinus mathesoni Rubin, Proc. Ent. Soe. Washington, vol. 48, No- 5, p. 121, figs. 1, 8, 10, 12, 1946. Type pata: USNM 57685. Holotype female from Citellus v. couchi (Baird) [Citellus variegatus couchi], at Nuevo Leén, Mexico, August 12, 1938, collected by H. Hoogstraal. The holotype female bears the following host and locality data: “Citellus v. couchi, Ojo de Aqua, Municipio de Galeand, N. L., Mexico, Aug. 11, 1938,” (not Aug. 12). The localities are properly Municipio Galeana and Agua del Oro. PRESENT status: A junior synonym of Neohaematopinus marmotac Ferris, 1923, NEw SsyNoNyMy. One would expect Neohaematopinus laeviusculus (Grube) rather than N. marmotae, which is a normal par- asite of Marmota and Cynomys, to occur on Citellus. However, the type and paratypes of N. mathesoni agree with Ferris’s figures and description of N. marmotae and with specimens from the normal hosts. Haematopinus montanus Osborn, U.S. Dep. Agr. Bull. 5, new. ser., p. 184, fig. 107, 1896. Type pata: On western gray squirrel, Fort Collins, Colo. (Baker). There is in the collection a slide containing two males and two females and labeled as follows: Left label: ‘Linognathodes [montanus Osb. (crossed out)] laeviusculus (Grube).’’ Right label: ‘Near Ft. Collins, Colo., 9-11-92, Stannard. From Spotted grey squirrel.” These labels are both in Ewing’s handwriting. Although the above specimens may be type material, the absence of a label in Osborn’s handwriting and the disparity in host names and the difference ia what may be the collector’s names do not allow them to be considered as such. Apparently no type material of H. montanus is in existence. Present status: A junior synonym of Neohaematopinus laevius- culus (Grube), 1851. Neohaematopinus patiki Rubin, Proc. Ent. Soc. Washington, vol. 48, No. 5, p. 121, figs. 2, 5, 7, 11, 1946. TypE pata: USNM 57686. Holotype female, 11 paratype females, from Citellus (Ammospermophilus) sp., Delta, Utah, Apr. 27, 1938, Nual Walter collector. Eight paratypes are in the Cornell collection, the rest are in the U. S. National Museum (No. 57686). 44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The type slide is in the collection, but there are two female speci- mens on it and no designation as to which should be considered the holotype. Therefore, both female specimens on this slide must be considered syntypes. The female nearest the right-hand label on the slide marked ‘‘type’’ is here designated lectotype, and circled with a diamond point pencil. Present status: A junior synonym of Neohaematopinus laevius- culus (Grube), 1851, NEW SYNONYMY. Neohaematopinus traubi Rubin, Proc. Ent. Soc. Washington, vol. 48, No. 5, p. 120, figs. 3, 4, 6, 9, 13, 1946. Type pata: USNM 57684. Holotype female from Citellus adocetus (Merriam), Michoacan, Mexico, Aug. 3, 1941, R. Traub collector. The holotype female bears the additional locality data: Apatzingan, Michoacan; and the date is given as “21 Aug. 1941,’ not Aug. 3, 1941. Present status: As originally described. Genus Pecaroecus Babcock and Ewing Pecaroecus javalii Babcock and Ewing, Proc. Ent. Soc. Washington, vol. 40, No. 7, p. 199, figs. 1-3, 1938. Type pata: USNM 52758. Males, females, nymphs, and eggs from Pecari angulatus [Pecari tajacu angulatus], collared peccary, western Texas, between Juno and Pecos River, Jan. 29, 1932, O. G. Babcock collector. In the collection are numerous specimens with the data similar to the above. Only two males and two females bear the type number (USNM 52758). A well-preserved male bearing the following data has been chosen and labeled as lectotype: Left label: ‘““Peccary, Juno, Texas, 1-29-1932, Bishopp No. 14093’; right label: ‘“‘Pecaroecus javalii Babe. & E., Type No. 52758 U.S. N. M., Long-nosed Peccary- louse, O. G. Babcock coll.” Present status: As originally described. Genus Pediculus Linnaeus Pediculus (Parapediculus) atelophilus Ewing, Proc. U. S. Nat. Mus., vol. 68, No. 19, p. 9, figs. 44, 5, 1926. Typr pata: USNM 28105. Holotype male from Ateles geoffroyi [Ateles geoffroyi subsp] (gray form, or melanochir type of coloration), type locality ?. Present status: Ferris (1951, Mem. Pacific Coast Ent. Soc., vol. 1, p. 273) presumes this species to be the same as Pediculus lobatus Fahrenholz, 1916, but does not actually synonymize it. Pediculus (Parapediculus) chapini Ewing, Proc. U. 8S. Nat. Mus., vol. 68, No. 19, p. 13, text figs. 2, 48, 5, pl. 1, figs. 3, 4, 1926. Typr pata: USNM 28106. Described from males and females from Ateles ater [Ateles paniscus paniscus], locality unknown. JOHNSON 45 TYPE SPECIMENS OF LICE IN USNM The only slide with the above data contains two syntype specimens, a male and a female. The male is designated lectotype. Present status: Probably the same as Pediculus atelophilus Ewing, 1926. Pediculus (Parapediculus) pseudohumanus Ewing, Journ. Parasit., vol. 24, No. 1, p. 23, figs. 3, 5a, 6a, 1938. Tyrer pata: USNM 51451. Type specimens taken from a saki monkey (Pithecia monachus) [Pithecia monacha subsp.] that died at the National Zoological Park, Washington, D. C. Original home of the type host is the Upper Amazon. There are four slides in the collection labeled essentially as above and containing (1) 4 eggs, (2) 4 nymphs, (3) 4 males, and (4) 4 females. A well-preserved female has been chosen lectotype. This female is the one farthest from the type label (i. e., farthest to the right) on the slide containing four females and has been circled with a diamond point pencil. Present status: Pediculus pseudohumanus Ewing, 1938. Pediculus (Paenipediculus) simiae Ewing, Proc. Biol. Soc. Washington, vol. 45, p. 117, 1982. Type pata: USNM 44327. Holotype female from chimpanzee, Pan sp., London Zoological Gardens. PRESENT sTaTus: A junior synonym of Pediculus shdéffi Fahrenholz, 1910. Genus Phthirpediculus Ewing Phthirpediculus propitheci Ewing, Journ. Washington Acad. Sci., vol. 13, No. 8, p. 150, 1923. Type pata: USNM 23762. Two females and one male (on the type slide) from female skin (USNM 63352) of Propithecus edwardsi [Propithecus diadema edwardsi] taken at Ambodiasy, eastern Mada- gascar, and two males from male skin (USNM 63354) of same host, taken at same place. The above syntype specimens are in the collection. A well-pre- served male is chosen lectotype. The lectotype is on the type slide which bears data as in the original description and a USNM type label. PRESENT status: As originally described. Genus Phthirus Leach Phthirus gorillae Ewing, Proc. Ent. Soc. Washington, vol. 29, No. 5, p. 120, 1927, Type pata: USNM 40161. First stage nymphs and eggs from two Gorilla beringeri [Gorilla gorilla beringei] (USNM 239883, 239884), eastern Belgian Congo. 46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 In the collection are five slides labeled ‘“‘Phthirus gorillae.”” Three of the slides contain only eggs, and since Ewing described just the first stage nymph the eggs need not be considered in type selection. Of the two nymphs, one bears the following data: Left; “Type No. 40161 U. S. N. M., Phthirus gorillae n. sp., Ist nymph”; Right: “Beloian Congo, 1923, by Benj. Burbridge, from young gorilla, U. S. N. M. 239884.”’ The other slide has, on the left, ‘“Phthirus gorillae Ewing, Ist nym.,” and, on the right, ‘Belgian Congo, Rec’d Jan., 1924, Benj. Burbridge, from skin of young ¢@ gorilla (U.S. N. M. 239884).”” Although this second specimen is not marked as a type, Ewing mentions “nymphs” in his description, making the selection of a lectotype necessary. The nymph with the type label USNM 40161 is selected as lectotype. PresENT status: The taxonomy of the genus Phthirus is in a chaotic state. Ferris (1951, Mem. Pacific Coast Ent. Soc., vol. 1, p. 281) has pointed out that the description of this species merely demonstrates the occurrence of a species of Phthirus on the gorilla. Genus Polyplax Enderlein Polyplax alaskensis Ewing, Proc. Ent. Soc. Washington, vol. 29, No. 5, p. 118, 1927. Typr pata: USNM 40159. Holotype male from Microtus sp. from Alaska. PRESENT stratus: As originally described. Figures of the holotype of P. alaskensis have been published elsewhere (Scanlon and Johnson, Proc. Ent. Soc. Washington, vol. 59, p. 282, fig. 3, 1957). Polyplax dentaticornis Ewing, Proc. Biol. Soc. Washington, vol. 48, p. 207, fig- c, 1935. Typr pata: USNM 44905. Holotype male and a last nymph from the skin of Cricetulus andersoni [Cricetulus longicaudatus anderson] (USNM 172610), Shansi, China. The holotype male and last stage nymph are on separate slides and both bear the above data. PRESENT status: Until more specimens from the type host are examined, the status of this species must remain in doubt. In the holotype, most of the paratergal plate setae are missing, and the third pair of legs is missing. The shape of the thoracic sternal plate and the fact that one seta on paratergal plate four is longer than the plate suggest that P. dentaticornis is closely related to P. serrata (Burmeister) and might be an aberrant specimen of this species. It should be noted that Ewing’s figure of the third antennal segment of the holotype (loc. cit., p. 203, fig. c) pictures the apical spine as a lobe rather than a spe. This mistake was probably due to the fact that the entire head is flattened, changing normal relationships of the various parts. TYPE SPECIMENS OF LICE IN USNM—JOHNSON 47 Eremophthirius eropepli Ewing, Proc. Biol. Soc. Washington, vol. 48, p. 209, fig. B, 1935. Tyre pata: USNM 44906. Two male and two female cotypes from Eropeplus canus (USNM 219711), Celebes. The four syntype specimens are mounted in a horizontal line on a single slide, and the label bears the data given above. All the speci- mens are in poor condition; however, one of the two males has most of the leg segments and the majority of the paratergal plate setae are still present. I select as lectotype this male (second from the right on the slide) and have circled it with a diamond poimt pencil. The third antennal segment of the lectotype was figured by Ewing (fig. B). Ferris (Mem. Pacific Coast Ent. Soc. vol. 1, p. 207, 1951) pre- sumed the host to be Eriopeplus incanus, which he states is now known as Cricetulus longicaudatus, and ememded the species name of eropepli to eriopepli. However, Ewing’s description properly lists the host (USNM 219711) as Hropeplus canus. Present status: Polyplax eropepli (Ewing), 1935. This species would key to couplet 19 of Ferris’ (Mem. Pacific Coast Ent. Soc., vol. 1, p. 204, 1951) key to Polyplaxy species. A redescription and figures of eropepli will be published elsewhere. Polyplax tarsomydis Ewing, Proc. Biol. Soc. Washington, vol. 48, p. 206, fig. b, 1935. Types pata: USNM 44904. Holotype male from Tarsomys apoen- sis, Mindan4o, Philippine Islands. Present status: As originally described. Genus Proenderleinellus Ewing Proenderleinellus africanus Ewing, Journ. Washington Acad. Sci., vol. 13, No. 8, p. 148, 1923. Typa pata: USNM 23760. Holotype male from Thryonomys gregor pusillus [Thryonomys gregorianus pusillus] (USNM_ 184180), taken at Maji-ya-Chumvi, British East Africa. PRESENT status: A junior synonym of Proenderleinellus calvus (Waterston), 1917. Genus Prolinognathus Ewing Prolinognathus ferrisi Fahrenholz, Ztschr. Parasitenk., vol. 11, No. 1, p. 12, 1939. Prolinognathus leptocephalus (Ehrenberg), Ferris, Stanford Univ. Publ. Biol. Sci., vol. 2, No. 5, p. 142, figs. 2508, 251c (err. det.), 1932. Typr pata: USNM 184247. Holotype female from Procama brucer rudolfi [Heterohyraz syriacus rudolfi|, British East Africa. Fahrenholz used Ferris’ (1932) figures and description of ‘“‘lepto- cephalus” as a basis for his name ferrisi, and did not see the specimens. 48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ferris (Mem. Pacific Coast Ent. Soc., vol. 1, p. 251, 1951) accepted the type of ferrist as being the female from Procavia brucei rudolfi, and stated that this specimen should be in the U. S. National Museum. There is in the collection a female with the following data on the label in Ferris’ handwriting: ‘Ferris Col. 474, Prolinognathus lepto- cephalus (Eh.), from Procavia brucei rudolfi, Marsabit Road, B. E. Africa, U.S. N. M. 184247.” This specimen is the holotype of Pro- linognathus ferrisi Fahrenholz and has been so labeled. PRESENT status: As originally described. Genus Scipio Cummings Scipio longiceps Ewing, Proc. Helm. Soc. Washington, vol. 4, p. 81, fig. 29, 1937. Typz pata: USNM 49919. Holotype male from Thryonomys gregor pusillus [Thryonomys gregorianus pusillus] (USNM 184180), British East Africa, Maji-ya-Chumvi. PRESENT status: A junior synonym of Scipio aulacodi (Neumann), 1911, New synonymy. Ferris (Mem. Pacific Coast Ent. Soc., vol. 1, p. 154, 1951) expressed the opinion that S. longiceps Ewing was probably a synonym of Scipio aulacodi (Neumann). The holotype male of S. longiceps and a female with the same data as the holotype, and a second male from the same locality and host, differ in no way from Ferris’ (Stanford Univ. Publ. Biol. Sci., vol. 2, No. 3, p. 170, figs. 113, 1144, B, c, 1922) description and figures of aulacodi. Ewing’s supposed differences were due partly to the fact that the abdomen of longiceps holotype is telescoped and therefore smaller in relation to the length of the legs. The head is not longer than in Ferris’ drawings, nor is the shape of the tarsal claws different. In Ewing’s specimen the claws are more extended than in Ferris’ drawings, leading to his mistaken impression. Geographically, the distribution of aulacodi is such that one would expect to find it in the type locality of longiceps. (Maji-ya-Chumvi is in southeast Kenya.) Genus Solenopotes Enderlein Linognathus panamensis Ewing, Proc. Ent. Soc. Washington, vol. 29, No. 5, p. 119, 1927. Typrre pata: USNM 40160. ‘Described from a few females which are a part of a lot of six specimens,” from Odocoileus chiriquensis [Odo- coileus virginianus chiriquensis] (origin, Panama), which died at National Zoological Park on Jan. 28, 1925 (USNM 240843). There is in the collection a slide with six females, all of which are accepted as being syntypes, since there is no way of knowing which “few’’ females of the lot of six Ewing referred to in his original de- scription. These syntype females are all the same species. ne vi tary Re Lo direkt iW a sik st ne fis weap i“ me oui pwr iH Sie Aine a fain. a i Pak oy ea Pasines (oily AY ane la ie | “ ah, uy [wating wy Hi (ee Aj Wey , tay oe pio Vy Ae ae si Vin aes ‘gintdont a meee a roa agi (G8 of ray sine ig _ Ae “qs el ugha Be | Gt ph SO SE | i ai tigenha, mg bs 9 a mn trian visio | a Uh LOB i ad Pe aah. abiauutt : ie AP DERG Se wn oes yu it is) i ut , ) ney) a j Thi , Ub ccer}er yahline ' t : 7 us Bs , sad 1 |, eters g ue h i i. ae 4 rv ae ~ i PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM issued #4 SMITHSONIAN INSTITUTION U, S. NATIONAL MUSEUM Vol. 108 Washington : 1958 No. 3394 A NEW NORTH AMERICAN BIRD-FLEA By F. G. A. M. Suir! Dr. Phyllis T. Johnson, of the Entomology Research Division, U.S. Department of Agriculture, Washington D. C., most kindly invited me to describe a new bird-flea, material of which was found in the collections of the U.S. National Museum. The specimens of the new flea were collected from the nest of a barn swallow and are part of a series which constitutes the first record of fleas from the North Amer- ican subspecies of this host. It is a subspecies of Ceratophyllus affinis Nordberg, 1935, and the host record is a valuable indication as regards the true host of the little-known nominate subspecies. I am much indebted to the authorities of the U.S. National Museum for the donation of one pair of paratypes of the new flea to the British Museum collection of fleas at Tring. Ceratophyllus affinis neglectus, new subspecies Figures 1, 2, 5-10 TYPE MATERIAL: Male holotype, female allotype, and 207, 2 2 para- types from Smithfield, northern Utah, U. S. A., from nest of barn swallow Hirundo rustica erythrogastra, June 6, 1951, collected by 1 British Museum (Natural History), The Zoological Museum, Tring, Herts. 453664—58 51 52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 T. Tibbetts.? Holotype, allotype and two paratypes (167, 19) in the collection of the U.S. National Museum, and two paratypes (1 o, 1 9) in the British Museum collection of fleas at Tring. Draanosis: The new subspecies differs from all known representa- tives of the genus Ceratophyllus by the great development of the post- spiracular area of the spiracular fossa on terga 1—vui (figs. 1, 2); in the nominate subspecies the postspiracular areas are somewhat less i Ficures 1-4.—Spiracular fossa of tergum vii of: 1, Ceratophyllus affinis neglectus, new sub- species, male, holotype; 2, C. a. neglectus, female allotype; 3, Ceratophyllus niger C. Fox, male, from Essington, British Columbia; 4, C. niger, female, from Essington, British Columbia. 3 a 2 Cross and Knowlton (1953) recorded material with virtually identical data, but determined as Cerato- phyllus garei Roths., “collected from nests of the barn swallow, Jdiruwndo erythrogaster, by T. Tibbets and L. Dale Haws at Smithfield, Utah, June 7, 1951.’’ Although C. garei, a flea of damp birds’ nests, is known to occur in Utah, it is very likely that the specimens recorded by Cross and Knowlton will also prove to be representatives of C. affinis neglectus. One pair of the USNM specimens was misdetermined as Cerato- phyllus niger, while the other two pairs had not been determined. NEW NORTH AMERICAN BIRD-FLEA—SMIT 53 well developed, though still very conspicuous and larger than in any other species of the genus. The new subspecies differs from the nominate form in the male by the longer movable process of the clasper (fig. 5) and the much more weakly developed spiculose area of tergum vi (fig. 7); in the female by the slightly less prominent lateral lobe of sternum vir (figs. 9, 10) and a somewhat zigzag-shaped sclerotized portion of the ductus obturatus (if this latter characteristic proves to be constant, which seems doubtful). For figures of pertinent struc- Figures 5-8.—Ceratophyllus affinis neglectus, new subspecies, male: 5, clasper, sternum 1x and sternum vu, holotype; 6, aedeagal crochet, holotype; 7, dorsal part of tergum vit, holotype; 8, processes of clasper, paratype. 54 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 tures of C. a. affinis see Darskaya (1950) and Smit (1956). The polytypic species C. affinis appears to be related to Ceratophyllus idius Jordan and Rothschild and, more distantly, to C. niger C. Fox. Description: Head with a moderately developed frontal tubercle; preantennal region with a frontal row of six or seven setae in the male, in the female the frontal row is vestigial and consists of only a few minute setae; ocular row with three setae in both sexes; postantennal region of head with three or four setae above the antennal fossa and a posterior row of five or six setae each side. The long setae of the antennal pedicel reach to about the seventh segment of the club in the male, and well beyond the apex of the club in the female. Labial palp not quite reaching the apex of the fore coxa. Pronotum with a ctenidium of 31-33 spines, the most dorsal of which are about as long as the pronotum anterior to the upper spines (as in C. idius; in C. niger the pronotum is dorsally much longer than the pronotal spines). Mesepimeron with one or two setae on the middle of the surface though fairly close to the mesopleural rod, apart from the few ventral and posterior setae (these median setae are usually also present in C. idius, but absent in C. niger). Pleural arch of metathorax well developed; squamulum of metasternum short and broad. Fifth tarsal segment of all legs with numerous minute setae scattered over the plantar surface. Terga I-v in the male with the following numbers of marginal spinelets per side: 2, 3 (4), 2 or 3, 2, 1 (2); in the female: 2 or 3, 3 or 4, 2 (3), 1 or 2, and 0 respectively. Spiracular fossae of terga u—vir usually with large postspiracular areas, the largest for any species in the genus (figs. 1, 2; ef. figs. 3, 4), although there is considerable variation in the size of these areas on individual segments. In other respects the structure and chaetotaxy of the head, thorax, legs, and unmodified abdominal segments are as in related species. Male (figs. 5-8): Tergum vin (fig. 7) with a weakly developed and rather narrow spiculose area, a row of 9-13 strong setae along the dorsal margin and two or three setae on the lateral surface. Sternum vul (fig. 5) long and narrow, distinctly curved upwards, with a triangular basal portion, two slender apical setae, and a long and nar- row apical membranous process which bears a posteriorly directed, rather narrow, spiculose basal lobe, as in the nominate subspecies. Manubrium of clasper fairly broad basally; fixed process almost quad- rangular, with a broadly rounded apex; the two acetabular setae are situated not far below the middle of the posterior margin of the clasper (fig. 5). Movable process (figs. 5, 8) elongate, its smoothly convex posterior margin bearing several slender setae along the upper half, while the anterior margin is angulate at about the middle. Apical lobe of the distal arm of sternum rx (fig. 5) narrowing very NEW NORTH AMERICAN BIRD-FLEA—SMIT 55 little towards the truncate apex; chaetotaxy of this arm as shown in figure 5. Aedeagal crochet with a fairly long, narrow, and blunt tipped apical portion (fig. 6). Female (figs. 9, 10): This sex resembles that of C. a. affinis very closely and the main differences, apart from the average size of the postspiracular areas, are that in the new subspecies the lateral lobe of sternum vir projects to a lesser degree and the margin below the lobe is almost straight (figs. 9, 10), not markedly concave as in C. a. Ja Ficures 9, 10.—Ceratophyllus affinis neglectus, new subspecies, female: 9, sternum vm and spermatheca, allotype; 10, sternum vi and spermatheca, paratype. affinis, and the heavily sclerotized basal part of the ductus obturatus is somewhat zigzag-shaped in the three females studied (this may be an abnormality), while this duct is smooth (as is normal) in C. a. afinis. The length of the cylindrical bulga of the spermatheca (figs. 9, 10) appears to be fairly variable, but the shorter of the two drawn (fig. 10) is like that of the nominate subspecies and is therefore presumably the normal type. Length: Male, 3 mm; female, 3.5 mim. Remarks: We know as yet extremely little about the range of distribution or true host of C. a. affinis; the specimens (207, 49) from southwest Finland are from an unknown host (Smit, 1956), but it is most significant that the specimens (11, 209) recorded by Darskaya (1950) from the Vologda oblast (roughly 450 km. north of Moscow) came from the nest of Hirundo rustica [rustica], for C. a. 56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 neglectus was collected from the nest of another subspecies of the Holarctic H. rustica, namely H. r. erythrogasira. Both subspecies of barn swallow build similar nests in similar places, and it may well be assumed now that H. rustica is the true host of C. a. affinis. The differences in structure between these two forms of barn swallow fleas are fairly pronounced, and some workers would perhaps consider them to be full species. However, the differences between, e. g., Ceratophyllus styx siyx Rothschild and C. s. jordani Smit, which are certainly subspecies and are parasites of the sand martin Riparia riparia in Europe, are even more pronounced than those between the two subspecies of C. afinis. Both barn swallow and sand martin build very isolated nests and they live in a considerable degree of isolation from other hosts. It is an acknowledged fact that isolation favors evolution and this may explain why the differences between subspecies of fleas which live in identical isolated ecological habitats (e. g., the fleas of the swallow and the sand martin) are greater than those between subspecies of fleas infesting hosts which occupy less isolated ecological habitats (e. g., the two subspecies of the tree- squirrel flea Tarsopsylla octodecimdentata, T. 0. octodecimdentata (Kolenati) in the Palaearctic region, and T. o. coloradensis (Baker) in the Nearctic region). Thus the degree of differences between C. a. affinis and C. a. neglectus might be interpreted as being the result of the occurrence of these fleas in isolated habitats and therefore strengthens the assumption that C. a. affinis, as well as C. a. neglectus, is a parasite of Hirundo rustica. If the true host (and the ecological factors concerning its nest) of one of the subspecies should prove to be different, it would be necessary to regard affinis and neglectus as full species, for, apart from considerations of geographical, morpho- logical, and reproductive isolation, it seems that in fleas we must usually also stipulate that subspecies of a certain species must occupy almost identical ecological niches, since it seems extremely probable that if two fleas have diverged so much as to be able to occupy sub- stantially different niches they will also have become too distant to have retained the capacity to interbreed without impairment of fertility. Thus, though the structural differences between Xeno- psylla ramesis (Rothschild) (a parasite of gerbils) and X. cunicularis Smit (infesting the rabbit in Morocco) are very small, and the two forms appear not to be sympatric, I regard them as full species be- cause of the very different nesting habits of their hosts. It may be that another form, more or less intermediate between C. a. affints and C. a. neglectus, occurs in eastern Siberia. Since barn swallows are widely distributed and travel over great distances, it seems likely that the distribution of the subspecies of C. affinis will NEW NORTH AMERICAN BIRD-FLEA—SMIT 57 also be found to cover a large area, and it is to be hoped that collectors will turn their attention to the usually easily collected nests of barn swallows. In Europe the nests of H. rustica, which are now usually built inside barns and other outbuildings, are obviously too dry to form a suitable environment for the development of flea larvae; the same may apply to the nests of the North American subspecies. Ceratophyllus affinis must have become associated with the barn swallow long before this bird adopted man-erected shelters for nesting sites, and it seems possible that it has not been able to adapt itself to this relatively new macro-habitat, that it survives mainly in the rela- tively few nests which are still built in natural sites (e. g., in roofs of caves, in little rock-pockets in wild ravines, in sea-washed caverns) and that from these natural sites occasional specimens are introduced to nests in barns. ‘This is a rather speculative suggestion and it is only by collecting fleas from large numbers of barn swallow nests from natural and man-made sites, both in the Nearctic and Palaearctic regions, that we shall learn something about this intriguing question. After the above was written I looked through those tubes in the alcoholic portion of the Tring collection which were labeled as having been collected from Hirundo rustica, and to my delight found in a tube containing four males and ten females of Ceratophyllus gallinae (Schrank) and one male of C. garet Rothschild (the occurrence of all of which must be regarded as casual) a single female of C. a. affinis Nordberg. The fleas in the tube in question were collected from Hirundo rustica (presumably from a nest) at Wulfsdorf, 7 km. south of Liibeck, Germany, in 1923 by W. Blohm, and the record gives some support to my assumptions about the host and distribution of the flea. Unfortunately the Wulfsdorf specimen is somewhat abnormal (e. g., it has only two antesensilial setae each side instead of three, several setae of one midtibia are malformed, and the anal tergum seems to be shortened), so I do not feel justified in figuring any part of it, but its sternum vii and spermatheca agree perfectly with those of Finnish specimens and with the figure published by Darskaya. References Cross, E. A., anD Knowuton, G. F. 1953. Fleas on flying squirrel and barn swallow. Bull. Brooklyn Ent. Soce., vol. 48, p. 73. Darsxaya, N. F. 1950. Key to bird fleas of the genus Ceratophyllus. Materialy k Poznaniyu Fauny i Flory SSSR, new ser., Otdiél Zoologicheskil, No. 15, Ektoparazity, No. 2, pp. 85-105, figs. 1-19. Smit, F. G. A. M. 1956. Redescriptions of fleas described by Nordberg in 1935. Opusc. Ent., vol. 21, pp. 1382-146, figs. 1-17. U. S. 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OS 18 2 tn walk Pamgeht) heal Ce Be b lod f [ ats if ti vache ont aa rival dio bky vat ed) tages Al OY atch a We). 16 Mote ifn moviaye.odt). btice eahaly o. @igaiat aliyirie, s peo" f stoallon. ater e ef doe ar YA rohetley ey, ta ( PCChCey Rett bynes aetoanih bia wali AY, ‘ed. lo sont slintei by: baat) old Ty ahs STEOs nla ti by) ena aig ensinage. | ' Bi p, ae eels te bh. hagigrth shia diag oe 108: i aoe baa eee anirgyted fhe ont | Os tra see, one siphtin 90 ig, than yA eieaken eg feats ais, bag ator ob I Lom | ieg bs exoult cpbve uhiooteag OOTY, i ca digces ie ho aay se 1 gg exaCl, edd bin le ih ish Orit wait, nid ese 40 iT ; ree at Phe : =i Qaonin rclail. ‘ bata i Laie a 4 _ ei vidnre 00d) Jikan f joott pli malta aah hy vip i ify fu Sal Ree . WAS f "i ylmokeT ty late ‘lg a usm yy el ov thdan Mfdsiyoloos ‘aba i.e a NA ‘0 Ss 1 sey OR PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM issued {6 ( SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington: 1958 No. 3395 A REVIEW OF SOME GALERUCINE BEETLES WITH EXCISED MIDDLE TIBIAE IN THE MALE By Doris H. BLAKE Unlike the Alticini, many of the Galerucini do not have outstanding characters. This is particularly true of such genera as Drabrotica, Neobrotica, and Luperodes, and has led to many really diverse beetles being included under these generic names. Yet any attempt to create a better classification is bound to result in a somewhat artificial arrangement because these beetle groups overlap in certain peculiari- ties and in other ways possess traits common to most Galerucini. There is one approach that can be used, which Horn (1893, p. 124) suggests when he remarks concerning Phyllecthrus: ‘Too many genera of Galerucini have been described from uniques without any pub- lished reference to sexual peculiarities which are often a guide to rela- tionships when other characters cause doubt by their double indication.” There is a somewhat miscellaneous series of Galerucine beetles in which the male character is a cut-out area on the inner margin near the apex of the middle tibiae similar to the emargination in the front tibiae of Carabidae. Some of these beetles have been allocated to that all-embracing genus, Diabrotica, in spite of not having bifid claws, others to Neobrotica, others to Phyllecthrus and Oroetes, and I 59 60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 have described under the name Ectmesopus a group of West Indian beetles—two of which had been described as species of Luperus—that have this character in the male. In the case of the majority of these beetles not only is there the male character of the cut-out middle tibiae, but also there is an abnormality of the antennal joints. All over the world there are Galerucini with queer male antennae, but the combination of these leg and antennal abnormalities in the male serves to make these particular beetles stand out from the bulk of the Galerucini closely resembling them in the female sex, in which the males and females are essentially identical. This paper is an attempt to bring together these beetles and consider their likenesses as well as differences. In describing the genus Phyllecthrus, LeConte (1865) states that he took the name from the Dejean (1837, p. 406) catalog. He used Dejean’s form, Phyllecthris, in his key in the first mention of the name, but two pages later, in his description of the genus, he used Phyllec- thrus, and subsequently described other species under that spelling, and that form of the name has persisted. LeConte listed the old Olivier species, Galleruca dorsalis Olivier (1808), given under Phyl- lecthris in the Dejean catalog as Phyllecthrus dorsalis, and added Say’s Galleruca atriventris, now regarded as a color phase of dorsalis; he also described a smaller species, P. gentilis. In 1868 he added another species, Phyllecthrus nigripennis, now regarded as a color form of gentilis. In 1884 he described Phyllecthrus texanus, which is closely related to gentilis and considered by Horn as a variety of it. All of these species that LeConte described, as well as the original dorsalis of Olivier and Say’s atriventris, have one point in common—the antennae in the male are 10-jointed. In 1891 Jacoby described a new genus Luperosoma for an Ecua- dorean species, Luperosoma marginatum. This had thickened an- tennal joints of the normal number (11) in the male. It resembled LeConte’s species in the cut-out middle tibiae of the male. Later, Weise (1924) in the Junk catalog, synonymized it with Phyllecthrus. Meanwhile, Horn (1893, 1896) described three species under Phyl- lecthrus, parallelus, schwarzi, and subsulcatus, all with thickened antennal joints, 11 in number, and cut out middle tibiae in the male, as in Jacoby’s genus, Luperosoma. In 1940 I described a group of tiny West Indian beetles, all very closely related to each other, with cut out middle tibiae in the male, as Ectmesopus. All but two of these species had abnormal antennal joints in the male, sometimes only | thickened, but usually deformed in some way. One of the two that did not have the abnormal antennal joints had a slight difference in the length of the joints in the sexes, and in the other there seemed to be no difference at all, yet both of these species in color and structure were — GALERUCINE BEETLES—BLAKE 61 very close to the rest of this homogeneous little group, so that the lack of these sex characters seemed only incidental and not at all com- parable with the marked differences between the species of LeConte’s genus Phyllecthrus and Jacoby’s Luperosoma. H.S. Barber, in fact, suggested to me putting Horn’s much larger species, subsulcatus, schwarzi, and parallelus, into my West Indian Eetmesopus. But not only is there the much smaller size of Ectmesopus species, but also the shape of the prothorax is unlike that of LeConte’s, Horn’s or Jacoby’s genera, in that the West Indian beetles have a smoothly convex prothorax without any hint of transverse depression. They form a distinct little group of beetles probably endemic in the West Indies, and it seems best to keep them separate. As above indicated, Horn’s species fit better into Jacoby’s genus Luperosoma. In dealing with the Galerucini it has been my experience that in the sex characters of the males there is an infinite variety of forms, many so unusual as to merit generic recognition. Note the monotypic genus Oroetes described by Jacoby. In common with Phyllecthrus and Ectmesopus, Oroetes flavicollis has notched middle tibiae in the male, but the male antennae somewhat resemble those of Cerotoma in being cut out, but in a different way from Cerotoma. Like Cerotoma, too, is the excavation of the face. This excavation of the face is also found in certain beetles at present ascribed to the genera Neobrotica and Eucerotoma as well as in species of the Asiatic and African genus Palpozena. The prothorax in Oroetes in addition has a peculiar median tubercle in the male. Another peculiar beetle is Diabrotica cyanospila Suffrian, which has both enlarged antennal joints and cut out middle tibiae in the male, and does not have bifid claws, the one character that always distin- guishes the Diabrotica group. There is still another male character in cyanospila that I have not found in any of the others with notched middle tibiae, and that is a greatly enlarged hind femur in the male, so that it would seem almost like an alticid. I have found one other instance of such enlargement of the hind femora in the male, that is in Leptoxzena eximia Baly, a monotype from the Andaman Islands. In this species also the male antennae are dilated. There are no notched middle tibiae, however, and the beetle itself has a quite different aspect. Suffrian describes cyanospila as an Altica-like beetle, although he does not mention the enlarged femora nor the dilated antennal joints, leading me to believe that he had examined no males. In his description of Diabrotica semicyanea, which follows the descrip- tion of D. cyanospila, he stated that one specimen had a quite abnormal build of the last joints of the antennae in that the last joint is spoon- shaped and the preceding broadened. He believed it was a male of semicyanea. He did not mention in either case the excised middle 62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 tibiae or enlarged femora. Although I have not seen any male speci- men of this species, I think it is not unlikely another of this group, closely related, as Suffrian stated, to D. cyanospila, and possibly also with enlarged hind femora. On searching through the list of species under section “P” in Gahan’s (1891a, p. 418) table of Diabrotica, which is composed of species that may be distinguished by the peculiar structure of the antennae of the male, I have found two that have excised middle tibiae in the male. One of these, D. amplicornis Baly, seems to be another of the Lwperosoma group in which Horn and Jacoby described species. In the other species, D. dilaticornis Baly, both the anterior and middle tibiae are widely emarginate, the emargination being unlike that of the group under consideration, but it has bifid claws, as in a true Diabrotica, and therefore cannot be considered one of the group of genera under discussion. Among species of that heterogeneous genus Neobrotica occur several that differ from the type of the genus, Neobrotica variabilis Jacoby, in being smaller, oblong in shape, and more slender and with excised middle tibiae in the male. They do not have any peculiarity of the male antennal joints, in this respect being unlike most of the rest of this group. These species are Neobrotica ruatanae Jacoby and a closely related and undescribed beetle, and Neobrotica sexplagiata (Jacoby), the latter related to Diabrotica nymphaea Jacoby. The name N. sexplagiata covers a number of closely related species. Besides these are others from South America, so far undescribed, that fall into this group. Jacoby has described still another genus with at least one species, Platymorpha smaragdipennis, having excised middle tibiae. In his other species, P. variegata, I am unable to detect any such character. In both, however, the male antennae are peculiar in that the third joint is very short and truncate. Chapuis (1875, p. 163) included Phyllecthrus with Phyllobrotica in his treatment of Phyllobroticides, and separated them in his key by the epipleural characters. He stated that in Phyllecthrus the epipleura are very narrow. LeConte rightly described the elytra as distinctly margined with the epipleura narrow and not extending to the tip. In Phyllobrotica the epipleura are either very narrow or wholly lacking. The genus Phyllobrotica, in spite of its worldwide range in the temperate Northern Hemisphere, is exceedingly homogeneous, the species all being closely related and rather uniform in markings. The chief points of likeness between the two genera lie in the semicircular depres- sion across the prothorax and the appendiculate claws. In at least one species, Phyllobrotica limbata (Fabricius), the male antennal joints are thickened, but the middle tibiae show no emargination, and there is no thickening of the anterior tibiae in the male or short first GALERUCINE BERTLES—BLAKE 63 a.Phyllobrotica limbata (Fab) (RY fei By z @ | ie Leptonesiotes cyanospilatSulfs fan) ie Phyllecthrus dorsalis Oliv. Ficure 1.—a, Phyllobrotica limbata (Fabricius); b, Diabrotica dilaticornis Baly; ¢, Phillecthrus texanus LeConte; d, Phyllecthrus gentilis LeConte; e, Leptonesiotes cyanospila (Suffrian); f, Phyllecthrus dorsalis (Olivier). 64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 c.).sexpla rakon ge P sop bivittat colle (Baly) \V cHo » d. l.parviplagiatum n.ssp.teapense 2. Oroctes flavicollis Jacoby Ficure 2.—a, Iceloceras sexplagiatum (Jacoby); 6, I. parviplagiatum, new species; ¢, ng sexplagiatum bivittaticolle (Baly); d, I. paroiplagtatum teapense, new subspecies; ¢, Oroetes flavicollis Jacoby. GALERUCINE BEETLES—BLAKE 65 t. l. verbesinae n.sp. d. |. flavipes asp, c. |. maximum “sp. Ficure 3.—a, Iceloceras latiplagiatum, new species; b, I. verbesinae, new species; ¢, I. maximum, new species; d, I. flavipes, new species. 66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 c L.marginatum Jacoby d. Platymorpha albiventris nsp. 2. Lenigrum n.sp h. L.amplicorne (Baly) Ficure 4.—a, Luperosoma schwarzi (Horn); b, L. parallelum (Horn); ¢, L. marginatum Jacoby; d, Platymorpha albiventris, new species; ¢, Luperosoma nigrum, new species; f, L. subsulcatum (Horn); g, Iceloceras nymphaea (Jacoby); hk, Luperosoma amplicorne (Baly). GALERUCINE BEETLES—BLAKE 67 & \.vittatum nop d. Lruatanae (Jac) «.|.biplagiatum nsp Ficure 5.—a, Iceloceras flavocyaneum, new species; g, I. otttatum, new species; ¢, Platymorpha smaragdipennis Jacoby; d, Iceloceras ruatanae (Jacoby); ¢, I. biplagiatum, new species. 453666—58——2 68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 bs E. pallidus Blake c.E.tristis BlaKe a.Ectmesopus crassicornis Blake d.E.vitticollis Blake 2.E. leonardorum nsp. }.E.zonatus Blake q-E.occipitalis Blake nn oy 4, E.malachoides Guffrian) i.Eongusticollts Blake # E.darlingtoni BlaKe K. E.longicornts Blake Ficure 6.—a, Ectmesopus crassicornis Blake; b, E. pallidus Blake; ¢, E. tristis Blake; d, E. vitticollis Blake; ¢, E. leonardorum, new species; f, E. zonatus Blake; g, E. occipitalis Blake; h, E. malachoides (Suffrian); i, E. angusticollis Blake; j, E. darlingtoni Blake; k, E. longicornis Blake. GALERUCINE BEETLES—BLAKE 69 tarsal joint as in the group with the excised middle tibiae. In short, Phyllobrotica is not much more closely related to Phyllecthrus than is Diabrotica. I wish to acknowledge my indebtedness to the U. S. National Museum and the Museum of Comparative Zoology for the material on which this study is based. Mr. G. E. Bryant of the British Museum (Natural History) has kindly made comparisons of types there and loaned me other specimens. Dr. Floyd Werner has sent me specimens from Arizona, and Mr. H. J. Grant has sent me notes on the Horn types. Dr. J. A. Wilcox has sent me a specimen with excised tibiae that he found in the Cornell University collection together with notes on other species, and called my attention to others in the Bowditch collection. Key to genera of Galerucini here treated 1. Elytra without epipleura or the epipleura so narrow as to be inconspicuous, middle tibiae not excised in male .... . . .. . . Phyllobrotica Elytra with epipleura. Middle tibiae in male petigedl: Sa ae ae a a ee ee 2. Two anterior pairs of tibiae in male shallowly and broadly emarginate. Distal four joints of antennae in male greatly enlarged; claws bifid. Diabrotica dilaticornis Baly Middle tibiae alone deeply and narrowly excised. Claws appendiculate. . 3 3. Antennae 10-jointed in male. . . . s «6s 6 ee ee ee caer s Antennae ll-jointedin male. .... A ee 4, Face excavated in male, third antennal juin faaeh Reformed a nodule in middie of pronetum . <5 27k 2 OA a Wh ok oe spre 10 2 Oroetes Hacein mate notextavated). tts)... st dhetie lk BNW A. «he 8 SSS WS Me 5. Hind femora in male greatly swollen ......... .. . Leptonesiotes Hind femora in male not swollen. ..........2..888088. 6 6. Disk of pronotum not depressed... . . . .. . . Eetmesopus Disk of pronotum with transverse Searerealar ide preation elie ob. Se ey cee 7. Antennae in male with the distal joints thickened . ... . . Luperosoma Antennae in male with the distal joints not thickened. . ........ 8 8. Third antennal joint in male very short, cup-shaped, truncate at apex. Platymorpha Third antennal joint in male usually much longer than second, not cup-shaped at all, or truncate at apex. . .......+.4+6-+.+. +... + Iceloceras Genus Phyllecthrus LeConte Phyllecthrus LeConte, Proc. Acad. Nat. Sci. Philadelphia, p. 207, 1865. The original generic description is as follows: Body elongate, glabrous and nearly smooth above. Head transversely impressed between the eyes, and with a short median impressed line; acutely carinate be- tween the antennae, which are very long, 2nd and 8rd joints together shorter than the 4th, nearly equal in size in the female, 2nd connate with the 38rd, and nearly obsolete in the male. Maxillary palpi stout, the last joint shorter than the preceding, slender, subsulcate, acute at tip. Prothorax quadrate, truncate at the apex with a lunate dorsal impression more or less distinct. Elytra with 70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 10% the lateral margin distinct, epipleurae very narrow, not extending to the tip. Anterior coxae conical, contiguous; legs slender, tibiae not sulcate externally, middle tibiae of the male incised at the extremity on the inner margin; ungues with a large angular basal dilation. Abdomen with five ventral segments nearly equal in length and alike in both sexes. I have adopted the generic name proposed in Dejean’s Catalogue for Gall. dorsalis Oliv. Typs or Genus: Phyllecthrus dorsalis (Olivier). Key to species of Phyllecthrus t-"’ Large;'from’5 to 6.5 mnt, mn Jensth i. 0. 7.2 os .. dersalis (Olivier) Small} from*3S!to’ 4 mma Tenet hits Me Is EAs Nan oi) erat gs 2. Prothorax with wide dark lateral markings. . .. . . . .gentilis LeConte Prothorax entirely pale or with at most faint brown lateral spotting. TEXAS so, o creche ec Nal eee ay ete lav ee ey eibte en hese ee LOXBELEES Ente OO NLS Phyllecthrus dorsalis (Olivier) FicureE 1,f Galleruca dorsalis Olivier, Entomologie, vol. 6, p. 646, 1808. Phyllecthris dorsalis Dejean, Catalogue des Coléoptéres, ed. 3, p. 406, 1837. Galleruca atriventris Say, partim, Journ. Acad. Nat. Sci. Philadelphia, vol. 3, pt. 2, p. 461, 1824. Phyllecthrus dorsalis LeConte, Proc. Acad. Nat. Sci. Philadelphia, p. 207, 1865. Between 5 and 6.5 mm. in length, elongate oblong oval, alutaceous but moderately shiny, finely punctate; head, prosternum, breast and femora yellowish brown, the last with a dark streak; prothorax with wide dark piceous lateral markings, the lateral margin and middle of disk yellow-brown; in some Kansas, Nebraska, and Missouri specimens the pronotum entirely pale; scutellum pale, elytra, tibiae and tarsi piceous. In male the antennae 10-jointed and middle tibiae notched. Head with interocular space half width of head, smoothly rounded over occiput, finely and sparsely punctate, frontal tubercles well marked, a short bulging carina between antennal sockets, lower front moderately long, with scattered punctures. Antennae in male 10- jointed, 2d joint alone short, rest long, black and hairy and extending well below middle of elytra, the terminal joint often reddish brown. Antennae in female 11-jointed, 2d and 3d joints short, with the 3d a little longer, but together not so long as 4th, remainder a littler shorter than 4th and slender, not so long or wide as in male and deep brown, not black. Prothorax wider than long with only slightly curved sides and a shallow depression in lower half, in some specimens faint; shiny, impunctate; dark piceous with pale lateral margin and a pale median area varying in size and shape from a central roundish spot to an elon- gate pale line; in some Nebraska, Kansas, and Missouri specimens the pronotum entirely pale. Scutellum pale. Elytra widened toward apex with well-marked humeri and short intrahumeral sulcus, dis- GALERUCINE BEETLES—BLAKE 71 tinctly alutaceous although shiny, and finely punctate, entirely dark piceous. Epipleura disappearing after middle. Body beneath pale with abdomen dark, femora pale except for dark streak and apex of middle and posterior legs, tibiae and tarsi piceous. Anterior coxal cavities open, middle tibiae of male notched near apex, tibiae with inconspicuous spine, claws appendiculate, front tibiae in male thick- ened and first tarsal joint swollen with tympanum-lke under- part. Length 5-6.5 mm.; width 2.3-3 mm. Type: Perhaps in Paris, described by Olivier from “Caroline.” Disrrisution: District of Columbia: Washington, Hubbard and Schwarz; Maryland: Glen Echo, J. C. Bridwell; Plummers Island, W. L. McAtee, F. Knab; Great Falls, F. Knab. Virginia: Falls Church, Wm. Middleton; Rosslyn, F. H. Chittenden, F. Knab, Glencarlyn, F. Knab, J. C. Bridwell. Pennsylvania: Fall collection. Illinois: Southern Hlinois; Monroe County, H. Soltau. West Vir- gima: White Sulphur Springs. North Carolina: Raleigh. South Carolina: Near Charleston (Olivier’s type probably collected here by Bosc). Georgia: Fall collection. Missouri: Crevecoeur Lake, F. Knab. Nebraska: West Point; Kenosha, Elliott. Kansas: Topeka, Popenoe; Ashton; Onaga, F. F. Crevecoeur. Remarks: In Illinois, Kansas, Nebraska, and Missouri occur speci- mens with the pronotum entirely pale. However in a series of four specimens from Crevecoeur Lake, St. Louis County, Mo., all collected June 7, two specimens have the pronotum entirely pale, two are typ- ically dark marked. In a series of seven from Onaga, Kansas, five have entirely pale pronotum, two are dark marked. Similarly, in a series from southern Illinois, while the majority are with an entirely pale pronotum, a few have typically dark-marked pronotum. Hence this pale coloration would seem merely a color phase. Phyllecthrus gentilis LeConte Figure 1,d Phyllecthrus gentilis LeConte, Proc. Acad. Nat. Sci. Philadelphia, p. 208, 1865. Phyllecthrus nigripennis LeConte, partim, Trans. Amer. Ent. Soc., vol. 2, p. 58, 1868. Phyllecthrus gentilis var. nigripennis, Horn, Trans. Amer. Ent. Soc., vol. 20, p. 127, 1893. About 3 mm. in length, elongate oblong oval, shining although finely alutaceous, pale yellow with dark antennae except the three basal joints, and with wide dark lateral markings on pronotum, the elytra dark except for the lateral margins and sutural edges; in some western specimens (nigripennis) the entire elytra dark; tip of abdomen dark; in male the antennae 10-jointed, and middle tibiae notched. Head smooth and shiny over occiput with punctures above the distinctly marked frontal tubercles, interantennal area narrow and i2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 flat, the labrum wide, lower front paler than upper. Antennae in male 10-jointed, basal three joints pale with dark edges, the re- mainder piceous, joints 3-5 long, rest gradually diminishing in length. In female, antennae 11-jointed, joints 2, 3 short, subequal, remainder longer and subequal. Prothorax a little wider than long with nearly straight sides, disk with a faint semicircular depression in basal half, sides widely piceous, margins, base and median space pale yellow, shiny, punctate on sides. Scutellum pale. Elytra with small humeri, elongate and narrow, alutaceous but shiny, piceous except for pale lateral and sutural margins; in some western specimens (nigripennis) elytra entirely dark. Body beneath, except tip of abdomen, and legs pale; middle tibiae of male notched near apex, and tibiae with fine spine, claws appendiculate. Length 3-4 mm., width 1.4-1.8 mm. Typr: A male in the LeConte collection, bearing a pale scarlet round locality label signifying the Southern States. There are two others with similarly colored labels, a male and female. The type of Phyllecthrus nigripennis is a male from “‘Ks.’’ Two others, bearing a similar locality label, are females (?). Distrisution: New York: C. V. Riley. Pennsylvania: Alleghany, Wickham. Maryland: Beltsville, W. L. McAtee; Plummers Island, W. Li. McAtee; Chesapeake Beach, H.S. Barber. District of Columbia: Washington, Hubbard and Schwarz on Pinus virginiana, W. L. McAtee; Rock Creek Park, Heidemann. Vorginia: Glencarlyn, F. Knab; Kanawha, on Robinia, A. D. Hopkins. Ohio: Cincinnati, H. Soltau. Illinois: Heyworth, Wolcott; Starved Rock State Park, F. G. Werner. Towa: Iowa City, Wickham. Missouri: St. Louis County, F. Knab.—Distribution of P. nigripennis: Nebraska: Fillmore. Towa: Lake Okoboji, Buchanan. Kansas: Onaga, F. F. Crevecoeur; Wickham; Riley County, F. Marlatt. Remarks: Horn reduced P. nigripennis LeConte to a variety of P.. gentilis. Comparison of the two types in the LeConte collection shows that the two are very much alike except that nigripennis is darker in coloring, having the elytra entirely dark, and the abdomen dark, the breast not so dark, the epipleura pale. The aedeagus of a dissected specimen from Kansas appears like that of gentilis. This darker coloring of the western specimens may be analogous to that of western specimens of dorsalis, Say’s atriventris. Horn gives a record of P. gentilis being taken on Robinia pseudacacia by Dr. Hamilton. Phyllecthrus texanus LeConte FIGuRE l,c Phyllecthrus teranus LeConte, Trans. Amer. Ent. Soc., vol. 12, p.28, 1884. Phyllecthrus gentilis var. texanus Horn, Trans. Amer. Ent. Soc., vol. 20, p. 58, 1893. About 3 mm. in length, elongate oblong, shining although the elytra are alutaceous, finely punctate, prothorax with a semicircular depres- GALERUCINE BEETLES—BLAKE 73 sion in basal half, in the male the antennae 10-jointed and the middle tibiae notched near apex; antennae dark, head, prothorax, undersur- face, legs (except tarsi), and margin of elytra pale yellow-brown, elytra otherwise piceous. Head entirely pale, smoothly rounded over occiput down to the frontal tubercles with an area over tubercles on each side of distinct, closely placed punctures, tubercles not swollen but distinctly marked, interantennal area narrow, not much produced, labrum wide. Anten- nae dark and in male 10-jointed, joints 3-10 long and subequal; antennae in female 11-jointed, the 3d joint short and equal to 2d in length. Prothorax almost as long as wide with sides only slightly curved, a semicircular depression in lower half, entirely pale yellow- brown except in two or three specimens a faint brownish spotting on each side; shining. Scutellum yellow-brown. LElytra smoothly rounded, without depression, shining although alutaceous and with fine, not dense punctures becoming invisible towards apex; piceous with pale yellow-brown explanate margin from apical curve on becom- ing piceous. Epipleura indistinct after middle. Body beneath pale yellow-brown, lightly pubescent; legs pale with tarsi deeper in color; middle tibiae of male notched near apex; tibiae with fine spine at apex; claws appendiculate. Length 3.3 mm.; width 0.9 mm. Typs: Female, in the LeConte collection, and four paratypes (two males, two females) collected by Belfrage in Texas. In the collection of the U.S. National Museum are 30 more specimens labelled ‘Texas, Belfrage, Eutopotypes” by H. 8. Barber, which are evidently from the same series as the LeConte specimens. Remarks: This is one of the species described by LeConte and published after his death by Horn who later synonymized it with P. gentilis, as a color form like P. nigripennis. It is probably a distinct species as the aedeagus is somewhat different and the prothorax has no distinct dark markings as in gentilis, and the elytral markings are unlike either gentilis or nigripennis. I have seen no other specimens besides those taken by Belfrage. Genus Oroetes Jacoby Oroetes Jacoby, Biologia Centrali-Americana, Coleopt., vol. 6, pt. 1, p. 600, 1888. The original generic description is as follows: Head broad, the penultimate joint of the palpi incrassate; antennae filiform, deformed in the male, the second and third joints short; thorax transversely subquadrate, the surface deeply depressed; elytra irregularly punctured, their epipleurae obsolete below the middle; the posterior tibiae with a short spine (this is present on all tibiae); the intermediate tibiae notched near the apex in the male; the first joint of the posterior tarsi rather longer than the following three joints united; claws appendiculate; the anterior coxal cavities open. This new genus, which should be placed near Luperus, is separated from any of the genera composing this section by the broad head and the depressed thorax, 74 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 and by the elytral epipleurae being obsolete below the middle; the male insect is further distinguished by the structure of the antennae, and by the intermediate tibiae being notched near the apex. Tyrer or GENUS: Oroetes flavicollis Jacoby. Oroetes flavicollis Jacoby FIGURE 2,e Oroetes flavicollis Jacoby, Biologia Centrali-Americana, Coleopt., vol. 6, pt. 1, p. 600, 1888. About 5 mm. in length, oblong oval, shining, the elytra finely punc- tate; pale reddish yellow, antennae with basal joints pale, rest piceous, elytra a violaceous blue except for a pale margin in basal half. In male the 3d antennal joint long and with a rounded excision, 4th joint flattened and curved, in female the 3d joint short; in male the face excavated and a spine in the middle of the lower front, the pro- notum with a median tubercle; middle tibiae notched near apex. Head in female smoothly rounded over occiput and down front, with little sign of frontal tubercles or interantennal carina, lower front flat, interocular space more than half its width; in male, the lower front excavated between antennae and below this a transverse ridge with a median spine, two flat side pieces extending from below antennal sockets to labrum. Antennae nearly as long as beetle, in female filiform, slender, pubescent, first two joints pale yellow, 3d, also pale, and 4th short and subequal, rest much longer and dark. In male the first four basal joints pale, the first joint swollen, 2d short, 3d long and flattened below and broadened with a cut out ex- cision near the base, 4th joint flattened below and a little swollen and long, remainder long and hairy and dark. Prothorax almost rec- tangular with sides only slightly curved, in male the anterior margin roundly produced in middle over occiput of head, in female straight across; in male, a median tubercle on anterior half with a fovea in front of it; in both sexes a semilunate transverse depression in basal half. Surface shining, impunctate, pale reddish yellow. Scutellum pale. Elytra shining violaceous with the anterior explanate margin pale, distinctly and moderately densely punctate. Epipleura becoming indistinct before apical curve. Body beneath and legs entirely pale, anterior coxal cavities open, tibiae with fine spine, middle tibiae of male notched near apex, first tarsal joint of anterior legs in male slender and cylindrical with tympanum-like covering underneath; in posterior legs first tarsal joint long, claws appendiculate. Length 5-5.4 mm.; width 2.4 mm. Tyrer: Probably in British Museum (Natural History), described from specimens from Chontales, Nicaragua, and Bugaba and David, Panama. GALERUCINE BEETLES—BLAKE 19 OTHER LOCALITIES: Escocia (Costa Rica?), Nevermann; Piedras Negras, Costa Rica, Schild and Burgdorf. Remarks: The extraordinary development of the antennae, head, and pronotum in the male bewilders a taxonomist trying to place this beetle in a genus. The excavation of the head and abnormal antennae are suggestive of Cerotoma, but the depressed pronotum, the notched middle tibiae in the male, and the Luperus-like shape place it near Phyllecthrus. It deserves generic standing. Jacoby describes the Nicaragua and Panama specimens as having four pale basal antennal joints in the male. In the specimens I have examined from Costa Rica, only three basal joints are pale in the male and only the first two in the female. Otherwise the description fits. Leptonesiotes, new genus Elongate oblong oval, glabrous and without conspicuous punctation, the head with well-marked frontal tubercles, with a short carina between the antennal sockets, jaws large; antennae 11-jointed, not extending much below the middle of the the elytra, in the male, joints 10 and 11 enlarged, joint 3 barely twice as long as joint 2 and a little shorter than joint 4, joints 3-9 subequal. Prothorax a little wider than long with slightly curved sides, somewhat narrowed towards base, disk convex, without depressions. Elytra a little broader behind, the humeri well developed with an intrahumeral sulcus. Epipleura extending to about the middle of the elytra; anterior coxal cavities open; in the male a thickening of the front tibia, the first joint of the front tarsi short and thick with the undersurface having a flat tympanum-like appearance; middle tibiae with a notch on the inner margin near apex; hind femora much enlarged but narrowed at apex, no apical spine discernible, posterior first tarsal joint longer than the rest together; claws appendiculate. Type oF GENus: Diabrotica cyanospila Suffrian. Remarks: The name is derived from the Greek “leptos,’’ delicate, and ‘“‘nesiotes,”’ islander. Leptonesiotes cyanospila (Suffrian) FIGURE l,e Diabrotica cyanospila Suffrian, Arch. Naturg., vol. 33, No. 3, p. 318, 1867. About 5.5 mm. in length, elongate oblong oval, shining, the elytra conspicuously punctate, yellow (? reddish in life) with dark head and dark humeral and apical spot on each elytron shining with a purplish or violaceous lustre, last two abdominal segments deep violaceous, the femora and tibiae also dark, tarsi reddish brown, antennae pale, and in the male with the two distal joints enlarged, in the male also the middle tibiae notched near the apex. 453666—58——-3 76 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 Head smooth and shining, with a shallow depression over the well- marked frontal tubercles, space between antennal sockets slightly produced in a short carina down front; labrum wide, jaws large, lower front deep reddish brown, upper part of head dark with a violaceous lustre. Antennae entirely pale, not extending to middle of elytra, 3d joint shorter than 4th, joints 3-9 subequal, the two distal joints in the male enlarged and somewhat distorted. Prothorax a little wider than long, with slightly curved sides and oblique basal angles, moderately convex, the disk without depressions, shiny yellowish. Scutellum pale. Elytra shiny, pale with dark violaceous spot covering humerus and extending almost to scutellum, another larger dark spot below middle extending from the margin nearly to the suture, but not reach- ing apex. Body beneath pale except the last two ventral segments, which are deep violaceous. Femora and tibiae dark, the tarsi reddish brown. Middle tibiae of male with notch near apex, no spurs dis- cernible on tibiae. Claws appendiculate. Length 5.5 mm.; width 2.4 mm. Tyrer: Perhaps in Gundlach collection in Havana, Cuba. Collected in Cuba. OrHerR Localities: Cuba: Havana, Baker; Taco Taco, S. C. Bruner, A. Acufia, C. H. Ballou. Remarks: I know of no other galerucid beetle that has the enlarged hind femora, notched middle tibiae, and antennal enlargement in the male. Leptoxena eximia Baly from the Andaman Islands, also a galerucid beetle, has similar enlargement of the hind femora and dilated antennal joints in the three distal joints, but not the notched middle tibia. It is an entirely different looking beetle and in no way related to the present one. Iceloceras, new genus Elongate oblong oval, glabrous, confusedly punctate, antennae filiform, not enlarged in male, the 3d joint sometimes subequal to 2d, sometimes twice as long; prothorax subquadrate, wider than long, with slightly curved sides; a semicircular depression in lower half. Elytra little widened apically, epipleura vanishing at apical curve; anterior coxal cavities open; anterior front tibiae in male swollen and first tarsal joint short and wide with undersurface having tympanum- like covering; middle tibiae of male notched near apex; middle and hind tibiae with small spur; first tarsal joint of posterior legs long, nearly equalling the rest of joints together; claws appendiculate. TyPsE oF GENUS: Diabrotica sexplagiata Jacoby. The type of the genus Neobrotica, as has been so designated by Weise, is NV. variabilis Jacoby. The several species that have been confused under the name of N. sexplagiata (Jacoby) bear little GALERUCINE BEETLES—BLAKE Td resemblance to N. variabilis. They are differently shaped, being oblong instead of widened at the apical end. They have longer heads and smaller eyes. The sex character of the excised middle tibiae in the male, found in them, is not found in variabilis. In short, they fit better into the group with notched middle tibiae than into that mis- cellany of species called Neobrotica, although there is no antennal difference between the sexes. Besides sexplagiata are other species, one of which has been also described under Neobrotica, N. ruatanae Jacoby from Honduras. There is another undescribed species from Mexico closely related to ruatanae. In both of these the middle tibiae of the male are notched and the antennae not enlarged in the male. But they differ from sexplagiata in the length of the 3d antennal joint, which is short and subequal to the 2d, whereas in sexplagiata it is long. In this regard these two form a connecting link with the genus Luperosoma. Besides these is another undescribed species from South America that is more closely related to sexplagiata. All of these species are a little different from the other groups having for the male character the notched middle tibiae in that they do not have any enlarged or otherwise deformed antennal joints in the male; conse- quently, the genus Iceloceras, is here erected for them. The name is from the Greek “‘ikelos,” like, and ‘‘keras,”’ horn. Key to species of Iceloceras 1. Third antennal joint short . .... bits PST PROG ec Aieh-< Renee Third antennal joint fully twice as long as Tsecond: ar wee fa ie: Rte 2. Elytra with a wide dark sutural marking curving down and w denne to cover apex of elytra except margin ........ ... . Fruatanae (Jacoby) Elytra with only apical third dark ... . . . flavocyaneum, new species ee VETS MAGUH GOs ASR te BS a ke alin BS a a 2 et Sn Seed, ia Elytra not vittate. .... Hd eee tet 4. Elytra with a sutural and a leper ite mot feed x ence vittatum, new species Elytra with a wide median dark vitta. . . . . . nymphaea (Jacoby) 5. Elytra piceous with a single pale spot in the middle of each elytron. biplagiatum, new species Elytra piceous with three pale spots on each, sometimes confluent . 6. Legs entirely pale . ‘ Tibiae, tarsi and usually femor ae ee dari SS Sy te tal Swors 7. Orange-brown with dark brown or piceous markings. South America. maximum, new species Pale yellow with piceous markings .. . et IASI ES 8. Pronotum nearly impunctate, a dark spot on either side ide extending below middle. Costa Rica... . . . . . verbesinae, new species Pronotum distinctly and ihe dessely punters. the dark spot on either side extending nearly entire length. Panama . . . flavipes, new species 9. Femora entirely pale, tibiae and tarsi dark, the large darle spot on each side of pronotum extending nearly to middle. Panama. latiplagiatum, new species Femora with a dark streak at apex, tibiae and tarsidark. . ..... 410 orn 78 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 10. Pronotum with vittate piceous mark on each side. Panama, South America. sexplagiatum (Jacoby) Pronotum: witha spot.on eachside: oy... Gs ss Vsgiekiola vs. logue 11 11. Large, 5-6 mm., an inconspicuous dark spot on each side of pronotum, some- times lacking. Guatemala, Nicaragua, Honduras, Costa Rica. parviplagiatum, new species Smaller, 4-5 mm., a dark spot on anterior sides of pronotum usually con- spicuous,, . Mexico! (te cu ecu parviplagiatum teapense, new subspecies Iceloceras sexplagiatum (Jacoby) Ficure 2,4 Diabrotica sexplagiata Jacoby, Proc. Zool. Soc. London (1877), p. 151, 1878; Biologia Centrali-Americana, Coleopt., vol. 6, p. 527, 1887. Neobrotica sexplagiata (Jacoby), Gahan, Trans. Ent. Soc. London (1891), p. 524, 1891.—Jacoby, Biologia Centrali-Americana, Coleopt., vol. 6, Suppl. p. 331, 1892. Between 3.5 and 5 mm. in length, elongate oblong oval, the elytra densely and moderately coarsely punctate, shining, pale yellow with brown to piceous antennae and legs with a dark streak on the upper femora, dark tibiae and tarsi, and on the pronotum a dark vitta on each side, and the elytra with six yellow spots and a pale margin, the rest piceous black; middle tibiae of the male with a small notch near apex. Head with interocular space a little more than half width of head; deep orange on occiput, pale yellow on lower front, smoothly rounded with fine punctures on occiput and a fine median line extending down lower front between frontal tubercles and a thin carina down lower front. Antennae entirely dark, 3d and 5th joints a little shorter than 4th. Prothorax about a third wider than long, sides slightly curved and a semicircular depression on lower half of disk; shining although finely alutaceous and finely punctate; pale yellow with a lateral piceous vitta on each side extending from anterior margin nearly to basal margin. Scutellum pale. Elytra dark piceous with three pale yellow spots on each and a pale margin; surface densely and coarsely punctate, shining. Epipleura extending to apical curve; in the female the last abdominal segment dorsally conspicuously black-edged, and the one before with a dark spot on each side. Body beneath entirely pale, with pale pubescence; femora pale with a dark streak above, posterior tibiae and tarsi dark, the notching on the middle tibiae of the male very small. Length 3.5—-5 mm.; width 1.5—-2 mm. Tyr: Probably in the British Museum (Natural History). Distripution: Panama: Bella Vista, Nathan Banks, Aug. 8, 1924; Paraiso, C. Z., E. A. Schwarz, A. H. Jennings; Porto Bello, August Busck; Summit, N. L. H. Krauss; Miraflores, A. H. Jennings; Tobago Island, H. F. Dietz; Pueblo Nuevo, Papaya plantation, H. Morrison; GALERUCINE BEETLES—BLAKE 79 El Cermino, J. Zetek; Pedro Miguel, N. L. H. Krauss; Volcan de Chiri- qui, 2500-4000 ft., Champion. Remarks: In 1878 Jacoby described as Diabrotica sexplagiata a beetle whose habitat he gave as Peru and Panama. In 1887 he stated that the Peruvian specimens were a different species from the Panama ones and used the name sexplagiata for the Panama species. Gahan in 1891 called attention to the fact that the claws of D. sexplagiata were appendiculate and referred the species to the genus Neobrotica. ‘This emendation Jacoby (1892) accepted in his Supplement to the Biologia. Meanwhile, Baly had described as D. bivittaticollis a similar form from Magdalena River, Colombia, which Jacoby (1892) referred to synonymy under his Panama species. In the material examined from Panama that I have tried to match with Jacoby’s description of sexplagiata, | have found four distinct species. There are many of Jacoby’s specimens in the Bowditch collection of the Museum of Comparative Zoology, and Mr. G. E. Bryant has kindly sent me other material from the British Museum, but in all this I have not found specimens that Jacoby originally had from Panama for his first descrip- tion. In the Bowditch collection are three specimens from Volcan de Chiriqui, Panama, collected later by Champion, and the specimens from the British Museum are of this same series collected by Cham- pion. Possibly Jacoby had earlier specimens collected by Salvin. But these Champion specimens match Jacoby’s description. However, the two figures given later for this species in the Biologia Centrali- Americana Supplement are of beetles collected elsewhere, the first from Belize, British Honduras, with an entirely pale pronotum (in the original description the pronotum is given as having a lateral piceous vitta on each side extending from the anterior margin nearly to the basal margin), and the second from Tapachula, Chiapas, Mexico, which Mr. Bryant has sent me, with sutural and lateral elytral vittae, distinctly different from his description of the three large yellow spots on each elytron. It seems quite evident that Jacoby was confused by the wealth of his material, all of which he tried to include in his later account of sexplagiata in the Biologia. In reality there are a number of closely related species having the 6-spotted elytra, but differing in many other less obvious ways. These range from Mexico to Bolivia and Brazil. In my interpretation of serplagiatum I am considering these speci- men from the Volcén de Chiriquf as typical as they correspond with Jacoby’s original description. In Central America this species seems not to occur north of Panama. In South America, however, is found a beetle very similar to the Panama species in markings. Baly (18862) described specimens from Magdelena River, Colombia, as Diabrotica bivittaticollis. Mr. Bryant has sent me a paratype of Baly’s species 80 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 which closely resembles the Volcén de Chiriquf specimens and may be identical with them. The Baly specimen unfortunately is a female. The only other specimens that I have seen from that region are two fe- males collected by P. J. Darlington at Aracataca, Magdalena Province. Specimens closely resembling these have been collected in Venezuela, Trinidad, Brazil, Peru, and Bolivia. On the whole, these South American specimens may be a little longer, the pale spots on the elytra larger, and the last three antennal joints, instead of being piceous, are generally pale. Because of these and some other minor differences, including the aedeagus, I am resurrecting Baly’s name bivittaticollis for a South American subspecies. Future collections and study will contribute more knowledge to the subject. Iceloceras sexplagiatum bivittaticole (Baly) FIGURE 2,c Diabrotica bivittaticollis Baly, Journ. Linn. Soc. Zool., vol. 19, p. 257, April 1885. Neobrotica sexplagiata (Jacoby), Jacoby, in part, Biologia Centrali- Americana, Coleopt., vol. 6, Suppl., p. 331, 1892. The only differences between this and typical [celoceras sexplagiatum are in the somewhat larger size (length 4-5.5 mm., width 2-2.5 mm.), the larger pale spots on the elytra, and the wider and longer black vittae on the pronotum. The last thres joints of the antennae, instead of being black, are usually pale brown. The aedeagus (at least of Tingo Maria specimen) is slightly different. The tip of the abdomen dorsally is darker. The small notch of the middle tibiae of the male is like that in typical J. sexplagiatum, these two forms differing from all the rest by having a much tinier notching. Type: In British Museum (Natural History), collected on the Magdalena River, Colombia. Distrrisution: Colombia: Aracataca, Magdalena Province, P. J. Darlington; Rio Dagua, W. Rosenberg. Peru: Tingo Maria, Dieguez; Rio Charapa, C. H. T. Townsend. Venezuela: Caracas, Panduze; San Esteban, E. Simon, Ist Jacoby collection (? Colombia or Venezuela). Jrinidad: G.E. Bryant. Brazil: Bahia, P. G. Russell; Ceara; Independencia, Parahyba, W. M. Mann and Heath; Jataky, Goiaz Province; Bonito, Pernambuco Province, Dr. Brauns, 2d Jacoby collection; Santa Catharina. Bolivia: Rurrenabaque, Rio Beni, W. M. Mann; Guanay, Mapire River, 1500 ft., Stuart. Iceloceras maximum, new species FIGURE 3,c Approximately 7.5 mm. in length, elongate oblong oval, alutaceous, only faintly shining, head and prothorax densely and finely punctate, the elytra more coarsely and densely punctate; deep yellowish or -GALERUCINE BEETLES—BLAKE Sl orange-brown, the prothorax with a broad piceous spot on each. side coming about half way down, elytra piceous with six pale yellow- brown spots and pale margin, legs and undersurface entirely pale and antennae pale with the three distal joints a bit deeper brown in color. Head with interocular space about half width of head, alutaceous and finely punctate, a well-marked narrow carina running from be- tween antennal sockets down lower front. Antennae pale with the last three joints brownish. Prothorax approximately one-third wider than long with somewhat rounded sides, disk with a semi- circular transverse depression; alutaceous and distinctly and rather densely punctate; yellow-brown with a broad piceous spot on each side extending just below the middle of pronotum. Scutellum pale. Elytra distinctly alutaceous, and densely and shallowly punctate with a tendency to ridging, producing a semicostate appearance; three large, dull, yellow-brown spots and pale margin on piceous background. Epipleura extending to apical curve; legs and under- surface and last two abdominal segments dorsally entirely pale. Length 7-7.5 mm.; width 3.5-3.7 mm. ' Type: Female, USNM 63355, Territorio Federal Amazonas Randal Salas, Aug. 15, 1951, Exp[edicion] F[ran]co Vene[zolana], Alto Orinoco. OTHER LOCALITIES: One female from King Frederick William Falls, Couratryne River, Dutch Guiana, J. G. Myers; one female from “Culebra, N. Duida Territory, Amazons, 1950, J. Maldonado Cap- riles’’; one female from Bogoté, Colombia. Remarks: This is the largest of the sexplagiatum group of beetles, and, unlike the rest, is dull deep orange-brown in its pale markings instead of pale yellow. Unfortunately, no male specimen has been collected. Iceloceras flavipes, new species Figure 3,d From 5-6.5 mm. in length, elongate oblong, somewhat shiny although distinctly alutaceous; prothorax rather densely punctate, elytra densely punctate with semicostate ridging; head, antennae, legs, and body beneath entirely pale, prothorax with wide piceous spot on each side wider anteriorly and extending nearly entire length; elytra piceous with three large pale spots on each elytron, the margin pale. Head with interocular space about half width of head, shining, pale yellow, the occiput finely punctate, frontal tubercles distinctly marked, a narrow carina running down lower front from between antennal sockets. Antennae entirely pale, 3d joint shorter than 4th. Prothorax about one-third wider than long, with slightly rounded sides and semicircular transverse depression; surface shiny, more or 82 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 less alutaceous, and with moderately coarse punctures. Scutellum pale. Elytra faintly alutaceous and densely and coarsely punctate, with irregular ridges giving it a semicostate appearance; piceous with three large pale yellow spots on each elytron, margin pale. Epipleura vanishing at apical curve; last two abdominal segments pale above. Body beneath and legs pale. Length 5.2-6.5 mm.; width 2.4-3.2 mm. Type: Male and three paratypes (2 females, 1 male), USNM 63356 from Colima, Panama, collected by August Busck. OTHER LOCALITIES: Panama: Pedro Miguel, A. H. Jennings; Barro Colorado, Canal Zone, May 1929, P. J. Darlington; Ft. Davis, Canal Zone, Nathan Banks, July 5, 1924. Colombia: Rio Frio, Magdalena Province, P. J. Darlington. Remarks: Of the Central American species having pale legs and antennae, this one is the largest of the serplagiatum group, and is characterized by having longer spots on the side of the pronotum extending nearly to the base. In addition, the pronotum is dis- tinctly punctate. It approaches in size the South American species, I. maximum, but the coloring is different, and the mark on the pro- notum is longer. In the Bowditch collection at the Museum of Comparative Zoology is a specimen from the first Jacoby collection with the label Peru and a name label “D. bivittaticollis Baly’”’ Gn Jacoby’s handwriting?). The specimen is similar to the above speci- mens from Panama in its markings. It is a male but the abdomen is shrivelled and the specimen so old that I have not ventured to dissect it. Possibly this is the specimen from which in part Jacoby originally drew up his first description of Diabrotica sexplagiata, since the locality is Peru. Until further specimens from Peru can be obtained, I believe this specimen should be considered the same as [celoceras flavipes. Iceloceras verbesinae, new species FicurE 3,b Between 5 and 6 mm. in length, elongate oblong oval, shining, the elytra coarsely punctate and semicostate appearing; head, prothorax, body beneath, antennae, and legs pale yellow, the prothorax with a large piceous spot on each side in anterior half, elytra piceous with six pale yellow spots and margin pale. Head with interocular space approximately half width of head, pale yellow, shining, a few fine punctures over occiput, tubercles well marked, carina between antennal sockets narrow and short, not ex- tending down front. Antennae entirely pale, 3d joint shorter than 4th. Prothorax about one-third wider than long with slightly curved sides, a semicircular depression across disk, nearly impunctate, shining, pale yellow with a large piceous spot on each side in anterior half. Scutellum pale. Elytra shining, sometimes faintly alutaceous, GALERUCINE BEETLES—BLAKE 83 densely and coarsely punctate, the punctures forming irregular ridges, giving the elytra a semicostate appearance; piceous with three pale yellow spots on each elytron, the margin pale. Epipleura extending to apical curve, last two abdominal segments entirely pale above, legs and undersurface entirely pale. Length 4.8-6 mm.; width 2-2.5 mm. Type: Male, USNM 63357, and three paratypes, from Tucurrique, Costa Rica, Schild and Burgdorf; two paratypes in Museum of Com- parative Zoology (No. 29612). OrHER LocALitiEs: Costa Rica: Juan Vincas, collected by C. H. Ballou on Verbesina turbacensis H. B. K.; Turrialba, Schild and Burgdorf, 2d Jacoby collection; Port Limén, P. J. Darlington; San José, Schild and Burgdorf. Remarks: In this species with entirely pale legs and antennae, the pronotal spots are near the anterior margin of the prothorax and extend barely to the middle of the pronotum. Iceloceras latiplagiatum, new species Figure 3,a About 5.5 mm. in length, elongate oblong oval, shining although faintly alutaceous; head and prothorax finely and elytra coarsely punctate; pale yellow, the antennae brown with the four distal joints pale, pronotum with a large piceous spot on each side in anterior half nearly meeting in center; elytra dark with six pale yellow spots, margin pale; all femora pale, middle and posterior tibiae dark, anterior tibiae dark on one side; last two segments of abdomen entirely pale on dorsal side. Head with interocular space half its width; smooth and shiny with a few fine punctures on occiput, frontal tubercles distinctly marked, a short carina between antennal sockets. Antennae brown with the four distal joints pale. Prothorax a little wider than long, with slightly curved sides, disk with deep semicircular depression; shiny although alutaceous, finely punctate; pale yellow with a broad piceous mark on each side of anterior half of pronotum, in two of the four specimens nearly meeting in the middle. Scutellum pale. Elytra piceous with three large pale yellow spots on each elytron, the margin pale, densely punctate, faintly costate. Epipleura vanishing at apical curve; last two segments of abdomen dorsally entirely pale. Body beneath entirely pale, anterior tibiae dark on one side, middle and posterior tibiae and all tarsi dark. Length 4.8—-5.5 mm.; width 2.2—2.6 mm. Typr: Male and two paratypes, USNM 63358, collected at Porto Bello, Panama, Mar. 3, 1911, by August Busck. One paratype in Museum of Comparative Zoology (No. 29613). 84 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Remarks: This species is characterized by having bicolored an- tennae, entirely pale femora, dark tibiae and tarsi, and very large dark spots on the anterior half of the pronotum. Iceloceras parviplagiatum, new species FIGurRE 2,6 Between 5 and 6 mm. in length, elongate oblong oval, somewhat shiny, although finely alutaceous, the head and pronotum finely and the elytra densely and coarsely punctate, pale yellow-brown, the spots and margin of piceous elytra pale yellow, the pronotum with a very small dark spot near lateral margin anteriorly, sometimes lacking; antennae dark with the basal and three distal joints tending to be paler, tibiae and tarsi more or less dark, and femora with a dark | streak. Head with interocular space more than half width of head, pale yellow in lower front, deeper in color on occiput, the occiput finely punctate, frontal tubercles somewhat swollen, a short carina not ex- tending more than half-way down front. Antennae brownish or piceous with the three basal and three terminal joints tending to be paler. Prothorax approximately one-third wider than long, with slightly curved sides and a transverse semicircular depression ; surface alutaceous and finely punctate, bright yellow with a small lateral piceous spot near anterior margin, not extending beyond the middle, sometimes very faint or even lacking. Scutellum pale. Elytra alutaceous, densely and coarsely punctate, the punctures often con- fluent and with semicostate ridging; piceous with three large pale spots on each elytron, the upper two sometimes confluent, the margin pale. Epipleura vanishing at apical curve, last ventral segment usually entirely dark on dorsal side. Body beneath pale, legs with anterior tibiae streaked with dark on upper surface, femora pale with a dark streak, middle and posterior tibiae and tarsi dark. Length 5-6 mm.; width 2—2.5 mm. Tyrer: Male, USNM 63359, and 22 paratypes, from El Salvador, E. J. Hambleton, collector, on yams and corn. OrueEr LocALities: Guatemala: San Geronimo, Champion; “‘Salagar, El Chico, Ven. Jicama.” British Honduras: Belize, Blancaneaux, 2d Jacoby collection. Honduras: Tela, W. M. Mann; La Ceiba, F. J. Dyer; Rio Hondo, Blancaneaux, 2d Jacoby collection. Hl Salvador: La Ceiba, K. A. Salman. Costa Rica: Port Limén, P. J. Darlington, May 9, 1929. Rewhnee: Of the species with dark antennae, tibiae, and tarsi, this one is distinguished by the spot, sometimes lacking enetrely. on GALERUCINE BEETLES—BLAKE 85 the anterior side of the pronotum being small and near the anterior margin. This spot, located on each side of the pronotum, is half hidden when viewed from above and is usually much smaller than in the other closely related species. The antennae tend to have pale brown tips. This is one of the two species figured by Jacoby (1892) under Diabrotica sexplaqiata, the drawing having been made from a specimen from Belize, British Honduras, having no pronotal spots. Iceloceras parviplagiatum teapense, new subspecies Fiaure 2,d About 4.5 mm. in length, elongate oblong oval, somewhat alu- taceous, faintly shining, head and pronotum almost impunctate, elytra with dense coarse punctures; pale yellow, the pronotum with a black spot (usually small) on anterior lateral half on each side, the spot never extending as much as half-way down, in one specimen entirely lacking; elytra dark with three pale yellow spots and a pale margin on each, the pale spots sometimes confluent; antennae entirely dark, middle and posterior tibiae and tarsi dark, and femora dark at apex. Head with interocular space half its width; smooth, impunctate, frontal tubercles distinct, a very short elevated line between antennal sockets not extending down front. Antennae usually entirely dark, but in two specimens the last two or three joints paler, 3d joint shorter than 4th. Prothorax wider than long with slightly curved sides and semicircular transverse depression across disk; alutaceous, very finely punctate; pale with a dark piceous spot on each side, in anterior half barely reaching half-way down, and usually small, in one specimen entirely lacking. Scutellum pale. Elytra densely and coarsely punctate with faint sign of longitudinal ridging; piceous with three large pale spots, often confluent, on each elytron, the margin pale. Epipleura disappearing at apical curve; last two ventral segments entirely dark on upper side; femora pale with dark apex; tibiae of anterior legs dark on one side, middle and posterior tibiae and tarsi entirely dark. Length 4.3 mm.; width 1.8 mm. Tyre: Male, in Museum of Comparative Zoology (No. 29611), and six paratypes, from Teapa, Tabasco, Mexico, H. H. Smith, in 2d Jacoby collection, Bowditch collection. Also one specimen taken at Teapa by Salle, 1st Jacoby collection. Remarks: This subspecies is smaller and darker than the more southern race, having entirely dark antennae and usually larger side spots on the pronotum, although they are sometimes lacking. In both races is found the tendency for the elytral pale spots to be confluent. 86 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Iceloceras vittatum, new species Ficure 5,6 About 4 mm. in length, elongate oblong oval, faintly shining, head and pronotum smooth, very finely punctate, elytra coarsely and densely punctate; pale yellow, the pronotum usually with a piceous vittate lateral marking on each side, variable in length, sometimes lacking, elytra with sutural and lateral deep brown vittae not joined at apex, femora pale with a dark streak, tibiae and tarsi in middle and posterior legs dark, tibiae of anterior pair with dark line, first three or four antennal joints pale, remainder dark. Head smooth, shining, with distinctly marked frontal tubercles and short elevated line running down from between antennal sockets; entirely pale. Antennae with four basal joints paler than the piceous or brown outer joints, 3d joint shorter than 4th. Prothorax a little wider than long; with sides slightly curved; a semicircular transverse depression on disk; surface faintly alutaceous and finely punctate; pale yellow with reddish brown or piceous lateral vitta, sometimes lacking. Scutellum pale. Elytra pale with sutural and lateral brown vittae not united at the apex; surface densely and coarsely punctate, with no sign of costation. Epipleura vanishing at apical curve, last two ventral segments dorsally black with a pale median area. Femora pale with dark streak, anterior tibiae dark on one side, middle and posterior tibiae and tarsi entirely dark, and tibiae with tiny terminal spine. Middle tibiae in male notched. Length 44.5 mm.; width 1.7-2 mm. - Typr: Male, USNM 63360, and nine paratypes, from San Marcos, Nicaragua, C. F. Baker collection (through Knab). OruER LocaLitres: Nicaragua: Chinandega, Baker collection. Mexico: Jicaltepec, Vera Cruz; Tapachula, Chiapas, Hége; San Rafale; “Mexique.”’ Remarks: This species is characterized by having vittate elytra, with the vittae not united at the apex. There is no sign of the elytral costation so common in the 6-spotted species. The specimen (one of two) figured by Jacoby (1887) under Diabrotica serplagiata and sent to me by G. E. Bryant from the British Museum is this species, but it does not have the usual dark piceous vittate marking on each side of the pronotum, although there is a faintly marked vitta there. Iceloceras nymphaea (Jacoby) FIGuRE 4,9 Diabrotica nymphaea Jacoby, Biologia Centrali-Americana, Coleopt., vol. 6, pt 1. p. 541, 1887. GALERUCINE BEETLES—BLAKE 87 About 4 mm. in length, elongate oblong, shining, the elytra strongly and densely punctate, pale yellow brown with brown antennae having three apical joints pale; pronotum with a dark brown vitta on each side, elytra with a broad median dark brown vitta, apex of femora, tibiae and tarsi deeper brown. Head shining, entirely pale brown, with well-marked frontal tubercles and carina extending down lower front. Antennae very long and slender, 3d joint half as long as 4th, brown with the three terminal joints pale. Pronotum a little broader than long with nearly straight sides, only shallowly depressed on each side in lower half; shining, impunctate, pale yellow-brown with a wide dark brown vitta on each side extending nearly the entire length. Scutellum pale. Elytra narrow and long, with small humeral prominences, densely and strongly punctate; pale yellow brown with a broad median dark brown vitta on each, wider at base and curving inwards toward the suture near apex. A few scattered hairs near apex of elytra. Body beneath entirely pale, femora with dark streak or apex dark, tibiae and tarsi deep brown. Anterior coxal cavities open, tibiae with ter- minal spine, first tarsal joint on hind legs about equal to remaining joints together, middle tibiae of male very deeply excised, claws appendiculate. Length 3.8-4.4 mm.; width 1.7—1.8 mm. Typr Locauities: Panama: Bugaba, Volcén de Chiriqui, Boquete, Caldera, collected by Champion. Remarks: This is another species described as a Diabrotica that has appendiculate claws and in the male deeply excised middle tibiae. It belongs near the Jceloceras sexplagiatum group. Iceloceras biplagiatum, new species FIGureE 5,e About 5 mm. in length, narrowly oblong oval, head and prothorax smooth and shining, elytra densely and coarsely punctate and with faint median costae, prothorax with semicircular transverse depression; pale yellow with head from tubercles upwards piceous, antennae piceous except the two pale distal joints, the tips of these being dark, prothorax pale with a large median dark spot, elytra piceous with pale margin and a roundish pale median spot on each elytron, body beneath with breast and abdomen brown, femora dark on upper side and at apex on posterior femora, tibiae and tarsi piceous, middle tibiae of male with notch near apex. Head with interocular space about half width of head, eyes large, antennal sockets contiguous, tubercles distinctly marked, a shallow semicircular depression above tubercles, this upper portion of head shining piceous and finely punctate, below antennal sockets the front 88 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 broad, moderately long and pale, a shallow depression in middle, coarsely and densely punctate, labrum short. Antennae long and slender, with joints 3-11 subequal with 4th and 5th joints longest, piceous except the two distal joints and these with a darker tip. Prothorax approximately one-third wider than long with nearly straight sides, only slightly tapering apically; a semicircular transverse depression on disk, shining, impunctate, pale yellow with a large median dark brown spot nearest anterior margin and extending to below middle. Scutellum dark. Elytra widening slightly towards apex, densely and coarsely punctate, with faint costae not reaching either base or apex; piceous with a pale margin, wider at apex, and a large, pale, roundish spot in middle of each elytron. Epipleura pale, vanishing at apical curve. Body beneath with breast and abdomen brown, the former with pale sides and the latter having tip of each | segment pale, lightly pubescent. Legs with the pale femora bearing a dark streak on top, and hind femora dark at apex, tibiae and tarsi dark; middle tibiae of male with a notch near apex; claws appendic- ulate. Length 4.8 mm.; width 2 mm. Typr: Male, USNM 63361, from Isiamas, Bolivia, W. M. Mann, Mulford Biological Expedition, 1921-1922. Remarks: This is clearly closely related to I. sezplagiatum, although with only one pale spot on each elytron, and a median dark spot on the pronotum. Iceloceras ruatanae (Jacoby) Ficure 5,d Neobrotica ruatanae Jacoby, Biologia Centrali-Americana, Coleopt., vol. 6, Suppl., p. 335, 1892. About 6 mm. in length, elongate oblong oval, shining, the elytra obsoletely punctate, prothorax with a wide transverse depression; elytra with deep incurving intrahumeral sulcus extending nearly a third of the way down; pale yellow, the antennae deepening in color towards apex, elytra with a violaceous brown median marking, beginning about scutellum and curving down to cover whole width of elytra before apex, leaving the margin pale; middle tibiae of male notched. Head with interocular space half width of head, shining, finely punctate on occiput, frontal tubercles distinctly marked, a slight median elevation in lower front, labrum wide; entirely pale except the brown tips of mandibles. Antennae long and slender, the 2d and 3d joints short and subequal, rest long, the joints gradually becoming a little shorter near apex and deeper in color. Prothorax somewhat wider than long with only slightly curved sides, not much narrower anteriorly; a wide transverse depression across disk; surface GALERUCINE BEETLES—BLAKE 89 entirely pale, shining, impunctate. Scutellum pale. Elytra a little wider towards apex, with wide explanate margin, a well-marked intrahumeral sulcus curving towards suture; surface obsoletely punctate, shining, pale with a violaceous brown sutural mark that is wide about suture, curving down to a narrow stretch in middle of elytra and again widening beyond the middle to cover all but margin in apical third. Epipleura vanishing before apical curve. Body beneath and legs entirely pale. Middle tibiae in male notched, a fine spine on tibiae; claws appendiculate. Length 6 mm.; width 2.4 mm. Typs: In British Museum (Natural History), from Ruatan Island, Honduras. OrHER LocaLitizs: Honduras: Puerta Castilla, Jos. Bequaert, April (MCZ); ‘“‘Roatan Id.,” F. J. Dyer, Mar. 28, 29, 1916. Remarks: In the short third antennal joint this species resembles LIuperosoma, but there is no enlargement of the antennal joints. Iceloceras flavocyaneum, new species FicureE 5,a Between 4 and 5 mm. in length, elongate oblong oval, shining, nearly impunctate, the elytra very finely punctate, yellow with posterior part of elytra violaceous; antennae pale with the last five joints a little darker; middle tibiae notched in male. Head with interocular space half its width, smooth, shining, im- punctate, pale yellow; frontal tubercles not very distinct, a median line from between antennal sockets. Antennae with 2d and 3d joints short, 4-11 long and subequal, the five basal joints pale, the remainder reddish brown. Prothorax a little wider than long with slightly curved sides, shallowly depressed in lower half, shining, impunctate, pale yellow. Scutellum pale. Elytra shining, very finely punctate, pale yellow with a violaceous spot covering apical third, leaving margin pale (in one specimen from Panama the suture also pale). Epipleura vanishing before apical curve. Body beneath entirely pale, tibiae with tiny terminal spine; claws appendiculate, middle tibiae notched inmale. Length 4-5.3 mm.; width 1.8 to 2.2 mm. Type: Male, USNM 63362, from Santa Lucretia, Tehuantepec, Mexico, W. M. Mann; one paratype in Museum of Comparative Zoology. OTHER LOCALITIES: Santa Lucretia, Vera Cruz, Mexico, F. Knab; XX Plantation, Panama. Remarks: This species is closely related to I. ruatanae (Jacoby) with the same short 2d and 3d antennal joints, and is intermediate in the matter of generic placing between Jceloceras and Luperosoma. 90 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Genus Luperosoma Jacoby Luperosoma Jacoby in Whymper, Travels amongst the Great Andes of the Equator, Supplementary Appendix, p. 87, 1891. The original generic description is as follows: Male. Body elongate; eyes entire; third joint of palpi robust, swollen; frontal tubercles very swollen, transverse; antennae subfiliform, incrassate towards the apex, second and third joints very short, subequal, fourth as long as the two preceding joints together; thorax square-shaped, transversely depressed at the disk; apex of the scutellum obtuse; elytra irregularly punctured, their epipleurae indistinct below the middle; tibiae without spine ‘‘(not true),’’ the intermediate emarginate at the apex; the inner margin produced in shape of a spine; posterior first tarsal joint as long as the two following ones united; claws appendiculate; anterior coxal cavities closed. Female. Intermediate tibiae without emargina- tion, prosternum not visible. Type (of genus) Luperosoma marginata. I am obliged to erect this genus for the reception of a small species of Galeruca having the appearance of Diabrotica or Luperus, and distinguished from either and other genera by the unarmed tibiae [there is a small spine on the middle and posterior tibiae]; short second and third joints of antennae and the other char- acters given above. The curious structure of the tibiae in the male is another peculiarity of the genus which would enter Chapuis’ 26th group, the Platyxanthi- nae. Key to species of Luperosoma ToNorth Americans co.) 62 io ve. bay peak agua kay Picea tie csi er bcos, eee ob oigkuend vex CROCS South Americans visi. sci scapes Core ea gate le Cahier Well i ogar FSR coke etapa eee: 2. Elytra distinctly costate .......... .. . subsuleatum (Horn) Elytra’ notidistinctlycostates cir a 2 sow (oN oh een le ene gotten cen eee en 3. Elytra piceous with pale margin. ...... . . . . parallelum (Horn) Elytra entirely piceous ......2.2.4..... =... sehwarzi (Horn) 4. Thoraxentirely dark. .... oe + we we ee) Migrum, new species Thorax pale with dark caren cad oats ava aide a Ab eae 5. Large, 5 mm., last three antennal joints! in anne eAteapedt mronoure with heavy interal dark markings... . . . . amplicorne (Baly) Smaller, 3-3.5 mm., joints 8-10 in male palareed: pronetar pale with median Garkispoty 6) sascha Vie Se ere es Tree _. . marginatum Jacoby Luperosoma marginatum Jacoby Figure 4,c Luperosoma marginata Jacoby, in Whymper, Travels Amongst the Great Andes, Supplementary Appendix, p. 87, 1891. Phyllecthrus marginatus Weise, in Junk, Coleopterorum catalogus, pt. 78, p. 109, 1924. From 3-3.5 mm. in length, oblong oval, shining, the elytra very faintly punctate, pale yellow-brown with piceous antennae and deep brown occiput of head, a dark brown median area on pronotum, the elytra sometimes with a wide brown median vitta on each elytron, in other specimens this vitta so pale as to be nearly indistinct; body beneath dark, legs for the most part pale, middle tibiae in male notched, and antennal joints 8-10 swollen. GALERUCINE BEETLES—BLAKE 9] Head with interocular space half its width, frontal tubercles swollen, and interantennal area carinate, upper part of head deeper brown than lower. Antennae fully half as long as beetle, the basal joints a little paler, rest piceous, 2d and 3d joints short and together barely as long as 4th, remainder shorter than 4th and subequal, joints 8-10 swollen in male. Prothorax wider than long with nearly straight sides, a faint semicircular depression in lower half, nearly impunctate, shining, dirty yellow-brown with a roundish piceous median area in anterior half. Scutellum dark. Elytra wider towards apex and with small humeral prominences and short intrahumeral sulcus; shining, very faintly punctate, dirty yellow-brown, in some specimens a wide piceous vitta, in others this vitta pale reddish brown and indistinct. Epipleura extending to middle of elytra. Body beneath dark, legs pale, the two anterior tibiae with a pale reddish brown area in middle, in the male the anterior tibiae somewhat swollen and the first tarsal joint cylindrical and short, the middle tibiae notched near apex, the middle and posterior tibiae (contrary to Jacoby’s description) with a very fine, inconspicuous spine; claws appendiculate. Length 3-3.5 mm.; width 1.5 mm. Typ: Probably in the Bowditch collection, Museum of Comparative Zoology, where there are three specimens (named in Jacoby’s descrip- tion) from Quito, Ecuador, alt. 9500 ft., Ed Whymper collector, bearing an old label “LZuperosoma marginata,’’ written by Jacoby, 2d Jacoby collection. Remarks: These three specimens in the Bowditch collection are probably the three original ones described by Jacoby (1891) although in the text of his description the locality is given as ‘‘Panecillo Quito, 10,000 ft.” Luperosoma nigrum, new species FicureE 4,¢ Approximately 3 mm. in length, oblong oval, shining, the elytra distinctly punctate; entirely black except for a narrow pale sutural and marginal edging on the elytra; antennal joints 8-10 enlarged and the middle tibiae notched in the male. Head with interocular space a little more than half its width, polished, impunctate, entirely black, the frontal tubercles distinct and a broad carina between antennal sockets not much _ produced. Antennae entirely black, in male joints 8-10 swollen, joints 2, 3 short and together about equal to4. Prothorax wider than long with nearly straight sides, obsoletely and faintly punctate, shining, a semicircular depression across disk; entirely black. Scutellum black. Elytra moderately densely and distinctly punctate, shining black with pale yellow sutural stripe and pale margin. Epipleura pale, extending to 92 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 apical curve. Body beneath entirely black and shining; anterior tibiae in male somewhat thickened, middle tibiae widely emarginate with prominent hook near apex, tibiae with small spine at apex. Length 3 mm.; width 1.3 mm. Type: Male in Museum of Comparative Zoology (No. 29610); one paratype, from ‘““Colombien’” (USNM 63363). Remarks: The wide emargination of the middle tibiae in the male is greater than in any of the rest of the species with excised tibiae, but it is only a matter of degree. The enlargement of the antennal joints is the same as in L. marginatum Jacoby. In fact, except for the wider notching of the tibiae and the darker color, which has the same pattern on the elytra as in marginatum, this species closely resembles L. marginatum and might be only a color phase. Luperosoma amplicorne (Baly) FiaurE 4,h Diabrotica amplicornis Baly, Trans. Ent. Soc. London, p. 446, 1886. About 5 mm. in length, oblong ova], the pronotum and elytra distinctly punctate, shining, pale yellow-brown, the head with two dark spots on occiput and dark labrum, antennae with joints 9-11 dark and in the male thickened, prothorax with broad dark lateral areas, elytra with median and lateral dark vittae joined at apex, legs and undersurface, except prosternum, dark. Middle tibiae of male notched. Head with interocular space half its width; occiput impunctate, with wide dark areas on either side narrowly separated by pale median line; front brownish, the frontal tubercles much swollen, the inter- antennal area with a carina down front, labrum dark. Antennae in male with joints 9-11 dark and enlarged in the male, no female examined. Prothorax wider than long with slightly curved sides and semicircular depression in lower half, surface shining and distinctly punctate, yellow-brown with wide black area on either side extending from anterior nearly to basal margin. Scutellum brown. Elytra moderately shining, quite densely and rather coarsely punctate, a depression from intrahumeral sulcus down about one-fourth the length of the elytra; yellow-brown with a wide dark median and lateral vitta uniting shortly before the apex, leaving the suture and the lateral margin narrowly pale, and a slightly broader median pale vitta. Epipleura disappearing before apical curve, body beneath (except prosternum) and legs deep brown, middle tibiae of male notched near apex; tibiae with fine terminal spine; anterior tibiae swollen in male, first tarsal joint of anterior legs cylindrical, first tarsal joint of hind legs as long as remaining joints. Claws appendiculate. Length 4.8 mm.; width 2.2 mm. GALERUCINE BEETLES—BLAKE 93 Typr: In British Museum (Natural History), collected at Parand, Brazil. Remarks: What appears to be a paratype of this is in the Bowditch collection, 1st Jacoby collection, Museum of Comparative Zoology. No females have been examined. Luperosoma parallelum (Horn) Figure 4,6 Phyllecthrus parallelus Horn, Trans. Amer. Ent. Soc., vol. 20, p. 126, 1893. About 5 mm. in length, elongate oblong oval, rather flat, moderately shiny although alutaceous, and with shallow, not dense elytral puncta- - tion; head reddish brown, deeper brown over occiput, prothorax pale yellowish brown, elytra piceous with pale margin, legs and antennae pale, the former with dark streaks, abdomen and part of breast piceous; antennae in male enlarged at end, and middle tibiae notched near apex. Head with deeper reddish brown color over occiput becoming pale yellow down front, alutaceous and obsoletely punctate, with swollen frontal tubercles and slightly produced carina between antennal sockets. Antennae pale except basal joint, with a deeper brown on upper surface, 2d and 3d joints short, 4th longest of all, thence gradually diminishing in length to terminal joint, which in male is enlarged. No female examined. Prothorax about a third wider than long with sides nearly straight, only slightly curved, anterior angles toothed, basal angles obliquely cut, disk with semicircular depression in basal half and a smaller one in middle of anterior half; pale yellow, often with spot or widely brown or piceous areas on each side. Seutellum reddish brown. Elytra somewhat depressed, shining although distinctly alutaceous and with scattered shallow punctures, deep piceous except the pale lateral (not apical) margin, sometimes slightly paler along sutural edges. Epipleura pale. Body beneath with breast brownish, abdomen even deeper brown, legs pale with a dark streak along upper side; middle tibiae of male notched near apex, first tarsal joint of anterior legs in male short; middle and posterior tibiae with fine spine at apex. Claws appendiculate. Length 4.3-5.2 mm.; width 1.8—2 mm. Typre: Not examined by the writer, but H. J. Grant of the Phila- delphia Academy of Sciences writes as follows: “Type No. 3792. Labeled holotype. Sex not indicated. Locality: ‘Tex.’ In the Horn collection there are two additional specimens—one with no data; one labeled simply ‘Tex.’ ” be hia OrnerR xLocauities: Teras: Round Mountain (Fall collection) ; Douglass, H. S. Barber; College Station, W. D. Pierce. Oklahoma: Ardmore, C. R. Jones. Kansas: West Kansas, Popenoe; Onaga, 94. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 F. F. Crevecoeur, Wickham; Topeka, Popenoe; Riley County, Popenoe. Remarks: The characters described by Jacoby (1891) for the genus Luperosoma so well fit the species described from North America by Horn (1898, 1896) as Phyllecthrus parallelus, P. schwarzi, and P. subsulcatus that I am transferring Horn’s species, leaving to Phyjllec- thrus the original ones described by LeConte (1865, 1868, 1884), who took as type of that genus Phyllecthrus dorsalis (Olivier). Luperosoma subsulcatum (Horn) Ficure 4,f Phyllecthrus subsulcatus Horn, Trans. Amer. Ent. Soc., vol. 20, p. 126, 1893. About 3.5 mm. in length, elongate oblong oval, shining although somewhat alutaceous, elytra with numerous longitudinal ridges be- - tween which are semistriate, moderately coarse punctures; head, antennae, prothorax, abdomen, tibiae, and tarsi reddish brown, elytra piceous; antennae in male flatly widened towards apex, middle tibiae notched near apex. Head shining reddish brown, smoothly rounded over occiput with some punctures in the depression over the frontal tubercles, tubercles swollen, a faint median line; antennal sockets depressed but the area between not prominently produced into a carina; eyes widely sepa- rated and from above not showing their comparatively large size. Antennae stout in male, the 2d and 3d joints short and robust, 4th longest, 5th and 6th shorter and equal, 7th cylindrical and a little shorter, 8th and 9th short and broad, 10th and 11th very broad and long; in the female, 3d and 4th joints longer than 2d and subequal, 4th longer than succeeding joints which are not enlarged. Prothorax almost as long as wide, with only slightly curved sides, a small tooth at apex, shining reddish brown sometimes with a median round brown- ish or piceous area; without much sign of semicircular depression on most specimens, a few inconspicuous punctures along base. Scutellum dark. Elytra usually entirely piceous (in one specimen the base and side from humerus down reddish brown, the rest of elytra pi- ceous); shining although faintly alutaceous, a little wider than pro- thorax and narrowly elongate; humeri moderately prominent, nu- merous irregular longitudinal ridges between rows of punctures, the punctures moderately coarse and irregularly striate, both punctures and ridges becoming obsolete near apex. Epipleura disappearing at apical curve; body beneath with breast dark, prosternum and abdomen pale, femora more or less dark, tibiae and tarsi paler brown. Middle and posterior tibiae with a fine spine, and in male, the anterior Ist tarsal joint cylindrical and twice as broad as the next; middle tibiae notched, claws appendiculate. Length 3-4.3 mm.; width 1.3-1.7 mm. GALERUCINE BEETLES—BLAKE. 95 Typs: Not examined by writer but H. J. Grant of the Philadelphia Academy of Sciences writes as follows: ‘““Type No. 3794. Lectotype. Male. Locality: Yuma. There are no other specimens of this species in the Horn collection.”” Horn wrote: “It occurs in New Mexico, Texas, and Arizona, in the latter region near Yuma and Santa Rita Mts.” OTHER LOCALITIES: Arizona: Bear Valley, Atascosa Mts., F. G. Werner; 9 miles west of Benson, F. G. Werner; Brown’s Canyon, Baboquivari Mts., F. G. Werner and G. D. Butler; Huachuca Mts.; Marana, Butler and Werner; Nogales, Santa Cruz County, Nunen- macher; 6 miles north of Nogales, 3500 ft., Werner and Nutting; Oracle Junction, Werner and Nutting; 6 miles east of Peace, F. G. Werner; Pima, 3000 ft.; 2 miles east of Ruby, F. G. Werner; Sabino Canyon, Santa Catalina Mts., Werner and Nutting; 8 miles east of San Vicente, F. G. Werner, A. A. Nichol, Tucson Mts., G. D. Butler. Texas: Hl Paso. New Mexico: Mesilla Park. Remarks: The semicostate elytra readily distinguish this species. Luperosoma schwarzi (Horn) FiGguRE 4,a Phyllecthrus schwarzi Horn, Proc. California Acad. Sci., ser. 2, vol. 6, p. 377, 1896. About 5 mm. in length, elongate oblong oval, moderately shiny, although somewhat alutaceous, antennae heavy in male with the last three joints wider, middle tibiae notched near apex in male; prothorax with a depressed spot on each side; head and prothorax reddish brown, the latter with piceous lateral darkening; elytra, legs, and under- surface dark; elytra with faint irregular costae; the punctures with a suggestion of irregular striation in basal half. Head smooth and shiny above the swollen frontal tubercles, a slightly produced short carina between antennal sockets, deep orange- brown with darker labrum. Antennae of male with the three basal joints shining piceous, remainder densely pubescent with grayish brown hairs, 2d and 3d joints short, 4th longest, last three joints heavier and wider; in female, the joints longer and more slender, 2d and 3d joints together equalling 4th. Prothorax approximately one- third wider than long, nearly rectangular, with only slightly rounded sides, deep yellow-brown with the sides piceous, on each side and in middle of base a shallow depression; alutaceous and finely and incon- spicuously punctate near base. Scutellum dark. Elytra long and narrow with slight humeral prominences and faint irregular costae near base, punctation with a tendency to becoming striate near base, not very dense or coarse and becoming much finer near apex; surface alutaceous but shiny, deep piceous. Epipleura vanishing at apical curve. Body beneath entirely dark, legs dark except a little paler at 96 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 knees; middle tibiae of male notched near apex, tibiae with fine spine, claws appendiculate. Length 4.5-5.0 mm.; width 2 mm. Typx: Not examined by writer, but H. J. Grant of the Philadelphia Academy of Sciences writes as follows: ‘“Type No. 3793. Holotype. Male. Locality: Tex. In the Horn collection there are seven para- types, these being from Texas, Arizona and New Mexico, and four other specimens from Arizona, Kansas and California.” OTHER LOCALITIES: Houston, Tex., on Helianthus; Shreveport, La., F. V. Mally. Remarks: Horn originally mistook this for Phyllecthrus dorsalis (Olivier), and three years later corrected this error, which E. A. Schwartz had pointed out to him, and described it again as new. Genus Ectmesopus Blake Figure 6,a—-k Ectmesopus Blake, Proc. Ent. Soc. Washington, vol. 42, p. 95, 1940. Small, slender beetles 2-4 mm. long, with long slender legs, usually pale yellow or reddish with lustrous green, blue or violet elytra. Head smoothly rounded, without depressions or protuberances, frontal tubercles not pronounced. Antennae usually not much over half the length of the beetles, in one species nearly the length. Usually some abnormality in the apical antennal joints, 7, 8, 9, 10 or 11 of the male. In one species, the one with the long antennae, this abnormality in the 3d joint. Prothorax from one-fourth to one-third wider than long, with the sides only slightly curved, often nearly straight, narrowly margined and with a small seta-bearing pore at the corners, hind margin nearly straight; the disk not at all depressed or deeply punc- tate, but usually smoothly rounded and polished. Elytra wider than the prothorax with a small humeral prominence, and usually a little wider in the apical half. Beneath, the epipleura gradually disappear towards the apex. Anterior coxal cavities open. In the male, the front tibiae frequently stout and the middle tibiae with a deep notch on the inside near the apex. Tibiae with a tiny spur, extremely hard to detect in some specimens. First joint of the hind tarsi equal to or longer than the following. Claws with a small basal tooth. Type or Genus: Ectmesopus darlingtoni Blake. Key to species of Ectmesopus 1. Prothorax entirely dark without any pale areas, joints 8-10 of male antennae Geformed..,, : JAmaieas..:c)' oxiiec sisters ssupayes, Nets see sis eo LER ley tristis Blake Prothorax entirely pale or pale with dark markings .......... 2 2. rrothorax pale with dark markings) 3) <0 .a)e0 coe ene Al eee te ee 3 Prothorax entirely pales) CARP hie le Mea eae ie tree, cheney caer anata 7 GALERUCINE BEETLES—BLAKE 97 3. Elytra shining green with a broad irregular pale band. Puerto Rico. zonatus Blake Elytra entirely dark... . . oe A hae, 4. Antennae extending much below fie Wade of fhe alytrs, in male the 3d joint very tiny, in both sexes the 4th joint very long. Haiti. longicornis Blake Antennae not reaching beyond the middle of the elytra, mostly not so far as the middle. .... Sings a aeipie 5. Prothorax with a broad, eee Genet piedian aeons extendas nearly the whole length; 10th joint of male antennae slightly enlarged. Cuba. ye Blake Prothorax with lateral dark marks. . .. . ce ae eG 6. Prothorax with a vitta on each side, this sorhetinies very ghost: sometimes well marked and extending down the whole side; 10th joint of male antennae much enlarged. Puerto Rico... . . . . Vitticollis Blake Prothorax with sides near margin somemies an ee antennal joints in male not deformed or enlarged. Dominican Republic. angusticdllis Blake Prothorax with lateral margin piceous black. Tortue Island, Haiti. leonardorum Blake 7. Notch in middle tibiae not deep, little more than an emargination; male antennae not deformed or enlarged or unlike those of the female. Domini- can Republic. ... . . . . angusticollis Blake Notch in middle tibiae of male Aon Paras a "ehaup tooth on upper inside margin; last joints of male antennae either deformed or enlarged. . . . 8 8. Antennae of both sexes very similar up to the last joint and in male the last joint enlarged. Cuba..... . . . + malachiodes (Suffrian) The 10th antennal joint and Somaiimnes) Benes of the male deformed. . . 9 9. Antennae dark with the last 2 or 3 joints pale, joints 6-9 in male triangular, 10th much enlarged. Haiti. .... . .. . . darlingtoni Blake Antennae not bicolored, joints 6 to 9 in mele not at all triangular. . . 10 10. Head, thorax, legs and antennae pale yellowish; 10th antennal joint in male much enlarged. Dominican Republic. ..... . . . pallidus Blake Head, thorax, and legs reddish, antennae with dark outer joints; 10th joint in male constricted and not any longer than 9th. Dominican Republic. crassicornis Blake Since all but one of the species given in the above key have been described by the writer (Blake, 1940), it does not seem necessary to repeat the descriptions in detail, as the key and illustrations are sufficient for purposes of identification. Ectmesopus leonardorum, new species FIGURE 6e Approximately 2.7 mm. in length, elongate oblong oval, shining, inpunctate, yellow-brown with piceous lateral margin to the pronotum and metallic green elytra; middle tibiae of male widely and shallowly emarginate; antennae with distal joints missing. Head with interocular space more than half width of head, entirely pale, smoothly rounded over occiput, a slight depression over the bulg- ing frontal tubercles, a short carina between antennal sockets running 98 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 down the Jower front. Antennae with the four distal joints missing, pale yellow-brown, 3d joint longer than 2d and shorter than 4th. Prothorax subquadrate, nearly as long as wide, with only slightly curved sides, pale yellow-brown, the lateral margins piceous black, shining, impunctate; a dent in middle of disk (probably from im- mature softness of specimen), the surface in life probably without depression of any sort. Scutellum yellowish brown. Elytra shining metallic green, impunctate. Undersurface and legs entirely pale yellow-brown; anterior tibiae thickened in male, middle tibiae widely and shallowly emarginate near apex. Length 2.7 mm.; width 1 mm. Tyrer: Male, USNM 63364, from Bassin Bleu, Tortue Island, Haiti, collected by E. C. and G. M. Leonard. Remarks: Although the specimen is immature and badly shrivelled, and the distal joints of the antennae are missing, this can readily be - seen as a distinctive little beetle with its heavy dark markings on the lateral margin of the prothorax. In addition, the middle tibiae of the male are widely emarginate, in this respect resembling EF. angusticollis Blake from the Dominican Republic. It differs from that species in the coloration of the margin of the prothorax as well as in having metallic green instead of blue elytra, and also in being even narrower. It is unfortunate that the end joints of the antennae, usually so ab- normal in the male, are lacking. Genus Platymorpha Jacoby Platymorpha Jacoby, Biologia Centrali-Americana, Coleopt., vol. 6, pt. 1, p. 602, 1888. The original generic description is as follows: Body elongate; antennae longer than the body, the second and third joints ex- tremely short, the other joints elongate-triangular; thorax subquadrate, the disc depressed; elytral epipleurae continued below the middle; the posterior tibiae mucronate, the first joint of the posterior tarsi as long as the following three joints together; claws appendiculate; the anterior coxal cavities open. Anterior tibiae and the first joint of the anterior tarsi strongly dilated in the typical species in the male. Type Platymorpha variegata. In general appearance Platymorpha agrees with Chthoneis; the third joint of the antennae, however, is extremely small, the posterior tibiae are armed with a spine. The last named character proves the affinity of Platymorpha with Luperus and its allies. I probably have only male specimens of P. variegata before me; these are at once distinguished by the curious dilation of the anterior tibiae and the first joint of the anterior tarsi. The two species I refer to this genus inhabit Mexico and Guatemala. To Jacoby’s description may be added that it is possible these two species he has assigned to the genus are not congeneric. He originally described smaragdipennis in the genus Chthoneis. The anterior tibiae and first tarsal joint in the male in Platymorpha variegata are much GALERUCINE BEETLES—-BLAKE 99 swollen and resemble that of Cerotoma dilatipes Jacoby, while the beetle itself bears a striking resemblance to Phyllecthrus dorsalis (Olivier). Although J. A. Wilcox has written me that Platymorpha variegata has excised middle tibiae in the male, I have been unable to detect it in the mounted specimens in the Bowditch collection. P. smaragdipennis, on the other hand, has no such dilation in the front legs, the middle tibiae are clearly excised, and the shape of the beetle is different. In one point both agree: in the short, truncate 3d anten- nal joint. I am including in this genus, purely because of the similarity of its excised middle tibiae and the short third antennal joint with its _ truncate apex, a third species from Peru. It is not closely related to either P. variegata or P. smaragdipennis and someday will doubtless be referred elsewhere. Platymorpha smaragdipennis Jacoby Fiaure 5,c Pialymorpha smaragdipennis Jacoby, Proc. Zool. Soc. London, p. 786, 1879. About 6 mm. in length, elongate oblong, shining, the elytra densely and coarsely punctate; head, prothorax, femora, and undersurface yellow, elytra shining metallic green, tibiae and tarsi deep brown. Head with a few minute punctures, carina and encarpae ill-defined. Antennae long, black, the 2d and 3d joints very small, truncate at apex; remainder long, densely covered with short hairs. Prothorax subquadrate, wider at middle, contracted near the base; impunctate, very shiny, yellow, disk in basal half with transverse depression. Scutellum pale. Elytra bright metallic green, closely and coarsely rugose punctate. Body beneath and femora pale, tibiae and tarsi dark; middle tibiae of male deeply excised; tibiae with spine, claws appendi- culate. Type: Male in Bowditch collection from Capetillo, Guatemala, and two paratypes (one male, one female). Platymorpha albiventris, new species Figure 4,d About 4 mm. in length, oval, parts of the prothorax and the entire elytra densely punctate, shining black with pale abdomen and the 8th and 9th antennal joints (possibly also the terminal ones which are missing in the single male specimen examined) pale, prothorax with a transverse scooped-out depression; middle tibiae of male excised. Head with interocular space half width of head, occiput polished, frontal tubercles distinctly marked, a small carina between antennal sockets, lower front somewhat bulging. Antennae in male with 3d joint very short, about half length of 2d, remainder long and slender; 100 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 only nine joints of one antenna and the basal joint of the other left; joints 1-7 black, 8 and 9 pale. Prothorax not quite twice as wide as long with nearly straight sides and scooped-out transverse depression in basal half; except in this depression the surface punctate; shining black. Scutellum black. Elytra widened toward apex with wide explanate margin, very shiny black with moderately dense puncta- tion; no suggestion of costae. Body beneath shining black with pale abdomen; anterior coxal cavities open, anterior tibiae in male thick- ened, middle tibiae in male excised near apex; first tarsal joint of hind legs longer than succeeding joints together; claws appendiculate. No terminal spurs discernible on tibiae. Length 4 mm.; width 2.2 mm. Tyre: Male in Cornell University collection, from LaChorerra, Putumayo District, Peru, collection Aug. 17-20, 1926. Remarks: This is another galerucid that has characters similar to - the group with excised tibiae but differs in the male sex characters, being unusual in the shortness of the 3d antennal joint. In this case the 3d joint is unlike the short 3d joint found often in Diabrotica, Luperosoma, or Iceloceras in that it is not so long as the second, truncate at the apex, and cup-shaped. It most closely resembles the antennae of the two species of Platymorpha, and I am tentatively placing the species in that genus although it is not very closely related. References Baty, JoSEPH SUGAR 1886a. The Colombian species of the genus Diabrotica, with descriptions of those hitherto uncharacterized. Pt. 2. Journ. Linn. Soc. Zool., vol. 19, pp. 230-259. 1886b. Descriptions of uncharacterized species of Diabrotica. Trans. Ent. Soc. London, pp. 443-455. BLAKE, Dorts HOLMES 1940. A new genus of Galerucinae (Coleoptera) from the West Indies. Proc. Ent. Soc. Washington, vol. 42, pp. 95-104. Cuapuis, FELICIEN 1875. Histoire naturelle des insectes. Genera des coléoptéres . . . [cont. of Lacordaire]. Vol. 11, 420 pp. DEJEAN, P. M. M. A. 1837. Catalogue des coléoptéres. . . . Ed. 2, livre 5, 503 pp. GAHAN, CHARLES JOSEPH 1891a. On the South American species of Diabrotica. Pt. 2. Trans. Ent. Soc. London, pp. 415-472. 1891b. On the South American species of Diabrotica. Appendix. Trans. Ent. Soe. London, pp. 521-524. Horn, Grorce HENRY 1893. The Galerucini of boreal America. Trans. Amer. Ent. Soc., vol. 20, pp. 57-136. 1896. Coleoptera of Baja California (supplement II). Proc. California Acad. Sci., ser. 2, vol. 6, pp. 367-381. GALERUCINE BEETLES—BLAKE 101 Jacosy, Martin 1878. Descriptions of new species of phytophagous Coleoptera. Proc. Zool. Soc. London (1877), pp. 144-153. 1879. Descriptions of new species of phytophagous Coleoptera. Proc. Zool. Soe. London, pp. 773-793. 1887, Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae, vol. 6, pt. 1, pp. 497-584. 1888. Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae, vol. 6, pt. 1, pp. 585-625. 1891. Coleoptera. Jn Whymper, Travels amongst the Great Andes of the Equator, supplementary appendix, pp. 82-88. 1892. Biologia Centrali-Americana, Insecta, Coleoptera, Supplement to Phytophaga, vol. 6, pt. 1, Suppl., pp. 313-348. LEeContTE, JOHN LAWRENCE 1865. On the species of Galeruca and allied genera inhabiting North America. Proc. Acad. Nat. Sci. Philadelphia, pp. 204-222. 1868. New Coleoptera collected on the survey for the extension of the Union Pacific Railway, E. D. from Kansas to Fort Craig, New Mexico. Trans. Amer. Ent. Soc., vol. 2, pp. 49-64. 1884. Short studies of North American Coleoptera. Trans. Amer. Ent. Soc., vol. 12, pp. 1-82. OLiIvieR, ANTOINE GUILLAUME 1808. Entomologie. . . . Coléopteres, vol. 6. Say, THOMAS 1824. Descriptions of coleopterous insects collected in the late expedition to the Rocky Mountains. . . . Journ. Acad. Nat. Sci. Phila- delphia, vol. 3, pt. 2, pp. 239-282, 298-331, 403-462. SuFFrRIAN, E. 1867. Verzeichniss der von Dr. Gundlach auf der Insel Cuba gesammelten Chrysomelinen. Arch. Naturg., vol. 33, pp. 283-328. WEISE, JULIUS 1924. Chrysomelidae: Galerucinae. In Junk (ed.), Coleopterorum cata- logus, pt. 78, pp. 1-225. U. S, GOVERNMENT PRINTING OFFICE: 1958 w dt , 4 J y rr ; ‘ i pK ¢ rT eR y i he 2 , + \ ’ tees i 4 r yee 7 . A > { : Pde bee ’ i , Py , b . ‘ > ‘ \ - Tee 34! : , . : “nh 7 ope f ; nh ratere se be : Ca uw ( i ; 7 ep, + Ags i | 5 ‘ ‘ ; ‘ i ip tr : é : es ex ‘ z * 3 a ry ws + ’ ; t = "i ; i i PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM MM aky SA Ng a anak VAN a Sa MNO oy SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM ooo Vol. 108 Washington : 1958 No. 3396 ee A REDEFINITION OF THE BUTTERFLY GENERA TATOCHILA, PHULIA, PIERCOLIAS, AND BALTIA, WITH DESCRIPTIONS OF RELATED, GENERA AND SUBGENERA By Witur1aM D. FIetp The genera of butterflies treated in this paper are those labeled by Klots (Ent. Americana, vol. 12, new ser., pl. 13, fig. 100, 1933) as the “‘Tatocheilae-Phulia” branch of the family tree. These genera probably will be considered as constituting a subtribe of the Pierini when that tribe is subdivided. The history of revisionary work in the Rhopalocera shows that the systematics of the group has progressed by three main stages. The earliest workers studied the habitus (shape, color, and pattern) of the wings almost exclusively. Before the middle of the 19th century the veins of the wings were discovered to be of great value and were thereafter used extensively in revisionary work. A few years later the male genitalia were found to be of great systematic value and slowly were used by more and more workers throughout a large part of the Rhopalocera. Surprisingly, studies of the female genitalia were almost never attempted, partly because the females were frequently harder to dissect and because of an erroneous contention of early workers that the female genitalia were of little taxonomic value. We know today that this is not true, and it has been demonstrated numerous times 453667—58 103 104 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 that the female genitalia are often as useful as the male genitalia in ascertaining the relationship of butterfly groups. Concerning the genera now under scrutiny, the soundest early works were those of Butler (Cist. Ent., vol. 1, pp. 83-58, 1870) and Grote (Proc. Amer. Philos. Soc., vol. 39, pp. 4-67, 1900) and were based, according to the practice of that time, mainly upon venational characters. To these studies Klots (loc. cit., pp. 217-220) added a study of the male genitalia, arriving at a fairly satisfactory generic treatment which was superior to all similar generic studies in the Rhopalocera and was certainly an outstanding landmark. In preparing the present paper, the male genitalia, venation, and habitus were thoroughly studied and evaluated. Also, the legs, including the claws (which proved to be of great value), and the female genitalia were studied. The complexities of relationship among the genera studied were found to be much greater than heretofore believed. The use of any one character (or kind of character) alone was found to produce a different alignment than when some other character was employed. Thus, a consideration of the uncus alone shows Theochila, new genus; Tatochila; Phulia (Infraphulia, new subgenus); Piercolias; and Baltia to be quite distinct from one another, while Hypsochila and Phulia (Phulia) would be considered very close to each other and quite like Tatochila. Considering the aedeagus only, one group of the genus Tatochila and the genera Hypsochila, Phulia, Prercolias, and Baltia would certainly seem to be nearly identical, while Theochila and four of the species groups of Tatochila would appear to be quite different from each other and from those mentioned above. On the other hand, Hypsochila, Phulia, Piercolias, and Baltia differ greatly from Theochila and Tatochila in lacking the tibial spurs of the mid and hind legs. Again, if we placed too much emphasis upon the claws, paronychia, and pulvilli, we would place Hypsochila and Theochila an unreasonable distance from the other genera and from each other. A consideration of the number and fusion of the radial veins taken by itself would give us the unnatural alignment of having Theochila, Tatochila, Hypsochila, and the nominotypical subgenus of Piercolias at one end (with four radial veins), one species of Phulia Unfraphulia) at the other end (with two radial veins), and Phulia (Phulia), a second species of Phulia (Infraphulia), and Pirercolias (Pierphulia, new subgenus), in the middle (with three radial veins). It became quite obvious, if we were going to attain a nearly natural classification, that all of these characters had to be considered and used. The result was that sometimes special importance was given to a single structure in one genus (or in several genera) and that | much less importance had to be given that same structure in another | genus (or in several other genera). CERTAIN BUTTERFLY GENERA—FIELD 105 Two of the genera treated below are each divided into two subgenera upon characters that certainly would be considered of generic instead of subgeneric value if it were not for the fact that they could not be separated upon habitus. It did not seem logical, for example, to treat the subgenera Phulia and Infraphulia as separate genera in spite of the several really important differences in male and female genitalia and venation— simply because they look so very much alike, Thus, habitus was regarded as being of great importance and genera were not divided unless differences in habitus supported the other reasons for such division. On the other hand, habitus differences alone were not regarded as sufficient to divide genera. Thus, the two species of Baltia, although differing greatly in habitus (one species resembling Pzercolias and the other somewhat resembling Phulia) were retained together in the genus Baltia because other characters did not support a division based upon habitus differences either on the generic or the subgeneric level. The species here transferred to Piercolias (Pierphulia) were taken from Phulia, where they have hitherto been placed, because of the differences in the uncus of the male, in the inner genital plates of the female, and in other characters which were supported by habitus differences. On the other hand they could not be regarded as typical Piercolias (except on habitus characters) because of some differences in these same structures and in venation. All of the genera treated here are Neotropical in distribution except Baltia, which occurs only in the Himalayas of central Asia. Klots (loc. cit., p. 219) says “whether there is a real relationship between Baltia and Phulia or whether the resemblances are merely to be regarded as similar developments in the same type of environment is a matter of doubt.”” Klots had the false notion that Phulia and Baltia both lacked pulvilli and paronychia. These structures are present in all of the genera here treated but are sometimes so greatly reduced in Phulia, Peircolias, and Baltia that they cannot be seen without dissection and great magnification. Baltia, however, in the majority of its characters, certainly seems more closely related to these Neotropical genera with which Klots has placed it than to any of the Old World genera. Of the Neotropical genera, Theochila is found in the southern nontropical parts of Brazil and eastern Argentina; Tatochila in Colombia, Ecuador, Peru, Bolivia, Argentina, and south to Tierra del Fuego; Hypsochila in the Andes of Bolivia and Peru, south to the high elevations in central Chile and west-central Argentina, and south from there to Tierra del Fuego; Phulia in the high Andes of Bolivia and Peru, south to the high elevations in central Chile and west- central Argentina, and Piercolias in the Andes of Bolivia and Peru, south into northern Chile. 106 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 In the treatment below, the species belonging to each of the genera are neither described nor discussed, but are merely assigned to their proper place. Subspecies are not listed at all in this paper. Com- plete specific accounts will be offered at a later date by both the present writer and by Prof. José Herrera-G. of the University of Chile. Key to the genera and subgenera treated in this paper 1. Tibial spurs present on mid and hind legs ......4..24e08-6 3 tates Tibial spurs absent on*mid ‘andhind legs 's0'5 2°52). ee Ete Se Sat} 2. Male with uncus distinctly shorter than its cranteat width (dorsal or ventral view) (fig. 9); female with inner genital plate having a finger-like process that is nearly parallel to the anterior lobe (fig. 34); claw short and thick, only slightly more than twice as long as its width at base (fig. 25). Theochila (p. 106) Male with uncus at least as long as its greatest width (dorsal or ventral view) (fig. 10); female with inner genital plate having a finger-like process that is not parallel to anterior lobe (fig. 35); claw nearly three times ag long as its width at base (fig. 26). . . . .. . . . . Tatochila (p. 108) 3., Korewing with four radial veins . 21 .- is. «se wens), euisuseld afi tale iogihiees 4 Forewing with two or three radial veins . . . asters Set 4. Forewing with vein M, from apex of cell or om sae near nen of cell (con- nate or nearly connate with base of R3-+ Rays+ My) (fig. 19). Hypsochila (p. 110) Forewing with Mg; from distinctly below apex of cell (fig. 23) (see third GUOIGE) si aieia 4, PHULIA (PHULIA) 2. TATOCHILA “5. PHULIA (INFRAPHULIA) Ficures 1-5.—Explanation on facing page. 124 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 EXpLaNaTION OF Ficures 6-16 Figures 6-8 are of male genitalia in lateral view with aedeagi removed. In figure 7 the subscaphium is shown both fastened to anal tube in lateral view and detached and flat. Figures 9-16 are ventral views of the uncus and tegumen. Ficure 6.—Ptercolias (Pierphulia) nysiella (Rober). Type species of Pterphulia, new subgenus. Drawn from Preparation 5343. Ficure 7.—Piercolias (Piercolias) huanaco (Staudinger). Type species of Piercolias Staudinger. Drawn from Preparation 155. Ficure 8.—Baltia shawi (Bates). Type species of Baltia Moore. Drawn from Prepara- tion 5411. Ficure 9.—Theochila maenacte (Boisduval). Drawn from Preparation 107. Ficure 10.—Tatochila autodice (Hitbner). Drawn from Preparation 22. Ficure 11.—Hypsochila wagenknechti (Ureta). Drawn from Preparation 31. Ficure 12.—Phulia (Phulia) nymphula (Blanchard). Drawn from Preparation 299. Ficure 13.—Phulia (Infraphulia) illimani Weymer. Drawn from Preparation 169. Ficure 14.—Piercolias (Pierphulia) nysiella (R6ber). Drawn from Preparation 5343. Ficure 15.—Piercolias (Piercolias) huanaco (Staudinger). Drawn from Preparation 5411. Ficure 16.—Baltia shawi (Bates). Drawn from Preparation 5411. CERTAIN BUTTERFLY GENERA—FIELD 125 6. PIERCOLIAS (PIERPHULIA) Z PIERCOLIAS ieconia. Sa Voss eso niun « FACACHEA HYPSOCHILA PHULIA. ae BALTIA i) ee) PHULIA NF. IA (INFRAPHUL PIERCOLIAS (PHULIA) p IERCOLIAS (PIERCOLIAS) (PIERPHULIA) = Ficures 6-16.—Explanation on facing page. 126 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 ExpLaNaTIon oF Ficures 17-24 The figures are of wing venation and are slightly larger than natural size. Ficure 17.—Theochila maenacte (Boisduval). Drawn from Slide 1148. Ficure 18.—Tatochila autodice (Hiibner). Drawn from Slide 1151. Ficure 19.—Hypsochila wagenknechti (Ureta). Drawn from Slide 1159. Ficure 20.—Phulia (Phulia) nymphula (Blanchard). Drawn from Slide 114. Ficure 21.—Phulia (Infraphulia) illimant Weymer. Drawn from Slide 169. Ficure 22.—Piercolias (Pierphulia) nysias (Weymer). Drawn from Slide 180. Ficure 23.—Piercolias (Piercolias) huanaco (Staudinger). Drawn from Slide 155. Ficure 24.—Baltia shawi (Bates). Drawn from Slide 3810. CERTAIN BUTTERFLY GENERA—-FIELD 127: Sc R, Ro Rz Ry S¢ $ R 34, 24,4 Cup IZ THEOCHILA 18. TATOCHILA BS 20. PHULIA WS (PHULIA) 2), PHULIA (INFRAPHULIA) 19. HYPSOCHILA LS 22. PIERCOLIAS (PIERPHULIA), Ficures 17-24.—Explanation on facing page. a 4 24. BALTIA 23. PIERCOLIAS (PIERCOLIAS) 128 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 EXPLANATION OF Ficures 25-33 Figures 25 through 32 are greatly enlarged outline drawings of the claws, paronychia, and pulvilli. Setae and other armature are omitted, except on the paronychium in figure 27. All drawings are to the scale shown in figure 28. Figure 33 is of the female genitalia showing lateral view, except for the bursa copulatrix, which is turned to show the signum in ventral view. The 7th tergite and the outer genital plate are shown only in dotted outlines to allow other structures to show up more clearly. Ficure 25.—Theochila maenacte (Boisduval.) Drawn from Preparation 1148. Ficure 26.—Tatochila autodice (Hibner). Drawn from Preparation 1151. Ficure 27.—Hypsochila wagenknechti (Ureta). Drawn from Preparation 1160. Ficure 28.—Phulia (Phulia) nymphula nymphaea (Staudinger). Drawn from Prepara- tion 1172. Ficure 29.—Phulia (Infraphulia) illimanit Weymer. Drawn from Preparation 1169. Ficure 30.—Piercolias (Pierphulia) nysiella (Reber). Drawn from Preparation 1165. Ficure 31.—Piercolias (Piercolias) huanaco (Staudinger). Drawn from Preparation 1167. Ficure 32.—Baltia shawi (Bates.) Drawn from Preparation 1168. Ficure 33.—Baliia shawi (Bates). Drawn from Preparation 5412. CERTAIN BUTTERFLY GENERA—FIELD 129 OL 25. THEOCHILA EO OcneS 27 HYPSOCGHILA = Se =< | a) | ee cc : 30. PIERCOLIAS Ce CC 28. PHULIA 29, PHULIA - (PIERPHULIA) (PHULIA) ~~" (INFRAPHULIA) (n 31. PIERCOLIAS (PIERCOLIAS) a ( eG 32. BALTIA Ficures 25-33.—Explanation on facing page. 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GOVERNMENT PRINTING OFFICE: 1958 ~ “w - PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM issued WA SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1958 No. 3397 THE SYSTEMATIC POSITION OF THE BIRD GENUS APALOPTERON By H. G. Deignan The Ornithological Society of Japan’s (1942) list of Japanese birds indicates that, by 1942, 100 species of birds had been reported from the Bonin Islands, an oceanic group of volcanic origin lying about 500-600 nautical miles southeast of Yokohama. Of these, only 19 species (four of them by then extinct on the islands) were known to have bred. Of the 19, eight pelagic forms (two albatrosses, four shearwaters, one gannet, and a tern) may be disregarded at this time. Of the land birds, seven are mere races of species common in the Japanese Archipelago (one hawk, one pigeon, one bulbul, one crow, one thrush, one warbler, and one ereenfinch), while two (a pigeon and a hawfinch), extinct and not examined by me, may be presumed to have had similar origin. An anomalous element in the avifauna is found in the former presence of a night heron characteristic of the coasts and islands of the southwestern Pacific and otherwise not occurring north of the Palaus and the Philippines. Finally, there is the genus Apalopteron Bonaparte, the subject of these remarks. First named I[zos familiaris by its discoverer, F. H. von Kittlitz,' and considered a bulbul, it was removed to the ‘“Timalidae’’ by Bary tesa aap. Sci. St. Petersbourg, vol. 1, pt. 3, p. 235, pl. 18, 1830. 136 134 PROCEEDINGS OF THE NATIONAL MUSEUM—DEIGNAN vot. 108 Bonaparte? in 1854 as a monotypic genus, Apalopteron. Sharpe (1882, p. 120, footnote) restored it to the bulbuls as a member of the type genus Pycnonotus, and the most recent Japanese writers, using the name Apalopteron, have retained it in that family. Delacour (1946, pp. 21, 29), on the other hand, has considered it to be a timaliine closely related to the genera Actinodura and Minla. It is my conten- tion, however, that Apalopteron is in fact a fairly typical genus of the Australasian Meliphagidae or honey-eaters. The true meliphagid tongue, fringed and quadrifid at its distal end, has been most recently discussed and portrayed by Scharnke (1931, pp. 454-466) in the genera Myzomela, Myza, Melidectes, Orodytes, Philemon, Xanthotis, Ptiloprora, and Toxrorhamphus; by the same author (Scharnke, 1932, pp. 117-119) in Promerops; and by Dorst (1952, pp. 185-214) in Meliphaga, Gliciphila, Melithreptus, Zantho- miza, Meliornis, and Melipotes. Comparison of the flattened tongue of Apalopteron (pl. 1) with the drawings of the tongues of Myzomela and Philemon (Scharnke, 1931, pp. 456, 457) and the schematized drawing of the tongue of Meliphaga (Dorst, 1952, p. 187) will show that all are modeled upon a common pattern. The Meliphagidae are unusual, if not unique, among oscinine birds by their pervious nostrils. I have noted this character in freshly col- lected specimens representing the Australian genera Melithreptus, Entomyzon, Ramsayornis, Conopophila, Myzomela, Meliphaga, Lich- mera, Myzantha, and Philemon. Specimens of Apalopteron in the U. S. National Museum that quite certainly have never had the nostrils pierced by a needle are similarly devoid of the narial septum. Such unspecialized genera of the Meliphagidae as Myzomela, Lichmera, Ramsayornis, Conopophila, Meliphaga, et al. have the tarsus in the adult so obscurely scutellate as to appear booted (bila- minate behind), although scutellation is usually apparent in the young. The tarsus of adult Apalopteron shows the same quasi-booted aspect, and it is interesting to note that, according to a recent communication from Dr. Yamashina, scutellation cannot be seen even in the newly hatched chick. In its gross external features, Apalopteron certainly more nearly resembles the unspecialized honey-eaters than it does any member of either the Pycnonotidae or the Timaliinae. Its but slightly decurved bill, with strongly operculate nostrils, is not strikingly different from that of Liehmera, while its general coloration and pattern about the head are reminiscent of those found especially in certain species of Meliphaga (e. g., M. chryseps and M. melanops). ‘The short, some- what recurved, bristle-like feathers that appear on the front and 2 Comptes Rendus, Acad. Sci. Paris, vol. 38, pp. 54, 59, 1854. PROC. U.S. NAT. MUS. VOL. 108 DEIGNAN—PLATE 1 E i l E Prate 1.—Dorsal view of tongue of subadult Apalopreron familiares. Magnification, < 15. BIRD GENUS APALOPTERON 135 throat of Apalopteron seem to be identical with those seen in the two above-named species and many others. Yamashina (1930, pp. 332-334) has reported upon six nests of Apalopteron, each of which contained either two brown-spotted, greenish blue eggs or two naked nestlings. Five of the nests were con- structed principally of fibers of Pandanus and Liwvistona, one (in a Livistona flower) wholly of Livistona fibers. The cup-shaped nest, the materials used, the number of eggs, and the nakedness of the chicks all would, if the bird were native to Australia, be used to confirm the view that it is a typical honey-eater. (It should be mentioned, however, that the eggs of the Australian species of the unspecialized genera seem always to have a pinkish or buffy rather than a greenish blue ground color.) The food habits of Apalopteron, and its probable role as a pollinator of one or more species of ornithophilous plants, are of prime im- portance. Dr. Yamashina has written to me that his visits to the Bonins, in winter and spring, did not coincide with the true season of flowers, and that he found the birds then feeding on insects and fruits: ‘. . . when we broke a large fruit of Pandanus, Apalopteron gathered to eat its seeds.” He adds that Apalopteron (in captivity) likes sweet food and delights in sugar-water or honey added to its diet. Parenthetically, it may be noted that insects and fruit seem to be the primary foods of many of the Meliphagidsae, with pollen and nectar only secondary ones. At discovery the Bonins were described as wholly overgrown with a dripping forest of tall trees. The endemic flora was of mixed proven- ience, some of the genera having originated in temperate eastern Asia, while others, perhaps a majority, were unquestionably of Oceanic origin and here reached their northernmost outpost from the Papua- sian matrix. Hosokawa (1934, pp. 201-209, 657-670) has discussed the phytogeography of the Bonins and has pointed out that, among the botanical curiosities of the islands, are a palm, Cyphokentia (a genus otherwise restricted to New Caledonia and the Samoan Islands), and an arborescent lobelioid (whose nearest relatives are restricted to the Hawaiian Islands). Endemic plants of Oceanic origin that may have had a special relationship with Apalopteron are Myoporum boninense (northernmost representative of a Papuasian family highly attractive to the Meliphagidae) and the Lobelia (whose allies are favored feeding flowers for the Hawaiian Drepaniidae). While scttlement of the islands has brought about immense alteration of the flora, it may be supposed that Apalopteron has benefitted by the introduction of such fruits as the banana, orange, mango, papaya, guava, longan, persim- mon, and passionfruit, to say nothing of the ornamental, often nectar- iferous, plants common to subtropical gardens. 136 PROCEEDINGS OF THE NATIONAL MUSEUM—DEIGNAN vot. 108 ACKNOWLEDGMENTS: For comment and advice, I am indebted to Dr. Ernst Mayr of the Museum of Comparative Zodlogy and Dr. Austin L. Rand of the Chicago Natural History Museum, both of whom have had broad experience with the Meliphagidae in the field and in the laboratory. To the Marquess Yoshimaro Yamashina I owe thanks for information on specimens in his collection and un- published observations on the living bird, and to Dr. F. Raymond Fosberg of the National Research Council and Dr. Egbert H. Walker of the U.S. National Museum for guidance in the botanical literature. Finally, I am indebted to Mrs. Patricia Isham of the U.S. National Museum for the drawing of the tongue of Apalopteron. References Drracour, JEAN 1946. Les timaliinés. L’Ois. Rev. Francaise d’Orn., new ser., vol. 16, pp. 7-36. Dorst, JEAN 1952. Contribution a l’étude de la langue des méliphagidés. L’Ois. Rev. Frangaise d’Orn., vol. 22, pp. 185-214, 14 figs. Hosokawa, TAKAHIDE 1934. Phytogeographical relationship between the Bonin and the Marianne Islands .... Journ. Soc. Trop. Agr., vol. 6, pp. 201-209, 657-670. ORNITHOLOGICAL Society or JAPAN 1942. A hand-list of the Japanese birds, ed. 3, 238 pp. ScHARNKE, Hans 1931. Beitrige zur Morphologie und Entwicklungsgeschichte der Zunge der Trochilidae, Meliphagidae und Picidae. Journ. f. Orn., vol. 79, pp. 425-491, 54 figs. 1932. Ueber den Bau der Zunge der Nectariniidae, Promeropidae und Drepanidae nebst Bemerkungen zur Systematik der bliiten- suchenden Passeres. Journ. f. Orn., vol. 80, pp. 114-123, 8 figs. SHARPE, RicHarp BowpLuR 1882. Passeriformes. Coliomorphae. Pt. 3, Timeliidae. Catalogue of the birds in the British Museum, vol. 6, xiii+420 pp., 18 pls. YAMASHINA, Y. 1930. Birds of Parry-Group, Bonin Islands. Tori, vol. 6, pp. 323-340, illus. U. S. GOVERNMENT PRINTING OFFICE: 1958 PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM aN by the SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1958 No. 3398 A REVIEW OF THE COPEPOD GENUS RIDGEWAYIA (CALA- NOIDA) WITH DESCRIPTIONS OF NEW SPECIES FROM THE DRY TORTUGAS, FLORIDA By Mitprep Stratton Witson* The calanoid copepods discussed herein represent a small number of incompletely known, closely allied species for which three generic names have been proposed. ‘To this list are added two new species from collections made among the keys of the Dry Tortugas, Florida, by Mr. Clarence Shoemaker of the United States National Museum in the course of an amphipod survey of that region (Shoemaker, 1933, 1956). The debris washed from rocks and corraline algae taken at very shallow depths of a few inches to a few feet has yielded not only amphipods but other Crustacea among which were many small copepods. These included mostly Harpacticoida and Cyclopoida characteristic of such a habitat, but some Calanoida were also present. Among these were adults of the two new species and a few subadult males of an unknown species that are herein assigned to the genus Ridgewayia. This rather unusual genus has been known since 1903 when Thomp- son and Scott described the female of R. typica from Ceylon. Since then, closely allied species have been described as the types of new genera without comment on their obvious relationship to Ridgewayra. 1Collaborator, SmithsonianfInstitution, 137 138 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Through a study of literature and comparison of descriptions with the Tortugas material, it is apparent that Lampoidopus marki Esterly from Bermuda and Suezia canalis Gurney from the Suez Canal are very closely allied to Ridgewayia typica. Of these species, only marki is known from both sexes, and has been listed in literature beyond the original record. Only the female of typica has been described; canalis is known only from the male. A single male speci- men, representing either an unknown species or the male of typica, has been briefly described from Madras as Suezia sp. by Krishnaswamy (1953). Both this Madras male and that of canalis are relatable to the female of Ridgewayia typica through knowledge of both sexes of marki and the new Tortugas species. On the basis of present know]l- edge, there seems little justification for maintaining three generic names for this small group of species, and they are all herein referred to Ridgewayra (R. typica, R. marki, and R. canalis). Specimens of only the new species have been available for study. The specific diagnoses included here are all based upon the original descriptions (text and figures); note is made of characters omitted in these descriptions. Generic synonymy The results of study of the literature and of the available specimens do not permit an absolute, unqualified synonymy of the generic names that have been proposed. Therefore this study is presented with the recognition and suggestion that Lampoidopus and Suezia may with increased knowledge be fully and satisfactorily separable generically from Ridgewayia. That this is not now possible is due largely to the fact that in Ridgewayia and Suezia only the one sex is known. If I were presenting a paper dealing only with literature, or with speci- mens of the same sexes as were originally available for the described species, 1t would seem sufficient to point out the hitherto unsuspected relationship of Ridgewayia typica, Lampoidopus marki and Suezia canalis. Since, however, there exists the primary problem of placing two new species as accurately as possible, it has been necessary to consider critically the question of whether there is any valid objective reason for not placing them in the genus of prior date. The generic diagnosis given for Ridgewayia by Thompson and Scott (1903) confused the exopod and endopod of the antenna, and gave no information for the third leg of typica. It is also possible that the presence of an inner seta on the first basipod segments of legs 1 and 2 may have been overlooked. It is doubtful if the setation shown in the figure of the antennule is entirely exact. Otherwise the descrip- tion is complete and accompanied by excellent figures. It affords therefore an adequate basis of support for inclusion in the genus Ridgewayia of the females of the new Tortugas species and of Esterly’s COPEPOD GENUS RIDGEWAYIA—-WILSON 139 (1911) Bermuda species marki. Among calanoids, the modification of the female fifth leg is distinctive for these species. That of R. typica differs from that of the three American species only in having a shorter inner apical spine on the third exopod segment. This and other differentiating characters are either interpretable as specific, or concern structural features that have been sketchily presented, or could have been overlooked. These characters and features are pointed out in the specific diagnoses and discussion. When the similarity of the basic characters is considered, it does not seem possible to me that there is any choice other than to refer the - Tortugas material to Ridgewayia. It seems inconceivable that Esterly would have erected a new genus for his Bermuda speci- mens had he been aware of and considered the description of R. typica. Since the male of typica is still unknown, there exists no more justification for the genus Lampoidopus today than there did in 1911. Gurney (1927) appears to have been unaware of Esterly’s paper, since he failed to point out not only the similarity of his Suez Canal specimens with the males described by Esterly, but in the same paper he named a new species of Pseudocyclops without noting its striking similarity to P. magnus described in Esterly’s Bermuda paper. It also seems inconceivable that if Gurney had referred in his study to both Esterly’s and Thompson and Scott’s papers that he would have failed to note the relationship of his specimens to Ridge- wayia, as shown through Esterly’s description of both sexes. Krish- naswamy also makes no mention of Ridgewayia or of Lampoidopus. Thus, all the authors who have reported species of this group since the original description of Ridgewayia typica have failed to relate their material to it. Sewell (1929, 1932, 1948), in summaries of copepods of Indian waters, has not included R. typica, although Suezia is men- tioned. The genus Ridgewayia has therefore in effect become buried in the literature. The generic diagnosis given herein for Ridgewayra is a composite of the characters of both sexes of all the known species. It is detailed, contains exceptions and alternatives, and points out wherein know!l- edge is not complete for some species. Such a diagnosis is intended to form a working basis for the present report and also to be useful in future studies. In searching for the characters that indicate rela- tionship of a group of calanoid species to one another, either on the generic or familial level, it is necessary to consider every appendage. In the species dealt with here, the evidence for their very close rela- tionship is found in all appendages. Particularly noteworthy is the correlation of all oral appendages, the maxilliped and the first and fifth legs. The characteristics of these appendages in large groups may define a family, but they are also highly pertinent at the generic level 140 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 in the Calanoida. It remains for future studies and accurate, detailed knowledge of both sexes of all the species to determine whether we are dealing here with more than one genus. Most of the differences that have been found to exist on the basis of literature, need, in my opinion, to be verified by further examination of specimens. The determination of the proper generic status of all these species is primarily dependent upon knowledge of the male of typica. Its discovery should serve to differentiate Ridgewayia and Lampoidopus if they are truly generically distinct. It may or may not solve the status of Suezia canalis, which may be further dependent upon knowl- edge of the female and reexamination of the male. The two new Tortugas species would be referable to Lampoidopus if it is established as a separate genus. Subclass COPEPODA Order CALANOIDA Genus Ridgewayia Thompson and A. Scott Ridgewayia Thompson and A. Scott, 1903, p. 245. Lampoidopus Esterly, 1911, p. 219, new synonym. Suezia Gurney, 1927, p. 457, new synonym. Dracnosis (emended): Species small, approximate range of length between 0.6 and 1.0 mm. Metasome stout, 5—6-segmented, the somite of leg 1 distinctly or indistinctly separated, or fused with cephalic segment. Urosome less than half the length of metasome, 3—-4-segmented in female; if present, the fourth segment reduced to width of caudal rami; 4—5-segmented in male, fifth segment, if present, reduced as in female. Paired genital openings of female closely set (known only for the new species). Caudal rami longer than wide, with four terminal setae longer than rami, of these the second from the inner the longest (an outer spine also present in marki and in the new species). Rostrum downturned, broad at base, rounded or pointed distally; filaments lacking. Cephalic appendages all of primitive calanoid type, without re- duction, excessive modification, or sexual differentiation. Antennule reaching from near end of metasome to caudal rami; 25-26-segmented in female (21-22-segmented in canalis?); the three apical segments elongate. Left antennule male like that of female (where known); the right of 21-24 segments, with a moderately de- veloped geniculation, segmentation beyond this specialized joint varying from three to four segments (geniculation reportedly not present in canalis and Ridgewayia sp. (Madras)). COPEPOD GENUS RIDGEWAYIA—WILSON 141 Antenna: Outer ramus a little longer than inner, 7—8-segmented, only the apical segment elongate. Inner ramus 2-segmented, the first segment longer than the second, bearing 1-2 lateral setae; the second with terminal portion expanded into two setiferous lobes, the outer produced beyond the inner. Mandible: Masticatory blade not conspicuously expanded, pro- duced or grooved, with about 7-10 shallowly incised teeth not sepa- rated by any large gaps. Palp with broad, unsegmented basipod bearing 3-4 lateral setae. Exopod 4-segmented, the two distal seg- ments more or less defined. Endopod 2-segmented, with numerous apical setae. Maxillule (unknown for canalis): Basal portion well developed, with indistinct segmentation or none; its greater proximal part con- sisting of an expanded inner lobe (gnathobase) bearing short, spini- form setae; outer portion an unexpanded setiferous plate with 9-10 mostly elongate setae. Distad to this outer plate a reduced lobe (epipodite) bearing (in the new species) a single seta (no seta shown in illustrations for typica, marki). Inner side beyond gnathobase with two narrow laciniae, each bearing 4-5 setae. Beyond these laciniae, the basis elongated and produced on inner proximal side into setif- erous lobe. Endopod attached just outside this lobe, 2-segmented (apical segment not shown as separated in marki) ; proximal segment greatly enlarged with several (6-8) lateral setae; apical segment re- duced, with 5-6 setae. Exopod rather well developed, arising from the basis nearly opposite the inner laciniae (clearly separated in the new species, but not shown as demarcated in marki, typica), with lateral and apical groups of setae (exopod entirely unsegmented in typica and in the new species, but apex shown as demarcated in marki). Maxilla: The whole equal to or a little longer than the basal segment of the maxilliped, with three broad primary divisions ex- panded into small setiferous lobes, and a reduced terminal portion which may or may not be distinctly separated or segmented. Lobes 5-7 in number, of which 1-3 are on the proximal division; the other two divisions each with two lobes. Setae of lobes long and slender, none conspicuously more developed than the others. Maxilliped: Not conspicuously enlarged, but its length 3-4 times greater than that of the maxilla and longer than the first leg. The basipod of 2-3 segments (a proximal, nonsetiferous segment not shown in figures of other species, but present in the new species); second segment with four lobes or groups of setae. Endopod shorter than total basipod, of five well-defined segments; intercalated between it and basipod an incompletely separated segment bearing two setae. Legs 1-4 slender, with narrow connecting pieces; biramous, both rami 3-segmented. Inner seta on basipod segment 2 of leg 1. (Infor- 142 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 mation on setae of basipods otherwise incomplete in literature; the new species and marki have inner setae on both basipod segments of leg 1, and on basipod 1 of legs 2-4; legs 3 and 4 entirely unknown for canalis; leg 3 unknown for typica.) Exopod segments 1 and 2 with one outer spine and inner seta. Exopod segment 3, total number of outer and terminal spines (so far as known): three on legs 1 and 2, four on legs 3 and 4; these spines mostly without serrations or mem- branes; total number of inner setae: four on leg 1; five on legs 2-4. Endopod segment 1 with one inner seta on all legs; endopod segment 2 with two setae on legs 1 and 2, and on leg 3 where known; one or two setae on leg 4. Endopod segment 3, total of six setae on leg 1; eight on leg 2; five to eight on leg 3; six or seven on leg 4. Most of the setae divided into two joints, consisting of a stiff basal cylinder and a longer, flexible distal part, densely plumose. Leg 5 of female showing only slight specific differences, slender, symmetrical, with well-developed 3-segmented exopod and reduced 2-segmented endopod. Exopod modified; segment 3 constricted basally and set into narrowed, well-defined socket of segment 2; the outer, distal spine-bearing portion of segment 2 enlarged and consid- erably produced beyond this insertion. Exopod segment 3 with four spines and four inner marginal setae. HEndopod segment 1 reduced, without inner seta. Endopod segment 2 at least twice the length of first segment and usually longer; with seven setae (two outer, two apical, three inner). All setae with jointed bases. Leg 5 of male: Right and left basipod segment 1 fused or with definable connecting plate. Both rami modified and strongly asym- metrical. Right exopod 2-segmented, sometimes with imperfectly separated apex; second segment tending to elongation, with two outer marginal spines, or with one proximal spine and more distally placed spinous points; the segmental portion beyond proximal spime nar- rowed and more or less incurved. Left exopod 3-segmented, or third segment not entirely separated from second (as in Gurney’s figure for canalis); the whole third segment or apical portion considerably modi- fied, with a short but stout basal portion from which may extend spines, complex ornamented processes and fragmented membranes of irregular length. Endopods unsegmented; the right elongate, nearly as long as or longer than exopod; the left much shorter than the right (tending to be about half as long or less); either endopod entirely unarmed, or with setae, spines or processes. Typ spectus: Ridgewayia typica. Ridgewayia typica Thompson and A. Scott Ridgewayia typica Thompson and A. Scott, 1903, p. 245, pl. 1, figs. 1-13. Diacnosis (after Thompson and Scott): Female: Length about 0.85 mm. Metasome of six well-defined segments, the somite of leg 1 COPEPOD GENUS RIDGEWAYIA—WILSON 143 clearly separated; ventral margin of last segment deeply incised and showing in lateral view a hooklike process. Rostrum broad and pointed. Urosome 4-segmented; genital segment with distal hooklike process on right side. Caudal ramus twice as long as broad, with four long apical setae (details unknown). Antennule reaching to near end of metasome, 25-segmented. Antenna, exopod 8-segmented. Mazxilla with seven well-developed setiferous lobes, of which three belong to basal division. Leg 1 (from figure): Exopod 2, outer distal part with inner, narrow, serrate process about half length of outer spine. Inner apical spine of exopod 3 nearly as long as total exopod (about 17:20). For armature of legs 1-4, see under ‘‘Discussion”’ (p. 168). Leg 5: Exopod 3 with all four spines shorter than segment, the innermost apical spine of about same length as outer apical spine. Endopod 2, first inner seta placed above middle of segment, at point about 23 percent of total length of segment; first outer seta at point about 54 percent of length of segment. Male: Unknown. Distrisution: Ceylon, from the Muttuvaratu pearl oyster wash- ings. Ridgewayia sp. (Madras) Suezia sp., Krishnaswamy, 1953, p. 127, figs. 7-9. This reference is to a brief description of a single, damaged male specimen found in plankton collected on the Madras coast of India. The record is particularly interesting because it is from the same geographic region as R. typica, and raises the question of whether it may represent the unknown male of that species. Unfortunately, the description given is too incomplete to allow for any decision other than the relationship traceable through the species in which both sexes are known. The textual description is brief and so may be repeated here exactly as given (with correction of obvious printing errors). ‘Size: Male 0.679 mm. Colour: Formalin fixed specimens appear yellow. Salient features: General body shape cyclopiform. The last thoracic segment with a small projection. The basal 2 of the endopod of first leg with a curved spine. Antennule not prehensile. Fifth leg highly modified and biramous.”’ Under ‘‘Remarks’”’ there is a short comparison with Gurney’s figures of the fifth leg of Suezia canalis: “. . . the left leg resembles Gurney’s figures exactly while the right exopod differs from it in the second joint being shorter.” One of the figures given is of the last thoracic segment, and shows a spinous point somewhat similar to that of typica, but smaller. The 144 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 other figures are of the first and fifth legs. Both call for comments on detail of structure. In the figure of the first leg, both spines and processes are solidly inked, so that they cannot be distinguished from one another. The exopod would have the same number of spines as in the other species, if distinction had been made between the processes and spines. The long inner apical spine of exopod 3 is shorter than that shown for typica, being equal to the length of the third segment plus about half of segment 2, and may thus indicate that the two are not conspecific. In both the text and the figure there is some confusion as regards the distinction between the produced inner portion of the second basipod segment and the endopod. ‘The text refers to the spine of the ‘‘basal 2 of the endopod.’’ Obviously the spine referred to belongs to the second basipod segment and not to the endopod; it appears to be similar to the stout, curved seta of the other species. No inner setae are shown on endopod segments 1 and 2; this is probably due to incomplete delineation rather than actual lack of setae. Endopod segment 3 differs from all the other species in that there is a stout apical spine in the position of the spinous process, though because all processes and spines are solidly inked, the actual nature is not deter- minable from the illustration. There seem to be five inner setae, but these are somewhat indefinitely portrayed. The structure of the fifth leg is for the most part clear from the figure. As further pointed out in the discussion (p. 171), the left exopod is relatable to the American species and is not identical to that of canalis. The important characteristics of the leg are: Right and left basipods about equal in length; the first segments not fused; left segment 2 with inner seta. Right second exopod segment only little longer than first, with one prominent, proximally placed outer marginal spine; beyond this spine the margin with a pair of closely set points beyond which the segment is abruptly shortened, narrowed, and inwardly directed. Right endopod reaching beyond exopod, with three inner marginal setae. Left exopod shortened but 3-segmented, not reaching to end of right exopod; segments 1 and 2 with prominent outer spines about as long as total exopod and closely set to one another; exopod segment 3 reduced (detail not shown in figure). Left endopod reaching to end of exopod, unarmed. There is little doubt that this Madras specimen represents a different species from canalis or from any other in which males are known. Its occurrence near the type locality of typica makes it imperative that consideration be given to the possibility that it may be the male of that species. This point may or may not be establishable from a complete comparison of appendages. COPEPOD GENUS RIDGEWAYIA—WILSON 145 Ridgewayia canalis (Gurney), new combination Suezia canalis Gurney, 1927, p. 457, fig. 109. Diaenosis (after Gurney): Female: Unknown. Male: Length (2 specimens), 0.72, 0.74 mm. Body form “cyclopoid,’’ metasome 6-segmented, lateral wing of last segment with small, backwardly directed tooth. Rostrum pointed. Urosome 4-segmented, fifth seg- ment said to be “scarcely distinct.” Antennule reaching to about end of metasome; segmentation said to be “indistinct,” of 21 or 22 segments; the right not prehensile. (No direct reference made to left antennule.) Exopod of antenna 7-segmented. Mandible palp as for genus. Maxillule undescribed. Maxilla shorter than segment 1 of maxilliped; with five inner setiferous lobes, of which one belongs to basal division; apical portion of two segments. Maxilliped as for genus (setae of basal segment appear to be incompletely shown in illustration). Leg 1 (from fig. 109%): No inner seta shown on basipod 1; present on basipod 2. Exopods 1 and 2, outer margin with distal, serrate process, that of second segment about half length of outer spine. Exopod 38, inner apical spine shorter than segments 2+3, about 14:18. Endopod 1, outer margin with stout, partially separated lobe (as in new Tortugas species). Spines and setae as given for the genus. Leg 2 with inner seta on basipod segment 1; setation of exopods and endopod as given for the genus (see also p. 169). Legs 3 and 4 unknown, except that the rami are 3-segmented and the exopod spines lack hyaline membranes. Leg 5 (from fig. 109H,1): No spines or setae shown on basipods; first basipod segments at least partially separated by connecting plate; right and left basipods of nearly equal length. Right exopod 2, outer margin with one spine set near proximal fourth of segment, the inner margin incised at this point and the segment narrowed and tapered, forming an incurved prolongation; outer margin with two minute spinous points at middle and near tip of segment. Left leg shorter than right. Outer spine of exopod 1 reaching to near end of leg, evenly tapered, with wide, serrated fringe. Exopod 2 produced into stout lateral spinous point inside of which is a modified spine ornamented with outer hyaline flange and inner basal process (com- parable to outer spine of other species?). Inside this modified spine, a shorter produced portion of the segment shown as partially sepa- rated on the anterior side, divided into an outer, seemingly flattened structure with bifid tip, and an unmodified segmental inner portion (this appears to represent a reduced third segment, less complex in structure and armature than that of the other species). Endopods modified as in other species of the genus. The right elongate, reach- 455057—58——_2 146 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 ing to near end of exopod, inner margin with four short, thick setae disposed along its length, the distal the largest and armed marginally with a hyaline flange. Left endopod much shorter than right, but nearly as long as the shortened left exopod, its outer distal margin with two short lobed processes. DistrrpuTIon: Suez Canal. Original record from two specimens, taken in separate plankton collections at night, from Kabret and Ismailia. Remarks: Since only the female of Ridgewayia typica and the male of ?. canalis are known, the question of their possible conspeci- ficity has been considered in this study. It has been concluded from the following differences in the first leg and the cephalic appendages, exclusive of the antennule, that R. canalis is a distinct species and does not represent the male of R. typica: ANTENNA: Endopod 8-segmented in typica, 7-segmented in canalis; last seg- ment with 3 apical setae in typica, with 4 in canalis. Maxiuua: Seven lobes in typica, of which three belong to the basal division: five lobes in canalis, of which one belongs to the basal division. Lec 1: Exopod segment 1 without distal process on outer margin in typica, with process in canalis. Apical inner spine of exopod segment 3 longer than last two segments in typica, shorter in canalis. These characters have been taken from illustrations given in the descriptions of the two species. The characters are easily determined for the antenna and maxilla, and it is probable, unless immature specimens were originally studied, that examination of the species when again collected will show them to be as given. With the exception of the spine length, the characters of the first leg need most careful study from an advantageous view. Mere comparison of the figures may not necessarily give exact detail of the armature of the exopods. All of these differences should also help in identifying the female of canalis. Ridgewayia marki (Esterly), new combination Lampoidopus marki Esterly, 1911, p. 219, pl. 1, fig. 4; pl. 2, figs. 18, 14, 20, 21; pl. 8, figs. 25, 26, 28-31, 34; pl. 4, figs. 35, 38, 42.—Pinney, 1933, p. 142. Draenosts (after Esterly): Length of both sexes about 1.0 mm. Metasome 5-segmented, somite of leg 1 fused with cephalic segment. Rostrum broad and rounded at apex. Urosome female 3-segmented; male 4-segmented; caudal rami about 3.5 times as long as broad in both sexes. (See also p. 162.) Antennules reaching to end of caudal rami; female and left male 25-segmented. Right antennule male geniculate, 23-segmented, “terminal portion 4-jointed.’”’ Antenna, exopod 8-segmented. Max- COPEPOD GENUS RIDGEWAYIA—WILSON 147 illa with six well-developed lobes and unsegmented apical portion; basal division with two lobes. Leg 1 (from fig. 42): Exopod 2, outer distal part with flattened, serrate process, a slender spine (or process?) between it and outer spine, both shorter than outer spine. Inner apical spine of exopod 3 a little longer than exopod 2+3, equal to about 75 percent of total exopod length. (See p. 168 for detail of setal armature of legs 1-4.) Leg 5, female: Exopod 3 with the three outer spines shorter than segment; inner apical spine subequal to segment and about twice the length of outer apical spine; basal joints of all inner setae reaching beyond point of insertion of succeeding seta. Endopod segment 2, first inner seta placed a little above middle, at point about 44 percent of total length of segment; first outer seta placed below middle of segment, at point about 69 percent of total length of segment. Leg 5, male (rami separated in Esterly’s illustrations, and the basal segments of left leg incompletely shown; not possible to judge relative length of left and right sides): Right exopod, outer spine of segment 1 reaching to about middle of segment 2; tip of exopod 2 truncated, without lappet or partial division, with two outer spines, relative length of segment and first and second spines about 38:20:15. Left exopod 3-segmented; segment 1 with long, narrow, setiform outer spine that reaches a little beyond the second segment. Second seg- ment about twice the length of and much broader than first segment, its outer distal spine stout basally, tapered apically, its length only little more than half that of segment. Segment 3, membranes and processes elongated, length from base to tip exceeding that of segment 2 and reaching far beyond end of its outer spine. Endopods unarmed; the right elongate, club-shaped, reaching to beyond middle of last segment; the left shorter, length about twice its own width, reaching to near end of exopod 2. Distrrisution: Bermuda Islands. In cave on small ledge-like island across from Agar’s Island, at high tide (Esterly); in night plankton haul in Grasmere Cove, near shores of Bermuda (Pinney). Ridgewayia gracilis, new species Figures 1—27 SPECIMENS EXAMINED: 31 females, 40 males. Vicinity of Logger- head and Bush Keys, Dry Tortugas, off the southwestern coast of Florida, July 23—-Aug. 12, 1926, Clarence R. Shoemaker. Typrs: Holotype female (alcoholic) USNM 99511; allotype male (alcoholic) USNM 99512. Paratype specimens (slides and alcoholic) in U.S. National Museum collections. 148 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 DiaGNnostic CHARACTERS: Urosome female 4-segmented, male 5- segmented; the two middle caudal setae jointed basally. Antennules, female and left male, 26-segmented; segments 13-22 with partial rows of spinules. Right antennule male 23-segmented, with three segments beyond geniculation; segment preceding geniculation with distal longi- tudinal comblike row of spinules. Maxilla with six lobes, two on basal division. ‘Total number setae, endopod 3, legs 1-4: 6, 8, 8, 7. Leg 1, inner apical spine of exopod 3 subequal to segments 2+3. Leg 5 female, inner apical spine of exopod 3 longer than outer spine, about as long as segment. Leg 5 male, right exopod 2, first outer spine about twice the length of second spine; left exopod 3-segmented, outer spine of segment 2 reaching beyond segment 3; left endopod with movable basal process as long as the endopod. FEMALE Length, dorsal midline, 0.83-0.90 mm.; the greater number of specimens 0.86-—0.87 mm. Body slender and of distinct calanoid shape. (In a few specimens, the fore part is curved downwards so that the body does not have the usual erect appearance shown in fig. 2. Although the normal flexibility of the body segments or effect of the preservative might account for some of this, there is a real though small difference in the amount of the curvature of the fore part of the body in individual specimens.) Metasome 2.7—3 times the length of the urosome, with its greatest width at beginning of second segment; 6-segmented, the division be- tween the cephalic segment and that of leg 1 not so distinct as those of the other segments. Cephalic segment, in dorsal view, rounded anteriorly and tapered sharply outwards so that beyond the middle the segment is nearly as wide as the second segment; its length, in midline, a little greater than that of the other segments combined. Length of segment 2 a little less than that of segments 3 and 4 together (relative lengths, segments 2-4: 35:20:20). Segments 2—5 with lateral, apically acute, hyaline flanges. Last segment not expanded laterally, in dorsal view the ‘‘wings’”’ narrowed and slightly pointed; in lateral view, the wings show on the inner edge, three notches bearing minute hairs, the notch nearest the outer edge the largest and easily visible, the others seen only at high magnification (fig. 3). Urosome (fig. 1) 4-segmented, the genital segment the longest; the fourth segment very short, reduced to the width of the caudal rami, with which it is more or less fused medially, but clearly distinct outwardly. Relative lengths of the segments and rami (dorsal): 1 2 3 4 CR 28 13 10 4 16 COPEPOD GENUS RIDGEWAYIA—WILSON 149 Surface of all segments and of caudal rami, both dorsally and ventrally, covered by irregular groups of minute spinulose scales (not illustrated). Segments with nonserrate, inconspicuous fringe; second and third segments with a prominent proximal sclerotization on each side (fig. 1). In lateral view, the genital segment appearing rounded and only a little produced ventrally. External portion of the genital field simple (fig. 8), more or less defined by a cuticular sclerotization which is heavier in the posterior area; the distal half with a crosswise, asym- metrical opercular flap drawn out on the right side into a pointed process; the slit formed by the flap noticeable in lateral view (fig. 2); when turned semilaterally, the process of the right side prominent. Paired genital openings set close together, rather large, filling most of the area defined by the external sclerotization (outlines visible with oil immersion objectives, but structurally indistinct). Caudal ramus with its inner portion somewhat expanded proximally; the distal inner margin armed with fine hairs. In most specimens the rami a little divergent, but parallel in some specimens and closely set so that the inner expansions of the basal part cross over one another (the rami thus apparently with a somewhat flexible attachment). Caudal setae consisting of an outer, subterminal spine shorter than the ramus, and four long, plumose, terminal setae, the outer of which is shorter than and the inner subequal to the urosome. The two middle setae with thickened, jointed bases; both longer than the urosome; the innermost of these two setae the longer, jointed second- arily near its distal third and without hairs beyond this joint. A short seta with very long marginal hairs inserted dorsally between the bases of the innermost setae. Ventrally, two flat spinules overlying the bases of the setae (fig. 7). Rostrum (fig. 4) of the broad form characteristic of the genus, not demarcated at base, tapered to a rounded point. A pair of minute frontal hairs present above the base of the rostrum. Antennule reaching to about the end of the metasome; comprised of 26 clearly defined segments (fig. 14). The two proximal segments subequal in length to one another (fig. 16), wider and longer than most of the succeeding segments except the four apical segments, which are progressively narrowed and lengthened (fig. 21). Two setae on every segment except segments 1, 21, and 22 which have only one each, segment 2 which has four setae, and segment 26 which has one lateral and five terminal setae. On many segments, particularly in the midportion of the antennule, the proximal seta short and _ hairlike. Elongate setae (reaching at least beyond the succeeding four segments) on segments 4, 8, 10, 13, and 22; the longest of these on segment 4 (reaching to segment 12) and on segment 22 (reaching to end of 150 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 antennule; fig. 21). Some of the setae of apical segments modified by division into one or more joints (fig. 21). Aesthetes present on most of the segments, stouter than the setae and of uniform width throughout their length; those of the proximal segments (fig. 16) stouter than those beyond the midportion of the antennule; the longest that on segment 10 (reaching to the middle of segment 15). Segments 138-22 and 24-25 with small groups of surface spinules (figs. 15 and 21). Summary of setation of individual segments as follows (s=seta; a=aesthete): Dpto y Pon inet (uO Nis Oeiy a pyri Oli Mey) One Hl pclieyi aha. bogs eno: Scag 23 Qo seas. OOS. 628 Derr 28a 2s" e 2s 2s MSs a a Aa Wat ay Seah eka Wy aay ay an a ee fa 16T7 1S PIS 20" POhig 22 83 BA 2b, 26 2s) (28° 28.128 28) 8 8 . 25) 280 425.0 6S a oe a a a Antenna (fig. 20) with basipod of two well-defined segments, the proximal with a plumose inner seta, the distal with two unarmed setae. Exopod reaching beyond inner ramus by about half the length of its apical segment; 8-segmented, the first seven segments differing little in length from one another, but progressively narrowed from the basal segment, each with a single inner seta; apical segment about 3 times the length of the other segments and with four terminally placed setae. Endopod of two stoutly developed segments, the apical (measured to end of outer lobe) about three-fourths the length of the first; two setae at distal inner third of first segment; inner lobe of second seg- ment with eight setae graduated in length from outer to inner edge; the outer lobe with four setae. Apex of mandible blade (fig. 11) only slightly expanded and with shallowly incised teeth (fig. 10). Palp with expanded basipod armed with three inner setae; the four segments of exopod well defined, the proximal three with inner seta, the apical with three setae of which the outer is much shorter than the others; first segment of endopod shorter and broader than the apical, with four inner, distally placed setae; apical segment of endopod with two groups of terminal setae, six in the outer group and three in the inner. Ficures 1-13.—Ridgewayia gracilis, new species, female: 1, Distal part of metasome and urosome, dorsal; 2, habitus, lateral; 3, detail of metasomal wing, lateral; 4, outline of rostrum, ventral, with frontal hairs; 5, leg 5, detail of endopod; 6, leg 5; 7, distal part of urosome and caudal rami, ventral; 8, genital segment, ventral, with detail of operculum; 9, leg 1; 10, detail of edge of mandible blade; 11, mandible blade, with palp; 12, leg 2, exopod; 13, leg 4, with detail of marginal armature of inner apical spine. 151 COPEPOD GENUS RIDGEWAYIA—WILSON facing page. ayia gracilis, new species, female. Explanation on 1-13.— Ridgew FIGURES 152 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Mazxillule (fig. 17) with well-developed basipod but without defina- ble lines separating segments from one another. The first inner lobe (gnathobase) comparatively large, prominently produced, oval in out- line, bearing eight stout spines and five subapically placed setae. Just distal to this lobe two narrow laciniae, each bearing four apical setae. The proximal outer portion an unexpanded plate bearing nine setae, of which the distal six are greatly lengthened. Between this group of setae and the basal attachment of the exopod, a protrusion (epipo- dite?) bearing a single setae. Exopod and endopod borne on the dis- tal narrowed portion of the basipod, which has its inner part produced as a small lobe bearing four setae. The endopod 2-segmented; its proximal segment comparatively enlarged, with eight lateral setae; the apical segment reduced, with six setae. Exopod constricted be- yond its middle so as to form two setiferous portions, the proximal bearing five lateral setae; the distal somewhat expanded and bearing three lateral and three apical setae. Mazxilla (fig. 19) with six distinctly developed lobes of which two belong to the basal division, which is incompletely demarcated from the second. An accessory seta on a short stalk at the proximal base of the first lobe. The fifth lobe (the proximal of the third division) the largest. The number of setae on the lobes as follows (lobes num- bered from proximal to distal): Lobes: 1 2 eo 4 5 6 Apical setae: 4 2 2 3 4 3 Basal setae: 1 2 The reduced terminal portion of the maxilla indistinctly separated and segmented, with three setae. Maxilliped (fig. 18) with short, nonsetiferous basal segment (present in all dissections). Second segment with four lobes, the number of setae from proximal to distal lobe, 1, 2, 4, 3. The longest setae that of the first lobe and the proximal of the second lobe; both of these naked. Two of the setae of the distal lobe extremely reduced. Third segment with three plumose setae, between it and the endopod an incompletely separated segment, distinct neither from the endopod nor from the basipod. Endopod of five distinct segments bearing setae as follows: Ficures 14-27.—Ridgewayia gracilis, new species. 14-21, Female: 14, outline of anten- nule, showing segmentation; 15, antennule, segments 13-15; 16, antennule, segments 1-4; 17, maxillule, greatly enlarged; 18, maxilliped; 19, maxilla; 20, antenna; 21, anten- nule, apical segments 22-26. 22-27, Male: 22, right antennule, apical segments; 23, right antennule, segments 9-15; 24, leg 5, detail apex of left exopod, anteromedial view; 25, same, posterior view; 26, leg 5, detail right exopod and endopod, anterior view; 27, leg 5, posterior view. COPEPOD GENUS RIDGEWAYIA—WILSON 153 Ficures 14-27.—Ridgewayia gracilis, new species. 455057—58——3 f f ff $ f i eae Explanation on facing page. 154. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Segment: 1 2 3 4 5 Apical setae: 4 4 3 3 3 Basal setae: 1 1 Legs 1-4: Both basal segments of leg 1 with inner seta; only first segment of legs 2—4 with inner seta. First two segments of exopod of legs 1-4 with outer spine and inner seta; segment 1 of endopod with inner seta, segment 2 with two inner setae. Armature of third segments of both rami as follows (sp=spine; s=seta): Leg 1 Leg 2 Leg 8 Leg 4 Exopod: 3sp 4s 3sp 5s 4sp 5s 4sp 5s Endopod: 6s 83 8s 7s Leg 1 (fig. 9) with several distinctive modifications. Outer spines of exopod with marginal flanges and tipped with a hair. Distad to the spine a flattened process; the processes of segments 1 and 2 with serrate edges; that of segment 1 very small; that of segment 2 larger, about half the length of the outer spine, between it and the spine an unarmed spiniform process; the process of segment 3 extremely small, nonserrate and placed at the outer distal corner of the segment. Inner apical spine of third segment of exopod subequal in length to segments 2+3. Seta of basipod segment 2 long and characteristically curved over the first two endopod segments. Distal outer portion of endopod segment 1 with an enlarged, partially separated, lobelike process or extension, densely setose on its anterior side. Segment 3 of endopod reduced in width, its terminal outer margin extended into spinous process. Excepting the seta of endopod 1 of leg 1, all the setae of legs 1-5 jointed, usually below the middle. The proximal portion of these modified setae appear stiff and rodlike, and are sparsely plumose; the terminal flexible portion densely plumose. Spines of outer margins of exopods of legs 2 and 3 bladelike and unarmed (fig. 12), those of segments 1 and 2 elongate, reaching to near the base of the next spine or beyond, set into deeply defined sockets. The distal outer spine of segment 3 set in nearly terminal position with a prominent segmental process between it and the longer inner apical spine; inner spine about as long as its segment and unarmed. Spines of leg 4 (fig. 13) differing from those of legs 2 and 3 in having marginal flanges; the flange of the long inner apical spine of segment 3 armed with marginal hairs; near its distal fourth, these hairs cross over the surface of the spine to the edge of the inner margin. Leg 5 (fig. 6): Basipod segments without inner setae; segment 2 with small outer spinous process and submarginally placed seta, usually inwardly directed; surface of these segments with groups COPEPOD GENUS RIDGEWAYIA—WILSON 155 of small spinules. Exopod segments 1 and 2 with stout outer spines armed marginally with serrate flanges; segment 2 with an inner, jointed seta, the stiff basal portion of which reaches more than halfway to the base of the first seta of segment 3. Exopod segment 3 with a group of three outer, serrate spines, all shorter than the segment; the two proximal spines placed marginally, the distal apically; the fourth spine (inner apical) with outer hyaline flange, about as long as the segment and nearly twice the length of the distal outer spine (about 60:35). The basal portions of the jointed setae of the inner margin of exopod 3 all reach beyond the base of the succeeding seta. Segment 1 of the endopod without seta; its outer distal margin pro- duced to point. Segment 2 with seven jointed setae, the basal portion of each of the three inner setae reach beyond the base of the succeeding seta. The first outer seta placed just below the middle of the segment (at a point representing about 60 percent of the length of the inner margin of the segment); the first inner seta placed above the middle of the segment (at about 36 percent). Apical outer spinous process and basal portion of apical setae long, the process 20 percent, the setal bases 54 percent of the length of the inner margin of the segment. (See also p. 170.) MaLE Length, 0.8mm. Habitus asin female. Urosome 5-segmented, the first four segments subequal to one another in length, the fifth reduced as in the female. Caudal rami and setae exactly like those of female. Rostrum enlarged as in female, with broader tip and separated from forehead by a distinct surface demarcation across its entire base. All appendages except right antennule and fifth leg like those of female. Left antennule 26-segmented and with setation as in female. Right antennule with proximal segments as in female; midportion modified in that segments 13 and 14 are fused outwardly and im- perfectly divided from one another by a deep cleft beginning at the inner margin and extending irregularly into the middle of the segment (fig. 23). The antennule with 23 apparent free segments (counting 13 and 14 as two segments). The four apical segments elongated and of similar length (fig. 22); the point of geniculation between the proximal two of these segments (segments 20 and 21) so that there are three segments beyond the geniculation. Segment 20 with a longi- tudinal row of spinules arranged as a comb along the distal inner half. (See p. 163 for interpretation of segmentation.) Leg 5 (figs. 24-27): First basal segments fused, forming a narrow crosswise bar to which the enlarged second basal segments are at- tached. Right basipod 2 longer than left, each with outer distally placed spinous process and submarginal seta. Right exopod: Seg- 156 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 ments subequal in length, the second narrowed. Outer spine of segment 1 with serrate flange, reaching to end of segment 2. First outer spine of segment 2 similar in size and armature to that of seg- ment 1, second outer spine about half length of first; both spines reach beyond end of exopod. Apical part of segment 2 modified, an imperfectly separated, rounded serrate lappet on the inner margin opposite the placement of the second outer spine; beyond this lappet, the segment slightly incised on the anterior side (the possibility that this tip is at least partially movable is suggested by the various positions found in different dissections; the tip may or may not be interpretable as an imperfectly separated third segment). Left exopod: Segment 2 much stouter than segment 1, broadened and stout at its apex into which is set the highly modified third seg- ment; its outer distal spine narrowed and tapered, reaching beyond the tips of the processes of the third segment, armed only with outer marginal spinules. Third segment consisting of a shortened seg- mental portion clearly separated on the anterior side from the second segment (fig. 24) but imperfectly separated on the posteromedial side. The segmental portion of the surface of the anterior side deeply incised medially with heavy marginal sclerotizations and form- ing in part on its posterior side the base for the attachment of a set of thin, apically and irregularly fragmented membranes (or a single folded membrane) and three heavier, exceedingly flexible processes. Of these, the innermost simple in structure (setiform). One process, with an irregularly serrate, flared tip, has a broadened base which is set into a socket of the posteromedial portion of the segment (fig. 25). The other process is deeply widened at its base and attached inside the segment on the anterior side (fig. 24), below its attachment it is abruptly contracted into a long narrow setiform process with a slightly widened tip which is split near its end. These processes appeared in dissections separated as shown in figure 25, or twined around one another as in figure 24. In interpreting the figures given here, it must be remembered that the membranes, though perhaps representing only one single struc- ture, are irregularly fragmented and folded, and their appearance in any one dissection may be different from any other dissection. In figure 24, the membrane shown is only the expanded outer portion of that shown in figure 25; the edge appeared in all dissections to be strengthened by a heavy band. Right endopod elongate-narrow, reaching to near end of exopod, the tip partially split; anterior side set basally with groups of surface spinules (fig. 26). Left endopod short, reaching only little beyond exopod segment 1; inner margin with two movable processes; the COPEPOD GENUS RIDGEWAYIA—WILSON 157 distal process short; the basal process nearly as long as the endopod (in dissections, this process found expanded as shown in fig. 27, or entirely “pulled in” and lying along the margin of the endopod). Ridgewayia shoemakeri, new species Figures 28-35 SPECIMENS EXAMINED: 2 females, 13 males. Occurring with R. gracilis in the vicinity of Loggerhead and Bush Keys, Dry Tortugas, Fla., July 23-Aug. 12, 1926, Clarence R. Shoemaker. Typss: Holotype female (slides) USNM 99517; allotype male (slides) USNM 99518. Paratype specimens (slides and alcoholic) in U.S. National Museum collections. DraaNnostic cHARACTERS: Urosome female 4-segmented, segments with distal serrate fringe; male 5-segmented; the two middle caudal setae not jointed basally in either sex. Antennules, female and left male, 26-segmented; segments 13-22 with row of spinules extending across distal edge of segment. Right antennule male 24-segmented, with four segments beyond geniculation; segment preceding genicula- tion without lateral comblike row of spinules. Maxilla with six inner lobes, of which two belong to basal division. Total number setae, endopod 3, legs 1-4: 6, 8, 8, 7. Leg 5 female, inner apical spine exopod 3 longer than outer spines, about as long as segment. Leg 5 male, right exopod 2, first outer spine only little longer than second spine; left exopod 3-segmented, outer spine of segment 2 reaching to end of modified processes of segment 3; right endopod with slender outer seta; left endopod unarmed. FEMALE Length, dorsal midline, 0.67-0.68 mm. Body slender, but lacking the erect appearance of gracilis, due to the strong curvature of the forepart of the cephalic segment (as shown for male, fig. 32). Pro- portions and segmentation of metasome very similar to those of gracilis. Cephalic segment longer than rest of metasome (about 80:71). Segments 5 and 6 both reduced in midline; wings of last segment like those of gracilis, except that the outer notch of the posterior edge is not enlarged. Urosome segmented as in gracilis, the fourth being likewise reduced to the width of the caudal rami. Relative lengths of the segments and rami: 2 3 4 CR 20 8 6 2 13 158 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Posterior edges of the segments with a complete dorsal fringe, that of segments 1-2 indistinctly serrate, that of segment 3 deeply serrate. External genital field defined by sclerotization, the distal edge of operculum a flaplike opening, the right side rounded (fig. 31). Caudal setae exactly like those of gracilis, except that the two long middie setae are not jointed at their bases. Appendages differing very little from those of gracilis. Antennule longer, reaching to the end of the genital segment; clearly 26-seg- mented, the proportions of the segments and numerical setation exactly like gracilis except that the last four segments tend to greater elongation, and segment 24 is nearly subequal to 25 and 26; the long setae of segments 4, 8, 10, 138, and 22 comparatively shorter; the surface spinules of segments 13-22 arranged in single rows extending across the entire distal edge of the segment, size of spinules varying from segment to segment. Segmentation and setation of antenna and mandible palp like gracilis. Maxillule like gracilis except that the first of the two laciniae just distad to the gnathobase has five ‘instead of four setae. Maxilla also with six large lobes, of which two belong to the basal portion; the apical portion clearly 2-segmented, a difference from gracilis that might be an individual variation or due to position in mounting. Maxilliped exactly like that of gracilis. Legs 1-4 identical to gracilis in arrangement, structure, and number of setae and spines. Leg 1 differing slightly in that the spinous process between the outer spine and flattened process of exopod segment 2 is as long as the distal process instead of shorter; the inner apical spine of segment 3 about as long as segments 2+3. Leg 5 (fig. 28) very similar to that of gracilis. Basal portions of all jointed setae of exopod and endopod comparatively shorter; that of exopod 2 hardly reaching beyond segment, first two setae of exopod 3 reaching about to point of insertion of succeeding setae. Endopod segment 2, first outer seta set below middle of segment, at point about 64 percent of total length; first inner seta set a little above the middle, at a point about 42 percent of total length; spinous process of outer margin short, only about 10 percent of total length of segment. (See also p. 170.) Ficures 28-37.—Ridgewayia shoemakeri, new species, and Ridgewayia sp. (Tortugas). 28-35: R. shoemakeri, new species: 28, female, leg 5; 29, male, right antennule, apical segments; 30, male, right antennule, segments 9-16, with armature of segments 10, 13, 14; 31, female, genital operculum; 32, male, habitus, lateral; 33, male, leg 5, detail apex of left exopod, anterior view; 34, same, posteromedial view; 35, male, leg 5, anterior view, with detail apex of right exopod. 36, 37, Ridgewayia sp. (Tortugas), male copepodid stage V: 36, leg 5, posterior view; 37, left antennule, segments 8-13, with armature of segment 9. COPEPOD GENUS RIDGEWAYIA—WILSON 159 Ficures 28-37.—Ridgewayia shoemakeri, new species, and Ridgewayia sp. (Tortugas). Explanation on facing page. 160 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 MALE Length, 0.63-0.68 mm. Forepart of body strongly curved (fig. 32). Urosome 5-segmented, the fifth segment reduced. Antennules reach- ing beyond the metasome; the left like that of the female, the right (fig. 29) with 24 free segments, four beyond the point of geniculation. (See pp. 163-167 for comparison of segmentation with gracilis.) Segments 13 and 14 modified as in gracilis but completely separated (fig. 30). Segment preceding the geniculation without a comblike group of spinules. Cephalic appendages and legs 1-4 as in female. Leg 5 (figs. 33-35): First basal segments reduced in size and com- pletely fused. Second basal segments of nearly equal size. Right exo- pod: Segment 1 with medial inner expansion and a very fine acces- sory seta; outer spine reaching just beyond base of first spine of second segment, with a broad, serrate hyaline flange. The two outer spines of segment 2 subequal to one another in length, both reaching beyond the end of the segment, and with serrate flanges. Beyond insertion of second spine, the inner margin of segment with a cleft and a serrate lappet which appears hardly separated from the anterior surface; tip of segment minutely serrate. Left exopod: Segment 1 reduced in both width and length, with a stout outer spine reaching beyond segment 2 and armed with hyaline flange. Segment 2 broadened and thickened at its apex; its outer spine thick and irregular, without marginal flange or serrations, longer than its segment and reaching to about the same point as the longest of the processes of segment 3. Segment 3 distinctly separated from segment 2 on anterior side and very nearly so on posterior side. Membranes folded and irregular in length and fragmentation as in gracilis (figs. 33, 34). In addition to membranes, a plumose seta as in gracilis, and two modified processes. Of these, one simple in structure with a relatively small base, set into the segment near the outer edge, not reaching beyond the membranes. The other much longer, set into a well-defined socket on the inner portion of the segment; protruding from near its distal end a large flattened structure with hairy margins (from its position and mode of attachment this process comparable to the one with flared serrate tip in gracilis). Right endopod elongate, reaching to near base of distal spine of exopod 2, without evidence of segmentation, armed only with a slender plumose seta on the outer edge. Left endopod swollen, not reaching quite to end of exopod 2, without ornamentation or processes. Riégewayia sp. (Tortugas) Fiaures 36, 37 SPECIMENS EXAMINED: 6 males, copepodid stage V. Occurring with R. gracilis and R. shoemakeri, off Loggerhead Key, Dry Tortugas. COPEPOD GENUS RIDGEWAYIA—WILSON 161 Description: Length, 0.665-0.69 mm. Somite of leg 1 separated from cephalic segment. Urosome 4-segmented; caudal rami with more or less distinct division of the outer margin near the base (representing incipient division of a reduced fifth segment ?); fourth segment with distal row of fine spinules ventrally and on the lateral areas of the dorsal side; caudal rami covered with hairs on both sides. Caudal setae of same relative lengths as in gracilis and shoemakeri, none jointed basally, but the longest jointed near the end as shown for gracilis in figure 1. Antennules alike, reaching just beyond metasome; with 25 free segments; segment 9 somewhat elongate and partially divided by a line running from the inner margin to the middle of the segment, with two setal groups (fig. 37); segments 1 and 2 subequal in length; the three apical segments elongate, the last two subequal to one another. All cephalic appendages weakly developed, but fully segmented. Exopod of antenna 8-segmented. Maxilla with six lobes, of which two are on the basal division. Legs 1-4 fully segmented; segmentation and setation exactly as in gracilis and shoemakeri; all setae jointed. Leg 1 resembles the adult of gracilis and shoemakeri, but the processes of the outer margin of the exopod segments are not fully developed. Leg 5 (fig. 36): First basipod segments well developed, connected by medial plate. Both exopods 2-segmented, the distal segments elongate and of similar length, the left broader than the right. Right exopod 2 with an outer medially placed spine that reaches to end of segment; distally with two shorter spines and a long apical spine equal to about three-fourths the segment length; distal inner portion with three closely set, jointed setae. Left exopod segment 2 with medially placed spine on outer margin, spine reaching beyond apex of segment; apex with three membranous spines of graduated length, increasing from outer to inner; the innermost with a stout inner marginal scle- rotization; at inner apex of segment, a conspicuous hyaline projection overlying base of inner spine. Right endopod longer than left, its proximal part partially segmented; the apex armed with setae and processes; of these, the outermost a stout spiniform process arising laterally; setae four in number, all jointed basally. Left endopod broad, armed near distal part on anterior side with a slender surface seta; otherwise closely resembling the unarmed endopods of the adult shoemakert and marki. Remarks: It does not seem possible to relate this subadult male to either of the Tortugas species. In many calanoid families the develop- ment of the fifth leg is progressive, and, if such were known to be true in Ridgewayia, it might be assumed that these copepodids represent a third unknown species in which the right exopod may have inner apical setac, and the right endopod is also armed with setae. How- 162 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 ever, there are often striking changes in this appendage between the recognized stage V and the adult, such as is known for Ceniropages (Gurney, 1931). It is therefore impossible to say with absolute certainty that this does not represent the subadult stage of one or the other of the two species. The left antennule in the adult could be 25- or 26-segmented, depending upon whether the partially divided ninth segment becomes fused or separated. There is no indication of modification of the right antennule, either in the middle or distal portions. One interesting point brought out by examination of this copepodid is that the modified processes of the first leg are not fully developed until the adult stage, although the leg is otherwise like that of the adult. Discussion The name Ridgewayia was proposed in honor of Sir West Ridgeway, governor of Ceylon. Attention is drawn to this in order to emphasize that the spelling of the generic name is correct. A genus of birds, Ridgwayia Stejneger 1883, named for the ornithologist Robert Ridg- way differs in the spelling by one letter. For purposes of brevity in the following discussion, the new Tortugas species (gracilis, shoemakeri) and the Bermuda species (marki) are referred to collectively as the American species. SpEeciric DIFFERENTIATION Hasitus: In general appearance, the species are alike. Only for marki is there a recorded lack of separation of the cephalic and first thoracic somites. Since the separation of this segment in the Tortugas material was not always as distinct as that of the other segments, specimens in future collections of marki should be carefully examined for indistinct or partial separation. There is also need to determine the possible presence of the reduced last urosomal segments in both marki and canalis. Esterly’s (1911) drawing of the female shows the caudal rami united basally and suggests the presence of this reduced segment. The urosome of canalis (male) as illustrated is very like that of the subadult male listed herein as Ridgewayia sp. (Tortugas). Gurney (1927) says “Abdomen of four somites” but adds ‘‘the 5th somite scarcely distinct.” The possibility that Gurney’s specimens were in the subadult stage is dismissed on the basis of the apparent complete development of the first leg, and of Gurney’s wide experience in study of developmental stages of copepods, precluding the possi- bility that he would err in this regard, even with an unfamiliar genus. COPEPOD GENUS RIDGEWAYIA—WILSON 163 The specific difference noted in the two Tortugas species in the jointed basal portions of the two middle caudal setae is a valuable character for distinguishing whole specimens, and particularly the females of associated species. It is not mentioned for any of the other species, but should be recorded for all species encountered in future studies. In all of the species the rostrum appears to be a large, somewhat expanded structure, without filaments. ANTENNULE: The antennules of the two new species have been studied in detail at high magnification with oil immersion objectives. The material has been critically checked and rechecked, partly because the antennules of the female and that of the left side of the male are 26-segmented, differing thus from the segmentation recorded for typica and marki, or recognized for any other calanoid species. The greatest number of segments that has been conceded to be present in the antennule of existing calanoids is 25. The few instances in which a 26-segmented antennule has been reported are thought to be due to the fact that the observer included the surface eminence to which the antennule is attached. Whole specimens of both species as well as dissected antennules have been examined with this in mind, so that such an error would not be repeated in the case of these two species of Ridgewayia. Gurney (1931, pp. 40-48; 1933, pp. 46-61) has discussed the inter- pretation of the armature and development of the antennule in rela- tion to its evolution, and points out (1931, p. 42) that the primitive antennule of calanoids probably consisted of 27-28 segments, or even of 30 or 31 segments. It seems apparent in all species that some of the fusion leading to reduction has taken place in the proximal part of the antennule, particularly in the usual second segment of a 25- segmented appendage. This segment is usually comparatively long, and bears more than the two setae and aesthete considered to repre- sent the archetypical grouping for each segment. ‘The second segment shown in the illustration of the antennule of typica (Thompson and A. Scott, 1903, pl. 1, fig. 3) is elongate and appears to have two or even three groups of setae. On the basis of length it is comparable to segments 2 and 3 of gracilis and shoemakeri, indicating that the differ- ence in segmentation of these congeners may be due to fusion in these proximal segments. Reference to the summary of setation given herein in the description of gracilis, and found to be identical in shoemakeri, shows that what is considered a primitive armature, as well as segmentation, has been largely retained—most segments have two setae, and aesthetes are abundantly distributed. 164 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 Although admittedly an obvious point, it does seem apropos to stress the desirability of including results of detailed critical examina- tions of the antennules in published records of any specimens of Ridgewayia or of allied genera. The discovery of two species of Calanoida with 26-segmented antennules is a matter of considerable systematic interest. Such an unusual segmentation might be an important part of a generic definition. In the present instance, it cannot separate the Tortugas species generically from Ridgewayia typica because the otherwise obviously related Bermuda. species marki is said to have a 25-segmented antennule. If this is actually the case, then the difference in segmentation must be considered specific. However, on the basis of their descriptions, it is not in- appropriate to suggest that there is need to verify whether the anten- nules of typica and marki are really 25-segmented, and, if so, how their armature compares with that of gracilis and shoemakeri. Esterly (1911) gave no detail of the female or left male antennules in his ac- count of marki. The antennule is figured for typica; it shows a very elongate second segment, and three elongate distal segments. It is personally considered doubtful if the setation shown is entirely exact. There are two setae on nearly every segment but there is no distinc- tion between setae and aesthetes, and many of the setae shown are too similar in length to have been based on exact observation. Considered critically, it cannot be judged from the text of Gurney’s description of Swezia canalis whether or not the segmentation given applies to both of the antennules or only to the right. No direct reference is made to that of the left side. If the statement “Ist antenna of 21 or 22 joints” refers to both antennules, then the seg- mentation within this group of species varies over the considerable range of from 21 to 26 segments. In the specimens observed, the geniculation of the male right antennule is only moderately developed. It was noted while working with the Tortugas material that the antennule could be turned or mounted in such a way that the jointing becomes obscured. ‘There exists, however, as illustrated for the two Tortugas species and as shown by Esterly for marki, a real constriction between two of the distal elongate segments. In whole specimens (most advantageously observed in alcohol), the distal part of the antennule is frequently bent upwards or outwards at this joint as is characteristic of geniculate antennules. This modification as it occurs in these species of Ridge- wayia, though weak, is obviously a specialized joint, giving to the distal portion of the antennule a unit flexibility and freedom of move- ment not present at any other part of the appendage. Such a spe- cialized joint may be presumed to be functionally and structurally COPEPOD GENUS RIDGEWAYIA—WILSON 165 comparable to the variously developed geniculations found in many calanoid genera. Gurney (1927) and Krishnaswamy (1953) have recorded non- geniculate antennules in their specimens. In reviewing their ac- counts in light of knowledge of these other species, it is difficult to know how to assess their records. Both worked with limited material (one or two specimens) and both presented their observations in a very brief fashion, Gurney’s description can only be considered in- definite inasmuch as he referred to the joints as “rather indistinct.” In view of the observation made in my study that the geniculation may easily be obscured in mounting, it does not seem unreasonable to consider this character as inadequately known in these two species. Since it may be of generic significance, it is an exceedingly important character to reaffirm by critical observation of both mounted and un- mounted material. The middle region of the male right antennule is not enlarged in gracilis and shoemakeri, but there are modifications of some segments. Segment 10 is shortened on the outer side. Segment 13 is even more reduced on the outer side, and while remaining distinctly separated in shoemakeri (fig. 30), it seems to have become partially fused with seg- ment 14 in the specimens of gracilis that were critically examined. In both species, segment 14 is elongate on the outer side. In gracilis there is on the inner side a medial incision with sclerotized edges; the sclerotization appears to extend into the internal part of the segment. Beginning at this point in gracilis, and at a similar position in shoe- makeri, there is a longitudinal muscle band that extends through seg- ment 19. There is indication in Gurney’s illustration of a modification at the same point of the antennule of canalis, involving a reduction of one segment and elongation of another, but no detail is given. Esterly did not mention such a modification of these segments in the right antennule of marki. The antennule is described simply as ‘23- jointed” with a ‘‘4-jointed terminal portion.’”’ The modified geniculate portion of the antennule is illustrated, but unfortunately the figure does not include all the succeeding terminal segments. There is a question as to whether the ‘‘4-jointed terminal portion” was meant to include only the segments beyond the geniculation, since the two Tortugas species differ in having three or four segments beyond this joint. This difference is apparently due to a fusion in gracilis of the two segments immediately distad to the geniculation. This is shown by comparison of the terminal segments of the two species (figs. 22, 29). The two distal segments correspond to one another in elongation and in the number, placement, and length of the setae and aesthetes, and so would appear to be of identical origin. In gracilis, the segment 166 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 preceding these has the appearance of at least two coalesced segments comparable to the two distinct segments of shoemakeri. The two groups of setae on the fused segment of gracilis correspond to those of the two separated segments of shoemakeri not only in placement but also in modification. It therefore appears that the geniculation occurs at exactly the same point, although the number of free segments beyond the joint is different. In both gracilis and shoemakeri there are 20 free segments preceding the geniculation. ‘There would appear to be no question that the modified segments of the midportion counted as segments 13 and 14 represent two segments (fig. 30). It therefore seems impossible to fix the position of the hinge at a point comparable to that presumed to be identical for all calanoids. It has long been held, as Gurney (1931, p. 47) emphasizes, that the “.. . . position of the hinge may be regarded as a fixed point. In the Calanoida, this point is always between segs. 18 and 19.”’ In these two species of Aidgewayia it seems correct to assume that the 26-segmented unmodified antennule of the male and female has resulted from a lack of fusion of two segments of the proxi- mal area and may differ from the closely allied species typica in the division of these segments. The modified antennule agrees with the unmodified antennule in relative length and armature of the proximal segments. In comparing this 26-segmented antennule to the basic calanoid 25-segmented appendage from whose study the hinge position has been derived, it would seem necessary only to add one segment to arrive at a comparable position. The hinge in these two species of Ridgewayia should therefore fall between segments 19 and 20, but this is not the case. Not only are there 20 countable segments preceding the geniculation but the elongation and armature of the 20th segment suggests that it may have resulted from fusion of at least two or even three of the segments of the preceding stage. Comparison of the armature of the unmodified antennule with that of the modified, may not indicate what segments are included in the fused distal region of the right appendage. But attention should be drawn to similarities that characterize certain areas of each in these two species. The proximal and the two distal segments are alike in armature and length. The groups of surface spinules occur on identical segments of the left and right side in both species (13-22 and 24-25). Segment 22 has a particularly long seta on the unmodified antennule; such a long seta is present on the segment preceding the geniculation in both species. Jointed setae are present on the unmodified anten- nule beginning with segment 23; similar setae are present beyond the geniculation point of the right antennule. Thus, on the basis of com- parison of armature, the distal part of the segment preceding genic- ulation (visible segment 20) is comparable to segment 22 of the un- COPEPOD GENUS RIDGEWAYIA—WILSON 167 modified antennule. If this is a true correlation, then the elongate segment preceding the geniculation represents a fusion of three seg- ments. Such an interpretation is easily supported in shoemakeri (fig. 29), which has three groups of spinules and three setal groups; in gracilis, the comb of spinules obscures any middle setal group. These two Tortugas species are thus not only unusual among Cal- anoida in the segmentation of the antennule but also in the position of the specialized hinge. It is not too surprising to find calanoid cope- pods with 26-segmented antennules, but the difference in the hinge position is unexpected. This seeming departure from what has been considered a basic pattern raises the question as to how well the facts are known. In studying the literature it is apparent that knowledge of detail of antennule structure and armature is lacking for many species and genera. Most of the available data of worth come from the observations of early workers (Claus, Schmeil, Giesbrecht) ; among the most important examples are the incomparable, detailed figures of Giesbrecht (1892). Since then few workers have given more than the rudiments of antennule structure in their descriptions. This is unfortunate since it is apparent that some very exact patterns of segmentation and armature have been established in the evolu- tionary development of this appendage in the Calanoida, and it therefore has high significance at all taxonomic levels. AnTENNA: This appendage is figured for all the species except marki, for which Esterly (1911) describes the exopod as 8-segmented. This agrees with the other species except canalis, which Gurney (1927) shows as having a 7-segmented exopod. All segments have an inner, lateral seta except the last, which has four apical setae in all except typica, for which three are shown (not known for markt). ORAL APPENDAGES: Where known, the mandible and maxilliped show no significant differences. The maxillule is unknown for canalis, but agrees closely in the other species. A single epipodal seta is present in gracilis and shoemakeri but is not shown for typica and marki, though the lobe is present in the illustration of the latter. This point should be checked in future studies of these two species as it may be of taxonomic importance. The number of setae shown in the figures of typica and marki on the various lobes and laciniae, and on the exopod and endopod, show slight differences from one an- other and from the new species. As some of these may have specific value and should be checked in future studies, they are summarized here (table 1). The maxilla appears to have the most taxonomic importance of all the oral appendages in species differentiation, inasmuch as the number of lobes varies from five in canalis to seven in typica. The reduction 168 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 is in the proximal portion of the appendage. The American species (including the subadult male) agree with one another in having two lobes on the basal division, and also have a seta at the base of the first lobe, not shown in either typica or canalis. TABLE 1,—Setation of mazillule in Ridgewayia (Unknown for R. canalis, and Ridgewayia sp. from Madras) Basal portion Endopod Species Inner Outer Exopod | Seg- Seg- ment 1 ment 2 Gnath- Lacinia Lacinia Lobe of| Coxa Epipod obase 1 2 Basis typica 10 5 4 5 9 0 129 WAS?) paivk(62) markt 13 5 4 5 10 0 aes 6 gracilis 13 f Bs 4 9 1 nie ee hs: 6 shoemakeri 13 5 4 4 9 i ens 6 Lees 1-4: Complete information on the armature of legs 1-4 is available for only the American species. It is alike in the Tortugas species, including the subadult male of unknown identity (Ridgewayia sp., Tortugas). Since the specific pattern is probably established by this stage, it is included in the summaries of armature. Esterly (1911) illustrated only leg 1 of marki, but he gave a table of setation which, if correct, shows that the number of setae on the endopods of legs 3 and 4 differ from the Tortugas species. Information is incomplete for the other species, being known for only some of the legs of a single sex of each species. No summary of setation is given in the text, and only some of the legs are illustrated. These are: typica 2 legs 1, 2, 4 canalis 3 legs 1, 2 sp. (Madras) # leg 1 An inner seta is known to be present on basipod segment 2 of leg 1 in all the species; so far as known, it is absent on all the other legs, but the information available in the literature is complete only for the American species. The same incompleteness of information applies to the inner seta of the first basipod segment. The presence or absence of this seta may be a basic character on all taxonomic levels in calanoid copepods, and it is important that it be accurately deter- mined for every leg of each species. As now recorded in the literature the inner seta of basipod 1 is as follows (+ present; — absent; ? un- known): COPEPOD GENUS RIDGEWAYIA—WILSON 169 Leg 1 Leg 2 Leg 8 Leg 4 typica 9 — _ ? + sp. (Madras) J — ? te ? canalis a = + ? ? markt oS Oot oH _ fs gracilis et + + + + shoemakeri od + + ae -+ sp. (Tortugas) J 4 a. -p -+- It is noteworthy that the figures given in the literature for the first three forms do not show this seta on leg 1, but that it is uniformly present in the American forms. This may be a character of signifi- cance on the generic level if it is correlated with other differences, and its presence or absence should be carefully determined in re- examination of the first three species. The presence or absence of this seta in the other legs is inconsistent in typica and canalis, so that it is difficult to accept the evidence as incontrovertible. Here again, future examination of specimens should be critical in this regard. Where known, the armature of the first two segments of the exopod of each leg of the incompletely known species agrees with that of the TaBLE 2.—Total number of spines and setae on segment 3 of exopod and endopod, legs 1-4 of Ridgewayia Leg 1 Leg 2 Leg 3 Leg 4 Species and known sex typica Q € 6 8 8 ? ¢ 9 7 sp. (Madras) J 7 7(6?)| ? ig ? ? ? ? canalis a 7 6 8 8 am ? ? ? gracilis ‘a Z 6 8 8 9 8 9 7 shoemakeri og ih 6 8 8 9 8 9 7 markt Set a 6 8 8 9 5 9 6 sp. (Tortugas) J 7 6 8 8 9 8 9 7 American species; that is, a single outer spine and single inner seta. With the exception of the first leg of the Madras specimen, the endo- pods likewise agree for legs 1-3; that is, one inner seta on segment 1 and two on segment 2. The complete absence of setae as shown for the Madras specimen would be very unusual, and, as suggested above, is probably due to incomplete delineation. Leg 4 is unknown for the Madras specimen and for canalis; typica agrees with the Tortugas species, but marki differs from the others in having only one seta on endopod segment 2. Where known, the total number of spines and setae on the third exopod segment are like those of the new species, but there is some 170 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 difference in the total number of setae on the third endopod segment of legs 3 and 4, as shown in table 2. Lec 5, Femaue: This appendage is so similar in the four species in which the female is known that only very precise examination reveals the small differences that do exist. The leg of typica differs notice- ably from the American species only in the shortness of the inner- most apical spine of the third exopod segment (see key, p. 173). Differences in the three American species are found only in the com- parative lengths of the basal jomts of the setae of the exopod and endopod and in the placement of the proximal inner and outer setae of the second endopod segment. These latter differences have been expressed in the text of the descriptions of the new species as the percentage of the inner margin of the segment. The length of the outer apical spinous process and of the basal portion of the outer apical seta of the endopod are expressed as a similar percentage. There is a striking difference between some of these points in the two Tortugas species (figs. 5, 28 and table 3). From the figure given for marki, the placement of the setae is very similar to shoemaker. Esterly’s (1911) figure of marki does not show the outer spinous proc- ess. &. typica seemingly differs from the others in the much closer placement of the proximal inner seta to the base of the segment. Taste 3.—Female leg 5, Ridgewayia. Ratio of certain characters of endopod segment 2, expressed as percentage of total length of inner margin of segment (KEY: A, distance between base of inner margin and placement of first inner seta; B, distance between base of inner margin and placement of first cuter seta; c, length of outer spinous process; D, length of basal joint of outer apical seta) Species A B C D typica 23 54 23 60 gracilis 36 60 20 54 marki 44 69 ? 50 shoemakeri 42 64 10 oo Since these characters of the endopod may be useful in differenti- ating species, a comparison has been made in table 3 between the two new species and the other species. Measurements were made with a millimeter rule on the illustrations given for typica and marki. These, of course, do not represent exact measurements, but in the absence of specimens they serve very well for comparative purposes. COPEPOD GENUS RIDGEWAYIA—WILSON 171 They are included in table 3 as percentage figures determined in the same way as those for gracilis and shoemakeri. Lec 5, Maus: An important part of the generic definition as now constituted is found in the modification and asymmetrical development of the endopods of the male fifth leg. This is expressed in the elongation of that of the right side in contrast to the shorter, broadened endopod of the left side and in the varied armature of the different species. The armature seemingly is specific in nature, although with increased knowledge it may be found, in part, to define groups of species or even genera. In the known species, segmenta- tion is suppressed in the adult, but since the subadult male (Ridge- wayia sp., Tortugas) shows partial segmentation of the right endopod, it may be that some species may be found in which at least the right endopod is distinctly segmented. The right exopod is 2-segmented in all the five known adult males (including Ridgewayia sp., Madras). The first segment is very similar in all the species, but the second segment shows definable differences. The three American species are noticeably most similar to one another not only in the shape and length of the segment but in the presence of two similarly placed outer spines. In canalis and the Madras male there is only one well-developed spine, the second or even a third spine being suppressed or broken off in the available specimen. Gurney’s (1927) figure shows two spinous points along the extended outer margin of canalis, and the exceedingly shortened segment of the Madras male has a projection of closely set points very suggestive at least of the remnants of the cuticular points defining the placement of a spine. With this exception, this segment of the leg of the Madras male, though shortened, is more similar to the American species than it is to that of canalis. The left exopod is separable into three distinct segments in the American species. The third segment is highly modified and is set into a centrally recessed area of the expanded, strongly built apex of the second segment. There is a higher degree of segmental develop- ment in shoemakeri than in gracilis. In marki, the apical processes and membranes are much more elongate than in either of the Tortugas species, but in other respects the whole left lez shows more similarity between shoemakeri and marki than exists between shoemakeri and the associated Tortugas species. Although shortened, the left exopod of the Madras male is easily correlated with that of the American species. The figure given by Krishnaswamy (1953) is reduced in size and allows for little detail; however, its outline is entirely credible and its structure interpretable in light of knowledge of the American species. The outer spines of the 172 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 first and second segments are present and strongly developed, being nearly as long as the exopod itself. These spines are placed close to- gether, the second segment being much reduced on the outer side. The apical portion appears to be structurally comparable to the modi- fied third segment of the American species. Its simplicity may be due to reduction or lack of some of the processes and fragmented membranes that complicate the structure in the other species, but it is certainly to be correlated with them. Indeed, it appears much like this portion of the exopod in the Tortugas specimens whenever they were viewed under relatively low power (fig. 35). Examination at high magnification with oil immersion objectives might well reveal complex detail in the Madras species such as has been found to exist in the American species. Krishnaswamy’s (1953) unnamed specimen is therefore seemingly relatable to these species through the fifth leg. In the presence of setae on the right endopod, in the lack of the second outer spine of the right exopod 2, and in the seeming reduction of the left exopod it represents a possible link between the American species and canalis. In the case of this latter species, however, the structure of the apical portion of the left exopod may be somewhat different. As drawn by Gurney (1927) it has a flattened appearance and is dif- ficult to reconcile exactly with the observed species. Such a differ- ence may be entirely graphic in nature. Here again there is need for further examination, and probably also comparison with actual specimens of some of the other species. Both of the Tortugas species have the first basipod segments fused and comparatively reduced. Unfortunately, Esterly (1911) has not shown or described the basipods completely for marki. In his illustra- tions the legs are entirely separated. A reduced but separated seg- ment is shown for the right leg, but only a portion of the left second basipod segment is included. It would be instructive to know the exact condition, since marki and the Tortugas species are obviously congeneric. Gurney shows well-developed segments joined by a center connecting plate very similar to that of the subadult Tortugas male (fig. 36). Apparently the first basipod segments are well de- veloped and separated in the Madras specimen. Correlated with other differences, the lack of fusion may have generic or other taxo- nomic significance, and it is a character that should be carefully noted. It is difficult to interpret the fifth leg of the subadult male (Ridge- wayia sp., Tortugas) in relation to the appendage in the adult. If it does represent a copepodid stage of one of the two known Tortugas species, then considerable change must take place between stage V and the molt to the adult. This would involve fusion of the basal segments, loss of one spine of the apex of the right exopod, and loss of the inner setae of the exopod and of the endopod. However, since COPEPOD GENUS RIDGEWAYIA—WILSON 173 two species are known in which the setae are present on the right endopod, it might be predicted that an unknown species with this character is present in Tortugas waters. The apical membranous spines and hyaline processes of the left exopod may or may not be the beginnings of development of the complex armature of the third seg- ment of the adult. Again, they may represent armature belonging to a species that has a simpler development of this part of the exopod, such as shown for canalis. The significant characters of the fifth leg have been summarized in table 4. The segmentation and spinal armature of the left exopod of canalis is listed as uncertain (see diagnosis of canalis, p. 145). TaBLeE 4.—Comparative characters of male leg 5, Ridgewayia (KEY: (+) character present; (—) character absent; (?) unknown) BasiIpoD RIGHT LEFT EXOPOD RIGHT LEFT EXOPOD ENDOPOD | ENDOPOD SPECIES First seg- | No. spines} No. seg- Armed with ments fused] on seg- ments No. spines | Inner setae} lobes or ment 2 processes markt im 2 3 2 — — shoemakeri a 2 o 2 — — gracilis + 2 3 2 — —- sp. (Madras) _ 1 3 2 ok — canalis — 1 2(3?) 1(2?) + + Key to known species of Ridgewayia The following key has been devised to include both sexes and to summarize and emphasize, in part, similarity of known basic charac- ters that may point up generic or species-group distinctions. Un- fortunately, the Madras male cannot be included in the key because of insufficient information. It differs strikingly from all other known males in that the right endopod of the fifth leg reaches beyond the apex of the relatively shortened right exopod. Characters of the caudal setae and antennules used in couplets 3 and 4 need verification in marki. 1. Antenna, exopod 7-segmented; maxilla with total of 5 lobes, of which one belongs to basal segment. (Female unknown.) ... . . . R. canalis Antenna, exopod 8-segmented; maxilla with total of 6-7 lobes, of which more than one belongs to basal segment... . . Dae pe 2. Maxilla, basal segment with 3 lobes; leg 5 female, ezaped 3, drneROst spine not longer than other spines. (Male unknown.) ... . . . R. typica Maxilla, basal segment with 2 lobes; leg 5 female, exopod 3, innermost spine longer than other spines, about equal to length of segment. . .... 3 174 PROCEEDINGS OF THE NATIONAL MUSEUM VuL. 108 3. Middle caudal setae jointed at bases; leg 5 female, endopod 2, first inner seta placed near the proximal third (equaling about 36 percent of total length of inner margin); leg 5 male, left endopod with a proximal, inner, movable process about as long asendopod. . ... . . . R. gracilis, new species Middle caudal setae not jointed at bases; leg 5 female, this seta placed below the proximal third (at about 42-44 percent of margin); leg 5 male, left endopod unarmed. .... Py hes oan ne 4, Antennules, female and left maiet Bicormnent eae lee A. puseced segment 2 with 2 inner setae, segment 3 with 7 setae; leg 5 mies, left exopod 2, outer spine Jonger than its segment, py ae Ss to same point as longest apical’process. . . . ... . . KR. shoemakeri, new species Antennules, female and left foal! '25-bepmenteds leg 4, endopod 2 with 1 inner seta, segment 3 with 6 setae; leg 5 male, left exopod 2, outer spine shorter than its segment, reaching only to about middle of extended apical membranes,.and processes: sje 5) eye een ee SR mark Systematic position Ridgewayia (as represented by the female of FR. typica) was referred to the Calanidae by Thompson and Scott (1903) on the basis of the resemblances of the cephalic appendages and legs 1-4. The lack of agreement in the fifth legs was noted. As now known, the highly modified fifth legs of both sexes and the geniculate antennule exclude the genus from the Calanidae, but Thompson and Scott are correct in their implication that the unreduced, simply modified oral append- ages and the first four pairs of legs are essentially primitive in structure. In his comments on Lampoidopus marki, Esterly (1911) did not place the genus systematically. Gurney (1927) doubtfully referred Suezia canalis to the Pseudo- cyclopidae because of its ‘general resemblance” to Pseudocyclops. He pointed out, however, that canalis differs ‘‘very materially from Pseudocyclops” but that he felt that “it must either be included in the Pseudocyclopidae or have a new family instituted for its reception,” a course “‘hardly justified without more complete knowledge of the two sexes.”’ Sewell (1932) and Krishnaswamy (1953) have listed Suezia in the Pseudocyclopidae without comment. The demonstrated relationship of the species considered herein makes it possible to evaluate more critically their possible relationship to the Pseudocyclopidae. There is some merit in Gurney’s sugges- tion, though the strongest resemblances between Ridgewayia and Pseudocyclops, the only known genus of the family, are the superficial ones of body form and habitat. Both have the stout body that is seemingly characteristic of extremely littoral and bottom-living calanoids. The segmentation of the body (including the reduced anal segment), the caudal setae, and the large, unfilamented rostrum COPEPOD GENUS RIDGEWAYIA—WILSON 175 are similar. The segmentation and armature of legs 1-4 are alike. There is also some similarity in the fifth legs of the females, though the distinctive modification of the exopod of Ridgewayia is not found in Pseudocyclops, and the endopod of Pseudocyclops varies considerably in segmentation and armature from species to species. If other char- acters showed strong correlation, the differences in the female fifth leg are such that they could well be considered generic rather than familial. It is more difficult, however, to correlate the highly com- plex male fifth leg of Pseudocyclops with that of Ridgewayia, though relationship of the two genera might be assumed through the modified endopods and the apical armature of the left exopod. In what are more easily comparable and, in part, more fundamental characters, the two genera differ more widely. Important among these is the contrast in the location of the paired genital openings of the female. Those of Pseudocyclops are widely separated; those of Ridgewayia, as exemplified by the two Tortugas species, are closely set as in most other Calanoida. The antennule of Ridgewayia is longer than that of Pseudocyclops, which has the segments reduced both in number and length so that the antennule is usually shorter than the cephalic segment. In Pseudocyclops the right antennule of the male is also more strongly geniculate and otherwise modified. The rami of the antennae differ in their segmentation pattern. There are strong resemblances in the mandible—the blades being very much alike in the two genera and the palps differing principally in the elongation of the basipod in Pseudocyclops. The maxillule has the same number of elements, but in Pseudocyclops the distal portions tend to elongation, and some parts have much more reduced armature than found in Ridgewayia. There are very striking dif- ferences in the structure of the maxillae and maxillipeds of the two genera; in Pseudocyclops, the maxilliped is much reduced. In general, the several differences of the cephalic appendages do not indicate an extremely close relationship between the two genera. This, com- bined with the difference in the location of the genital openings, seems to exclude placement in the same family. Qn the other hand, the likeness of habitus and habitat and the similarities of the legs may indicate some degree of phylogenetic relationship. The group of species referred herein to the single genus Ridgewayia have been shown to be in part inadequately known. The lack of knowledge, however, is specific or generic. The species are quite obviously referable to the same family. In the complete absence of any other family in which they can be placed, and in light of more complete knowledge than was available to former authors, it is appropriate to propose for these species a new family, as follows: 176 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 RIDGEWAYIIDAE, new family Copepoda, Calanoida. Detailed diagnostic characters as given above for the unique genus Ridgewayia. Characterized principally by the combination of unreduced and little-specialized cephalic appendages, weakly geniculate right male antennule, fully segmented legs 1-4, and distinctively modified but biramose fifth legs in both sexes. TYPE AND ONLY KNOWN GENUS (as herein defined): Ridgewayia Thompson and A. Scott (1903). Gurney (1931, p. 84) has outlined a grouping of the calanoid fam- ilies that is taxonomically very useful. For the most part, the new family Ridgewayiidae is taxonomically referable to the definition of the first group, the Centropagina. Gurney considered this group to represent the most primitive of the Calanoida and to be closely allied to another group which included only the Calanidae. Within the Centropagina there are some genera with highly modified fifth legs and reduced endopods (such as Jsias), so there would appear little reason to exclude Ridgewayia because of the complexity of the left exopod and modification of the endopods of the male fifth legs. The more primitive segmentation of the antennule found in the new Tor- tugas species emphasizes rather than negates relationship with this group. However, when the phylogenetic position of this family is considered it is probable that important significance must be given to the segmentation of the geniculate antennule in relation to the seemingly unusual position of the hinge. Too little is known to eval- uate this at present, and Ridgewayia appears as a highly singular genus exhibiting a combination of primitive characters with others of unique or specialized modification. For taxonomic purposes, how- ever, the Ridgewaylidae may be currently placed with Gurney’s Centropagina or considered allied to that group. Similar taxonomic considerations may apply, with some qualifica- tions, to the Pseudocyclopidae, placed by Gurney in an undefined group of ‘uncertain position.”” Gurney’s concept of the Pseudocyl- opidae may have been somewhat in error inasmuch as he appears to have considered it as including the very anomalous genus Platycopia, which he spoke of as being “related to Pseudocyclops’’ (1931, p. 82). As has been pointed out (M. S. Wilson, 1946), Platycopia is unique among known calanoids and cannot be closely related to any known genus. Nor is there any known allied family as implied by Lang (1948, pp. 24, 26) in his reference to ‘‘Platycopiidae and closely allied families.”” Lang has placed the Platycopiidae in a suborder separate from the Calanoida. Nomenclaturally, Lang’s system has the regret- table and inconvenient effect of eliminating entirely the much-used COPEPOD GENUS RIDGEWAYIA—WILSON ia term Calanoida, which, though equivalent to Giesbrecht’s term Gym- noplea, has long been preferred and extensively used. Lang’s com- ments on the systematics of the Copepoda are brief and in part in- conclusive, but the concepts presented are worthy of consideration by systematists. Attention is drawn here to Lang’s paper because it is one that may easily be overlooked by specialists in the Calanoida. Habitat and distribution The existing records of Ridgewayia are all from tropical or subtrop- ical coastal areas. Although they suggest that the species may be somewhat localized, they are too few to verify such a suggestion. They more strongly support the indication that the genus is not a pelagic form. This indication is not refuted by the instances in which it occurred in plankton hauls because the records of occurrence are so few and only one or two specimens were captured. Hauls reported by Gurney and Pinney were made at night when bottom copepods may ascend to higher water levels. Gurney interpreted Suezia canalis as “a species living on the bottom during the day” and included the record in the report on the littoral rather than the pelagic Copepoda of the Suez Canal. ‘The four species represented in collections by several specimens were all found in comparable situations. Specimens of R. typica were found in sediment from oyster shells taken on a reef; marki occurred in an island cave dominated by the coral Agaricia gracilis; and the two new species from the Dry Tortugas were associated with corraline algae. The warm, shallow waters of tropical and subtropical reefs and rocky shores, particularly among islands, is therefore suggested as a common habitat of the genus. This is unusual for Calanoida, but it is such a little-investigated habitat of Copepoda that our in- formation concerning the calanoids that may occur in such situations is very Meager. That the genus may also be a bottom-living form of deeper coastal waters is suggested by the records in which the species occurred in plankton hauls. Aside from the investigations made by Thomas Scott and G. O. Sars on the north European coast, the bottom-living calanoids are practically unknown. Since Scott and Sars found several genera and species not closely related to one another or to known pelagic genera and families, it would appear safe to hazard the guess that there may exist other species, genera, or even families of Calanoida that are as yet undiscovered. There are published records of only five genera of this habitat group from American waters. Pseudocyclops has been recorded from northern Canadian waters by C. B. Wilson (1936) and from Bermuda by Esterly (1911). A new 178 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 species of Stephos was found in collections from James Bay by Willey (1923). Two new species of Platycopia were described from the Maine coast by M.S. Wilson (1946). Ridgewayia was found in Bermuda by Esterly (1911) and to this is now added the Dry Tortugas records. In addition, Fleminger (1957) has described new species of Stephos and Bradyidius from the Gulf of Mexico. The distribution of the species of Ridgewayia emphasizes the known faunistic affinity between the Indo-West-Pacific region and the American tropical Atlantic (West Indian) region (Ekman, 1953; Hyman, 1955). Sewell (1948) has listed many species of pelagic copepods common to the two areas and has also (1940, p. 354) pointed out the similarity of the littoral copepod fauna of the Suez Canal with that of the coasts of India and Ceylon. The littoral copepod fauna of the West Indian region is scarcely known, but Willey (1930) has shown the Bermudan harpacticoid fauna to be related to that of the Suez Canal. Nicholls (1944) has pointed out the striking similarity of Suez Canal and Bermudan species of Pseudocyclops (P. magnus Esterly, 1911, and P. latens Gurney, 1927). The closely allied Tortugas and Bermuda species of Ridgewayia emphasize the relationship of these two areas of the West Indian region, and, through their demon- strated relationship to species of the Suez Canal and the Indian coast, are another example of littoral animals zoogeographically linking the Indo-West-Pacific and West Indian regions. Literature cited EKMAN, SVEN 1953. Zoogeography of the sea. xiv+417 pp. EstEeRLY, CALVIN O. 1911. Calanoid Copepoda from the Bermuda Islands. Proc. Amer. Acad. Arts Sci., vol. 47, No. 7, PR. aaa 4 pls. FLEMINGER, ABRAHAM 1957. New calanoid copepods of the? enattion Aetideidae, Euchaetidae, and Stephidae from the Gulf of Mexico. Fish. Bull. U. 8. Fish and Wildlife Service, vol. 57, No. 117, pp. 355-363, pls. 1-3. GIESBRECHT, WILHELM 1892. Systematik und Faunistik der pelagischen Copepoden des Golfes von Neapel und der angrenzenden Meeresabschnitte. Fauna und Flora des Golfes von Neapel, monogr. No. 19, 831 pp., 54 pls. GuRNEY, ROBERT 1927. Zoological results of the Cambridge Expedition to the Suez Canal, 1924. Report on the Crustacea: Copepoda (littoral and semi- parasitic). Trans. Zool. Soc. London, vol. 22, No. 4, pp. 451-577, figs. 108-168. 1931. British fresh-water Copepoda. Vol. 1, 238 pp., 344 figs. Ray Society, London. 1933. British fresh-water Copepoda. Vol. 3, 384 pp., figs. 1196-2061. Ray Society, London. COPEPOD GENUS RIDGEWAYIA—WILSON 179 Hyman, Lippie H. 1955. Le, ZL. oe Sora SE ea OZ n e O77, £7 “4 Te BAAN ne Lanes +) f => LIM fax Np ied U7 \ pore: nM ; oa < SV fos RS . aii te ‘ t \ VTA . . \ : . \ KN ie \ \ PWN "y-¢F cS Ze oe ¢ af / MI Ficure 7.—Mesial aspect of male gonopods of Pachydesmus: a, P. crassicutis crassicutis from Port Hudson, La.; b, P. c. duplex from Grenada, Miss.; ¢, P.c. incursus from Easley, S. C.; d, P. ¢. denticulatus from Atlanta, Ga. 202 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 as long, the caudal edge evenly curved, the setiferous tubercules small and set back from the edge. Sternites smooth and glabrous, sternal area of metazonites raised between legs into a platform higher than level of prozonites, seg- ments 8-10 each with a distinct cruciform impression, segments 11-16 with a large transverse knob between the second legpair. Interzonal furrow broad and well defined on sides, curving in front of spiracles, and reduced to a narrow suture across sternal area. Spiracular opening large and reniform, its edges low and rounded. Pleural areas finely granular, those of midbody segments swollen and more tuberculate just above the legs. Caudal edge of metazonites margined down the sides. Prozonites smooth and polished ventrally. Legs massive, the coxae without distal armature, prefemora with short but acute distal spines and a characteristic knob on the dorsal side. Femora robust, about twice as long as thick and little longer than the other leg joints. Tarsal claw long and slightly curved, with several very fine parallel ridges on the dorsal side. Legs, especially anterior pairs, covered on the ventral sides with very thick pubes- cence, the dorsal surfaces almost glabrous. Sternites of anterior segments without modification except fer very low knobs between the 4th legpair. Gonopod aperture broadly transverse, about twice as wide as long, with a raised margin in front of each coxite and caudally bounded by the elevated sternal area between the 7th legpair. Coxae of gonopods heavy and subyclindrical, with short coxal apodemes. Coxal apophysis small and distally indented. Telo- podite about same length as coxa, its prefemoral division 60 percent of length, and densely setose, prefemoral process rather long and slender, evenly curved instead of medially geniculate. Tibiotarsus slender and laminate, the leading edge finely fimbriate or serrate distally. Secondary tibiotarsus slender, nearly straight, a little longer than primary, its subterminal process normally smaller than the tip, and oblique to it, both distally divergent. VaRIATION: The few specimens that have been available for study afford practically nothing tangible in the way of variation. The total length ranges from 53 to 62 mm. in three intact specimens. In general the gonopods are remarkably alike in all three males, except for what is apparently individual variation affecting the end of the secondary tibiotarsus. As shown in figure 9, one of the prongs tends to be longer than the other, but the type specimen of P. simulans departs slightly in that the prongs are more approximate in size and length. Aside from this, the remainder of the gonopod and the rest of the animal is so similar to the other specimens of crassicutis that simulans can scarcely be maintained as a distinct entity. The speci- MILLIPEDS-——-HOFFMAN 203 men came from a locality directly between those from which the other two illustrated millipeds originated. Synonymy: Of the early references to crassicutis, only those of Wood and Brolemann seem to apply to the typical subspecies. The latter author described and illustrated material from Louisiana, and his account was abridged by Attems (1938) for his treatment of the species. Ficure 8.—Lateral aspects of male gonopods of Pachydesmus, the same specimens de- picted in figure 7: a, P. crassicutis crassicutis; b, P.c. duplex; ¢, P.c. incursus; d, P. c. denticulatus. 204 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Fontaria louisiana Chamberlin is included as a doubtful junior synonym partly on the basis of its type locality and partly because there is nothing in the original description to preclude such an associa- tion. The name was proposed without diagnosis, comparison to any other species, or illustration, and, to make matters worse, I have been unable on two occasions to locate the type at the Museum of Com- parative Zoology. The description of the gonopods of lowsiana is quoted in full: Characterized by the structure of the gonopods of the male. In these the posterior or principal limb of the telopodite is bifid, the branches long, slender, and subequal and cross those of the other gonopod; of the two branches of prongs the mesal one is geniculate near tip with the latter acute, while the ectal one is straight, its tip also acute. The proximal, undivided and less chitinous stalk is thick, densely hairy, and is prolonged along the mesal side of the mesal prong. The anterior or lesser spine is much shorter than the bifid branch, it is much nar- rowed distad, is moderately sigmoidally flexed, and crosses that of the other gonopod. According to this verbal characterization, both tibiotarsal branches are distally simple. I find, however, that in crassicutis the subapical Ficure 9.—Distal end of the secondary tibiotarsus of three speci- mens of Pachydesmus crassicutis crassiculis from Louisiana. Left to right: from Port Hudson, East Baton Rouge Parish; Gonzales, Ascension Parish; and New Orleans Parish. process of the secondary tibiotarsus is usually concealed when the gonopods are viewed in place. Until the holotype, or additional specimens from the type locality, can be examined and the status of louisiana settled conclusively, I feel that the evidence is entirely on the side of treating the name as a junior syonym of P. c. crassicutis. Through the kindness of Dr. Chamberlin; I was priviledged to study the type specimen of Pachydesmus simulans.. This name was based upon a male from the vicinity of Gonzales, Ascension Parish, La., a locality bracketed by the known localities for crassicutis. Chamberlin’s (1942) original description stated that simulans is— ; MILLIPEDS—-HOFFMAN 205 A species apparently very close to P. retrorsus, known from Knox County, Tennessee. It seems distinct, however, in the details of the gonopods, although these are of the same type. The two branches are more slender, and relatively longer and closer together. The distal prongs of the outer branch form an acute angle with each other instead of a very obtuse one or a semicircle, and the bent portion at end of inner branch is shorter. The differences stipulated above are quite valid ones to distinguish simulans from retrorsus, but no consideration was made, apparently, of the pachydesmid previously known from southern Louisiana. The three specimens I have seen from that area are all identical in gonopod structure except for a slight variation affecting the secondary tibio- tarsus in the type specimen of simulans (figure 9). This difference, probably only an individual variation, is hardly of sufficient magnitude to warrant recognition of simulans as a valid name. DistTriBuTIoN: Insofar as presently known, P. ¢. crassicutis is restricted to the Gulf Coastal Plain in the vicinity of the Mississippi Delta country. There are no definite localities known for Mississippi, although the form certainly must occur there. Material has been seen from the following localities: Louisiana. East Baton Rouge Parish: Port Hudson, 1 o&, Dr. Leavitt (MCZ)- Ascension Parish: 4 miles north of Gonzales, 1 o&, Aug. 31, 1940, Stanley and Dorothea Mulaik (RVC). Orleans Parish: New Orleans, 1 o&, H. E. Hubert (MCZ). Mississipp1. ‘BE. Miss.’’ with no further data, 2 #@@ (USNM). Pachydesmus crassicutis denticulatus Chamberlin Fiaurss 7,d, 8,d. Fontaria crassicutis Bollman, 1888, p. 344. Pachydesmus denticulatus Chamberlin, 1946, p. 152, figs. 8, 9—Chamberlin and Hoffman, 1958, p. 43. Type sPECIMENS: Male holotype, female allotype, and two male paratypes (RVC); from the vicinity of Atlanta, Ga., collected at in- tervals between 1939 to 1943 by the late Prof. Perry W. Fattig. Diacnosis: A medium-sized subspecies of the crassicutis phratry characterized primarily by the slender, sinuous secondary tibiotarsus, which terminates in two equal-sized processes, and by the simple, rather elongated coxal apophysis. The color in life is not known, but is probably brown with yellow paranotal maculae. VaRIATION: I have seen but a single specimen definitely referable to this form, the paratype kindly loaned for study by Dr. Chamberlin. The gonopods of this milliped appear to be identical in every respect with those figured in the original description. The length of 47 mm. cited for the holotype seems somewhat short in relation to the width of 12.5 mm., suggesting that perhaps the caudalmost segments are telescoped. The paratype examined measures 61 mm. in length and 13 mm. in width. The specimen recorded from Indian Springs, Ga., 206 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 by Bollman (1888) measures 70 mm. in length and 15 mm. in width. Distripution: P. c. denticulatus is known only from the Piedmont region of north-central Georgia. Aside from the type locality, the only known station is Indian Springs, Bibb County, Ga., where a single male was collected by Prof. Lucien M. Underwood. Unfortu- nately the gonopods of this individual were removed and lost at some time in the past, making a conclusive determination impossible. How- ever, considering the proximity of Indian Springs to Atlanta, and the similarity of size and structure in specimens from the two places. I think that my allocation will eventually be verified by collections made in the region concerned. Pachydesmus crassicutis duplex Chamberlin FicureEs 1,d, 5,e,f, 7,b, 8,5, Pachydesmus duplec Chamberlin, 1939, p. 5, fig. 8—Chamberlin and Hoffman, 1958, p. 43. TypE SPECIMEN: Male holotype (RVC), from Grenada, Grenada County, Miss., collected in July 1910 by Dr. Chamberlin. Diagnosis: A medium-sized member of the crassicutis phratry, characterized by the long, slender, simple secondary tibiotarsus, which completely lacks any remnant of the subapical process and which equals or slightly exceeds the primary branch in length. The cypho- pods are distinct from those of members of the daticollis phratry in that the receptacle is roughly triangular in mesial aspect, its distal end acuminate and considerably exceeding the ends of the valves. In life the color is dilute black, with clear yellow paranotal maculae. VaRIATION: Four adult specimens studied showed the normal range of variation in size. The largest is the male holotype, with a length of 72 mm. and a width of 15 mm. The other three specimens, a male and two females, measure 61 and 59 mm. in length, respectively. The male, from Memphis, Tenn., differs slightly from the holotype in that the secondary tibiotarsus is slightly sinuous instead of nearly straight, but this can at present be considered little more than within the normal range of variability. DistriBuTion: P. c. duplex appears to be endemic to the upper half of the Cretaceous embayment region in northern Mississippi and western Tennessee. It seems probable that collecting in the future will reveal the presence of the subspecies in the western tip of Kentucky. Specimens have been examined as follows: Mississippi. Grenada County: Grenada, 1 &@ (holotype), July 1910, R. V. Chamberlin (RVC). Marshall County: Holly Springs, 1 9, Sept. 17, 1955, L. Hubricht (RLH). MILLIPEDS—-HOFFMAN 207 TENNESSEE. Shelby County: Overton Park, Memphis, o& and 9, Oct. 1, 1955, Hubricht (RLH). Pachydesmus crassicutis incursus Chamberlin FIGURES 7,c, 8,c Pachydesmus incursus Chamberlin, 1939, p. 5, fig. 7—Chamberlin and Hofiman, 1958, p. 43. Pachydesmus retrorsus (not of Chamberlin 1921) Hoffman, in Wray, 1950, p. 44 (misidentification). TYPE SPECIMEN: Male holotype (RVC) from Taylors, Greeneville County, 8. C., collected Aug. 3, 1910, by Dr. Chamberlin. DiaGnosis: a moderate to large member of the crassicutis phratry characterized particularly by the shape of the secondary tibiotarsus of the gonopod, in which the subapical process is reduced to a rather small marginal spur, and the main termination somewhat exceeds the tip of the primary tibiotarsus. The leading edge of the latter is finely denticulate as in the three other members of the phratry, and the coxal apophysis is low and bilobed or trilobed. Paranotal markings apparently pinkish in life. VARIATION: A range in body length of from 51 to 63 mm. has been noted, the width varying from 10 to 13.5 mm. The gonopods of the three males at hand exactly match each other and the original illus- tration given for this form, except that in one, possibly only aberrant, the subapical process of the secondary tibiotarsus is enlarged at the expense of the terminal tip, so that the effect is a complete reversal of the appearance here figured as typical for the race. The original description states that the paranota of the living animal are yellow. The specimen from Easley, S. C. (not far from the type locality), is accompanied by a label indicating that in life the speci- men was “grayish brown with pinkish carinae.’”’ It will certainly be of interest to learn the living colors of material of incursus obtained in the future. Distrispution: This form, the easternmost representative of the genus, seems to be an upcountry and foothill milliped, as the known collection stations are in or closely adjacent to the southeastern end of the Blue Ridge range. I have examined three male specimens from the following localities: Sourn Carouina. Pickens County: Easley, 1 &@, November 1913, C. 8. Marshall (USNM). Grorara. Habersham County: Tallulah Falls, 1 #@, August 1887, Lucien M. Underwood (USNM). Nortu Carouina. ‘Mtns. of N. C.,” 1 &, without date, Roland Thaxter (USNM). This locality is probably either near Tryon, in Polk County, or in Macon County between Franklin and Clayton, Ga. 208 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The laticollis phratry Pachydesmus crassicutis laticollis (Attems) Figures 5,a,b, 10,d, 11,d Fontaria laticollis Attems, 1899, p. 258, pl. 13, fig. 312. Pachydesmus laticollis Pocock, 1909, p. 189.—Attems, 1938, p. 154, fig. 176.— Chamberlin and Hoffman, 1958, p. 44. Pachydesmus retrorsus Loomis, 1943, p. 403. TYPE SPECIMEN: Male, in the Berlin Museum, labeled only ‘“‘Tllinois.”’ Diacnosis: A medium-sized subspecies of the laticollis group, in which the tibiotarsus is only 30 percent of the telopodite length, with a small angulation on its leading edge; prefemoral process conspicu- ously thickened at the geniculation and nearly straight in its distal half; secondary tibiotarsus long, slender, and sinuous, with both its tip and subterminal process slightly recurved proximad. The distal margin of the receptacle of the cyphopod is straighter than in females of other known forms, and the valves longer in proportion to their height. Color in life medium to dark brown, with yellowish or testa- ceous paranota. VaRIATION: Despite the fairly large range occupied by this form, there is no perceptible variation in gonopod structure or in other details of body form. In the material studied the length varied from 62 to 70 mm., width from 12 to 14 mm. There is apparently some geographic variation in size, with a decrease from south to north. Specimens from the Tennessee River valley, in Marion County, Tenn., and Jackson County, Ala., range from 66 to 70 mm. in length; those from the Cumberland Plateau of central Tennessee from 62 to 65 mm. Attems’ type specimen is stated to be only 55 mm. long and 10 mm. wide, considerably smaller than any I have examined. Remarks: Fortunately, although Attems’ original drawing of the gonopod of Jlaticollis is made from low magnification and a difficult aspect, the characters peculiar to the subspecies are clearly shown. It is to be regretted that nothing is known of the history of the type specimen, for it seems unusual that a scarce milliped from a relatively remote area such as central Tennessee would find its way to the Berlin Museum. The specimen, according to Attems, was originally labeled “Fontaria virginiensis,” although by whom is unknown. DistripuTION: This subspecies is known to occur from Jackson County, Ala., north to Cumberland County, Tenn., chiefly through the Cumberland Mountain Range but also to the west of it in the Highland Rim area of central Tennessee. Presumably the great bend of the Tennessee River marks the southern limits of the range. Specimens have been examined from the following localities: TENNESSEE: Cumberland County: Grassy Cove, 12 miles southeast of Cross- ville, 1 @, Sept. 15, 1957, P. C. Holt (RLH). White County: Sparta, 1 o, July 15, MILLIPEDS—HOFFMAN 209 1933, W. J. Gertsch (AMNH); 2 miles west of Bon Air, 1 #, May 9, 1951, L. Hubricht (RLH). Warren County: McMinnville, 1 9, May 9, 1951, L. Hubricht (RLH). Marion County: west fork of Pryor Cove, 2 miles northeast of Jasper, 2 of, June 27, 1957, L. Hubricht (RLH). Coffee County: 4 miles southeast of Manchester, 1 o, June 26, 1957, L. Hubricht (RLH). ALABAMA. Jackson County: Russell Cave, 1 @, May—August 1957, Carl F. Miller (USN M); near Blowing Cave, 5 miles southeast of Limrock, 1 7, June 19, 1957, L. Hubricht (RLH). Ficure 10.—Mesial aspects of the left male gonopods of Pachydesmus: a, P. crassicutis hubrichti from Tuscaloosa, Ala.; 6, P. c. retrorsus from Knox County, Tenn.; c, P. c. adsinicolus from Mobile, Ala.; d, P. c. laticollis from Sparta, Tenn. 210 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The Tennessee specimens agree closely with the subspecies as defined by the drawings. Those from Alabama, however, tend to approach P. ¢. retrorsus both in size and gonopod characters, and the specimen taken by Hubricht near Limrock is so thoroughly inter- mediate in characters that it might easily be considered closer to retrorsus than to laticollis. It seems probable that the specimens reported (Loomis, 1943) as retrorsus from Madison County, Ala., are actually referable to laticollis on the basis of their provenance, somewhat west of Jackson County. Pachydesmus crassicutis hubrichti, new subspecies Fieures 5,c,d, 10,a, 1l,a TypE sPECIMENS: Male holotype (USNM 2272) from Tuscaloosa, Tuscaloosa County, Ala., collected Mar. 30, 1948, by George E. Ball; two female paratypes (USNM 2272) collected at Woodstock, Bibb County, Ala., May 4, 1953, by Leslie Hubricht. Diacnosis: A large form (more than 70 mm. long) characterized by the short, distally subdeltoid tibiotarsus of the male gonopod, as well as by the very large subterminal tooth of the secondary tibiotarsus. DescripTION OF HOLOTYPE: Length of body, 71 mm., width of midbody segment, 15.5 mm. Segments 4-16 of about equal width. Front of head smooth and polished, evenly convex except for a distinct subtriangular flattened clypeal area. Genae large, extending well beyond lateral edge of head capsule, each with a prominent broad median impression. Vertigial groove distinct, becoming more so anteriorly, terminating in a small bipunctate depression centered between the antennae. Two paramedian frontal setae, two sub- antennal setae, and four supra-antennal setae present. Antennae widely separated at base by a distance slightly greater than length of 2d antennal article. Antennae long, extending back to middle of 3d tergite. The 2d, 3d, and 6th articles similar in size and shape, 4th and 5th slightly shorter. The 7th article hemispherical, with four sensory cones. Collum large, 5.0 mm. long at middorsal line, almost 2 mm. wider on each side than following tergite, the lateral ends strongly narrowed and bent cephalomesiad, entirely concealing base of mandibles when head is depressed. Surface of collum polished, the disk with very fine scratches, becoming coriaceous toward the ends. Anterior edge straight, margined up as far as base of mandibles. Posterior edge straight across back, thence abruptly and sinuously swept forward. Paranota of first eight segments swept forward, their corners broadly rounded, those of segments 8-15 transverse, those of last segments becoming increasingly produced caudad. Paranota narrow, MILLIPEDS—-HOFFMAN 2t1 € d Ficure 11.—Lateral aspects of left male gonopods of Pachydesmus, from the same specimens depicted in figure 10: a, P. crassicutis hubrichti; b, P. c. retrorsus; c, P. c. adsinicolus; d, P. c. laticollis. 212 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 less than one-third the diameter of body cavity, almost horizontal but tilted cephaloventrad. Tergites of both prozonites and meta- zonites finely coriaceous, becoming more so on the paranota. Peri- tremata swollen and polished, but not set off by a submarginal groove, their surfaces thus continuous with rest of paranota except in texture. Anterior edges of paranota set off by a deep marginal groove, the posterior edges acute, not margined, and on the caudalmost segments are convex, meeting the caudal edge of the metatergite at a slight re-entrant angle. Interzonal furrow very distinct across dorsum, and finely costulate. Paranota of 18th segment forming acute subtriangular lobes, those of 19th more broadly rounded. Telson short, finely granular, its sides evenly converging, the median process distally truncate with the terminal setae set in a small median impression. No distinct setif- erous knobs near the base of telson on each side. Anal valves moderately convex, vertically striate, with a small swelling near the base of each, the mesial margins with strongly compressed, obliquely striate edges. Preanal scale large, about half as long as broad, its caudal edge evenly curved without distinct median projection. Lateral setiferous tubercules very small and indistinct, well removed from edge. Anterior edge of preanal scale slightly overlapping onto caudal edge of the preceding segment. Sternites smooth and completely glabrous, sternal areas of meta- zonites raised between legs into a platform, the podosternum, which is appreciably higher than level of prozonite, those of segments 8-10 each with a distinct cruciform impression and those of segments 11—16 with a large transverse knob between the caudal pair of legs. Inter- zonal furrow broad and sharply defined down sides, curving forward in front of spiracles, but reduced to a mere suture across the venter. Spiracular opening large and auriculate, with raised and flared edges. Pleural areas finely granular, those of midbody segments swollen and more tuberculate immediately above legs. Caudal edge of meta- zonite with a distinct raised rim down the sides. Prozonites smooth and polished. Legs stout and massive but rather long, the coxae without traces of ventral armature, prefemora with short but acute distal spines and a characteristic knob on the dorsal side. Femora very robust, less than twice as long as thick and but little longer than any other leg joint. Tarsal claw long and slightly curved, with four to six distinct parallel ridges on its dorsal side. Legs covered on the ventral side with very thick short pubescence, dorsal sides, except of tarsal joints, almost glabrous. Sternites of anterior segments without knobs. Gonopod aperture very broadly transverse, about 2% times as wide as long, laterally with a short raised rim in front of each coxite, and MILLIPEDS—-HOFFMAN 213 caudally bordered by the greatly elevated sternal area of the 7th legpair. Coxae of gonopods rather massive, somewhat longer than the telopodites, the coxal apophysis large but not elevated, its margin nearly smooth. Prefemur of telopodite large, 70 percent of the length, densely setose, prefemoral process slender and bisinuate, poorly sclerotized. Tibiotarsus short and stout, constricted at its midlength, distally expanded and subdeltoid. Secondary tibiotarsus shorter than primary, nearly straight, terminally directed mesiad and slightly hooked, the subterminal process large, perpendicular to the main axis, and directed laterad. FEMALE: Similar to the male in external structure, aside from a slightly more arched dorsum. The pleural lobes of the 3d segment are very pronounced and conspicuous. Cyphopods large, the valves in lateral aspect being nearly as large as the mesial face of the recep- tacle. Latter strongly concave on its inner distal surface, and pro- vided there with a small cluster of prominent rounded tubercules and rugosities. Inner valve slightly shorter than outer. Cotor: The color in life is not known, but recently preserved spec- imens are light brown or tan, with the caudolateral halves of the paranota dilute yellow. VartiaTION: The three known specimens, a male and two females, measure 71, 72, and 73 mm. in length, respectively. The male appears to be slightly broader in proportion to its length, being 15.5 mm. wide in contrast to 14.3 and 14.5 mm. for the females. The latter are entirely similar in details of the cyphopods. DistripuTion: Known so far only from central Alabama, at the edge of the Piedmont. Specimens from slightly to the east, in Tal- ladega County, Ala. (2.7 miles north of Sylacauga, 3 oo, May 5, 1954, L. Hubricht; and 1 mile east of Renfro, @ and 2, May 7, 1954, Hubricht), appear to be intermediate in gonopod structure between hubrichti and retrorsus, and are so indicated on the map. These five specimens are very massive; the largest male, 78 mm. long and 16 mm. wide, is the largest North American polydesmoid thus far recorded. Pachydesmus crassicutis retrorsus Chamberlin Fieurss 1,c, 10,b, 11,6 Pachydesmus retrorsus Chamberlin, 1921, p. 232, figs. 3, 4.-—Chamberlin and Hoffman, 1958, p. 44. Typr sPECIMEN: Male (MCZ), from Knoxville, Knox County, Tenn., collected by C. N. Ainslie. Dracnosis: A very large subspecies of the laticollis phratry, char- acterized particularly by the rather long and slender primary tibio- 214 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 tarsus of the male gonopod, and the somewhat flattened and massive secondary tibiotarsus, its distal half twisted about 90 degrees to the basal half. The cyphopod resembles that of P. c. hubrichti but lacks the cluster of tubercules on the side of the receptacle. Variation: The male specimen from Mentone, Ala., agrees closely with the holotype in gonopod structure as well as other details. The Ficure 12.—Distribution of the known forms of Pachydesmus. Solid triangle, P. clarus; hollow triangle, P. crassicutis crassicutis; hollow dot, P. c. duplex; hollow square, P. c. denticulatus; inverted hollow triangle, P. c. incursus; solid square, P. c. adsinicolus; solid inverted triangle, P. c. laticollis; half-shaded dot, P. c. hubrichti; solid dot, P. c. retrorsus; symbol X, intermediate populations. apex of the secondary tibiotarsus is directed distad, instead of slightly oblique, and the subterminal process is concealed in mesial aspect. These minor departures, however, are little more than individual variations. Farther south in Alabama, the gonopods become more definitely intermediate between those as figured for retrorsus and hubrichtt. The male at hand is 74 mm. in length and 15 mm. in width; the female from Georgia is 69 mm. in length. MILLIPEDS—-HOFFMAN 215 DistrisuTion: P. c. retrorsus seems to be endemic to the middle portion of the Tennessee Valley, ranging from the vicinity of Knox- ville south into adjacent parts of Georgia and northern Alabama. Specimens from the following localities have been examined: TENNESSEE. Knox County: Knoxville, 1 @, C. N. Ainslie (MCZ, holotype). ALABAMA. DeKalb County: Mentone, 1 #@, August 1952, Lindsey S. Olive (RLH). Grorgia. No precise locality, but inferentially from the extreme northwestern corner, 1 9, Apr. 1, 1950, B. D. Valentine (RLH). Pachydesmus crassicutis adsinicolus, new subspecies FiaureEs 10,c, 11,c Type spEcIMEN: Male holotype (USNM 2273) from Mobile, Mobile County, Ala., collected Mar. 29, 1948, by William R. M. Mason. Draenosts: One of the smaller members of the Jaticollis phratry, judging from the dimension of the single known specimen. It is certainly closest to retrorsus, differing chiefly in the shape of the sec- ondary tibiotarsus. In adsinicolus this structure is quite slender and nearly parallel to the primary branch, the two converging distally. In retrorsus the terminal elements of the secondary tibiotarsus are strongly unequal in size and both are directed distad, whereas in adsinicolus the terminal tip is bent mesiad and proximad. The holotype, which is 69 mm. in length and 14.5 mm. in width, agrees in nearly every particular with specimens of retrorsus. DistriBuTIoN: Known only from the type locality. This form is of considerable interest in emphasizing the curious north-south distributional pattern of the laticollis phratry. The subspecific name is given in reference to the habitat of the form near Mobile Bay. RELATIONSHIPS GENERIC RELATIONSHIPS: The generic affinities of Pachydesmus are not as obvious as have been surmised by previous investigators. The superficial similarity of the gonopods to those of the species of Harpaphe misled both Attems and Verhoeff to combine the two genera, despite a marked structural dissimilarity in other respects. Actually, Harpaphe is a member of the Rhysodesmus-Boraria complex, it being dificult to separate females of Harpaphe from those of Boraria carolina except by locality. The gonopods in Harpaphe are distinctly diminutive, as is also the gonopod aperture, and I assume that the secondary tibiotarsal process of Harpaphe must be the result of inde- pendent convergent evolution. Attems (1938, p. 151) stated that the genera Takakuwaia (= Xysto- desmus), Rhysodesmus, and Pachydesmus are very closely related but 216 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 he did not mention the basis for his opinion. I suspect his associ- ation must have been founded upon species really not congeneric with Rhysodesmus limax and Pachydesmus crassicutis, although lumped with them in his rather unwieldly classification. In my own opinion, the existing genus most closely allied to Pachy- desmus is Epeloria (Chamberlin, 1939), at least the species related to E. ficta, E. dela, and E. leiacantha. In this genus the podosterna of the males are produced into distinct transverse ridges, and the collum is definitely broader than the following tergite. The gonopods appear bifid near their ends, an effect readily achieved by the basal coalescence of the primary and secondary tibiotarsi. From a generalized ancestral stock, perhaps similar to P. clarus, it is quite feasible to derive two specialized groups, one represented by the large races of P. crassicutis, the other by Epeloria talapoosa and its relatives, which have remained small and developed a simplified gonopod but which are otherwise rather similar in body form and coloration to P. crassicutis. Their distribution around the southeastern periphery of the crassicutis group adds a certain amount of credence to the possibility of a com- mon origin. For the present it is sufficient to indicate that if the telopodite of the gonopod of Epeloria ficta, for example, was bifid entirely to the base instead of only halfway, the result would be almost identical to the gonopod of P. clarus, and the similarity of body structure is so close that it need scarcely be changed to approximate that of a true Pachydesmus. SPECIFIC RELATIONSHIPS: As already implied, P. clarus seems to represent clearly the ancestral stock from which P. crassicutis and its subspecies evolved. It presents none of the specializations peculiar to the larger species, such as more complex gonopods, increase in size, broadened collum, and podosternal ridges of the male. Specific evolution has perhaps been accelerated and almost certainly facili- tated by the Mississippi flood plain, which so far as now known com- pletely isolates P. clarus in the hill parishes of western Louisiana. The eight recognized subspecies of P. crassicutis fall into two dis- tinct ensembles which have been defined above. Of these two phratries, that including crassicutis, duplex, denticulatus, and wncursus is the more primitive. The species are smaller, with less specialized genitalia, and occupy peripheral locations on the range of the genus. During the Cretaceous embayment of the Coastal Plain province, animal life which had occupied that region must have been forced to withdraw to inland areas. Subsequent invasion of the emerged coastal plain would provide the opportunity for the ancestral pachy- desmid stock to spread southeastward probably from Ozarkian and Appalachian refuges. The conservative elements remained west of the Mississippi in the form of P. clarus, the more aggressive produced MILLIPEDS—-HOFFMAN Oty first the progenitors of the present crassicutis phratry and then the group now recognized as Lpeloria. Of the existing pachydesmids, incursus and duplex have gonopods most like those of clarus, and are in fact very similar to each other. They occupy respectively the northwestern and northeastern corners of the generic range, suggest- ing slightly modified remnants of their ancestral stock. Near the center of the range, a population reinvaded the southern end of the Appalachians along the old north-south course of the Tennessee River. This population evolved a greater size and more complex gonopod structure, and now, as the Jaticollis phratry, separates the eastern and western subspecies of the older crassicutis phratry. Literature cited AtTEMs, CaRL GRAF 1899. System der Polydesmiden. II. Denkschr. K. Akad. Wiss. Wien, vol. 68, pp. 251-435, pls. 12-17. 1938. Fam. Leptodesmidae, Platyrhachidae, Oxydesmidae, Gomphodes- midae, in Das Tierreich, Lief. 69, pp. 1-487, figs. 1-509. BouLMAN, CHARLES HARVEY 1888. Notes upon some myriapods belonging to the U.S. National Museum. Proc. U. 8. Nat. Mus., vol. 11, pp. 343-350. BROLEMANN, HEnriI W. 1900. Myriapodes d’Amerique. Mém. Soc. Zool. France, vol. 13, pp. 89-131, pls. 6-8. Causey, NELtL BEvEL 1955. New records and descriptions of polydesmoid millipeds (Order Poly- desmida) from the eastern United States. Proc. Biol. Soc. Wash- ington, vol. 68, pp. 21-30. CHAMBERLIN, RaLtpH VARY 1918a. Two new diplopods from Louisiana. Canadian Ent., vol. 50, pp. 361-363. 1918b. Myriopods from Okefenokee Swamp, Ga., and from Natchitoches Parish, Louisiana. Ann. Ent. Soc. America, vol. 11, No. 4, pp. 369-380. 1921. Chilopods and diplopods from Knox Co., Tennessee. Canadian Ent., vol. 53, No. 10, pp. 230-233. 1939. Onsome diplopods of the family Fontariidae. Bull. Univ. Utah, biol. ser., vol. 5, No. 3, pp. 1-19, figs. 1-37. 1942. New southern millipeds. Bull. Univ. Utah, biol. ser., vol. 6, No. 8, pp. 1-19, figs. 1-40. 1946. On some millipeds of Georgia. Ent. News, vol. 57, pp. 149-152, figs. 1-9. 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Wie sieges eo ae ee ees snaiauro snalaurd 6FF | GET | 6ZF | GIF | GOF | 66E | 68E | 6LE | 69E | BSE | BFE | BEE | GE | GTE | 6OE | O09 (209 WNOY WP -OF>1-07 =|: OF |= OF-11F109-=| 09: | OF OF | OF. |) 09 | OF OFF | OSF | OZF | OLF | OOF | O6E | O8E | OLE | OYE | OSE | OFE | OSE | OSE | OTE | OO salgedsqns pu soledg Y33Ug] [B0} OY} JO SYJPURSNOY} UT syysue, Apog JeZuo0g snuab ayz ur snuv 07 ynous fo dy woif yjbua] ipog—'9 ATAV I, EELS OF THE GENUS CONGER—KANAZAWA 243 Fowler (1912, p. 9) described as new the species Leptocephalus caudalus, from the coast of California. I doubt the validity of this record and discuss this problem on page 228. Dr. E. A. Lachner has examined Riippell’s type of Conger cinereus (No. 766, total length about 338 mm.) deposited in the Senckenberg Museum at Frankfort, Germany. He noted that the sixth infra- orbital pore is above rictus of jaw, that the lateral line pores from anus forward are 39 in number, and that there may have been a dark streak under the eye. Conger triporiceps, new name Ficures 2, 3; Puatss 1,p, 4,a Conger brasiliensis, Kaup, 1856b, p. 115 (type locality, Brazil). Conger occidentalis (not De Kay) Kaup, 1856b, p. 114 (Martinique). Leptocephalus conger Nichols, 1921, p. 22; Fowler, 1944a, p. 436 (in part). SPECIMENS stuDIED: Eight specimens, ranging in total length from 622 to 950 mm., from the following localities: Brazil, holotype (PM 8427) of Kaup’s C. brasiliensis (examined by Dr. E. A. Lachner at Museum National d’Histoire Naturelle, Paris); Cuba (USNM 24936, 9792, 19799); Bahamas, Turks Island (AMNH 7240); Bermuda (CNHM 48606, 48339, 48945). Diacnosis: Three supratemporal pores; 11 preoperculo-mandib- ular pores; four supraorbital pores; greater number of compressed teeth in jaws than in other species (fig. 2); in young specimens vomer- ine teeth in a single irregular row; premaxillary tooth-patch squarish; ossification developed across sensory canal of preorbital bone, one or two post orbital pores. Description: Length of head 126 to 132; tip of snout to origin of dorsal fin 164 to 177; tip of snout to anus 301 to 366; snout length 24 to 35; diameter of eye 17 to 19; length of upper jaw (to rictus) 44 to 51; length of pectoral fins 36 to 42; pectoral fin rays 14 to 17; pores in lateral line 39 to 41; teeth in jaws in a single row, at times with a few scattered teeth on inner row; number of compressed teeth in upper jaw 72 to 90; in lower jaw 74 to 97; vomerine teeth in single irregular row, in large specimens in a traingular patch; premaxillary tooth-patch squarish, with 40 to 105 teeth; preoperculo-mandib- ular pores 11; supratemporal pores 3; supraorbital pores 4, infra- orbital pores 7 or 8; vertebrae 157 to 160; origin of dorsal fin 4.3 percent anterior to pectoral tip to 28.35 percent posterior to pectoral tip; ossification developed across sensory canal of preorbital bone (see pl. 1,p); pectoral fins colored black with lower one-third of fin pale. Measurements and counts are recorded in tables 1 to 3, 5, 6 and in figure 2. 463736—58——_4 944 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 GEOGRAPHICAL RANGE: Brazil, Bermuda, Bahamas, Cuba, Marti- nique, and Pensacola, Fla. Remarks: Ranzani (1840, p. 79, pl. 13, figs. 1,a,b) described a new species, Conger brasiliensis; however his species belongs with the muraenesocid eels. Kaup (1856a) described a new conger as Conger brasiliensis, which makes this name a junior primary homonym. According to Article 36 of the International Rules of Zoological Nomenclature, a rejected homonym can never be used again; there- fore, I give Kaup’s C. brasiliensis, a new name, C. triporiceps. This name refers to the three supratemporal pores on the head, which is a distinguishing character of this species. Kaup’s (1856b, p. 114) description of C. occidentalis states vomerine teeth 9, small and irregularly spaced. The only species that has such a few number of vomerine teeth is triporiceps; also, it has the greatest number of compressed teeth (fig. 2). The head of one specimen (USNM 30710) was obtained from Pensacola, Fla. It was identified as this species by the ossification bridging across the sensory canal of the preorbital bone and the numerous compressed teeth in the jaws. This is the only species in the Atlantic that has the preorbital bone with ossification bridging across the sensory canal. Conger myriaster (Brevoort) Figure 7; Puatss 1,4, 3,A Anguilla myriaster Brevoort, 1856, p. 282, pl. 11, fig. 2 (type locality, Hakodate, Japan). Congromuraena myriaster Giinther, 1870, p. 40, footnote. Ophiosoma? myriaster Bleeker, 1879, p. 26. Conger japonica Bleeker, 1879, p. 32, pl. 2, fig. 2. Conger myriaster Steindachner, 1896, p. 222, pl. 4, fig. 2. Leptocephalus myriaster Jordan and Snyder, 1901, p. 849.—Tanaka, 1911, pl. 19, figs. 69, 70; pl. 20, fig. 77; 1912, p. 78.—Fowler, 1912, p. 9. Astroconger myriaster Jordan and Hubbs, 1925, p. 194.—Mori, 1952, p. 64.— Matsubara, 1955, p. 337. [?]Leptocephalus kiiensis Ui, 1931 (not seen). SPECIMENS STUDIED: 152 specimens from 129 to 606 mm. in total length from the following localities: Japan: Mororan, SNHM 6798 (5 specimens); Hakodate USNM 48194, 49898, 71027, 163410 (4); Aomori Ken, Pacific coast UMMZ 165496, 165505; Sendai, Shiogama USNM 71823 (2), UMMZ 165512 (2); coast of Boshyu, Chiba Ken UMMZ 165528; Tokyo Bay UMMZ 165515 (3); Tokyo market, USNM 49977 (8), 38806 (2), 71870, 151844 (2), SNHM 6696 (21); Yokohama market, SNHM 30624 (41), 26779, USNM 151793; Sagami Sea, Shimoda USNM 57596; Misaki, USNM 71868, MCZ 29018 (2); Suruga Bay UMMZ 177297, USNM 163411, 163412; PROC. U. S. NAT. MUS. VOL. 108 KANAZAWA PLATE 1 Preorbital bone in 13 species of Conger. a, myriaster; 3, oligoporus; C, oceanicus; D, tri po- riceps; E, orbignyanus; ¥, esculentus; G, jordant; H, wilsont, 1, cinereus; J, erebennus; K, philippinus; L, conger; M, macrocephalus. Magnification: F, L, natural size; CE, H-J, M, times 2; A, G, times 3; B, kK, times 7. KANAZAWA PLATE 2 108 PROC. U. S. NAT. MUS. VOL. ‘Apoq uo Sieg 9pImM oy 3ulMmoys “WU FI¢ fo y1sus] [e107 B YIM evluOops[e7) - C D1° (. . r ‘a> TATCY ‘ > at 2 = j 3 MON WO} “607F9L WNSf “ULIOF SUNOA ®B “q Suu CI8 JO Yisuys] [e107 & YIM puRsy AoI90g WO] F7/F[T WNSf UIIOF JINP! USD LSNILIUTI SNILIULI 4IGU0T) cea eerie ETE AT ELON AL 3 KANAZAWA PLATE 108 S. NAT. MUS. VOL. PROG. U. U t Dd $10¢ {sx Lic 3 ‘ O Y1Budg] [e107 & YIM oys “wt ¢/¢ JO YIBus] [e101 B ym ‘uedel ‘uay INnyNy wolf ‘OFZTST INNS! Se 2A LE: NRA LAA LLB RO ae A . i 3 - Uy 9 S LS OM tee eh ng 45 a a a . - retinal te an lnemepy ‘Nnyedg wolf ‘7[S79T INNS ‘snzogos1jo jo adAjopoy 9y1 EAL REIRAA RS ARETE I ‘ a ) ‘4apsorskue ‘y -snsodo ‘saioads s I ee) 7 6110 on ‘Oo SIIO} pue 49]S (iu 49301) PROC. U. S. NAT. MUS. VOL. 108 KANAZAWA PLATE 4 Conger triporiceps and C. erebennus. A, triporiceps, USNM 9792, from Cuba with a total length of 950 mm.; B, holotype of erebennus, SNHM 6466, from Misaki, Japan, with a total length of 483 mm. EELS OF THE GENUS CONGER—KANAZAWA 245 Oigawa 75981; Kobe market USNM 151820 (4), SNHM 30611 (5); Wakanoura SNHM 6968; Nagasaki, UMMZ 165513 (2); Tsushima Straits UMMZ 165533, 165534; Hamada UMMZ 165491, 165536; off Miyazu UMMZ 165519 (3); Wakasa Bay UMMZ 165520 (2); Fukui Ken USNM 151740. Korra: Fusan USNM 163469 (2), UMMZ 165502, 165522 (18). Curina: Tsingtao, Shantung USNM 130468 (3); Ningpo USNM 130525 (2); Chusan Island SNHM 32422 (3). Diacnosis: Whitish areas surround the sensory pores and pits on head and body; origin of dorsal fin 8.14 percent behind to 43 percent of pectoral fin in front of pectoral tip; number of compressed teeth 25 to 45; least amount of ossification developed in preorbital bone; pores in lateral line 39 to 43, head length 133 to 160. Description: Distance from tip of snout to origin of dorsal fin 172 to 209; length of pectoral fin 51 to 70; vertebrae 144 to 147 (50 to 56 ++ 90 to 96); in five specimens dorsal fin rays 272 to 287 and anal rays 187 to 218; compressed teeth in upper jaw 25 to 45, in lower jaw 29 to 43; generally two rows of teeth laterally in jaws, inner row of teeth from a few teeth to five-sixths the length of outer row; pre- maxillary tooth-patch wider than long; origin of dorsal fin from 8.14 behind posterior tip of pectorals to 43 percent in front of pectoral tip (table 1); peritoneum silvery to silvery with black specks; ossification not bridging over sensory canal in preorbital bone; white areas around lateral line pores and sensory pits; preorbital bone shows the least ossification when compared to other species. Additional measure- ments and counts appear in tables 1-6. GEOGRAPHICAL RANGE: In Japan from Mororan and Hakodate on the Island of Hokkaido southward on the Pacific coast of Japan to Nagasaki, northward to Fukui Ken in the Sea of Japan, to Korea southward to Chekiang Province of China. Remarks: This species can be distinguished by the white areas surrounding the sensory pores and organs on head and body. In some preserved specimens these white areas are not discernible, thus making this species frequently misidentified with C. jordan. The two species can be distinguished by the number of compressed teeth in upper jaw (table 4) and by the position of the origin of dorsal fin (table 1). The number of pores in lateral line is 39 to 43, whereas in C. jordani there are 36 to 40 (table 2). The description of Conger japonica Bleeker (1879, p. 32, pl. 2, fig. 2) gives the number of dorsal rays as 260, anal 170; inner row of teeth of both jaws well developed; Blecker’s illustration shows the dorsal origin slightly in front of pectoral tip. These characters agree with C. myriaster; therefore, I have placed C japonica as a synonym of that species. YAG PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ficure 7.—Distribution of Conger myriaster. The paratype of Leptocephalus erebennus Jordan and Snyder (SNHM 6968) from Wakanoura is a young specimen of C. myriaster. Dr. Kamohara has been very kind in giving me the description of kiiensis Ui which was not available to me. Following is the transla- tion of the description as transmitted to me. Dorsal fin inserted slightly behind and above posterior end of pectoral fin; caudal long, tapering, acute, head about 5 in its length, head and trunk together a little less than 2; back of body pale darkish brown; minute white spots along lateral line; dorsal and anal pale, both fins margined with black posteriorly; total length about 40 cm., rare. EELS OF THE GENUS CONGER—KANAZAWA 247 I am placing this species with C. myriaster until the type can be examined. Conger orbignyanus Valenciennes FicureEs 2, 3, 5; PLatE 1,5 Conger orbignyanus Valenciennes, 1847, p. 10, Atlas, pl. 12, fig. 1 (type locality, South America).—Evermann and Kendall, 1906, p. 76.—De Buen, 1950, p. 66. Conger multidens Castelnau, 1855, p. 84, pl. 44, fig. 1 (Rio de Janeiro).—Giinther, 1870, p. 40. Conger vulgaris Ginther, 1870, p. 38 (in part), item ‘k’’ (South America).— Perugia, 1891, p. 656 (Montevideo).—Berg, 1895, p. 23 (Mar del Plata, Montevideo). Conger conger Giinther, 1870, p. 13 (La Plata).—Tortonese, 1939, p. 72 (Monte- video). Leptocephalus conger Jordan and Davis, 1892, p 684 (coast of Brazil). Leptocephalus orbignyanus Devincenzi, 1924, p. 183 (Banco Inglés, Maldonado, Montevideo).—Devincenzi and Barattini, 1928 pl. 16, fig. 4.—Carvalho, 1943, p. 43. SPECIMENS STUDIES: 12 specimens ranging in total length from 603 to 1125 mm., from the following localities: Uruguay: USNM 87737; Montevideo USNM 22625, SNHM 13492 (2); Rio Plata USNM 86726, MCZ 4701. ArcGentTina: Buenos Aires USNM 55580 (3), SNHM 31617 (2). Braziu: Rio de Janeiro MCZ 4711. Diaacnosis: Pores in lateral line from 44 to 46; vertebrae 160-161; origin of dorsal fin one-tenth to 1 pectoral fin length behind posterior tip of pectoral fin; diameter of eye 12 to 17. Description: Tip of snout to origin of dorsal fin 178 to 200, length of upper jaw (to rictus) 39 to 52; length of pectorals 40 to 51; number of compressed teeth in upper jaw 41 to 61; in lower jaw 51 to 59; vomerine tooth-patch triangular, 30 to 45 teeth; premaxilary tooth-patch wider than long, with 31 to 45 teeth; vertebrae 55 to 56 precaudal, and 106 to 104 caudal (in 2 specimens) ; 9 or 10 preoperculo- mandibular pores; a single supratemporal pore; in one specimen dorsal rays 368, anal 301; preorbital bone does not have ossification bridging over sensory canal, pl. 1,n. Additional counts and measurements are recorded in tables 1-3, 5, 6 and in figure 2. GEOGRAPHIC RANGE: South America, from Uruguay and Argentina northward to Brazil. Remarks: The distribution of C. orbignyanus may ovrleap with that of C. triporiceps and C. esculentus; however, it can easily be separated by the number of lateral line pores. Also, it may be fur- ther distinguished from OC. triporiceps by the single supratemporal pore and absence of postorbital pores. This species is more closely related to C. conger from Europe, but it can be distinguished by the following characters: 248 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Character C. orbignyanus C. conger Eye diameter 12 to 17 20 to 28 Head length 122 to 150 143 to 168 Eye into snout 2.06 to 2.90 times 1.36 to 2.11 times Pectoral rays 16 to 17 17 to 19 Origin of dorsal in rela- one-tenth to 1 pectoral one-fifth in front to one- tion to pectoral fin fin length behind third behind tip The origin of dorsal fin in relation to pectoral fins in the specimens I have examined are from one-tenth to two-thirds pectoral fin length behind posterior tip of pectoral fin; however, in literature it has been described as 1 pectoral fin length behind. Dr. E. A. Lachner has examined the type of Conger multidens Castelnau (in the Muséum National d’Histoire Naturelle, Paris, No. 8428), 704 mm. in total length, and states that lateral line pore counts from a vertical from anus anteriorly was 43 on the right side and 45 on the left side, which places this species in the synonymy of C. orbignyanus. Conger conger (Linnaeus) Ficures 2, 3; Puate 1,1 Muraena conger Linnaeus, 1758, p. 245 (based on Artedi; type locality, Med- iterranean Sea). Conger verus Risso, 1826, p. 201 (Nice). Conger vulgaris Yarrell, 1831, p. 158.—Bleeker, 1864, p. 26.—Ginther, 1870, p. 38 (in part).—Day, 1880-1884, p. 250.—Moreau, 1881, p. 565.—Stassano, 1890, p. 32 (Spanish Sahara).—Vinciguerra, 1893, p. 333.—Chabanaud and Monod, 1926, p. 249 (Port-Etienne).—Duncker et al, 1929, p. f.60. Congrus vulgaris Lowe, 1837, p. 192 (Madeira).—Richardson, 1844-1848, p. 107. Conger rubescens Ranzani, 1840, p. 82, pl. 12. Leptocephalus conger Fowler, 1912, p. 9 (in part).—Jordan and Gunn, 1899, p. 339. Conger conger Fowler 1923a, p. 33 (Funchal Bay).—Nobre, 1935, p. 383.— Cadenat, 1937, p. 437.—Navarro, 1942, p. 196; 1943, p. 125 (Cabo Bojadar, Cabo Blanco).—Fowler, 1936, p. 268 (in part). SPECIMENS STUDIED: 25 specimens, from 83.5 to 87.7 mm. in total length, from the following localities: Sardinia, USNM 143454 (2); Naples, Italy, USNM 48482 (2), 48470, SNHM 20614 (3); Nice, France, MCZ 9326, 9332 (2); Madeira, USNM 6522, SNHM 24032; Azores, USNM 94503; Canary Islands, SNHM 10566, 10586; Bergen, Norway, USNM 17488; Europe, USNM 431238, 43587, MCZ 2554 (4), 2468A (2). Draenosts: Pores in lateral line 44 to 47; pectoral rays 17 to 20; diameter of eye 20 to 28; origin of dorsal fin from 19.30 percent before to 33.74 percent behind pectoral fin tip. Description: Distance from tip of snout to origin of dorsal fin 186 to 222; length of pectorals 37 to 70; supratemporal pore 1; preoperculo- mandibular pores 9; compressed teeth in upper jaw 25 to 62, in lower EELS OF THE GENUS CONGER—KANAZAWA 249 jaw 24 to 58; 2 rows of teeth laterally in jaws; inner row from one- quarter to whole length of outer row; premaxillary tooth-patch wider than long; upper edge of gill opening opposite middle of pectoral fin base; ossification not developed over sensory canal of preorbital bone. Additional counts and proportional measurements are recorded in tables 1-3, 5, 6 and in figure 2. GEOGRAPHICAL RANGE: Norway southward to Cape Blanco, Africa, and the Canary Islands, throughout the Mediterranean Sea, and occurring in the western half of the Black Sea. Kyle and Ehren- baum (in Duncker, et al, 1929, p. f.62) state that this species is found as far north as Trondhiem Fjord, Norway, is seldom found in the Baltic Sea in the Kattegat and in the Skaggerak, and is found mainly south and west of the British Isles, in the Biscayne west of Spain, and in the Mediterranean Sea. Remarks: There may be some population divergence in the Atlan- tic and Mediterranean forms in regards to snout and head length; however, more specimens are needed to reach definite conclusions. The following tabulations show the comparative snout and head lengths (in thousandth of total length) of the Mediterranean and Atlantic forms: Snout length in thousandths of total length Area 30.0 | 32.5 | 35.0 | 37.5 | 40.0 | 42.5 | 45.0 | 47.5 oa | Mediterranean _-_---_-_- 2 2 | 5 | De ai Sct | ae gee eee | eee GAG ee ge See a ees ese erin! 5 | 2 |------ 4 | | Head length in thousandths of total length Area 140 | 145 | 150 | 155 160 | 165 | 170 | 175 | | ire Mediterranean_-_-__-__- 2 3 3 Dee ea Le _| Lee ote Son tes Atlantic_____________|.____- os } 1 | 1 | oe. 2 | ee | 2 | | This species is related to C. orbignyanus from the Atlantic coast of South America; however, in C. orbignyanus the diameter of eye is smaller, pectoral rays fewer, and origin of dorsal fin more posterior in position. See remarks under C. orbignyanus. Giinther (1870, p. 39) records under the species Conger vulgaris a specimen from St. Helena, but this specimen is identified tentatively as C. esculentus. 250 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Yarrell (1831, p. 158) and subsequent authors based their vulgaris on Anguilla vulgaris Cuvier (1817, p. 231), which is not a Conger but a freshwater eel. Conger jordani, new species Fiaure 4; Puate 1,a Leptocephalus erebennus Jordan and Snyder, 1901, p. 849 (in part). Conger japonicus Jordan and Hubbs, 1925, p. 194 (in part). [?] Conger vulgaris Barnard, 1925, p. 187 (Agulhas Bank, off South Africa). [?] Conger conger Smith, 1949, p. 392, fig. 1110 (Cape of Good Hope to Mozam- bique). Hotorypr: USNM 71844, 606 mm. in total length, collected in 1906 at Misaki, Japan, by the Albatross. PaRaTyPEs: JAPAN: USNM 26250; Misaki USNM 49866, 71843, 71715, 71716, 71845, 72003, 71963, UMMZ 14280, 165503-4, SNHM 12924—5, 23466; Kagoshima USNM 163467; Tokyo market USNM 71818, UMMZ 165537, 165521; Suruga Bay, Shimizu Market UMMZ 165508; Nagasaki UMMZ 165517 (8), 165535; Oxinawa: USNM 71819. Draenosis: One row of teeth in jaws; origin of dorsal from 11 to 80 percent of pectoral fin behind posterior tip of pectoral fin; verte- brae 141 to 144; pores in lateral line 36 to 40; diameter of eye 18 to 26. Description: In the following description the first figure is for the holotype; the figures in parentheses refer to paratypes. Length of head 144 (125 to 148); tip of snout to origin of dorsal fin 200 (180 to 217); tip of snout to anus 379 (352 to 388); snout length 35 (380 to 37); diameter of eye 21 (18 to 26); length of pectoral fin 49 (40 to 56); upper jaw (to rictus) 49 (36 to 51); pores in lateral line 37 to 38 (36 to 40); supratemporal pore 1; preoperculo-mandibular pores 9; vertebrae 141 to 143 (48 to 50 precaudal, + 92 to 94 caudal); in two specimens dorsal rays 301 and 307 and anal rays 225 and 240; pectoral rays 17 (15 to 18); compressed teeth in upper jaw 42 to 77; 1 row of teeth in upper jaw, occasionally a few teeth in inner row anteriorly; pre- maxillary tooth-patch wider than long. In large specimens the ossification of preorbital bone bridges over sensory canal; in small specimens ossification not bridged over (Pl. 1,¢). Origin of dorsal fin 28.58 percent of pectoral fin behind posterior tip of pectoral fin (14.65 to 76.95 percent behind pectoral tip). Color notes taken by Dr. C. L. Hubbs for specimen UMMZ 165504 is as follows: Purplish brown fins, blue gray at base and inky black in margin. GEOGRAPHICAL RANGE: From Tokyo Bay southward to Nagasaki, Okinawa, and Africa. Remarks: The paratypes of Leptocephalus erebennus Jordan and Snyder (USNM 49866, SNHM 12924 and 12925) are not the same as the holotype of L. erebennus, but are of this new species. EELS OF THE GENUS CONGER—KANAZAWA 251 The affinity of this species is with C. oligoporus from Hawaiian waters, but can be distinguished from it by the following characters: The eye smaller, and more vertebrae and more pores in lateral line (see tables 2, 3, 5). I am unable to distinguish the two specimens from Port Alfred, South Africa, from C. gordani, but they are doubtfully included with this species because its geographical distribution is broken consider- ably from that of C. jordani. The origins of dorsal fin of the two African specimens are more anterior in position than in C. jordani. The counts and proportional measurements of the two specimens are given below (one specimen had a mutilated head so some of the characters are not given): total length 409 and approximately 635 mm.; pores in lateral line 40; pectoral rays 17 and 16; vertebrae 144 (X-ray); compressed teeth in upper jaw 42 and 55, in lower jaw 40 and 56; one row of teeth in jaws; a few scattered teeth in inner row; preoperculo-mandibular pores 9; supratemporal pore 1; dorsal rays 282; anal rays 228; origin of dorsal fin 24.25 percent behind posterior tip of pectoral fin; preorbital bone bridged over sensory canal; length of head 134 and 144; tip of snout to origin of dorsal fin 176; tip of snout to anus 374 and 386; snout length 32 and 40; diameter of eye 16 and 21; interorbital 21; length of upper jaw to rictus 45; length of pectoral fin 40; premaxillary tooth-patch wider than long. ‘Some preserved specimens of C. myriaster that lose their white markings around the sensory pores are difficult to distinguish from C. jordant. 'To separate these specimens see remarks for C. myriaster. The new species is named for David Starr Jordan, who pioneered in studies of Japanese fishes. Conger oligoporus, new species PLATES 1,B, 3,B Conger wilsont Fowler, 1923b, p. 375 (Honolulu) ; 1928, p, 38, pl. 1,c. Hotorype: USNM 162512, 237 mm. in total length, collected Dec. 22, 1951, 5-20 feet in depth at poison station, 200 yards west of Diamond Head Light House, edge of cut in reef, Oahu Island, Hawaii, by Gosline and class. ParatyPes: USNM 163567 (3 specimens), 194, 226, and 245 mm. in total length, taken with the holotype. George Vanderbilt Equatorial Expedition Stations 15 and 26, 2 specimens, Laysan Island and Maro Reef, Territory of Hawaii, 1951. Diaenosts: Teeth in one row in jaws, pores in lateral line 35 to 36, vertebrae 137 to 139, diameter of eye 24 to 30. 25? PROCEEDINUS OF THE NATIONAL MUSEUM VOL. 108 Description: The description is based on the holotype and five paratypes. The counts recorded first are for the holotype; those in parentheses are for the paratypes. Pores in lateral line 35 and 36 (35 to 36); pectoral rays 15 (16); vertebrae (X-ray) 137 (139); teeth in upper jaw in one row; compressed teeth in upper jaw 40 (30 to 46), in lower jaw 39 (42 to 45), vomerine teeth 13 (12 to 17), premaxillary teeth 16 (12 to 21); preoperculo-mandibular pores 9 (9); supra- temporal pore 1(1). Gull opening near middle of pectoral base, rictus of jaws extends posteriorly to rear margin of pupil or about two- thirds of eye; origin of dorsal fin from 2.63 to 28 percent of pectoral fin length behind posterior tip of pectoral fin. See table 7 for various measurements expressed in thousandths of total length for the holotype and paratypes. GEOGRAPHICAL RANGE: Known only from the Hawaiian Islands. Remarks: Fowler (1923b, p. 375) recognized two species from the Hawaiian Islands and thought one of his species was the same as Waite’s C. wilsoni (=verreauat); however, it differs from verreauxi in number of lateral line pores and number of vertebrae. Its affinity is with C. jordani, but differs from it in that the eye is larger and the number of vertebrae is tewer. This new species can be distinguised from the two species of Conger in the Hawaiian Islands by the characters given in the key (p. 230). The name of this new species is in reference to its possession of few lateral line pores. Tas.Le 7.—Measurements of Conger oligoporus expressed in thousandths of total length Holo- Paratypes type Characters USNM || USNM |! USNM | USNM | GVF | GVF 162512 || 163567 | 163567 | 163567 | 15 26 Total length (in mm.)___-- 237 226 194 245 227 110 Head length is be 143 153 157 145 145 141 Snout to dorsal origin_- --- 203 208 219 200 197 200 Snout to anus._-.-------_ 346 356 379 353 352 341 Snoutilength=—--2 282 S25" 37 36 36 37 35 34 Diameterioleyesu2—s2s22— 20 30 28 28 25 24 Length of upper jaw to rictus2=s 22h" a 51 53 54 48 of 45 Pectoral fin length_______- 53 52 59 51 49 45 Depthvotibody22—. 2-224 49 50 55 43 46 49 Length of gill opening__-_-- 15 16 7 16 15 14 EELS OF THE GENUS CONGER—KANAZAWA 253 Conger esculentus Poey Figures 2, 3; PuatE 1,F Conger esculentus Poey, 1858, p. 346 (type locality, Cuba); 1867, p. 246; 1868, p. 424; 1876, p. 151.—Stahl, 1883, p. 80.—Rivero, 1938, p. 173. Conger niger Bean and Dressel, 1884, p. 169 (Jamaica). [?] Conger vulgaris Giinther, 1870, p. 39, item “w,’’? (St. Helena).—Metzalaar, 1919, p. 14 (Curagao). [?] Conger vulgaris var. niger Giinther 1869, p. 239 (St. Helena).—Melliss, 1875, p. LO: SPECIMENS sTUDIED: Three specimens, from 970 to 1030 mm. in total length, from the following localities: Cuba, MCZ 9328 (holo- type), 9330; Jamaica, USNM 32091. Diacnosis: Two rows of teeth in jaws; pectoral rays 18 to 19; length of head 170 to 183. Description; Tip of snout to origin of dorsal fin 222 to 236; upper jaw 62; length of pectorals 59 to 70; number of compressed teeth in upper jaw 53 to 63; lower jaw 54 to 58; vomerine teeth 45 and in a triangular patch; premaxillary teeth 36; tooth-patch wider than long; vertebrae 46 precaudal, and 86 caudal; teeth in jaws in two rows; supratemporal pore 1; preoperculo-mandibular pores 9; preorbital bone without ossification bridging across the sensory canal, pl. 1,F. Additional counts and measurements are recorded in tables 1 to 6, and fig. 2. (All counts and measurements of the holotype were made by William C. Schroeder of the Museum of Comparative Zoology at Harvard.) GEOGRAPHICAL RANGE: Cuba, Jamaica, and possibly to Curagao and St. Helena. Remarks: This species is related to C. erebennus from Japan and OC. macrocephalus from the Philippines. The differences distinguishing these species are given in the key (p. 230). Giinther (1870, p. 39, item “‘w’’) lists under Conger vulgaris an adult from St. Helena. Dr. Marshall of the British Museum kindly furnished the following data for this specimen: Total length 1027 mm.; head length 169 mm.; tip of snout to anus 428; tip of snout to origin of dorsal fin 259; snout length 41; eye 21; pectoral length 68; lateral line pores 37; supratemporal pore 1; and pectoral rays 16. The lateral line pore count of 37 for this specimen is much fewer than the usual 44 to 47 for the species C. conger from European waters (table 2). The pore counts and proportional measurements are closest to C. escu- lentus from Cuba. For comparison of the two forms see tables 1, 2, 5, 6. The only difference between the two forms seems to be in the number of pectoral rays—18 to 19 for C. esculentus and 16 for the St. Helena specimen. The number of vertebrae and the number of 254 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 teeth in jaws may show some difference. Until more specimens can be examined I am placing this specimen with C. esculentus. Posada Arango (1909, p. 305) lists Conger vulgaris from the Colum- bian Caribbean; however, this specimen is not a Conger since it is described with tentacles and barbels. Metzalaar (1919) lists Conger vulgaris from Curagao. I am unable to verify this specimen as C. esculentus, but I am placing it here until it can be verified. Conger macrocephalus, new species PLATE 1,M ? Conger vulgaris Bleeker, 1864, p. 26, Java, Celebes and Letti. ? Conger conger Weber and de Beaufort, 1916, p. 259. Hototypr: USNM 164334, total length 803 mm., Philippines, Verde Island passage, lat. 13°34’37’’ N., long. 121°07’30’’ E., Feb. 22, 1909, collected by the Albatross, at a depth of 180 fathoms. Diaanosis: Teeth in upper jaws in two rows laterally; pores in lateral line 35; pectoral rays 17; length of head 199. Description: Length of head 199; tip of snout to origin of dorsal fin 247; tip of snout to anus 416; snout length 50; diameter of eye 24 length of upper jaw 66; length of pectorals 51; compressed teeth in upper jaw 66; two rows of teeth in jaws, inner row two-thirds the length of outer row; dorsal rays 288; anal rays 214; premaxillary tooth-patch wider than long; origin of dorsal fin 10.41 percent behind pectoral tip; preorbital bone without ossification bridging across sensory canal; supraorbital pore 1; preoperculo-mandibular pores 9; vertebrae 137 (X-ray). Additional counts and _ proportional Measurements are recorded in tables 1, 2, 3, 5, 6. GEOGRAPHICAL RANGE: Philippines and East Indies. Remarks: This species is closely related to C. esculentus from the West Indies; however, it differs from it by having fewer pectoral rays and more compressed teeth in upper jaws (table 4). This species is named in reference to its long head. Conger erebennus (Jordan and Snyder) Puates 1,3, 4,8 Leptocephalus erebennus Jordan and Snyder, 1901, p. 849 (in part), fig. 3 (type locality, Misaki, Japan). Leptocephalus kiusiuanus Jordan and Snyder, 1901, p. 851 (Hakata, Kiushu, Japan). Conger japonicus Jordan and Hubbs, 1925, p. 194 (in part). SPECIMENS STUDIED: Six specimens, ranging in length from 429 to 776 mm., from the following localities: Japan: Misaki SNHM 6466 (holotype of erebennus); Hakata, Kiushu SNHM 6467 (type of kiusiuanus) ; 1dzu USNM 22563; Tokyo market USNM 49440; Suruga Gulf USNM 163468. Korra: Fusan USNM 143410. EELS OF THE GENUS CONGER—KANAZAWA 255 Diagnosis: Two rows of teeth in jaws; pectoral rays 19 to 21; origin of dorsal fin from 10 percent behind pectoral tip to 40 percent anterior to pectoral tip; length of head 163 to 184; ossification not bridging across sensory canal of preorbital bone. Description: Tip of snout to origin of dorsal fin 199 to 222; length of pectoral fins 48 to 60; upper jaw (to rictus) 52 to 59; dorsal rays 288 and 300, anal rays 217 and 229 in two specimens; vertebrae 137 to 147 (50 precaudal and 87 caudal in one specimen); two rows of teeth in jaws laterally, inner row from one-half to whole length of outer row; 39 to 53 compressed teeth in upper jaw, 40 to 52 in lower jaw; premaxillary tooth-patch wider than long; one supratemporal pore; upper edge of gill opening near midpoint of pectoral base, slightly closer to upper edge than lower; rictus of jaw below middle of eye; length of head into trunk 1.10 to 1.39, head and trunk into tail 1.48 to 1.60; ossification not bridging over sensory canal of preorbital bone. Additional counts and proportional measurements are recorded in tables 1 to 6. (Counts and measurements of the types were made by Dr. J. Béhlke at Stanford Museum.) GEOGRAPHICAL RANGE: From Sagami Bay southward to Hakata, Kiushu, Japan, to Fusan, Korea. Remarks: Jordan and Snyder (1901, p. 849) had two other species mixed with the type series of Leptocephalus erebennus: one specimen of Conger myriaster and three of C. jordani. Jordan and Snyder recog- nized the differences in the head and trunk proportions in their key; however, in their description the proportional measurements were mixed. This species is closely related to C. esculentus from the West Indies. For differences see key (p. 231). The number of pectoral rays distinguishes this species from the others found in Japanese waters. It differs further from C. myriaster by having a longer head and without white areas around sensory pores and pits. It differs further from C. jordani by having two rows of teeth in jaws, the origin of dorsal fin situated more anteriorly, and ossification not bridging over sensory canal of the preorbital bone. Conger philippinus, new species PuaTE 1,Kk [?] Conger vulgaris Giinther, 1870, p. 39 (in part), item ‘‘l’”’ (East Indian Archi- pelago). [?] Conger multidens Pohl, 1884, p. 40 (fide Fowler, 1931, p. 316, Viti Levu).— Whitley, 1927, p. 4. [?] Conger orbignyanus Fowler, 1931, p. 316. Conger conger Herre, 1934, p. 16 (Cebu, Dumaguete, and Oriental Negros in the Philippines). Hototype: USNM 134969, 234 mm. in total length, collected at a market in Cebu, Philippines, Mar. 22, 1909, Albatross. 256 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 ParatyPes: SNHM 27116, 223 mm., from Cebu, Philippines, Aug. 27, 1931, Herre; SNHM 26848 (2), 212 and 278 mm., Dumaguete, Philippines, June 18, 1931, Herre’s Philippine Expedition. Diagnosis: Two rows of teeth laterally, pores in lateral line 37 to 38; vertebrae 127 to 135; pectoral rays 15 to 16. Description: The counts recorded first are for the holotype; those in parentheses are for the paratypes. Length of head 124 (126 to 144); tip of snout to origin of dorsal fin 201 (202 to 224); tip of snout to anus 359 (363 to 401); snout length 27 (27 to 34); diameter of eye 20 (22 to 26); length of pectoral fins 46 (47 to 50); upper jaw (to rictus) 39 (41 to 50); supratemporal pore 1; pores in lateral line 38 (37 to 38); preoperculo-mandibular pores 9; vertebrae 130 (127 to 135); pectoral rays 16 (15 to 16); compressed teeth in upper jaw 38 to 45; two rows of teeth laterally in jaw; premaxillary tooth-patch wider than long; origin of dorsal fin 75 percent of pectoral fin length behind posterior tip of pectoral fin (71.43 percent to 85.00 percent); pre- orbital bone without ossification bridging across sensory canal at 278 mm. in total length. GEOGRAPHICAL RANGE: Philippine Islands, Kast Indies, and possibly Fiji Islands. Remarks: This species is related to C. wilsont from Australian waters; however, it differs from it by having fewer lateral line pores, pectoral rays and vertebrae. Pohl (1884, p. 40, fide Fowler, 1931, p. 316) records a specimen of Conger multidens from the Fiji Islands with origin of dorsal fin about a pectoral fin length behind posterior tip of pectoral fin; however, until a specimen can be examined from that area I am referring it to this species. Dr. Marshall of the British Museum has measured and made counts on a specimen listed under Conger vulgaris by Ginther (1870, p. 39, item ‘‘l’’.) The counts and measurements compare closely with philipprnus. Conger wilsoni (Bloch and Schneider) FIGURE 2,w; PLATE 1,H Gymnothorax wilsoni Bloch and Schneider, 1801, p. 529 (type locality, New Hol- land). Conger ne Castelnau, 1872, p. 193 (Hobart, Tasmania, and Hobson’s Bay, Victoria).—Waite, 1923, p. 72, fig. 54. Leptocephalus monganius Phillipps, 1932, p. 230 (Mongoniu, Doubtless Bay). Leptocephalus labiatus Griffen, 1936, p. 15. Leptocephalus wilsont hesperius Whitley, 1944, p. 25, fig. 1, (Leighton Beach, near Fremantle, Western Australia). SPECIMENS STUDIED: Four specimens, ranging in length from 289 to 1012 mm., from the following localities in Australia: Port Jackson, New South Wales, USNM 47765, 47830, and 47764; Strickland Bay, Rottnest Island, Western Australia, USNM 164518. EELS OF THE GENUS CONGER—KANAZAWA 257 Draenosis: Two rows of teeth in jaws, lateral line pores 38 to 40, pectoral rays 17 to 18, vertebrae 146 to 147. Description: Tip of snout to origin of dorsal fin 185 to 194; length of head 119 to 139; tip of snout to anus 349 to 379; diameter of eye 15 to 23; length of snout 26 to 34; length of upper jaw (to rictus) 40 to 46; length of pectoral fin 42 to 47; supratemporal pore 1; preoperculo- mandibular pores 9; vertebrae 146 to 147 (50 to 51 precaudal plus 95 to 96 caudal); vomerine teeth 25 to 46; premaxillary teeth 12 to 36; two rows of teeth in jaws laterally; premaxillary tooth-patch wider than long; preorbital bone has ossification developed across sensory canal (pl. 1,4); origin of dorsal fin from 26 percent to 71.5 percent of pectoral fin behind posterior tip of pectoral fin. Additional counts and proportional measurements are recorded in tables 1-6. GEOGRAPHICAL RANGE: From Moreton Bay, Queensland, southward to Tasmania and New Zealand, and westward to Fremantle, Western Australia. Remarks: This species is related to C. oceanicus but differs in the origin of dorsal fin being more posterior in position and by having a greater number of compressed teeth in upper jaw (fig. 2). Griffen (1936) in his revision of the eels of New Zealand, lists 2 species from that area and distinguishes the 2 species by the origin of dorsal fin and the thickness of the body. Conger verreauxi Kaup Conger verreauxt Kaup, 1856a, p. 72 (type locality Australia). Conger vulgaris Giinther, 1870, p. 39, items “‘x’”’ and “‘y,”’ (Tasmania). [?] Conger labiata Castelnau, 1879, p. 396 (Port Jackson, New South Wales). Leptocephalus conger Waite, 1911, p. 164 (between Timaru and Porangahan Bay on east cost of New Zealand). Leptocephalus labiatus Phillipps, 1932, p. 229, fig. 3. Leptocephalus verreauxi Griffen, 1936, p. 15. Conger wilsont Waite, 1921, p. 49, fig. 74; 1923, p. 72, fig. 54. SPECIMENS STUDIED: A specimen in the British Museum, examined by Dr. Marshall, and Kaup’s type from the original description. Diagnosis: Two rows of teeth in jaws; vertebrae 152 to 156; origin of dorsal fin about over the posterior tip of pectoral fin; diameter of eye small. Description: Length of head 131 to 143; tip of snout to origin of dorsal fin 202; tip of snout to anus 377 to 412; snout length 39; diame- ter of eye 11; length of pectorals 39 to 59; origin of dorsal fin 9.43 percent of pectoral fin behind posterior tip of pectoral fin forward; two rows of teeth in jaws laterally, 54 to 55 precaudal and 97 to 102 caudal vertebrae. Additional counts and _ proportional measure- ments are recorded in tables 1-3, 5, 6. 958 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 GEOGRAPHICAL RANGE: From southern Queensland southward to Tasmania and New Zealand. Remarks: In the English translation, Kaup’s (1856b, p. 115) locality of the type specimen of Conger verreauxt was omitted. Kaup’s description (1856b, p. 73) states ‘‘nach meinen verehrten Freunde Julius Verreaux genannt, welcher ihn in Australien sammelte.”’ Whitley (in Griffen, 1936, p. 15) thinks that the type probably came from Tasmania, where Verreaux collected. Castelnau (1879, p. 396) and Phillipps (1932, p. 229) mention a small tentacle on each side of the snout; if this is so, C. verreauwzi is the only species in the genus that has such tentacles. There are two species of Conger in Australian waters. The main difference between them appears to be in the number of vertebrae and the origin of the dorsal fin, which is in a more anterior position in C. verreauxi. Conger oceanicus Mitchell Fieurs&s 2, 3; Puate 1,c Anguilla conger Mitchell, 1814, p. 360. Anguilla oceanica Mitchell, 1818, p. 407 (type locality, off New York, near Bloch Island).—Storer, 1846, p. 234. Conger occidentalis De Kay, 1842, p. 314, pl. 58, fig. 172. Isognatha oceanica Gill, 1861, p. 56. Echelus conger Jordan, 1888, p. 90. Leptocephalus conger Jordan and Davis, 1892, p. 664 (in part).—Jordan and Evermann, 1896, p. 354 (in part), pl. 57, fig. 148.—Smith, 1907, p. 111, fig. 35.— Fowler, 1912, p. 9 (in part). Conger conger Bigelow and Welsh, 1925, p. 86, fig. 36.—Hildebrand and Schroeder, 1928, p. 116.—Jordan, Evermann and Clark, 1930, p. 79 (in part). Conger oceanica Bigelow and Schroeder, 1953, p. 154, fig. 70. SPECIMENS STUDIED: 48 specimens, ranging in total length from 152 to 1052 mm., from the following localities: Massachusetts, USNM 16961, 49682, 126121, 148152 (2) MCZ 35806; Connecticut, USNM 16027, 43098, 14053 (2); Rhode Island, USNM 20745; New York, USNM 48941; New Jersey, USNM 6827, 28717, 28765, 33053, 36908, 118205 thru 118210 (7); Virginia, USNM 32796-7-8, 90395; Chesapeake Bay, USNM 127218; North Carolina, USNM 131327, 134184, CNHM 39999, 40000, 40331, AMNH 4369 (3), SNHM 1880; South Carolina, USNM 7418, 25352, 25358, SNHM 10314; Florida (New Smyrna), USNM 62652; Mississippi, USNM 164239 (4); Atlantic Coast of United States, USNM 44341. Diacnosis: Pores in lateral line 39 to 44; pectoral rays 16 to 18; vertebrae 140 to 147; diameter of eye 17 to 28. Description: Tip of snout to origin of dorsal fin 185 to 230; length of pectoral fins 37 to 62; supratemporal pore 1; preoperculo- EELS OF THE GENUS CONGER—KANAZAWA 259 mandibular pores 9; vertebrae 85 to 96 caudal and 50 to 51 precaudal; in one specimen dorsal rays 273; anal rays 187; compressed teeth in upper jaw 27 to 65, in lower jaw 28 to 59; two rows of teeth in jaws, inner row conical; premaxillary tooth-patch wider than long; upper edge of gill opening near midpoint of pectoral fin base; rictus of jaw below middle of eye. Additional counts and proportional measure- ments are recorded in tables 1-3, 5, 6, and in figure 2. DistriBuTion: On eastern Atlantic coast from Massachusetts to Florida, and Gulf of Mexico to Pascagoula, Miss. Bigelow and Schroeder (19538, p. 156) record adults from North Truro, Cape Cod, and near Provincetown in Cape Cod Bay, Mass. The Oregon collected four specimens just south of Pascagoula, Miss., at a depth of - 260 fathoms in an 80-foot balloon trawl. Remarks: There is a clinal difference in the length of head and snout from north to south. The forms from New England have a shorter head and snout than those of Carolinas and Florida; those from the Gulf of Mexico have the longest. Length of head in thousandths of total length Area 130 | 135 | 140 | 145} 150 | 155 | 160 | 165 | 170 | 175 | 180 New England States____| 1 1 3 1 2| 3 To eet ese Sie New Jersey—Virginia- __-_|_-_-_- 1 3 5 2 3 1 Oy eee sees | aa North Carolina south- SV CELT Cl eaten el eh ee ee eee |e oe 2 1 3 3 3 5 a Cul Of MexicOns 2.2 sate tesla cece teteaee Dieses! Dees 1 | Length of snout in thousandths of total length Area Ge tate nie ae le et eee 27. 6-|80. O-|32. 6-|35. O-|37. 5-|40. O-|42. 6-|46. O-|47. 6- 29.9 |82. 4 184.9 \387. 4 |89.9 |42. 4 |44.9 |47. 4 149.9 New England States___-_ 2 3 1 3 1 Lf he ste 2.22 ees New Jersey—Virginia____ 1 4 4 2 4 A i eee ee North Carolina south- WAT s6- d= fos oe | i 2 i 4 3 Balt oatalns sees SEU Oi RVC RICO = ars Oe eles al eh ha | a Dei \ Seyi tia 2 This species is related to C. wilsona from Australia, but differs from it by having the origin of dorsal fin more anterior in position and a fewer number of compressed teeth in the upper jaw (fig. 2). 260 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 References Autis, E. P. 1903. The lateral sensory system in the Muraenidae. Inter. Monat. Anat. Physiol., vol. 20, pp. 125-170, plates. Aoyaal, Hyoz1 1943. Coral fishes. Pt. 1, xii+224 pp., 54 figs., 37 pls. BARNARD, K, H. 1925. A monograph of the marine fishes of South Africa. Pt. 1. Ann. South African Mus., vol. 21, 418 pp., 18 text-figs., 17 pls. BEAN, TARLETON H., aND DresseEt, H. L. 1884. 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An improved graphical analysis and comparison of series of samples. Syst. Zool., vol. 2, No. 2, pp. 49-56, 92. JENKINS, J. TRAVIS. 1925. The fishes of the British Isles. 408 pp. JorRDAN, Davip STARR 1888. Manual of the vertebrate animals of the northern United States. Ed. 5, iii+-375 pp. JORDAN, Davip Starr, AND Davis, BRADLEY Moore 1892. A preliminary review of the apodal fishes or eels inhabiting the waters of America and Europe. Rep. U. 8. Comm. Fish and Fisheries for 1888, pt. 16, appendix No. 10, pp. 581-678, pls. 73-80. JoRDAN, Davin Starr, AND EVERMANN, BARTON WARREN 1896. Fishes of North and Middle America. U.S. Nat. Mus. Bull. 47, pt. 1, 1240 pp. EELS OF THE GENUS CONGER—KANAZAWA 263 1905. The shore fishes of the Hawaiian Islands with a general account of the fish fauna. Bull. U. 8. Fish Comm. (1903), vol. 23, pt. 1, xxviii+ 574 pp., 73 col. pls., 65 pls., 228 figs. JoRDAN, Davin Starr; EVERMANN, BARTON WARREN; AND CLARK, HOWARD WALTON. 1930. Check list of the fishes and fishlike vertebrates of North and Middle America north of the northern boundary of Venezuela and Colom- bia. Rep. U. 8. Comm. Fish. (1928), pt. 2, 670 pp. JoRDAN, Davip Starr, AND GILBERT, CHARLES H. 1882. List of a collection of fishes made by Mr. L. Belding near Cape San Lucas, lower California. Proc. U.S. Nat. Mus., vol. 5, pp. 378-381. JoRDAN, Davip Starr, AND GUNN, JAMES ALEXANDER JR. 1899. List of fishes collected at the Canary Islands by Mr. O. F. Cook, with descriptions of four new species. Proc. Acad. Nat. Sci. Philadelphia (1898), pp. 339-346. JorpAN, Davip Starr, AND Husss, Carn LEAVITT 1925. Record of fishes obtained by David Starr Jordan in Japan, 1922, Mem. Carnegie Mus., vol. 10, No. 2, pp. 93-346, pls. 5-12. JORDAN, Davip STARR, AND SNYDER, JOHN OTTERBEIN 1901. A review of the apodal fishes or eels of Japan, with description of nineteen new species. Proc. U. 8S. Nat. Mus., vol. 23, No. 1239, pp. 837-890, 22 figs. KaMmouara, TOSsHIJI 1950. Description of the fishes from the Provinces of Tosa and Kishu, Japan. 3+288+5+48-+ 27 pp., 220 figs. Kavup, JOHANN JACOB 1856a. Uebersicht der Aale. Arch. Naturg., vol. 22, pt. 1, pp. 41-77. 1856b. Catalogue of apodal fish in the collection of the British Museum, London, 163 pp., 19 pls. KLUNZINGER, CARL BENJAMIN 1871. Synopsis der fische des Rothen Meeres. II. Verh. Zool. Bot. Ges. Wien, vol. 21, pp. 441-668. LINNAEUS, CARL 1758. Systema naturae ... , ed. 10, vol. 1, 824 pp. Lowe, R. T. 1837. A synopsis of the fishes of Madeira, with the principal synonyms, Portuguese names and characters of the new genera and species. Trans. Zool. Soc. London, vol. 2, pt. 3, pp. 173-200. MarsuBaRA, K1yoMaTsu 1955. Fish morphology and hierarchy. Ishizaki-Shoten, Tokyo, pt. 1, xi+ 789 pp. MEEK, SetH E., anp HILDEBRAND, SAMUEL F. 1923. The marine fishes of Panama. Field Mus. Nat. Hist., Zool. ser., vol. 15, pt. 1, 330 pp., 24 pls. MELLIss, JOHN CHARLES 1875. St. Helena .... Pp. 100-113, pls. 19-21. METZALAAR, J. 1919. Report on the fishes collected by Dr. J. Boeke, in the Dutch West Indies, 1904-1905, with comparative notes on marine fishes of tropical West Africa. Jn Boeke, Rapport betreffende ... toestand ... Visscherij . . . Zeeproducten in de Kolonie Curacao. . . 1904, 315 pp., 64 figs. 264 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 MitTcHELL, SAMUEL LATHAM 1814. The fishes of New York. Trans. Lit. Phil. Soc. New York, vol. 1, pp. 355-492, 6 pls. 1818. Description of three species of fish, Anguilla oceanica, Gadus albidus, Salmo amethystus. Journ. Acad. Sci. Philadelphia, vol. 1, pp. 407- 412, Moreav, E. 1881. Histoire naturelle des poissons de France. Vol. 3, 697 pp. Mort, TAMEZO 1952. Check list of the fishes of Korea. Mem. Hyogo Univ. Agr., vol. 1, No. 3, Biol. ser., No. 1, 228 pp. Mort, TAMEzO AND Ucuipa, K. 1934. A revised catalogue of the fishes of Korea. Journ. Chosen Nat. Hist. Soc., No. 19, p. 1-23. Navarro, F. dE P. 1942. Note preliminar sobre los peces de la Costa de Africa, desde el Cabo Bojador a la bahia de Tanit. Bol. Real. Soc. Esp. Hist. Nat., vol. 40, pp. 189-214, pl. 16. 1943. La pesca de arrostre en los fondos del Cabo Blanco y del Banco Arguin (Africa Sahariana). Trabajos Inst. Esp. Ocean. Madrid, No. 18, 228 pp., 38 pls., 33 figs. NicuHois, JoHN TREADWELL 1921. A list of Turk Islands fishes, with a description of a new flatfish. Bull. Amer. Mus. Nat. Hist., vol. 44, art. 3, pp. 21-24, pl. 3. Nore, A. 1935. Fauna Marinha de Portugal, descricao dos peixes de Portugal. Pt. 3, 574 pp., 77 pls. OarLBy, J. Douaias 1898. New genera and species of fishes. Proc. Linn. Soc. New South Wales, vol. 23, pp. 280-299. OKEN, L. 1817. Isis .’.. , vol..1, 1575 pp.; 12"pls: PrruGis, ALBERTO 1891. Appunte sopra alcuni pesci sud-Americani conservati nel Museo Civico de Storia Naturale di Genova. Ann. Mus. Civ. Storia Nat. Genova, ser. 2, vol. 10, pp. 605-657, figs. Puiuurpes, W. J. 1921. Notes on the edible fishes of New Zealand with a record of fishes exposed for sale in Wellington during 1918. New Zealand Journ. Sci. Tech., vol. 4, No. 3, pp. 114-125. 1932. Notes on new fishes from New Zealand. New Zealand Journ. Sci. Tech., vol. 13, No. 4, pp. 226-234, 5 text-figs. Puriuiprs, W. J. AND HODGKINSON 1922. Further notes on the edible fishes of New Zealand. New Zealand Journ. Sci. Tech., vol. 5, No. 1, pp. 91-97. PLAYFAIR, R. LAMBERT, AND GUNTHER, ALBERT C. L. G. 1866. The fishes of Zanzibar. xiv+153 pp., 19 pls. Pory, FELIPE 1858. Memorias sobre la historia natural de la Isla de Cuba. Vol. 2, 342 pp., 19 pls. EELS OF THE GENUS CONGER—KANAZAWA 265 1867. Peces Cubanos. Rep. Fis.-Nat. Isla Cuba, vol. 2., pp. 229-268, 2 pls. 1868. Synopsis piscium cubensium. Rep. Fis.-Nat. Isla Cuba, vol. 2, pp. 279-484, 4 figs. 1876. Enumeratio piscium cubensium. Anal. Soc. Espafiola Hist. Nat., vol. 5, pp. 131-218, 2 pls. Pout, JULIUS 1884. Journ. Jus. Godeffroy, Cat. 9 Pisces pp. 25-46. (Not seen, Fide Whitley, 1927.) PosapA ARANGO, ANDRES 1909. Estudios Cientificos, 432 pp. RANZANI, CAMILLO 1840. De novis speciebus piscium; dissertationes quatuor. Acad. Sci. Inst. Bonon, vol. 4, pp. 65-83, pls. 8-13. REGAN, TaTE 1912. The osteology and classification of the Teleostean fishes of the order Apodes. Ann. Mag. Nat. Hist., ser. 8, vol. 10, pp. 377-387, 2 text- figs. RICHARDSON, SIR JOHN 1844— Ichthyology of the voyage of H. M. 8. Hrebus and Terror. viiit+ 1848. 139 pp., 60 pls. Risso, ANTOINE 1810. Ichthyologie de Nice, ou Histoire naturelle des poissons du départment des Alpes Maritimes. 388 pp., 11 pls. 1826. Histoire naturelle des principales productions de l Europe méridionale. et particulierment de celles des environs de Nice et des Alpes Maritimes. Vol. 3, xvi + 480 pp., 16 col. pls. Rivero, L. HowEii 1932. The apodal fishes of Cuba. Proc. New England Zool. Club, vol. 13, pp. 3-26. 1938. List of fishes, types of Poey in the Museum of Comparative Zoology. Bull. Mus. Comp. Zool., vol. 82, pp. 169-227. RUppeLL, WILHELM PETER EpUARD SIMON 1828. Atlas zu der Reise im Nordlichen Afrika. ... Fische des rothen Meers, vol. 4, 141 pp., 35 pls. Scumipt, J. 1931. Eels and conger eels of the North Atlantic. Nature, vol. 128, No. 3232, pp. 602-604, 3 _ figs. 1932. Danish eel investigations during 25 years, 1905-1930. Carlsberg Foundation, Copenhagen, 16 pp. ScHuuitz, LEONARD P. 1943. Fishes of the Phoenix and Samoan Islands, collected in 1939 during the expedition of the U. 8. 8. “Bushnell.” U. S. Nat. Mus. Bull. 180, x + 316 pp., 27 figs., 9 pls. ScHuitz, LEONARD P., AND COLLABORATORS 1953. Fishes of the Marshall and Marianas Islands. U.S. Nat. Mus. Bull. 202, vol. 1, xxxii + 685 pp., 90 figs., 74 pls. Smiryu, HuaeH M. 1907. The fishes of North Carolina. North Carolina Geol. Econ. Surv., vol. 2, xi+ 453 pp., 21 pls., 187 figs. Smiru, J. L. B. 1949. The sea fishes of Southern Africa. 550 pp., 102 pls., 1232 figs. 266 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 STauL, AUGUSTIN 1883. Fauna de Puerto-Rico. 249 pp. Srassano. E. 1890. La pesca sulle spiagge Atlantiche del Sahara. Ann. Agricl. Roma, No. 172, 103 pp. STEINDACHNER, FRANZ 1896. Bericht iiber die wihrend der reise Sr. Maj. Schiff ‘‘Aurora’’ von Dr. C. Ritter V. Mieroszewski in den Jahren 1885 und 1896 gesammelton fische. Ann. Naturh. Hofmus. Wien, vol. 11, pp. 197-230, pl. 4. Storer, Davin HuMPHREYS 1846. A synopsis of the fishes of North America. Mem. Amer. Acad., vol. 2, 298 pp. TANAKA, SHIGEHO 1911. Figures and description of the fishes of Japan. Vol. 4, pp. 51-70, pls. 16-20. 1912. Figures and description of the fishes of Japan. Vol. 5, pp. 71-86, pls. 21-25. THompson, WILLIAM 1856. Natural history of Ireland. Vol. 4, p. 227. TorTONESE, E. 1939. Resultati ittiologici del viaggio di cireumnavigazione del globo della R. N. ‘Magenta’ (1865-68). Boll. Mus. Zool. Anat. Comp. Torino, vol. 47, ser. 3, No. 100, 245 pp., 9 pls., 17 text-figs. TREWAVAS, ETHELWYNN 1932. A contribution to the classification of the fishes of the order Apodes, based on the osteology of some rare eels. Proc. Zool. Soc. London, pp. 639-659, 4 pls., 9 text-figs. U1, N. 1931. The fishes of the Province of Kishi, Wakayama Prefecture. (Not seen, fide Kamohara, 1950, p. 52.) VALENCIENNES, A. 1841. In Eydoux and Souleyet, Voyage autour da Monde exécute pendant des années 1836 et 1837 sur la corvette ‘‘La Bonito.’ Poissons. Vol. 1, pp. 157-215, 10 pls. 1847. Poissons, Catalogue des principales espéces de poissons rapportées del’ Amérique méridionale. (Jn D’Orbigny, Voyage dans 1’ Amérique méridionale. . . . Vol. 5, pt. 2, pp. 1-11, pls. 1-16. VINCIGUERRA, D. 1893. Catalogo dei pesci della isole Carnaie. Atti Soc. Ital. Sci. Nat., vol. 34, pp. 295-334. WalreE, Epaar R. 1911. Scientific results of the New Zealand government trawling expedition, 1907. Pisces: Pt. II. Rec. Canterbury Mus., vol. 1, No. 3, pp. 157-272, pls. 24—42. 1921. Catalogue of the fishes of South Australia. Rec. South Australian Mus., vol. 2, No. 1, pp. 1-199. 1923. Fishes of South Australia. 2438 pp., illus. EELS OF THE GENUS CONGER—KANAZAWA 267 WEBER, Max 1913. Szboga-Expeditie. ... Monogr. 57, livr. 65, Die Fische. xii + 710 pp., 12 pls., 123 figs. WEBER, Max, AND DE Braurort, L. F. 1916. The fishes of the Indo-Australian Archipelago. Vol. 3, xv + 455 pp., 213 figs. WuitLey, GILBERT P. 1927. A check list of fishes recorded from Fijian waters. Journ. Pan- Pacific Res. Inst., vol. 2, No. 1, pp. 3-8. 1935. Studies in Ichthyology, No. 9. Rec. Australian Mus., vol. 19, No. 4, pp. 215-250, pl. 18. 1944, Illustrations of some western Australian fishes. Proc. Roy. Soc. New South Wales, pp. 25-29, 6 text-figs. YARRELL, WILLIAM 1831. On the anatomy of conger eel (Conger vulgaris Cuv.) and on the difference between the conger and freshwater eels. Proc. Zool. Soc. London, pt. 1, pp. 158-159. U. S. GOVERNMENT PRINTING OFFICE: 1958 ee i ed FD, : ; , ow) ae a cist ii ‘ed in ook ee sO ala 110 nm F | i / st teat Ge, o on ee ae hth ny, els ay h ae a By Rae ee cane ; tl : Brana Py een a ae a iF Iyer id Sn pate PAder MR saan aoc a tnd / Bley ee” ru wpe mab thoes ie dita ‘aboot He na e eal tle a | eed ercot, oy ae cen 0 Sia WS t iy" Ps Neh : ey | peeve Web PRA eb deen ART al ball Ss Sas Rd a dove: ‘iter iciaded “OU we ete beste, meee ‘a Pay Pe Tae n m7 hae: ~ ; Bt Ay Mis. 21S Sty ae nt 88 om! "arate PhetiselVec.W bobled Moneta Hep sonal navies si | Pgtolegs th OR GL ey aloe Hein, wert rata Pee yee rent Me et AD get Eee PLY GA AW Teyn yy Pa! eg Atall’ ih ‘ata af? ty Se toot copy't “sls ‘patito Lie in Newt woes. aah ante: ADE! 5 by “pA aae ant! bt ODA Hie ah ty 7 i a 3) ’ j Ab ] i. 7 Ag )) wee Fi { i : y) ; ») hi ty (ie iY | sah tthh a : i) Deine. 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Remarks on Helmintholepis vermiculatus Cockerell PLATE 1 The North American Cretaceous fish Helmintholepis vermiculatus Cockerell (1919) is based on a single disassociated scale (USNM 8677). The specimen was collected in 1914 by T. E. Willard, of the U. S. Geological Survey, 1% miles northeast of Milliken, sec. 13, T. 5 N., R. 67 W., Weld County, Colo., in exposures of the Fox Hills sandstone (Maestrichtian). The scale, presumably deriving from the left flank of a fish, is com- paratively large and, although composed of successive fine lamellae, is of remarkable over-all thinness. The major portion of the apical margin is missing. The outline of that part impressed in the matrix, however, indicates a roughly 5-sided form for the scale. Of these edges the broadly rounded apical margin has the greatest length. The dorsal and ventral borders converge slightly toward each other, anteriorly, to distinct but obtuse basal angles. The basal margin is divided into upper and lower segments by a low, forwardly convex, median lobe. The greatest length of the scale (21.4 mm.) passes along a longitudinal axis a little nearer the dorsal border than to the ventral 468792—58 269 270 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 one, and the perpendicular of maximum depth (20.9 mm.) is situated far posteriorly in the vertical plane of the dorsal and ventral apical angles. The nucleus is a concentrated locus on the longitudinal axis of the element, somewhat basal of the middle length of the scale. Surround- ing this area in concentric fashion, relatively fine circuli (about 34 in 1 mm.) cover the dorsal, ventral, and basal quadrants of the external surface. No basal radii are evidenced, but suggestive of such struc- tures are a series of sinuous folds in the courses of the circuli, arranged in an anteroposterior line across the median basal field. The circuli terminate, posterodorsally, along a diagonal extending from the nu- cleus to the middle point of the dorsal border and, posteroventrally, along a more acutely oblique diagonal extending from the nucleus to a point near the ventral apical corner of the scale. The apical quad- rant is, thus, a triangular field with a nuclear angle of approximately 120°. The preserved portion of this part is ornamented with short vermiculate markings which in the center of the field are directed more or less longitudinally and on either side of this median area in zones adjacent to the apical diagonals, vertically. Peripherally the anterior terminations of generally longitudinal apical radii, spaced 4.5 to 5in 1 mm., may be observed. From the comparisons discussed below, it seems feasible to postulate the passage of these radii posteri- orly across a missing marginal and hyaline zone to end in notches along the dentate apical margin of the scale. The scale of Helmintholepis vermiculatus cannot be distinguished from anterior flank scales of the upper Cretaceous Pierre shale fishes Pelycorapis berycinus Cope (1877) and Paleoclupea dakotaensis Dante (1942). From well-preserved specimens in the U. S. National Museum, the skulls, as well as the scales of the latter two genera, are identical. It must be concluded, therefore, that Helmintholepis vermiculatus and Palecoclupea dakotaensis are synonyms of Pelycorapis berycinus. Unfortunately, the generic term Pelycorapis does not appear available for the berycinus species. Examination was re- cently made in the American Museum of Natural History of the genotypic specimen, Pelycorapis varius, defined by Cope (1874) on the basis of a fragmentary fish from an undetermined horizon in the “Benton” of Kansas. There can be little doubt of at least close relationship, if not even conspecific identity, between P. varius and the North American elopid fish Thrissopater intestinalis Moodie (1911). Woodward (1901, pp. 353, 354, 616) early advocated affinity, at least at the family level, between the two species assigned to the generic name Pelycorapis (Cope, 1874, 1877), Syllaemus (Cope, 1875), Apsopeliz (Cope, 1871), and Leptichthys (Stewart, 1899). Apsopelix CRETACEOUS FISHES—DUNKLE 27h from the “Benton” of Kansas is apparently known only by the fragmentary type specimen. No differences can be observed between the scales and vertebrae of this specimen and the corresponding parts of Syllaemus and Leptichthys. Most authorities (for example, Hay, 1929; Romer, 1945) continue to treat Syllaemus and Leptichthys as distinct genera, but in addition to their similarity of scale and vertebral structure, no major variation has been detected as yet among available skulls of the two genera. Certainly the North American materials which have been variously referred to either Apsopeliz, Syllaemus, or Leptichthys constitute a basically homo- geneous group peculiar to a number of marine strata exposed in the Western Interior, which in geologic range seem restricted to the interval between either the late Cenomanian or early Turonian and the Santonian stages of the upper Cretaceous. Relationship between the Apsopelix-Syllaemus-Leptichthys complex of forms and the late upper Cretaceous berycinus species is denoted by the common possession of distinctive modifications of the bone pattern of the skull, and by general over-all similarities in body habit, including the structure and relative position of fins. As might be anticipated, on the other hand, certain regularly occurring variations are to be observed. Of these, the proportions of the parts of the visceral skeleton differ and these features may account for the curious apposition of the mandibles commonly displayed by specimens of Syllaemus and Leptichthys. Further, the scales of the latter two genera have a large and diffuse nuclear field mainly devoid of any ornamentation and, depending on body position, with the apical quadrant covered proximally, either with complete and entirely vertical circuli or with a median hyaline zone. Such comparisons prompt, in at least tentative solution of the problem, allocation of the berycinus species to the oldest generic name unquestionably asso- ciated with this group, Apsopeliz. To be demonstrated shortly in another connection, the compact assemblage of fishes under consideration is of elopine stock. The members, thus, display interesting specializations of the most primitive teleostean stage of morphologic organization. They comprise a distinctive component of upper Cretaceous marine faunas and would seem best interpreted as a line of pelagic plankton feeders. Sincere thanks are extended to Drs. Bobb Schaeffer, American Museum of Natural History, and Robert W. Wilson, Kansas Uni- versity Museum of Paleontology, for courtesies in making pertinent materials available. The photograph was made by Jack Scott of the Department of Geology, U.S. National Museum. 262 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 II. Reassignment of Petalolepis? fibrillatus Cockerell PLATE 2 The generic name Petalolepis first appeared in an unpublished catalog of fish remains from the upper Cretaceous Planer formation of Saxony, Germany, compiled by Prof. Moritz Steinla. Sub- sequently in formal description, Geinitz (1868) considered Petalolepis a synonym of the elopid genus Osmeroides (Agassiz, 1844) and proposed the name O. divaricatus for the identical Steinla specimens. Cockerell (1919, p. 173) interpreted the same scales as those of an albulid fish, and finding the name Petalolepis available for the reception of the hence reassigned divaricatus species, provisionally referred a new American species, fibrillatus, to the generic term. Heretofore known only by the one holotypic specimen (USNM 8662), two additional seales of Petalolepis? fibrillatus were recently recognized among the national collections of fossil fishes. The variety and individuality of fish scales have been amply demonstrated through the work of many investigations in all parts of the world. However, when attempts have been made to arrange these exoskeletal elements in developmental series, the resultant trends of structural variation, either parallel, convergent or divergent, appear to cross phylogenetic lines. In direct consequence, serious doubts are entertained by many ichthyologists not only of the ability but also of the use for identifying and naming fragmentary remains of such parataxial nature. Although disassociated fish scales probably never can constitute a completely satisfactory base for precise studies of broad scope, the future clarification of group relationships and corresponding refinement of fish classification, coupled with accumu- lated factual data concerning development at all ontogenetic and phylogenetic levels, will correct many of the difficulties surrounding the use of scales. Within restricted limits, on the other hand, scales have been proved effective tools. In the present case a new definition of Petalolepis? fibrillatus seems justified. The fossil specimens are entirely distinctive in fundamental structural characteristics and are not to be identified with the genotypic Petalolepis divaricatus (Geinitz) from Germany. Equally important, they are of potential strati- graphic utility as representative of practically unknown marine faunas from near shore and lagoonal environments for a seemingly short interval of the late upper Cretaceous in North America. Grateful acknowledgment is offered Dr. R. W. Brown, Dr. Lore R. David, and Mr. Shelton P. Applegate for advice and various other aids. The photographs accompanying this note are the work of Mr. Jack Scott, Department of Geology, U. S. National Museum. CRETACEOUS FISHES—DUNKLE 213 Family GADIDAE Paractichthys,' new genus Diacnosis: A fossil genus distinguished from living anacanthine fishes by the following combination of scale characteristics: propor- tionately large, subquadrangular form with central oblong nuclear field and single Jongitudinal median basal sulcus; peripheral basal circuli more or less longitudinal from apical diagonals to basal scale margin; more proximal basal circuli crescentic, paralleling the basal border and rather sharply recurved or acutely angulated on approach- ing the median sulcus; and short, sinuous apical circuli diverging slightly to the apical scale border from a faint median longitudinal axis. Type species: Paractichythys fibrillatus (Cockerell) (= Petalolepis Jibrillatus Cockerell, 1919). Paractichthys fibrillatus (Cockerell) Diagnosis: The same as for the genus (the only species). Typr: USNM 8662; one disassociated scale. GEOLOGIC AND GEOGRAPHIC OCCURRENCE: Upper Cretaceous Blair formation (middle Campanian, following Cobban and Reeside, 1952) in the NWMNE sec. 26, T. 21 N., R. 104 W., Sweetwater County, Wyo. REFERRED SPECIMENS: USNM 14517 from the topotypic horizon and locality in Wyoming; and USNM 21898 from the upper Cretaceous Fox Hills sandstone (Maestrichtian), near Milliken, in sec. 23, T. 4 N., R. 67 W., Weld County, Colo. Description: Seales generally of moderate size and ranging from examples with equal dimensions to ones longer than deep (measure- ments, in millimeters: 10-17 long by 9-12 deep). In outline, the elements are subquadrangular, although the apical portion was evi- dently weak and the posterior border of the scale is usually preserved with an evenly rounded profile. As here interpreted, the dorsal margin displays a lower convexity than does the ventral one. Anteri- orly these edges diverge slightly from each other to obtuse basal angles. The basal border is of low forward convexity but displays a prominent notched median lobe. The longitudinai axis is situated nearer the dorsal border than to the ventral, and the perpendicular of maximum depth lies anteriorly near the vertical plane of the basal angles. The oblong nucleus occurs adapical to the middle scale length, along the main longitudinal axis. Neither basal nor apical radii are present, although a median longitudinal sulcus extending between the 1 Derived from the Greek para, near; akte, shore; and ichthys, fish; referring to the inferred near-shore habitat occupied by the fish. 274 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 basal margin and the nucleus is a prominent structural feature of all known specimens. Circuli are relatively coarse (about 19 in 1 mm.). On the covered portion of the scale the central ones are crescentic, and paralleling the basal border are either sharply recurved or acutely angulated as their courses approach the median sulcus. Peripherally the circuli are more nearly longitudinal in their arrangement and intersect in very acute angles the basal scale margin after only slight median flexure. The exposed apical quadrant has a nuclear angle of from 75° to 80° and is covered with short, sinuous circuli, which are directed backward and slightly away from a faint median longitudinal axis. Remarks: The scale characteristics of Paractichthys are more com- parable to those of anacanthine fishes than to any other living group (David, 1956). While very probably not closely related to any extant member, all features exist in one or another genus of the Gadidae. It is upon this basis that the family assignment has been made. Heretofore, the codfishes and their allies have been recognized only from marine sediments of Tertiary age. Paractichthys, thus, extends the known geologic range of the gadid fishes into the upper Cretaceous, and the genus may prove, on recovery of more complete remains, a key to the establishment of the complete phyletic history of this dis- tinctive assemblage of fishes. Ifl. Comments on the Status of Cyclolepis stenodinus Cockerell PLATE 3 The genus Cyclolepis was named and described by Geinitz (1868). The genotypic species, C. agassizi, is based on unassociated scales from the Planer formation (Cenomanian and Turonian) of Saxony, which display a subcircular or ovate outline with nearly central nu- cleus, concentric circuli, and an absence of either radii or ctenoid structures. Although various opinions have been held regarding the affinities of the form, most current classification outlines list Cyclolepis with the salmonoid fishes (Romer, 1945; David, 1946). In 1919, Cockerell described an isolated scale from the upper Maestrichtian (Fox Hills sandstone) of Colorado under the name Cyclolepis stenodinus. 'The Cockerell type (USNM 8703) is exposed in internal aspect, displaying prominent annular growth rings, and as thus observed would appear to compare closely with the characteriza- tion of Cyclolepis Geinitz. However, recent examination under highly refractive liquids and the binocular microscope has revealed previously unreported features of the embedded external surface of the scale. The new information creates serious doubt as to the correctness of the generic reference and even as to the validity of the Colorado species. PROC. U, S. NATIONAL MUSEUM, VOL. 108 DUINKLE, PLATE. 1 Photograph of type scale Helmintholepis vermiculatus Cockerell (1919), herein reassigned to Apsopelix berycinnus (Cope), with missing margins restored. Reproduction approxi- mately * 4. Explanation of abbreviations: ant, anterior; and p, dorsal. PROC. U. S. NATIONAL MUSEUM, VOL, 108 DUINIKKEE;, REATE 2 Ant. Ant. Scales (A, USNM 8662 type; B, USNM 14517) from Paractichthys fibrillatus (Cockerell). Reproduction the upper Cretaceous Blair formation in Sweetwater County, Wyo. approximately 10. Explanation of abbreviations: ant, anterior; p, dorsal. PROC. U. S, NATIONAL MUSEUM, VOL. 108 DUNKLE, PLATE 3 from right flank of fish, drawn with anterior basal edge oriented to the right; external surface ornamentation somewhat diagrammatized, and small vertical striae between cir- culi omitted. (Magnification 9.) CRETACEOUS FISHES—DUNKLE 275 The original description of Cyclolepis stenodinus now may be ex- panded. A relatively thin scale of moderate size (measurements, in millimeters: 12.4 long by 9.7 deep). In outline, subcircular with broadly rounded and confluent dorsal, apical, and ventral margins; obtuse but distinct basal angles; and a centrally lobed basal edge. Perpendicular of maximum depth adjacent to the basal angles and thus situated far anteriorly. Nucleus a concentrated focus both in- ternally and externally, slightly apicad of central. No basal or apical radii, but suggestive of the former are several (three or more) indis- tinct folds or undulations in the courses of the circuli across the central part of the basal field, arranged in lines convergent posteriorly toward the nucleus. Circuli on anterior inserted portion of scale coarse (18 in 1 mm. centrally and 12 in 1 mm. peripherally above and below, and in their posterior zone of termination along the apical diagonals) ; essentially vertical in arrangement although slightly curved in an anterior direction around the nucleus and bent posteriorly both along the ventral basal diagonal and in an upper peripheral zone so that the dorsal ends of the lines intersect the scale margin at more acute angles than do the ventral ends. Interspaces between the more widely separated circuli of the dorsal and ventral quadrants marked by short, fine, parallel striae (10 to 12 in 0.1 mm.), directed at acute angles to the circuli and generally perpendicular to the anteroposterior axis of the scale. Apical field triangular with a nuclear angle of about 101°; ornamented with fine circuli (34-36 in 1 mm.) arranged in parallel but slightly undulating crescentic courses to meet the termina- tions of the basal circuli along the apical diagonals in angles of ap- proximately 90°. Internal surface of scale distinctly marked by sparsely tuberculous and concentric annular growth rings which are in places so closely set they resemble concentric circuli. No materials of the genotypic Cyclolepis agassizt are available for comparison. Notwithstanding, the characteristics of the scale of Cyclolepis stenodinus are not those either commonly or easily asso- ciated with the scales of salmonoid fishes. Rather, scales of its type with essentially vertical circuli which are coarser in the basal portion than in the apical one and with nearly central nuclei are better recog- nized as pertaining to the Synentognathi, Scombrcidea and Caran- goidea. During search among these latter groups for more closely comparable materials, striking and fundamental similarities were ob- served between C. stenodinus and Hemilampronites hesperius Cockerell (1919). In fact, only minor differences of shape, absolute size, and slight details of sculpture exist between the two species and these are well within the range of variation to be expected between scales from different parts of the body of one fish. Interestingly also, the 276 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 holotype of H. hesperius (USNM 8713) and that of C. stenodinus were both collected by T. E. Willard of the U. S. Geological Survey on Sept. 14, 1914, from the same exposure of Fox Hills sandstone in sec. 23, T. 4 N., R. 67 W., southwest of Milliken, Weld County, Colo. It is concluded that Cyclolepis stenodinus is a synonym of Hemilampro- nites hespervus. Cockerell (1919) referred Hemilampronites hesperius to the Hemi- ramphidae. Such a relationship is indeed supported by the centrally lobate basal border and the marked angle of meeting of the apical and basal circuli displayed by the scales. As remarked by David (1946) in another case, however, the proportionately long, broadly rounded outline and lack of discrete basal radii suggest relationship also with either or both the Scomberesocidae and Exocoetidae, among other Synentognathi. This distinctive combination of structural fea- tures, which serves adequately to separate the Cretaceous species from the Tertiary and living members of the order, may well indicate a primitive ancestral type. Unfortunately, the fossil record of the flying fishes and their allies is meager. Until available information permits reconstruction of the evolution of the order, no precise desig- nation of the relationships of H. hesperius seems feasible. References Aaassiz, L. 1844. Recherche sur les poissons fossiles. Vol. 5, pt. 2, p. 103. Coppan, W. A., AND Ruxsipe, J. B., JR. 1952. Correlation of the Cretaceous formations of the Western Interior of the United States. Bull. Geol. Soc. Amer., vol. 63, No. 10, pp. 1011-1044, 1 pl. CocKkERELL, T. D. A. 1919. Some American Cretaceous fish scales. U.S. Geol. Surv. Prof. Pap. 120, pp. 165-188, pls. 31-37. Corr, E. D. 1871. On the fossil reptiles and fishes of the Cretaceous rocks of Kansas. U. 8S. Geol. Surv. of Wyoming and portions of contiguous Terri- tories. 2d Ann. Rep., pp. 385-424. 1874. Review of the Vertebrata of the Cretaceous period found west of the Mississippi River. Bull. U. S. Geol. Geog. Surv., vol. 1, No. 2, pp. 3-48. 1875. The Vertebrata of the Cretaceous formations of the west. Rep. U.S. Geol. Surv. Terr., No. 2, pp. 1-303, 10 figs., 55 pls. 1877. Report on the geology of the region of the Judith River, Montana, and on vertebrate fossils obtained on or near the Missouri River. Bull. U. S. Geol. Geog. Surv., vol. 3 (art. 29), pp. 565-597, 5 pls. Dante, J. H. 1942. Description of fossil fishes from the upper Cretaceous of North America. Amer, Journ. Sci., vol. 240, pp. 339-348, 2 pls. CRETACEOUS FISHES—DUNKLE Die Davi, L. R. 1946. Upper Cretaceous fish remains from the western border of the San Joaquin Valley, California. Contr. Paleont., Carnegie Inst. Wash- ington Publ. 551, pp. 81-112, figs. 1-11, pls. 1-3. 1956. Tertiary anacanthin fishes from California and the Pacific North- west; their paleoecological significance. Journ, Paleont., vol. 30, No. 3, pp. 568-607, 27 figs., 4 pls. Gerinitz, H. B. 1868. Die fossilen Fischschuppen aus dem Planerkalke in Strehlen. Denk- schr. Gesell. f. Natur- und Heilkunde Dresden, pp. 31-48, 4 pls. Hay, 0. P. 1929. Second bibliography and catalogue of the Fossil Vertebrata of North America. Carnegie Inst. Washington Publ. No. 390, vol. 1, vii + 865 pp. Mooprg, R. L. 1911. sy he oe eran Ey chow ete! Pisuialnon ij, apn Vol eins eit iin ~venyathi Pvhias alt’ iy FRR Hosen t Wareinigd ay in nasa a ek: init Ppt oe “ina har, Ly hee wie Wipe ie yu Senter ripe. ‘Cvsnnsan Le ‘yet d ey, (nye Bes raheg dt 2 ions Wye ‘8 Any oe iow, Wik Vs, a tthe Boe, | oa ii dpa Vietbeheapt ® ‘hw! Placed Stony ab nai ‘apse, Beaty, "Oa SORp a ar Maer TP gua 4 pity cae ap er Sa ig SN as ade sh a tiie rlueee Or ee ra) is id ae way) Lag. Aa ot pe TAP. i » AG til | ee wt LAN, aeameeiy steel ivaalk Nelo pl AA NR a oe het ches | Dn oni Aintiny, Phe, Wee, ale, Bay, Op caput pone hi we ssi EEK UM SEN iden he ‘ne Wes \ yalon ee Bieraiey Yannnwavee 0/4) ee oe a oe 7 i? ‘ - ae : ; : oh i nue ; K i 4 : la 7 : 71 it f a ih . or eet + i ‘os i : a gist ce ‘ ; i ae oi bi wie a WA ey TFs UR 4 WY iss Cates Se NIE a Bik | MM nM hair ae ye. A. iy ea ty A tal ; a ae o Ria B ihe Lin ei rule a a ih aie i a ae ts lige Wi Be oh iN) i nite i Arash nal ee PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1958 No. 3402 TAXONOMY AND NOMENCLATURE OF THREE SPECIES OF LONCHURA (AVES: ESTRILDINAE) By Kennetu C. Parkes! In August and September 1956 I collected birds for a virus re- search project in central Luzon, Philippine Islands. The museum skins and skeletons prepared as a by-product of this project are now in Carnegie Museum, and represent the first modern collection of Asiatic birds in that institution. Virtually all of the work, therefore, of identifying these Philippine specimens has been done at the Ameri- can Museum of Natural History (AMNH) and the U. S. National Museum (USNM). In connection with these more-or-less routine identifications, a certain amount of revisionary work proved to be necessary for certain species, several with ranges extending far beyond the Philippines. It has been thought desirable to publish such findings separately rather than include them in the final report of the 1956 collection, where they would be more-or-less irrelevant. The present paper treats the three species of the large estrildine genus Lonchura whose ranges include the Philippines. The field work was carried out as part of a project of the Graduate School of Public Health, University of Pittsburgh, under the sponsor- ship of the Commission on Viral Infections, Armed Forces Epidemi- ological Board, and supported in part by the Office of the Surgeon General, U. S. Department of the Army. Specimens were collected and prepared with the invaluable assistance of Mr. Telesforo Oane and the generous counsel of Dr. Canuto G. Manuel, both of the ~1Carnegie Museum, Pittsburgh, Pa. 469348—58 279 280 — PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 National Museum of the Philippines. Much of the taxonomic work was carried out at the American Museum of Natural History, where I received many courtesies from Drs. Dean Amadon and Charles Vaurie, and at the U.S. National Museum, with the help of Mr. H. G. Deignan. Additional specimens were borrowed through the kind- ness of Dr. Raymond A. Paynter, Jr., of the Museum of Comparative Zoology, Dr. Austin L. Rand of the Chicago Natural History Museum, Dr. S. Dillon Ripley, Jr., of the Yale Peabody Museum, and Dr. Dwain W. Warner of the Minnesota Museum of Natural History. Lonchura leucogastra (Blyth) Three subspecies of the white-breasted mannikin are currently recognized: Lonchura leucogastra leucogastra (Blyth) of Siam, the Malay Peninsula, and Sumatra; L. 1. everetti (Tweeddale) of the Philip- pines and northern Borneo; and ZL. 1. castanonota Mayr of southern © Borneo. Delacour (1943, p. 82) has shown that Lonchura leucogas- troides (Horsfield and Moore) of Sumatra, Java, Bali, and Lombok, formerly considered to be a subspecies of L. leucogastra, is not closely related to the latter species. Tweeddale described the Philippine everetti (type locality Monte Alban, Luzon) in 1877, and later (1878, p. 622) commented that specimens from Palawan appeared to be intermediate between everetti and leucogastra. He assigned them to the former, working under a strictly binomial system. After having examined over 60 Philippine specimens (this species is by far the least common Philip- pine Lonchura), I have come to the conclusion that the name everetti should be confined to the birds of Luzon and a few adjacent islands. The remainder of the range hitherto ascribed to everetti is occupied by a form which, although intermediate in many respects between everetti and leucogastra, occupies such an extensive range that I feel it merits recognition as an additional subspecies, which may be called Lonchura leucogastra manueli, new subspecies Type: USNM 201278; adult male; “near Isabella’? [=Isabela], Basilan Island, Philippine Islands; Jan. 26, 1906; E. A. Mearns, collector (original No. 13903). Draenosis: Similar to L. l. everetti and L. l. leucogastra, but, in general, more deeply colored than the former, less deeply colored than the latter (details below); bill (except for Palawan birds; see measurements) averaging larger than that of either subspecies; wing averaging longer than that of lewcogastra. Although in some char- acters manueli displays a condition intermediate between everettt and leucogastra, in others it presents a combination of characters typical of one subspecies or the other with a remarkable constancy THREE SPECIES OF LONCHURA—PARKES 281 considering the size of its range. In detail, then, the features distinguishing the three races under consideration are as follows. The anterior underparts of everetti are chocolate brown, becoming deeper and richer on the chin and throat. In manueli this deep, rich color prevails, deepening almost to black on the chin and throat. Finally, in leucogastra the whole anterior underparts are virtually black. Dorsally there is a corresponding but less striking deepening of the general brown color of crown and back. In everettt and manueli the border between the dark breast and white abdomen includes many dark-tipped white feathers, giving this border area a distinctly spotted appearance. Such feathers are lacking in leucogastra, in which the border is more clear-cut. In everettt the upper tail coverts are plain brown, somewhat darker than the rest of the upperparts. In manuweli the upper tail coverts are decidedly blackish, contrasting quite sharply with the rest of. the back. Finally, in leucogastra the upper tail coverts are black and this dark area has advanced to include the rump as well. The under tail coverts and thighs show a corresponding progression from brown to blackish brown to black in the three subspecies. The white shaft-streaks of the dorsal surface are best developed in everetti, somewhat reduced in manuel, and decidedly reduced in leucogastra, particularly on the head. The tiny feathers along the metacarpal edge of the wing are pre- dominantly white in everetti, predominantly dark brown or black in manueli and leucogastra. This is a matter of shifting proportions of white and pigmented feathers rather than a progressive intensification of pigmentation as in other color characters cited. The yellow of the edges of the central rectrices, while variable, is a warmer, more golden color in leucogastra than in either manuelt or everett. Rance: Southern half of Philippine Archipelago (see list of speci- mens for specific islands) and highlands of northern Borneo. The Tutong River specimen mentioned by Mayr (1938, p. 45) has been examined and belongs with manueli as might be expected. Six other Borneo specimens examined also belong here; one from Laham, eastern Borneo, and five from Sarawak (Poeh Mountain, Kelabit Plateau, headwaters of Baram River). In the diagnosis above, no comparisons have been made with the rather isolated L. l. castano- nota Mayr of southern Borneo; this is a distinctive race with much richer coloring, being deep rufous chestnut above and jet black below. Remarks: The only islands of the Philippines from which I have examined specimens which were not immediately referable to either L. 1. everetti or L. l. manueli are Sibuyan and Culion; birds from these islands are almost exactly intermediate. It is possible that such 282 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 intermediate populations may also be found on others of the central islands. Although, as mentioned above, manueli is quite constant in its characters for a bird whose range encompasses so many islands, there is a certain amount of intraracial variation present. The most noticeable of these variations is a tendency for Palawan specimens to have smaller bills than those of the other islands within the range of manueli as here defined. The two specimens examined from the Sulu Archipelago are also small-billed. That this does not represent a stage in a cline toward the small-billed leucogastra is shown by the fact that the Borneo specimens have bills fully as large as those of typical manuel. This subspecies is named for Dr. Canuto G. Manuel, chief of the zoology division of the National Museum of the Philippines, whose assistance and hospitality added immeasurably to the success of our — expedition. A series of specimens of Lonchura leucogastra from Kuching, Sara- wak, sent to the U. S. National Museum for identification by B. E. Smythies of the Sarawak Museum, shows that a race is found in that vicinity which differs remarkably from other Borneo birds; no less than three races of this species are thus found in Borneo. The birds of the highlands of northern and eastern Borneo are L. 1. manueli; those of southern Borneo, L. 1. castanonota. The Kuching birds are actually closest in appearance to the geographically distant L. 1. ev- eretti of the northern Philippines. Mr. H. G. Deignan, who had sus- pected that the Kuching series differed from everetti as previously understood, has generously permitted me to include a description of this race in the present paper. I am happy to use the name which had been selected by Mr. Deignan, and designate this subspecies Lonchura leucogastra smythiesi, new subspecies Typr: USNM 461688; adult male; Tabuan, Kuching, Sarawak; Dec. 9, 1938; collected for Sarawak Museum. Diaqnosts: Nearest to the geographically distant L. 1. everetti of the northern Philippines, but white shaft-streaks of dorsal surface re- duced, practically absent on crown; upper tail coverts virtually con- colorous with back instead of a darker brown; anterior underparts deeper brown. The neighboring LZ. J. manueli of the highlands of northern Borneo has blacker anterior underparts and the upper tail coverts decidedly blackish, as mentioned above. The wing of JL. l. smythiesi averages shorter than that of typical manueli, but matches that of the slightly smaller Palawan population assigned to the latter race, THREE SPECIES OF LONCHURA—PARKES 283 RanGE: Known only from the vicinity of Kuching, Sarawak, Is- land of Borneo. SPECIMENS EXAMINED (adults only): L. l. everetti: Luzon, 5; Catanduanes, 2; Polillo, 5; Mindoro, 1. L. l. everetti x manueli intermediates: Sibuyan, 6; Culion, 2. L. l. manueli: Negros, 8; Cebu, 4; Bohol, 3; Siquijor, 1; Mindanao, 4; Basilan, 5; Palawan, 12; Tawi-tawi, 2; northern Borneo, 7. L. l. smythiesi: vicinity of Kuching, Sarawak, 11. L. 1. castanonota: southern Borneo, 5. L. l. leucogastra: Siam, 3; Malay Peninsula, 6; Sumatra, 2. Measurements: All measurements were taken to the nearest half- millimeter. The wing was pressed flat against the ruler. The “bill index”’ is the sum of the culmen length and the greatest width of the bill. This index is an attempt to indicate the general “largeness’’ of the bill rather than merely the customary length figure. Perhaps the most obvious difference in dimension between large-billed and small- billed birds is that of depth at the base, but no satisfactory comparable measurements could be made of this because of the great variation in degree of bill closure effected by preparators. The plus sign (++) indicates that the wing or tail was worn, and that the true measure- ment would be perhaps a millimeter or so longer. Measurements of excessively worn wings or tails were not taken. L. l. everetti Luzon: Wing, 51, 52, 52, 52, 53. Tail, 36, 37, 38, 39. Bill Index (B. I.), 19, 19.5, 20, 20, 20. CATANDUANES: Wing, 50, 51. Tail, 36, 38. B.I., 20, 20. Ponto: Wing, 50, 50,-51,.51--, 52. Tail, 35+-, 35-+-,-36, 37. B. 1., 19.5 19.5, 20, 20, 20.5. Minporo: Wing, 52.5. Tail, 39. B. I., 19.5. L. l. everettt x manueli intergrades Srpuyan: Wing, 51, 51.5, 52, 52, 52, 52.5. Tail, 36.5, 37, 37, 37.5, 38+, 39.5. Bet 91 9-0,,19.5, 2b, 216 2h: Cuxion: Wing, 51, 51.5. Tail, 35.5, 37.5+. B.IL., 20, 20. ZL, l. manueli NeEcrRos: Wing, 50.5, 51.5, 52, 52, 52.5, 52.5, 52.5+, 53. Tail, 36-+-, 38, 40, 41. B. I., 20.5, 20.5, 20.5, 20.5, 21, 21. Crsu: Wing, 52, 53.5, 53.5, 55. Tail, 39+, 40. B.I., 20.5, 21, 22.5, 22.5. Siquiyor: Wing, 52. Tail, 37. B.I., 20. Bouou: Wing, 52.5, 55,55. Tail, 37.5, 41. B.I., 20.5, 21.5, 22. Minpanao: Wing, 52.5, 53, 54. Tail, 37, 37.5+, 38.5+. B.I., 20.5, 21, 21.5, 22. Basiuan: Wing, 51, 51.5, 52.5, 53, 53.5. Tail, 35+, 36+, 36.5, 38+, 39. B. I., 19.5, 20.5, 21, 21.5, 22. PALAWAN: Wing, 49, 50, 50.5+, 51, 51, 51, 51, 51.5, 51.5, 52, 53, 53. Tail, 34+, 34+, 36, 37+, 37.5, 37.5+, 38. B. 1. 19, 19, 19, 19, 19.5, 19.5, 19.5, 19.5, 19.5, 20, 20, 20.5. Tawi-Tawi: Wing, 50.5, 52.5. Tail, 36+,37+. B.I., 19, 19.5. Borneo: Wing, 50.5+, 51, 54. Tail, 34+, 36.5. B.I., 20, 20, 21. 284. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 L. l. smythiesi Sarawak: Wing, 50, 50, 50.5, 50.5, 50.5, 51, 51, 52, 52, 52, 52. Tail, 35, 36, 37, 37, 37.5, 38. B.I., 19, 20, 20,:20.5, 20.5, 20.5,.21, 21. L. l. leucogastra Stam: Wing, 49, 49, 49.5. Tail, 33.5, 34, 37. B. I., 17.5, 17.5, 17.5. Matay PEeNniInsuLaA: Wing, 48.5, 50, 50, 50.5, 50.5, 50.5. Tail, 34.5, 35, 35+, 35.5, 35.5, 36.5. B.1., 17, 17.5, 18, 18, 18. Sumatra: Wing, 49.5, 52.5. Tail, 35.5, 37. B.I., 17.5, 17.5. Lonchura punctulata (Linnaeus) Linnaeus (1758, p. 173) originally described the nutmeg mannikin as Loxia punctulata, listing its range simply as ‘‘Asia.’””? His descrip- tion was based entirely on the description and plate of Edwards (1743, p. 40). In the 12th edition of the “Systema Naturae” (1766), Linnaeus added a reference to Brisson, who specifically mentioned Java. Kloss (1931, p. 364) considered that Linnaeus thus acted as his own “‘first reviser,” and that the type locality of Loxia punctulata should be restricted to Java. Hellmayr (1914, p. 60) had earlier come to the same conclusion. This is at variance with the treatment of Baker (1926, p. 91; 1930, p. 223) and other authors who have used the Linnaean name for the Indian rather than the Javanese race. The identification of Linnaeus’ name punctulata with a particular population of the nutmeg mannikin must rest entirely on Edwards’ description and plate, since this was the sole reference in the 10th edition of the ‘Systema Naturae.’”’ If this description and plate are identifiable as one of the subspecies of Lonchura punctulata, any sub- sequent restriction by Linnaeus or anybody else is superfluous. Examination of Edwards’ plate and accompanying description make it evident that it cannot possibly apply to the Javanese form, but must be that of India, as advocated by Baker. ‘The evidence is as follows: 1. «The bill is of the shape and bigness of our Green-finch’s bill . . .”.. The plate clearly shows a large-billed form of L. punctulata; the Indian bird is large- billed, the Javanese small-billed. Culmen length of Chloris chloris (England), 13 mm.; Indian Lonchura punctulata, 11.5 mm.; Javanese L. punctulata, 10 mm. 2. Although too much reliance should not be placed in Edwards’ statement that the bird is figured in its natural size, still the wing-length of the figure matches Indian specimens exactly, and is far larger than Javanese specimens. 3. “Head, neck, beginning of the breast, back, wings and tail, are of a dark reddish brown . . .” ‘True of all except the tail of the Indian race; the Javanese bird has the dorsal surface dull grayish brown, not dark reddish brown. 4. As for the tail, which is described and figured as ‘dark reddish brown,” this does not match any form of the species, and is certainly not the Javanese bird in which the tail is distinctly ashy gray. The shape of the tail in the plate is also inaccurate, unless the specimen portrayed lacked the long pointed central rectrices, which are yellow in the Indian bird. A specimen of the Indian race in which the bright yellow, pointed central rectrices were missing, and in which THREE SPECIES OF LONCHURA—PARKES 285 the remaining rectrices were worn (as they appear to be in Edwards’ figure) would appear to have a brownish tail. 5. “The forepart of the neck has something of a purplish cast’; true of the Indian race, not of the Javanese. 6. ‘The rump is of a lighter greenish brown.’”’ The rump of the Indian bird is decidedly yellowish or greenish, with faintly indicated darker markings as shown on Edwards’ plate. The rump of the Javanese bird is heavily barred with dark gray and white, with no trace of yellowish or greenish. 7. The edges of the white abdominal feathers are described and figured as black; in the Javanese form these edges are dark brown, not black. 8. ‘The middle of the belly, thighs, lower belly and covert-feathers under the tail, are light brown, or dirty white.” Exactly true of the Indian race; the thighs and under tail coverts of the Javanese race are white, heavily barred with dark gray. The name of the Javanese race thus reverts to Lonchura punctulata nisoria (Temminck), while the Indian form is L. p. punctulata (Lin- naeus). Stuart Baker gives the restricted type locality as Calcutta. Hodgson’s L. p. lineoventer from Nepal, used for the Indian race by Kloss (1931), Stanford and Ticehurst (1938, p. 611), and others, is a synonym of punctulata. L. p. topela (Swinhoe), with type locality Amoy, has customarily been considered the form which occurs in the mountains of Yunnan and northeastern Burma, although its status in the latter area has been questioned (Smythies, 1953, p. 231). La Touche (1927, p. 295), under topela, states: ‘‘In eastern Yunnan it is represented by another race, probably more closely akin to the Indian forms.” All of the literature on Yunnan birds I have been able to find, however, lists the resident race as topela. The form found in most of Burma except the northeastern section is L. p. subundulata (Godwin-Austin). In any case, the Yunnan-northeastern Burma bird does differ from topela, as might be anticipated on geographic grounds. The latter race is found in Taiwan and Hainan, and in eastern China and north- eastern Indochina. It is the second-palest of the races of L. punctu- lata, being exceeded in this respect only by the Philippine race L. p. cabanist. The Yunnan bird may be described as follows: Lonchura punctulata yunnanensis, new subspecies Type: AMNH 720698; adult female; “hills around Tengyueh” [=Tengchung], western Yunnan, alt. 6,000 feet; June 1924; George Forrest, collector (original No. 5027). Diacnosis: Similar to L. p. topela (Swinhoe), but coloration markedly richer; pencilling of ventral feathers darker and coarser; dark throat-patch warmer, more reddish brown; dorsal coloration in general browner, less gray, with, in many specimens, a greater con- trast between the whitish shaft streaks and the ground color of the 286 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 feathers. The yellow of the upper tail coverts and central rectrices averages deeper, less greenish. In juvenal plumage, L. p. yunnanensis is everywhere brighter, less grayish in color than topela. There is no significant difference in size between yunnanensis and topela. Measurements of type: wing (flat), 54 mm.; tail, 39 mm.; culmen, 11 mm.; tarsus, 13 mm. Differs from ZL. p. subundulata (Godwin-Austin) of Assam and southern Burma in having more conspicuous light shaft streaks on the dorsum, a browner (less blackish) tone ventrally, and ventral markings of quite a different shape; the breast and flanks of sub- undulata have a spotted appearance, while those of yunnanensis are pencilled like those of topela. The posterior border of the brown throat of subundulata is more sharply defined than that of yunnanensis. Actually the population which is currently known as subundulata is a variable series of intermediates between the well-defined punctulata to the west and the yunnanensis-topela group to the east. Rance: Mountains of Yunnan and northeastern Burma (for localities, see list of specimens examined). Remarks: Four specimens from Mengtz, in southeastern Yunnan, appear to be somewhat intermediate between L. p. topela and L. p. yunnanensis, although closer to the latter. Three of the four, how- ever, are in such worn plumage as to make color comparisons almost valueless. Smythies (1953, p. 231) writes: ‘The status of topela [in Burma] is uncertain; a single specimen obtained by the Vernay-Hopwood expedition at Singhaling Hkamti may have been a winter visitor, whereas birds seen at 5,000 feet or more in the Sadon hills and in the Ngawchang valley and not collected may have been resident topela; Stuart Baker claims that it occurs in the Shan States.” The Vernay-Hopwood specimen, taken on Mar. 4, 1935, is in the middle of the post-juvenal molt. A specimen (AMNH 720672) taken at Sinlumkaba, east of Bhamo, on Apr. 19, 1908, has almost completed the post-juvenal molt. Both are referable to yunnanensis. That this subspecies does, indeed, breed in northeastern Burma is suggested by a specimen (USNM 377788) taken 6 miles north of Myitkina on May 24, 1945, and one (AMNH 347218) taken at Myitkina on June 25, 1945. The specimen from Washaung (near Myitkina) mentioned by Stanford and Mayr (1941, 355) is, of course, yunnanensis rather than subundulata. It was taken on Apr. 12, 1939, and is in about the same stage of the post-juvenal molt as the Singhaling Hkamti specimen. Baker (1930, 223) lists Munia inglisi Hume in the synonymy of L. p. topela; however, inglisi was named from Cachar, and birds from this region are referable to L. p. subundulata; THREE SPECIES OF LONCHURA—PARKES 287 L. p. catervaria Koelz (1954, p. 19), described from the Khasia Hills, represents a population intermediate between punctulata and sub- undulata, and is best placed in the synonymy of the latter. It will be remembered that subundulata itself represents a rather variable inter- mediate population. SPECIMENS EXAMINED: L. p. yunnanensis: YUNNAN: Vicinity of Tengyueh, 23; Nantien, 1; Shweli- Salaween Divide, 6. Burma: Myitkina, 1; 6 miles north of Myitkina, 1; Singhaling Hkamti, 1; Washaung, 1; Sinlumkaba, 1. L. p. yunnanensis x topela intermediates: YUNNAN: Mengtz, 4. L. p. topela: Taiwan, 40; Hainan, 23; Fukien, 6; Kwangtung, 1; Tonkin, 5; An- nam, 2. L, p. topela x subundulata intermediates: BurMa: Pegu, 2. L. p. subundulata: Assam, 5; Khasia Hills (‘‘catervaria’’), 10. Also series of other races in USNM and AMNH. Lonchura malacca (Linnaeus) Delacour (1943, p. 83) united three groups of forms which had often been considered separate species: L. malacca, L. ferruginosa, and L. atricapilla. He used the name ferruginosa for the combined species, overlooking the fact that Loria malacca Linnaeus, 1766, antedates Lozia ferruginosa Sparrmann, 1789. The desirability of merging the three groups has been questioned by some authors, but I am not prepared to go into this question here. The present paper is con- cerned only with that portion of the combined species formerly known as Lonchura atricapilla, all forms of which lack white on the head and underparts. It is tempting to consider L. grandis of New Guinea as part of this species, since the adults closely resemble the atricapilla group. Dela- cour (1943: p. 83) considered grandis and ferruginosa (=malacca) to comprise a superspecies, giving the large bill and extensively black underparts of grandis as specific characters. EZXven more important, in my opinion, is the pattern of the juvenal plumage, in which grandis differs markedly from all races of malacca in being streaked on the throat. Salomonsen (1953, p. 265) pointed out that a revision of this species was badly needed, and he reviewed it briefly himself. The large area ascribed by Salomonsen and other authors to the race atricapilla involves a number of well-marked clines, which can be divided into several subspecies and intergrades. Salomonsen’s treatment of the Philippine populations is critically discussed (p. 290) after a review of the mainland forms. The distribution of L. m. sinensis is substantially as given by Salomonsen (1953, p. 266), namely Malay States, southern Siam, and lowlands of Sumatra. The diagnostic characters can, how- VOL. 108 PROCEEDINGS OF THE NATIONAL MUSEUM 288 ‘ajeulxoidde olv sanyeso] [JY *YOeq ay} Jo sal oy} YIM snosojoouoD ATenji1A st duni sy} YY ul ssoyi pur Jojoo yoeq ay} Wo pouysp Ajdieys dun uoolew ev yitM suonrndod oy} saptaip e[nsutueg Av[eyy Joddn oy} ssosov oul] oy], ‘juasoid uaym ‘yoied eulWopqe yeIq JY} JO ssaujUTISIP puv oZIS DAT] LJAI SY} UMOYS SI ZUlIN}}9] Ja[[vUIs UT *S99113991 Jo saspa Pue s1JOA0d Jv} Joddn jo 10[09 dy} 0} JoJoI ,-MOT[AA,, PUL ,.P2Y,, °VIIVIVU DANYIUOT Ul UOLIIeA d1yde13003 Ul spudl.— gsnasia a7LLiy ] zuno1y THREE SPECIES OF LONCHURA—PARKES 289 ever, be expanded. The black abdominal patch is absent. The back is paler (particularly noticeable in fresh plumage) than that of adjacent populations to the north and west. The rump is darker and more reddish than the back, but this color is much paler and more diffuse than in any of the races to the north. The upper tail coverts and central rectrices are yellow. The race L. m. batakana (Chasen and Kloss) from the mountains of northern Sumatra is similar in dorsal color (including rump and tail), but smaller, and with a well- defined black abdominal patch. Toward the northwest, sinensis intergrades with atricapilla, as suggested by Chasen and Kloss (1929, p. 23). A specimen from Pegu is precisely like sinensis above and below, except that the rump is a deeper maroon and quite sharply defined. Another Pegu speci- men is similar dorsally, but blackish on the lower abdomen and under tail coverts. Specimens from northern Burma and northwestern Yunnan resemble atricapilla in having the tail and coverts yellow, and a well-defined maroon rump patch; the Yunnan specimens are paler than those of Burma. Ventrally these birds are rather variable, but all have a more-or-less diffused blackish area in the lower abdo- men. In Assam and Bengal occurs true atricapilla, which has a well- defined black abdominal patch, deep maroon rump, and yellow tail and coverts. Salomonsen (1953, p. 266), Smythies (1953, p. 229), and others have synonymized Munia rubro-nigra Hodgson (Nepal) with atricapilla of Bengal. This is understandable in view of the distributions and distinguishing characters for the two claimed by Stuart Baker (1926, pp. 80-81). Both are incorrect. As pointed out by Vaurie (1949, p. 38) and others, rubro-nigra does not differ from atricapilla in wing length or color of underparts. Judging from a limited amount of material, however, the name rubro-nigra is available for a well-marked race of Nepal and adjacent India. It occurs at least as far east as Jainagar, on the Nepal-Bihar border (specimen, Yale Peabody Mus. 26520, slightly intermediate toward atricapilla), and as far west as Dehra Dun. It resembles aéricapilla in having a well-defined black abdominal patch and sharply defined maroon rump, but the long, dissected upper tail coverts are a rich maroon red, little different from the rump, whereas in atricapilla these feathers are golden yellow to orange. The rectrices themselves are washed with yellow in atricapilla, deep reddish brown in rubro-nigra. The latter race is also slightly paler dorsally. The four rubro-nigra seen have quite large and heavy bills, but accurately sexed material is insufficient to deter- mine whether there is any actual difference between the two races in this respect. 290 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The population of Chieng Rai, northernmost Thailand, mentioned by Deignan (1945, p. 559), is somewhat variable, but cannot be satisfactorily assigned to either a known race or an intergrade between races. Rather than to try to stretch the limits of well-defined sub- species to include these birds, it would seem advisable to recognize them as a subspecific entity, for which an appropriate name is Lonchura malacca deignani, new subspecies Tyrz: USNM 350467; adult male; Chieng Rai, northern Thailand (1,214 feet); May 8, 1936; H. G. Deignan, collector (original No. 1297). DiacGnosis: Differs from all other subspecies of the mainland, glossy black-headed group except rubro-nigra in that the long upper tail coverts and central rectrices in most (not all) specimens are deep orange-red rather than some shade of yellow. In a minority of individuals these feathers are rich yellow-orange, not unlike (but somewhat more intensely colored than) those of atricapilla. All individuals of deignani, however, may be distinguished from rubro- nigra or atricapilla by their more diffuse and generally less blackish abdominal patch, and by having dark brown rather than black under tail coverts. The general color of both dorsal and ventral surfaces is also a paler reddish brown in rubro-nigra than in deignani. Raneer: Chieng Rai Province, northern Thailand, and probably Indo-China. A single specimen from Hue, Annam, is referable to deignani. A series of specimens from Hainan in the American Museum of Natural History consists of cage birds, as shown by the condition of the claws; the species is not known to occur in Hainan in a wild state. However, these specimens may well have originated on the Indo-China mainland across from Hainan, since they agree in every respect with deignant. SPECIMENS EXAMINED (mainland and Sumatra races): L. m. rubro-nigra: Nepal, 1; Nepal-Bihar border, 1; United Provinces, 2. L. m. atricapilla: Assam, 5. L. m. atricapilla x sinensis intermediates: northern Yunnan, 5; northern Burma, 2; central Burma, 2; southern Burma, 2. L. m. sinensis: Malay Peninsula, 2; Sumatra, 1. L. m. batakana: Sumatra, 2. L. m. deignani: northern Thailand, 6; Annam, 1; ‘‘Hainan”’ (cage birds), 8. Leaving the glossy black-headed birds of the mainland, we turn now to the eastern populations inhabiting Taiwan, the Philippines, Celebes, and Borneo. There is a general cline in the direction of increased saturation of the pigment of head and breast running from north to south, from grayish brown in Taiwan to rich black in Borneo and northern Celebes; then, surprisingly, back to brown again in THREE SPECIES OF LONCHURA—PARKES 291 southern Celebes (LZ. m. brunneiceps). The pale northern extreme is L. m. formosana (Swinhoe). Salomonsen (1952, p. 354) has shown that specimens from northernmost Luzon are inseparable from Taiwan examples of formosana. The rest of the birds from central Luzon to Borneo have until lately been considered to be L. m. jagort (Martens), type locality restricted by Salomonsen (1953, p. 267) to Manila. As represented in the Philippines, jagorz is actually a tremendously variable series of intermediates between formosana and the almost consistently black-headed Borneo population. Salomonsen endeav- ored to express this by describing the black-headed birds as L. m. gregalis, with type locality Opol, Mindanao. This merely complicated matters, however, because birds from the range ascribed by Salomon- sen to his ‘‘gregalis’”’ are not consistent in having dull black rather than dark brown heads, as he claims. Three Sulu birds in the Ameri- can Museum of Natural History, for instance, have distinctly brownish crowns, and even some of the Borneo specimens (cf. AMNH 720576, Labuan) tend to have rather brownish heads. In a series of 18 specimens from Basilan, which should be “gregalis’? (University of Michigan Museum of Zoology), there is much variation; although mostly black-headed, several are distinctly brown on the crown and nape. Some of my own Luzon series (which is quite variable inter se) are as dark as any Borneo specimens seen. Peters (1940, p. 209) could detect no significant difference between Borneo and Philippine specimens of this species, calling both jagort. De Schauensee (1957, p. 11), on the other hand, upheld ‘“‘gregalis,” calling it “‘a very distinct race,” on the basis of comparing one adult male and two immature females from Mindanao with three Luzon specimens! Ideally we should probably use two names, one for the Taiwan- northern Luzon bird and one for the birds of Borneo and northern Celebes, with the rest of the Philippine birds considered as a variable intergrading cline. Unfortunately the name jagori was based on the highly variable population of central Luzon. In view of the fact that variability in this species is so high in this part of the world, so that few specimens could be named if their localities were unknown, it appears best to continue to use jagori in its traditional sense, considering gregalis Salomonsen as a synonym. ‘The subspecies gagort thus includes both brown-headed and black-headed birds, with a preponderance of the latter toward the south, but no reasonable geographic line can possibly be drawn between them. There seems to be no consistent geographic variation in Lonchura malacca within the Island of Borneo; this, of course, is in striking contrast to the situation in L. leucogastra. 292 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Literature Cited Baker, E. C. Stuart 1926. The fauna of British India, including Ceylon and Burma. Birds. Vol. 3, xx+489 pp. 1930. The fauna of British India, including Ceylon and Burma. Birds. Vol. 7, vii+484 pp. Cuasen, F. N., anp Kuoss, C. BopEN 1929. Two new Malaysian birds. Bull. Raffles Mus., vol. 2, pp. 22-238. Deienan, H. G. 1945. The birds of northern Thailand. U. 8. Nat. Museum Bull. 186, v+616 pp. DeELacour, JEAN 1948. 3 oe 7 ™ RHIZOCEPHALA—REINHARD 299 its course to open alongside the testes at the junction of the mesentery with the mantle. The single specimen from the Barbados (fig. 1,d) was not sectioned. It appears to be an animal that is undergoing degeneration after the release of its brood. The mantle cavity is empty and the anterior end of the sac has a conspicuous round orifice, presumably the birth pore. The host also bears the scar of a companion parasite no longer present. Remarks: Dr. Fenner A. Chace, Jr., curator of marine inverte- brates, U. S. National Museum, for whom this species is named, recorded the presence of abdominal parasites on three examples of Munida irrasa A. Milne Edwards obtained by the Atlantis off Cayo Coco, Cuba (Chace, 1942, p. 46). Two of these were loaned to the author through the courtesy of the Museum of Comparative Zoology. They provided part of the material described here. Cyphosaccus cornutus, new species Figure 2; Puate 1, Ficure 2 Syntypes: Off Playa Baracoa, Havana, Cuba, Atlantis Station 3305, lat. 23°05’30’’ N., long. 82°35’ W., 330 fathoms, Mar. 23, 1939, three specimens on one Munidopsis erinacea (A. Milne Edwards), Harvard-Havana Expedition, Atlantis. MCZ 11721. Diagnosis: Body broadly U-shaped, comparatively stout, uniform in thickness, with the stalk in the midregion. Testes fused. Description: The parasites are attached to the sternites of the third and fourth abdominal segments. Two are oriented with the anterior arm towards the left side of the host; the other is turned in the opposite direction. All three are of approximately the same size, 8 mm. in length and about 1 mm. in thickness. The shape of the sac resembles an ox-bow, although the anterior arm is misshapen. The tip of this arm bears a nozzle-like prominence enclosing a blind canal. The smooth external surface of the sac is covered with a thin cuticle through which an underlying meshwork of delicate longitudinal and circular muscle fibers belonging to the mantle can be seen. The longitudinal fibers are more widely spaced than the circular ones. When the animal] is examined with the anterior arm to the observer’s left, as in the drawings, the mesenterial surface lies uppermost. The mesentery, which extends along the entire length of the sac, supports a well-developed visceral mass. In the animals that were sectioned, segmenting eggs are also present in the mantle cavity. Whether they hatch as nauplii or as cypris larvae is not known. 300 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The testes occur near the posterior end of the mesentery. These elongated sacs are fused for approximately one-third of their length in one specimen, and completely fused in the other. Each gives rise to a vas deferens shaped like an inverted J, which runs in a posterior Ficure 2.—Cyphosaccus cornutus, new species: a, Syntype, external appearance (MCZ 11721); 6, diagram of internal anatomy; c, section through region of stalk. The scale refers toa and b. Abbreviations as in figure 1. direction, then curves in the opposite direction shortly before it terminates. The vasa deferentia are about as long as the testes. The colleteric glands are located in the region of the stalk. They are of simple design, with no more than two or three narrow channels leading to the opening into the mantle cavity. Remarks: Chace (1942, p. 90) mentioned the presence of “ab- dominal parasites” on a female of Munidopsis erinacea (A. Milne Edwards) from Playa Baracoa, Cuba. This is the parasitized specimen that provided the material for the present study. RHIZOCEPHALA—REINHARD 301 Boschmaia, new genus Genotype: Boschmaia munidicola, new species. Diagnosis: Gregarious; body elongate, bifurcate, with arms of unequal length. Mantle aperture covered. Stalk on right side; testes adjacent to stalk. On Galatheidae. The orientation of the mesentery with respect to the stalk is a distinguishing feature of this genus. In the previously known mem- bers of the family Peltogastridae the stalk arises from the dorsal or mesenterial surface, but in Cyphosaccus and Boschmaia it is placed in a lateral position. Whereas in Cyphosaccus the mesentery lies uppermost when the animal is viewed in an upright position with the anterior arm to the observer’s left, in Boschmaia the opposite is true. Here the mesentery lies on the far side of the animal when the sac is viewed in the same manner as above. Since the mesentery in Rhizocephala marks the dorsal surface of the animal, the stalk in this genus arises on the right side. The generic name Boschmaia is given in honor of Prof. H. Boschma, director of the Rijksmuseum van Natuurlijke Historie, Leiden, Holland, the author of more than 80 papers dealing with the Rhizo- cephala and the foremost authority on this group of animals. Boschmaia munidicola, new species FIGURE 3 Driacnosts: Anterior arm much longer than posterior one. Testes minute, vasa deferentia long and straight; colleteric glands near anterior end. Syntypss: Off Cape Lookout, N. C., Fish Hawk Station 7302, U. S. Fish Commission, 7% fathoms, July 24, 1902, 9 specimens on one Munida irrasa A. Milne Edwards. USNM 100945. ADDITIONAL SPECIMENS: Off Key West, Fla., Fish Hawk Station 7279, U. S. Fish Commission, lat. 24°21’55’’ N., long. 81°58’25’’ W., 98 fathoms, Feb. 14, 1902, 14 specimens on one Munida irrasa A. Milne Edwards. USNM 100946. Description: The parasites are attached by means of a short, thin stalk to the third and fourth abdominal segments of the host. They have a somewhat hook-shaped appearance. Each sac is strongly bent, with one arm, which is the anterior part of the animal, nearly twice as long as the other. The basal portion of the sac tapers in the direction of the stalk. At the summit of the anterior arm there is a knob-like prominence formed by the mantle opening. The aperture, however, does not communicate with the exterior since it is covered by the external cuticle. 302 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The syntype specimens are mature animals measuring about 6 mm. in length and 1 to 1.5 mm. in thickness. They contain developing nauplii. Those from Key West are younger specimens, more slender Ficure 3.—Boschmaia munidicola, new species: a, Syntype, external appearance (USNM 100945); b, diagram of internal anatomy; c, d, younger specimens (USNM 100946); ¢, visceral mass and colleteric gland in transverse section. The scale appearing alongside figure a refers to a,c and d. Abbreviations as in figure 1. in form, without eggs or embryos in the mantle cavity but with a visceral mass that fills up most of the space. They measure about 5 mm. in length and 0.5 to 0.75 mm. in thickness. There is relatively little difference in size between the various sacs found on the same host individual. RHIZOCEPHALA—REINHARD 303 The mantle and visceral mass, in general structure, are like those of Peltogaster paguri and exhibit no unusual features. The mesentery is broad, about equal in width to the visceral mass, and extends from one end of the sac to the other. At the base where the two arms meet there is an extension of the mesentery towards the right to connect with the stalk. The visceral mass accompanies the mesen- tery to its termination. The testes are exceedingly small, rounded bodies set close to the stalk. They measure only 35 to 40 microns in diameter. The straight and comparatively long vasa deferentia pursue an upward course away from the stalk along the surface of the basal extension of the visceral mass. Their length is about 250 microns. The colleteric glands are located near the anterior end of the sac, with the left gland slightly in advance of the right. In a cross section of the animal the gland appears as an elliptical mass com- prising a single canal surrounded by several layers of cells. Its depth is 60 to 80 microns and its extent in a dorsoventral direction is about 150 microns. Genus Tortugaster Reinhard Tortugaster fistulatus Reinhard FiacurE 4,e Tortugaster fistulatus Reinhard, 1948, pp. 33-37, fig. 1 (external appearance, internal structures), fig. 2 (transverse section), fig. 3 (vas deferens). MATERIAL EXAMINED: South of Cay Sal Bank, north coast of Cuba, Ailantis Station 2987, lat. 23°22’ N., long 79°53’ W., 280-300 fathoms, Mar. 13, 1938; one specimen on Munidopsis spinifer (A. Milne Edwards). Harvard-Havana Expedition. MCZ 11755. East of St. Augustine, Fla., Atlantis Station 3780, lat. 30°27’ N., long. 79°52’ W., 250-265 fathoms, Feb. 24, 1940; one specimen on Munidopsis bahamensis Benedict. MCZ 11734. These are new locality and new host records for a species that has previously been known only from the type specimens found on Munidopsis robusta A. Milne Edwards, off Tortugas, Fla. In their attachment and orientation with respect to the host and in their external appearance these examples of 7. jistulatus agree with the type. The specimen on Munidopsis spinifer measures 6 mm. in length and 3 mm. in width and is a mature animal with eggs present in the mantle cavity. The parasite on Munidopsis bahamensis is also fully developed and differs from the other only in size. Its length is 8mm. and width 4.5mm. Because of their poor state of preserva- tion these animals were not sectioned. They are the specimens (“abdominal parasites’) mentioned by Chace (1942, pp. 89, 91) in 304 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 his report on the Galatheidae obtained by the Atlantis expeditions to the West Indies. Genus Galatheascus Boschma Galatheascus minutus Boschma Ficure 4,a—d Galatheascus minutus Boschma 1933, pp. 476-478, fig. 1 (external appearance), figs. 2, 3 (transverse sections); 1947, pp. 2—4, fig. 1 (attached to host). Type specimen on Galathea nera Embleton, from Oban, Scotland. MATERIAL EXAMINED: Off Cayo Coco, north coast of Cuba, 180 fathoms, 1939; two specimens on one Munida stimpsont A. Milne Edwards. Atlantis Coll. MCZ 11344. Gulf of Mexico, Blake Station 45, lat. 25°33’ N., long 84°21’ W., 101 fathoms, 1877-78; one specimen on Munidopsis barbarae (Boone). Blake Coll. MCZ11719. (Chace, 1942, p. 81, mentions this specimen. The host is an intersex.) Off Cape Lookout, N. C., Fish Hawk Station 7302, U. S. Fish Commission, 74% fathoms, July 24, 1902; one specimen on Munida irrasa A. Milne Edwards. USNM 100944. Off Fort Pierce, Fla., Combat Station 235, lat. 27°27’ N., long. 78°58’ W., 180 fathoms, Feb. 2, 1957; one specimen on Munida schroedert Chace. USNM 100947. All five specimens are oriented with their long axis perpendicular to the main axis of the host and with the anterior end directed towards the right. The site of attachment is on the second or third abdom- inal segment. The largest parasite, found on Munda schroederi, measures 7 mm. in length, 4 mm. in width, and 3 mm. in thickness; the smallest, found on Munidopsis barbarae, measures 3 by 2 by 1.5 mm. All have developing embryos in the mantle cavity and are therefore mature animals. The mantle opening in three of the specimens is small, indistinct, and narrow, but in the parasites of Munidopsis barbarae and Munida schroederi it is wider and surrounded by an elevated rim. The short, thin stalk, which is oval in cross section, arises in the median dorsal part of the body. The smooth external cuticle is only about 4 microns thick except where it forms an inconspicuous shield around the base of the stalk. In order to study the internal cuticle, one of the specimens occur- ring on M. stimpsoni was macerated in KOH. Retinacula were found on the cuticle covering the visceral mass but not on the lining of the mantle. They consist of one to three spindles having a length of 7 to 10 microns and a width of 2 to 3 microns. The retinacula of Galatheascus striatus, which they resemble in all respects except size, are 20 to 25 microns long. RHIZOCEPHALA—REINHARD 305 WS i (fl 4 ate < QWiZe> eS AA wi i WZ d Ficure 4.—a-d, Galatheascus minutus Boschma: a, external appearance of specimen on “ Munidopsis barbarae; b, transverse section of the same specimen; ¢, testes and vasa defer- entia of specimen on Munida stimpsont; d, chitinous linings of vasa deferentia revealed by maceration of another specimen on Munida stimpsoni; e, Tortugaster fistulatus Reinhard, external appearance of specimen on Munidopsis spinifer. Abbreviations: sh, sheath; od, vas deferens; others as in figure 1. Serial sections of the other parasite occurring on M. stimpsoni and of the Blake specimen were prepared. No differences were observ- able in the details of their internal anatomy. The mesentery begins just behind the mantle opening and gradu- ally increases in width as it passes backwards. It reaches its maxi- 306 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 mum breadth in the region of the stalk, then tapers off and ends about half way between the terminations of the male genital organs and the posterior end of the sac. The testes are located in the region of the stalk. They consist of a fairly long, straight portion followed by a slightly coiled portion. The vasa deferentia are long and highly tortuous tubes, nearly as wide as the testes at their beginnings, but gradually diminishing in thickness. Each of the male genital systems is enclosed in a loose sheath or bag. In the sections one sees a noticeable space between the testis or vas and the envelope that surrounds it. The colleteric glands, located anterior to the stalk, occur in the same sections of the animal as those which include the testes. They lie in the dorsal half of the visceral mass, rather close to the mesen- tery, and, although simple in construction, are not indistinct. The base of the gland forms a cavity which contains a coagulum. A short duct reaches from this cavity to the surface of the visceral mass, and its wall, as well as that of the cavity, consists of a layer of tall columnar epithelial cells. The gland reaches to a depth of about 140 microns below the surface of the visceral mass and extends in a dorsoventral direction for a distance of about 180 microns. The specimens identified here as Galatheascus minutus agree with the description of the type material in size, position of the stalk, nature of the external cuticle, structure and position of the colleteric glands, and in the general morphology and location of the male genital organs. The fact that the mesentery of G. minutus is described as extending from the anterior to the posterior end, while in our specimens it is only about three-fourths as long as the sac, is not considered to be a significant difference. Boschma (1933, p. 478) stated that the testes of minutus have their closed ends directed ‘‘towards the posterior extremity” of the body, but the word “posterior” in this connection was a mistake (Boschma, personal communication). In our material and in the type specimens of minutus the blind ends of the testes point in an anterior direction. This is likewise the condition in G. striatus (Boschma, 1929). Although the sac that surrounds each of the male reproductive systems is not mentioned in the description of the type, Boschma’s figures of transverse sections of striatus and minutus both show this feature. The list of Peltogastridae occurring on the Galatheidae is now as follows: Galatheascus striatus, G. minutus, Tortugaster fistulatus, Cyphosaccus chacei, C. cornutus, and Boschmaia munidicola. A com- parison of the morphology of these animals fails to reveal any distinc- tive feature that sets this group apart from the other Peltogastridae, although in all except Galatheascus there is a pronounced tendency toward asymmetry. RHIZOCEPHALA—-REINHARD 307 Literature Cited Boscuma, HILBRAND 1929. Galatheascus striatus—a new rhizocephalan. Journ. Mar. Biol. Assoc. United Kingdom, vol. 16, pp. 73-79, 6 figs. 1933. The Rhizocephala in the collection of the British Museum. Journ. Linn. Soe. London, vol. 38, pp. 473-552, 58 figs., 1 pl. 1947. The European Rhizocephala in the collection of the Brussels Museum of Natural History. Bull. Mus. Roy. Hist. Nat. Belgique, vol. 23, pt 23, pp. 1-7, 2 figs. CuHacE, FENNER A., JR. 1942. Reports on the scientific results of the Atlantis expeditions to the West Indies, ete. The anomuran Crustacea. I. Galatheida. Torreia (Havana), No. 11, pp. 1-106, 38 figs. REINHARD, Epwarp G. 1948. Tortugaster fistulatus, n. gen., n. sp., a rhizocephalan parasite of Munidopsis robusta. Proc. Helminth. Soc. Washington, vol. 15, pp. 33-37, 3 figs. U. S. 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Alaa De 0 a ll alee 1) emdialls ee / a ‘Lparssgy 6 ead state ua a bite) AAR sects Hy, pie eu (i ° ©, Gigi Arne . a Rigi? at yh | +h Gar oral . _ & meres BAe Pi We oT vs): ; MS ve as me : b5ei isthe Batt ning enanyed aM PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1958 No, 3404 ADVANCES IN OUR KNOWLEDGE OF THE HONEY-GUIDES By HersBert FRIEDMANN In the following pages I have brought together new information on various aspects of the biology of the honey-guides that has come to attention since the publication of my book in 1955. Together with these additional data, I have made such comments as seem necessary, either for their proper evaluation or for their allocation with respect to earlier knowledge. Only two phases of honey-guide studies are not included: Purely systematic data, and my continuing investigations on wax digestion, the results of which will be presented separately. In addition to such notes as have appeared in print and which are here collated, I am indebted to the following for unpublished obser- vations: J. P. Chapin, W. R. Ingram, C. H. Jerome, D. W. Lamm, H. M. Miles, B. Neuby-Varty, R. H. Stevenson, V. G. L. van Someren, and J. M. Winterbottom. Additional Data on Eggs and Egg-Laying Because of the difficulties attached to study of ovulation in para- sitic birds, one must always be alert for incidental data bearing on this topic. The new information is as follows: 1. Greater honey-guide, Indicator indicator One observation bearing on the time of day of ovulation comes to me from J. M. Winterbottom. On Sept. 16, 1943, in the northwestern 469350—58 309 310 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 corner of the Livingstone District, Northern Rhodesia, he caught a female greater honey-guide on the nest of a hoopoe in a hole in an ant hill. The bird had already laid its egg. He watched it go to the nest, and walked up at once, so it was only a matter of seconds, half a minute at the most, that the bird was on the nest. In answer to my query, Winterbottom informs me that this took place during the hot part of the day, between 11 a. m. and 2 p. m. A female collected April 28 at Enugu, eastern Nigeria, by Serle (1957, p. 415) had two large yolked ova in the ovary and two ruptured ege follicles, indicating that at least four eggs would have been laid. It may be recalled that earlier data of the same type (Friedmann, 1955, p. 136) suggested, in one case at least, that eight eggs would have been laid by one bird. We still do not know what the usual number may be. Recently, H. A. Roberts (1956, p. 114) has stated that, when about to lay in a barbet’s nest, the hen honey-guide goes there accompanied by the male, and that the latter acts as a lure to draw away the poten- tial hosts from their nest, thereby giving the hen the chance to enter it and deposit an egg. Roberts writes that the female barbet rushes out of the nest hole as the honey-guides approach, and back into it again as they depart a short distance, this performance ‘‘being re- peated until she becomes rather exhausted. At this stage the female honey-guide conceals herself nearby, and as soon as both barbets pursue the increasingly bold male honey-guide, the female honey- guide makes a dash for the hole. Usually a short lull among the con- testants now follows which enables the female honey-guide to deposit her egg. Should the barbets try and return too soon, the male honey- guide at once takes action to lure them away... .” This account is comparable to one by Millar which I have previously described (Friedmann 1955, pp. 136-137), and as I wrote then, it is “‘difficult to believe that the male accompanies the female to the nests of the po- tential hosts, as there are no data suggesting anything comparable to mating in these parasites. That the male should act as a foil, to draw off the barbets on guard while the female deposits her egg, seems like too good a story not to have entered into the recording of what- ever may have actually transpired.’ In the case described above by Winterbottom, no male honey-guide was noted. However, we still have so few direct observations of the act of egg deposition that it is advisable to keep an open mind on this matter, even though it seems rather unlikely that the males attend the laying females. 2. Sharp-billed honey-guide, Prodotiscus regulus Serle (1957, pp. 416-417) collected a female at Enugu, eastern Ni- geria, on September 27 that had an egg in membrane in the oviduct HONE Y-GUIDES—-FRIEDMANN 311 and three large yolked ova and two ruptured follicles in the ovary, indicating that at least five eggs could have been laid. Additional Host Records Our knowledge of the life histories of all the honey-guides is still so imperfect that it is to be expected that additions to the lists of known victims will be made for a long time to come. Besides these new host species, further data on previously poorly documented ones have also come to attention in the past three years. The total supplementary data, presented here under the various species of honey-guides, do not change the present picture materially but merely add to the total recorded information and help to orient more properly some of the earlier information. 1. Scaly-throated honey-guide, Indicator variegatus One additional host has been recorded for this still infrequently observed honey-guide. The case is of sufficient interest to quote the original account in some detail. Jackson’s tinker-bird, Pogoniulus bilineatus jacksoni (Sharp) Van Someren (1956, p. 220) writes that . at another nest I knew to hold young, I noted fresh chippings on the ground and thought perhaps another hole was being started, but there was no such cavity. Sitting down in cover, I heard dull tapping coming from the nest hole, then noticed chippings coming out .,.. With a sharp knife, I cut a circular opening . . . and exposed the nest. The chamber and tunnel had been considerably enlarged and within was a three-quarters grown variegated honey- guide. It was remarkable that the hen honey-guide had been able to force her way in and lay her egg in the original small chamber, and moreover, how did the barbets come to appreciate that the chamber was too small to accommodate the chick! Yet, here they were, enlarging the chamber to ensure the comfort of their foster child! I replaced the circle of wood and sealed it in. The young honey- guide was seen in the forest two weeks later, attended by the foster parents . It is fortunate that, in this case, the young parasite was feathered sufficiently to make its identification certain. In my book (1955, p. 105) I listed one record for the Uganda race of this tinker-bird, Pogoniulus bilineatus nyansae (Neumann), and echoed Jackson’s (1938, p. 734) doubts that either Indicator variegatus or I. indicator could possibly get inside the small nest opening to lay there, or that the young parasite, when ready to leave, could get out through it. It now appears that our doubts were needless. Van Someren (1956, p. 221) writes that he has seen a “variegated honey-guide struggling into a hole scarcely large enough for her to enter.”’ 2. Greater honey-guide, Indicator indicator The new data on this, the best known of the honey-guides, whose recorded hosts now number 82 species, or, including subspecies, 38 forms, are as follows: 312 PROCEEDINGS OF THE NATIONAL MUSEUM Vor. 108 Striped kingfisher, Halcyon chelicuti chelicuti (Stanley) Previously known from a single instance near Marandellas, Southern Rhodesia, this kingfisher is listed as a host by Smithers, Irwin, and Paterson (1957, p. 89), possibly on the basis of the same record. Neuby-Varty, the discoverer of the first case, has recently written me of what may be asecond case. Early in December 1955, near Maran- dellas, he watched a striped kingfisher at a nest hole about 20 feet up in a tree. Suddenly, a freshly dead kingfisher nestling, about 4 or 5 days old, fell out of the opening and to the ground. He picked it up and noted tiny punctures anterior to the wings that looked very much like the bill hook wounds of a nestling honey-guide. Unfortunately, he was not able to get to the nest and so could not prove that there was a honey-guide chick present. It may be recalled that in the case of nestling ejection by the young honey-guide in a nest of the crested barbet (Friedmann, 1955, pp. 147-148) the young barbets were ejected alive and unharmed, possibly because the nest stump had been broken off and there was little depth left to the nest cavity, thereby making ejection easier. In the present instance, it may be that the depth was greater and the parasite did not attempt eviction until after it had immobilized its nest mate. Cinnamon-breasted bee-eater, Melittophagus lafresnayii oreobates (Sharpe) In his recent book, van Someren (1956, p. 190) writes that he has taken eggs of the greater honey-guide from nests of this species in Kenya Colony, but does not give any indication of how many such cases he found. In addition, W. R. Ingram informs me (in litt.) that he once found a nest of this bee-eater, also in Kenya Colony, containing one egg of Jndicator indicator as well as some pecked eggs of the host. This bee-eater was not known to be parasitized prior to these observations. Crested barbet, Trachyphonus vaillantii vaillantii Ranzani To the two records given in my book (1955, pp. 147-148) may be added two more, both of parasitized nests found in Southern Rhodesia by C. T. Fisher, and mentioned to me by Mr. H. M. Miles (in litt.) Banded sand martin, Riparia cincta cincta (Boddaert) One record for this hitherto unrecorded host, but the identification of the species of honey-guide involved is only inferential. Captain R. H. Stevenson informs me that a honey-guide’s egg was found in a nest of this swallow in the Selukwe Reserve, Southern Rhodesia, about the end of November or the first days of December 1955. It is possible that either Indicator indicator or I. minor might be involved. In the absence of details, such as notes on the status of the two in that locality, dimensions of the egg, ete., it is not possible to allocate the record. It would seem more likely to be J. indicator because that species makes use of nests in holes in the ground much more HONEY-GUIDES—FRIEDMANN 313 frequently than does J. minor, but both are known to do so. In their recent book on the birds of Southern Rhodesia, Smithers, Irwin, and Paterson (1957, p. 89) list this swallow as a host of the greater honey-guide. I do not know if this is based on Captain Stevenson’s record or if it is still another case. Kenya anteater-chat, Myrmecocichla aethiops cryptoleuca (Sharpe) Previously only two records were known to me, to which two more may now be added, indicating more definitely something of the regularity, if not frequency, with which this species is parasitized. W. R. Ingram informs me that he has found two nests of this bird, each containing a single egg of the greater honey-guide in addition to several pecked eggs of the host. Red-shouldered glossy starling, Lamprocolius nitens (Linnaeus) To the single host record recorded in my book (1955, p. 152) may be added another, unfortunately without exact data. Mr. C. H. Jerome informs me that he has been told of a second such occurrence, but did not have any explicit information about it. Blue-eared glossy starling, Lamproceolius chalybeus chalybeus (Hemprich and Ehrenberg) The southern subspecies of this starling, LZ. c. sycobius Hartlaub, was previously known to be parasitized by the greater honey-guide in the northern Transvaal. We may now note the same for the nominate race. Mr. W. R. Ingram writes me that at Serere, Teso District, Uganda, on Mar. 22, 1956, he found a nest of this bird containing a single egg of the greater honey-guide, together with some pecked eggs of the starling. 3. Lesser honey-guide, Indicator minor Recent data of interest involve three species of hosts, one of them previously unrecorded in this capacity. Striped kingfisher, Halcyon chelicuti chelicuti (Stanley) Previously (Friedmann, 1955, p. 193) I knew of two records, both in South Africa. To these may be added a third, somewhat indefinite one from Kenya Colony, where van Someren (1956 p. 221) found an egg attributed to the lesser honey-guide in a nest of this little kingfisher. Cinnamon-breasted bee-eater, Melittophagus lafresnayii oreobates (Sharpe) Van Someren (1956, p. 190) has found this bee-eater to be par- asitized by the lesser as well as by the greater honey-guide in Kenya Colony. This is an addition to the known victims. Pied barbet, Tricholaema leucomelan (Boddaert) Previously known as a frequent victim in South Africa, this barbet is now recorded in this capacity in Southern Rhodesia as well, where, according to H. M. Miles (in litt.), Irwin found a nest containing a young lesser honey-guide as the sole occupant. 314 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Although only of suggestive interest, mention may be made of the fact that in Ghana, between Accra and Kumasi, early in June, Donald W. Lamm (in litt.) watched a colony of brown barbets, Gymnobucco calvus, ready to begin breeding. At least four lesser honey-guides (Indicator minor ussheri) were present. They were very quiet, perching on the heavier branches of the trees, and showed no attempts to enter any of the nest holes. Two of them, a male and a female, were collected, both with well-developed gonads. This observation suggests that this species of barbet, as well as Gymnobucco bonapartei (already so recorded by Friedmann, 1955, p. 193), may be a host of the lesser honey-guide. 4. Slender-billed honey-guide, Prodotiscus insignis To the little known information about the hosts of this honey-guide previously compiled by me (1955, pp. 251-252) may be added further details of the cases there described, as well as one additional host species. Like the previous data, the new observations all stem from Dr. van Someren. Black-throated wattle-eye, Platysteira peltata peltata Sundevall. Of this host all I was able to report previously (Friedmann, 1955, p. 251) was the bare fact that van Someren had once found a para- sitized nest. He (van Someren, 1956, pp. 281+283) recently supplied more data, of which the following is a summary. When the nest was found, it contained just the young honey-guide, dark brown with a yellow gape, and quite naked. ‘The chick grew rapidly and on the fourth day of observation was well feathered on the head and back. It was then that I saw that... the plumage being gradually assumed was the olive green of a pigmy honey-guide.... The youngster was now receiving quite large moth larvae, imago moths, and dozens of small Diptera.’”’ Shortly afterwards some predator robbed the nest and ended the opportunity for further observation. Kikuyu green white-eye, Zosterops virens kikuyuensis (Sharpe) To the case I previously recorded (Friedmann, 1955, p. 252) merely as having two young slender-billed honey-guides in the nest, van Someren’s (1956, pp. 222-223) additional data provide the following details. He found one of the young parasites just out of the nest on the ground near his house. As he was looking to see what nest it may have come from, he saw one of the white-eyes fly with food to a chick in a nest directly above the spot where he had picked up the fledgling. When his son climbed to the nest, the chick in it fluttered to the ground; it was found to be another slender-billed honey-guide. Van Someren put the two young birds in a cage, to which both parent white-eyes came with food for the next two days. The next night it rained very heavily and, as a result, one of the chicks died. The HONEY-GUIDES—FRIEDMANN 315 other one continued to be fed by the foster parents and was last seen in a tree nearby with the attendant white-eyes. Van Someren concludes his account by stating that the original white-eyes’ eggs had been disposed of and that the honey-guide laid more than one egg in the white-eyes’ nest. In view of the relative scarcity of these honey-guides, it doesseem more probable that the two eggs in this nest were laid by one hen rather than by two, but this is only an inference. Similarly, we have no knowledge as to whether the host’s eggs were removed by the laying honey-guide or ousted by the nestlings. In another place in his book, van Someren (1956, p. 434) does state that the hen honey-guide removes the victim’s eggs one at a time when laying her own, but he gives no substantiating evidence for this statement. Amethyst sunbird, Chalcomitra amethystina kalckreuthi (Cabanis) This sunbird is an addition to the known victims of the slender- billed honey-guide. In November 1956, at Ngong, Kenya Colony, G. R. C. van Someren saw a newly fledged young slender-billed honey- guide being fed by both members of a pair of amethyst sunbirds. He watched them for some time at a distance of less than 10 feet. Mammalian Symbionts It is well established that the original foraging symbiont of the greater honey-guide is the ratel or honey-badger, Mellivora capensis, and that the human has deliberately become a substitute symbiont in its place. In my earlier account (1955, pp. 41-50) I gave some data to indicate that very occasionally baboons might be involved, and cited one instance of a bird apparently attempting in vain to evoke response from a mongoose, Myonaz cauut. Verheyen (1951, pp. 91-93; 1957, pp. 105-113), on the other hand, suggested that the honey-guide may attempt to “call” to any or all of the larger mammals, between which he assumed it could not or did not distinguish. I find it difficult to agree with Verheyen in this matter and look upon the mongoose incident as an occasional error on the bird’s part. However, the fact that as un-ratel-like a creature as the African human could become accepted as a symbiont shows that there may be a basic symbiont tolerance beyond what normally transpires. In this connection, it is of interest to record the following incident involving a greater honey- guide and a genet, kindly sent me by Bryan Neuby-Varty, who made the observation on his farm near Marandellas, Southern Rhodesia. One day he was out in the brush when he heard a honey-guide calling and wondered if it was calling to any creature. Moving carefully he got behind a large tree and could see that it was watching a genet (probably Genetta genetta mossambica) on the ground at a hole at the 316 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 base of a tree from which swarms of bees were flying. As long as the bird kept calling, the genet remained motionless, but soon it disap- peared up to its shoulders down the hole, apparently not minding the bees. Ina short while, it backed out with a small piece of beescomb which it proceeded to eat. Neuby-Varty then moved to a better posi- tion on a termite mound about 30 yards from the genet, from which spot he watched it for about half an hour as it repeatedly inserted its paw into the hole, pulled out pieces of comb, and proceeded to eat them. Then the wind changed and the genet must have scented the observer and it slunk off into the tall grass and was gone. Exam- ination of the hole convinced Neuby-Varty that it was not dug by the genet but may have been the work of a jackal or possibly a mongoose or aratel, although he has never seen the last named animal on his land. In the above incident, there is no evidence that the bird had guided the genet to the hive, which apparently had been opened previously. It is more likely that the bird was attracted to the spot because of the bees, and its interest was then transferred to the genet that had come there independently and which, by virtue of its feeding there, actually kept the bird from doing the same. After the genet had left, the bird called intermittently for about 10 minutes and Neuby-Varty waited another 20 minutes to see if the beast would return. In those 30 minutes, the bird hopped down to the ground only once and pecked at tiny bits of comb. We still have too few observations of associations between the greater honey-guide and various mammals, but it does seem that ob- servations such as this one, or the one involving the mongoose, and even the baboon incident recorded earlier (Friedmann, 1955, pp. 45- 46), hardly justify looking upon these creatures as definitely proved symbionts of the bird in the sense that ratels and humans are. The Termination of Guiding Behavior In my detailed account (Friedmann, 1955, pp. 25-71) I stated that the stimulus which appears to bring to a halt the guiding behavior, released earlier by the bird meeting with a potential foraging symbiont such as a ratel or a human, ‘‘is the sight or sound of bees. It is tempt- ing to expand van Uexkiill’s and Lorenz’s fruitful concept of the ‘kumpan,’ or companion, as the releaser of instinctive actions in birds and apply it to the honey-guides . . . . From this standpoint guiding may be looked upon as the result of the reactions evoked in the bird when the releasing agent is met with in the bird’s natural environment, away from the bees’ nest, for which it is the ‘kumpan,’ and the ‘guid- ing’ behavior is ‘satisfied’ or, at least, brought to a stop when the bird brings together the ‘bee companion’ and the bees.’’ While I still think HONEY-GUIDES—FRIEDMANN 317 this statement is as close to the actual picture as present understanding permits, it has seemed to me possibly a little forced in its reasoning. It was, therefore, with considerable interest that I recently came across, in Russell’s (1953) description of the drive character of instinctive behavior, some ideas that give further support to the applicability of the ‘“kumpan” concept to guiding activities. In the current general theory of instinct, we find that usually a chain of actions grouped under one heading as instinctive behavior tends to be divided into two phases, an introductory one of openly seeking, striving, appetitive behavior, and a directly subsequent one of essentially consummatory action, generally of a quasi-mechanical or quite stereotyped nature. The emphasis on the innate drive char- acter of instinctive behavior, developed largely by Lorenz and his colleagues, seems warranted. These investigators account for the specificity of the drives by hy- pothesizing what they term “action specific energy,” which is said to accumulate and to be discharged with and into highly specific appeti- tive behavior patterns. Lorenz (1950) further assumes ‘‘that some sort of energy, specific to one definite activity, is stored up while this activity remains quiescent and is consumed in its discharge.”’ Russell sees no real need to hypothesize energy when all that seems to be “accumulated”? may be described just as readily as “specific tension or unreleased tendency to carry out a certain course of action.” Regard- less of whether it is a specific energy or a specific tension, the fact remains that we have, in either case, a support for what I implied when I wrote that the guiding behavior was “satisfied’’ and brought to a halt when both the bees and the bee “‘kumpan”’ simultaneously came to be within the sensory range of the bird. The specific energy is discharged, or the specific tension is dispelled, when the guiding bird achieves the unison of the releasing agent, the foraging symbiont, and the thing with which the releaser is associated in the experience of the activated bird, the bees. The mode of termination of guiding is one more example of the increasingly obvious fact that it is the discharge of consummatory action and not the biological or survival value in- volved that is the goal of innate appetitive behavior as far as the indi- vidual bird is concerned. The Rustling Flight The rustling flight, recorded for both the greater and the lesser honey-guides (Friedmann, 1955, pp. 130-133, 184), is of interest not only for itself but as the possible root from which evolved the highly specialized performance of the lyre-tailed honey-guide, Melichneutes robustus. The evidence is somewhat divided as to whether the rustling 318 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 sound is produced by the wings or the tail in the greater and lesser honey-guide species, while in the lyre-tail it seems obvious that the sound is made by the highly peculiar outer tail feathers. Two interesting additional observations have come to me, both having to do with the greater honey-guide. Neuby-Varty writes me that about the end of May, on his farm near Marandellas, Southern Rhodesia, he heard a greater honey-guide giving its victor call from the top of dead branches of a tree. Then it flew towards him, and just as the bird came above him, it started to make a rattling noise, appar- ently with its wings. The tail may have been involved as well, as the bird spread it fanwise while making the noise. Neuby-Varty timed the performance and found the noise (written down as feet-wp) was given 10 times, with an interval of about a second between the sounds. W. R. Ingram, at Serere, Uganda, informs me that he has found the rustling or drumming flight to be given only towards evening and al- ways in the early dry season. He thinks it has no connection with courtship, as there are no suitable hosts nesting at the time of the year. He first heard it in December 1955, at about 7 p. m. and almost dark; the noise ‘‘was most eerie and seemed to come from different parts of the sky almost at once, showing that whoever or whatever was making the noise was moving very fast indeed.” Ingram describes the sound made by suggesting that if one blows out the word whukooo with a great expulsion of air on the first syllable and with strong emphasis on the K, and then emptying one’s lungs on the 000, a similar sound can be produced. He goes on to say that “the noise was heard occasion- ally at dusk during the whole dry season (December—March) and again in 1956, but the author was never discovered.” It was not until 1957 that he succeeded in seeing as well as hearing the performing bird. Early in December of that year, at about 6 p. m., he saw the bird . . traveling at a very high speed in a circling, dipping flight. It careened around the sky for about 30 seconds and then dived into a large tree. . Immediately, three or four victor notes came from this very tree... I knew this tree to be a popular stud-post, all the year round, but still I could not connect the bird in the sky with the honey-guide. However, I did not have to wait very many evenings before I got a repeat performance. I managed to pick up the bird in flight with the binoculars during its drumming flight, follow it round and into the same stud-post. At the moment it darted into the tree, if fanned its tail and the outer white feathers were very conspicuous; this is the only time it opened its tail in flight, so I assume the noise is done with the wings. It landed on a prominent perch and I got close enough for a positive determination .... This account indicates a similarity in habit to the aerial evolutions of the lyre-tail even more definitely than did our previous data. It also supports the observations of Ranger, Neuby-Varty, and myself that the sound is produced by the wings and not by the tail. HONEY-GUIDES—FRIEDMANN 319 Feeding Habits That fair numbers of honey-guides may occasionally gather to eat at a single bees’ nest is shown by an observation sent me by H. F. Stoneham, who heard noises coming from one of his domestic beehives. Thinking that a rat or some other creature was raiding the nest, he went to it, and was surprised to see eight honey-guides fly out in rapid succession, six greaters and two lessers. Kettlewell (1955, pp. 45-47) describes a nest of wild bees built in an abandoned wrecked automobile, the metal of which became hot from the sun and caused the honey inside to ferment. This apparently had intoxicated a greater honey-guide, which Kettlewell pulled out of the automobile. An addition to the known diet of the lesser honey-guide is reported by van Someren (1956, p. 221), who saw one taking the young larvae and pupae from the paper nest of an aculeate wasp. This recalls the old observation of Butler, Feilden, and Reid (1882, p. 208) who reported the greater honey-guide pecking at the comb of a wasp’s nest that had fallen to the ground. Chapin has recently sent me some observations on the feeding habits of the least honey-guide, /ndicator exilis, in the eastern Belgian Congo. He opened an old bees’ nest in which he found considerable quantities of.comb, practically empty of honey. The next day a least honey- guide came to it, and again two days later he saw one there. He placed a piece of the comb in a branch of a tall bush where he could watch it, and the bird came there and ate pieces of the comb. The bird was alone in each case, which fact seems to answer Chapin’s (1939, p. 540) earlier statement that since this species does not guide humans, it may have some other mammalian symbiont. Chapin observed not only the least honey-guide feeding at open bees’ nests, but also his newly discovered pigmy honey-guide, J/ndicator pumilio. In fact, most of his specimens of the latter were captured with a butterfly net as they emerged from a beehive. Recently Verheyen (1957) has taken objection to my conclusion that the primary interest of the honey-guides in bees’ nests is the wax of the comb rather than the honey, pollen, or bee larvae. It should be pointed out that I have described that the birds do eat the bee larvae and pupae, and, adventitiously, the honey, but it still remains that the wax is the one substance they are most eager to get from the hive, and the one substance they cannot obtain elsewhere. They are constantly catching insects on the wing, and are certainly not primarily wax feeders. As I pointed out, honey-guides grow to full size in their hosts’ nests without getting any wax in their food, but once they begin fend- ing for themselves they eat wax avidly, not as a substitute for some 320 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 other food but as an addition to their diet. By itself, beeswax is not a ‘“‘total” food as it lacks nitrogen, without which no bird could survive for more than a few weeks or a month. References BieRENS DE Haan, J. A, 1956. Het honingaanwijzen van de Honingaanwijzers. Ardea, vol. 44, pp. 289-293. Butter, Epwarp A.; Frinpen, Henry W.; anv Ret, SAvILye G. 1882. Ornithological notes from Natal. Zoologist, ser. 3, vol. 6, No. 66, pp. 204-212. CHAPIN, JAMES P. 1939. The birds of the Belgian Congo, II. Bull. Amer. Mus. Nat. Hist., vol. 75, 632 pp. FRIEDMANN, H@ERBERT. 1955. The honey-guides. U.S. Nat. Mus. Bull. 208, 292 pp. JACKSON, FREDERICK JOHN. 1938. The birds of Kenya Colony and the Uganda Protectorate, vol. 2, pp. 545-1134. KETTLEWELL, H. B. D. 1955. Note. Entomologist, vol. 88, No. 1101, pp. 45-47. Lorenz, KONRAD 1950. The comparative method of studying innate behaviour patterns in physiological mechanisms in animal behaviour. Soc. Exp. Biol. Symposium 4, pp. 221-268. Roserts, Huau A. 1956. Breeding tactics of the two honey-guides—Indicator indicator (Sparrman) and Indicator minor Stephens. Bull. Brit. Orn. Club, vol. 76, p. 114. Russet, E. S. 1953. The drive character of instinctive behaviour. Arch. Néerlandaises Zool., vol. 10, suppl. 2, pp. 110-120. SERLE, WILLIAM 1957. - less 9 |= 302 | ss sa |x Sleeps Koa |X) gules ee TUT TOUELL Seales be eee tae] raja ees eae eae ee Peete ese | Sees eee aes Wiles saa] eae ceaalhg pee atee ReaRsaG oC ls | ae eg USUI aa NCA |x aaa ad || eee Ck eee 1837DMS mee |e eee nekid Nina Mn lA © |0 Awa ao T TLE O tele eile cl cet | dete kG we |prare goer Gm a Smeal ees | Dake | LUGE TOG 1 TS 0 ilies Nast $nI13 -UD]2D SNUDIITaUL Bah ee ee Gen On GR cola so len D RE ete ae eee Sa) Se) eerie ae mle ea snuvo -129UL SNUDILTIUL See asf elas see ee etl nes meae|Ge—-| ge |anao| nant ~2eccen-[---===-""="9 05ND t lecre|--2-]---- Te cee lene ooo snoiuod ot DIT gz | 22 | 92 | 90 | | e2 | ze] 1% | 0% | BL | SE | AT | OT | ST | HUTT] SLT | Ores | TUTE] OTT] eTT | OT | 6 | 8 #1 | €T | Ot | XT| TIIA|I-I-IA [esioq sopods pus eiouer) [810990g jeuy Yoiv JSuy WO sJeyeI [ITZ jo zaquumMu [v4.0], shel uy Jo 1OqUINN Pee a oer ae ea a ae ead ee ee (01N4B19}1] 9} WOT popsodel vyep sojousp ,,X,, 12}997]) wNnysysioyg pup “ewodoaidory ‘soyxlg viauab ay} 07 a1qnsafas savads sof papsovas sJuN0Q—'T DTIAVL 326 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 snout 89, 88, 84, and 77; fleshy interorbital space 59, 69, 62, and 60; diameter of eye 79, 77, 84, and 86; postorbital length of head 208, 200, 199, and 192; least width of preorbital 25, 32, 26, and 25; length of maxillaries 164, 167, 166, and 176; least depth of caudal peduncle 127, 135, 133 and 137; distance from base of last anal ray to midbase of caudal fin 228, 231, 235, and 242; length of longest (third or fourth) dorsal spine 119; 123, ——, and 116; of longest soft dorsal (third from last) ray 228, 211, 208, and 239; longest anal (third) spine 114, 107, 89, and 102; longest soft anal (third from last) ray 189, 177, —-, and 179; longest pectoral ray 297, 304, 310, and 297; longest pelvic ray 173, 181, 186, aad 176; longest caudal fin ray 238, —-, 257, and 2a. Cotor IN ALcoHOoL: Plain straw-colored, with traces of black pig- ment at tips of outer caudal fin rays. Remarks: This new species is distinguished from P. cubensis by having 14 soft dorsal rays instead of 13. It may be distinguished from the other related species by means of the key. Pikea mexicanus atlanticus, new subspecies Houtotypr: USNM 185005, collected Nov. 8, 1957, off French Guiana, lat. 7°18’ N., long. 53°32’ W., Oregon station 2021, 1090 fathoms, standard length 108 mm. Paratypes: USNM 185008, taken with holotype and bearing same data, standard length 98 mm. USNM 185007, off Surinam, lat. 7°32’ N., long. 54°12’ W., 100 fathoms, Nov. 8, 1957, Oregon station 2014, 5 specimens, 84 to 109 mm. USNM 185006, off Venezuela, lat. 9°53’ N., long. 59°53’ W., 125 fathoms, Nov. 3, 1957, Oregon station 1983, 9 specimens, 59 to 107 mm.. USNM 185082, off Surinam, lat. 7°30’ N., long. 54°16’ W., 125 fathoms, Nov. 8, 1957, Oregon sta- tion 2013, 1 specimen, 56 mm. USNM 185080, off Venezuela, lat. 9°24’ N., long. 59°41’ W., 110 fathoms, Nov. 4, 1957, Oregon station 1988, 1 specimen, 77 mm. USNM 185081, off Venezuela, lat. 9°39’ N., long. 59°47’ W., 100 fathoms, Nov. 4, 1957, Oregon station 1986, 9 specimens, 59 to 92 mm. This new subspecies from off the Guianas and Venezuela is essen- tially the same (in all important characters) as Pikea mexicanus mexicanus of the Gulf of Mexico, except for the number of gill rakers on the first gill arch—18 to 21 on P. m. atlanticus and 21 to 23 on P. m. mexicanus. Because of the slight overlap in counts of gill rakers, I prefer to consider the form from off the coasts of Venezuela and the Guianas as a subspecies. NEW SERRANID FISHES—SCHULTZ 327 Key to genera and species related to Pikea la. Spiny dorsal fin continuous with soft dorsal by a ridge of scales along sides of connecting dorsal spines; no scaled area separating spiny and soft dorsal rays; 244 to 5% scales in a row from lateral line to base of last dorsal spine; 16 to 21 predorsal scales to occiput. 2a. Dorsal spines vu (genus Pikea) } 3a. Anal rays 111,10; dorsal rays v1t1,14; pectoral ii,14; pores in lateral line 47; zig-zag scales around caudal preduncle 40 to 42; outer edge, distally, of each caudal lobe white (Japan). . . . P. japonicus (Déderlein) ? 3b. Anal rays 11,9; dorsal rays vi11,13; pectoral ii,13; pores in lateral line 48 to 51, zig-zag scales around caudal peduncle about 45 (Hawaiian Islands). ...... .. . . P.aurera Jordan and Evermann ? 3c. Anal rays 111,8. 4a. Pores in lateral line 43 to 49, rarely 49. 5a. Dorsal rays vu1,12. 6a. Pores in lateral line 43; pectoral rays ii,11; gill rakers on first gill arch about 15; tips of outer caudal fin rays black (Bermuda). P. mowbrayi (Woods and Kanazawa) 4 6b. Pores in lateral line 46 or 47; pectoral rays ii,12; predorsal scales to occiput 19 to 21; gill rakers on first gill arch about 28 (Philip- pines). ...... . . . P.swalesi (Fowler and Bean) ® 5b. Dorsal rays vu,13 or 14; pectoral rays ii,13 rarely ii,12 or ii,14; pores in lateral line 46 or 47; tips of outer caudal fin rays usually black in adults. 7a. Dorsal rays vi11,13; gill rakers on first arch 20 or 21 (Cuba). P. cubensis, new species 7b. Dorsal rays vr11,14; gill rakers on first arch 21 to 23 (Gulf of Mexico). P. mexicanus mexicanus, new species and subspecies 7e. Dorsal rays vir,14; gill rakers on first arch 18 to 21, rarely 21 (off Venezuela and Guianas). P. mexicanus atlanticus, new subspecies 4b. Pores in lateral line 48 to 70; gill rakers on first gill arch about 18 to 20 (unknown for P. maculata). ! Pikea Steindachner, Sitzber. Akad. Wiss. Wien, vol. 69, p. 2, 1874 (type species, Pikea lunulata (= Grystes lunulatus Guichenot 1863). Labracopsis Déderlein in Steindachner and Déderlein, Anz. Akad. Wien, vol. 20, p. 49, 1883; Denkschr. Akad. Wien, vol. 47, p. 235, 1883 (type species, Labracopsis japonicus Déderlein). 2 Labracopsis japonicus Déderlein in Steindachner and Déderlein, Anz. Akad. Wien, vol. 20, p. 49, 1883; Denkschr. Akad. Wien, vol. 47, p. 235, pl. 6, fig. 3, 1883 (Japan). * Pikea aurora Jordan and Evermann, Bull. U.S. Bur. Fish. vol. 22, (1902), p. 178, 1903; Ibid., vol. 23 (1903) pt. 1, p. 220, pl. 14, 1905 (Hilo). 4 Liopropoma mowbrayi Woods and Kanazawa, Fieldiana, Zool., vol. 31, No. 53, p. 633, fig. 134, 1951 (Ber- muda). ' Chorististium swalesi Fowler and Bean, U.S. Nat. Mus. Bull. 100, vol. 10, p. 186, 1930 (Gulf of Tomini, Celebes). 328 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 8a. Pores in lateral line 48 to 55. 9a. Pectoral rays ii,13 or 14; dorsal rays vi11,12; pores in lateral line 48 to 52; zig-zag scales around caudal peduncle about 40 to 52; body with numerous black spots (Reunion Island and Mauritius). ays. eae) oa yes 4c os bunulatay(Guichenot) ss 9b. Pectoral rays i,13; dorsal vi11,18; pores in lateral line 54 or 55 (Panama)! s'50 cay eens . .. . . P. longilepis (Garman) 7 8b. Pores in lateral line ibout 65 to 70; dorsal rays v111,12; pectoral ii,13; sides of body spotted (Japan). P. maculata Steindachner and Déderlein 8 2b. Dorsal spines 1x (genus Liopropoma)® 10a. Dorsal rays 1x,12; pores in lateral line 45 (Cuba). L. aberrans (Poey)?? 10b. Dorsal rays 1x,14; pores in lateral line 58 (my count of drawing is 62 or 63) (Pernambuco, Brazil)... ... . . L.roseus (Giimther)!! 1b. Spiny dorsal and soft dorsal fins completely separated externally by a scaled area of 5 to 7 rows across back between fins; dorsal rays vi-1-1,11 or 12; the seventh dorsal spine usually embedded below scales; pores in lateral line 45 to 48; 2% to 5 scales in a row from lateral line to last dorsal spine; zig-zag scales around caudal peduncle 30 to 32; gill rakers 5 to 7+1-++ 11 to 14 on first arch; anal rays1m,8. . . . (genus Chorististium Gill)” lla. Coloration plain pale, no stripes or dark pigment marks on sides or on fins; pectoral raysii,14;predorsal scales 12 to occiput; dorsal rays vi-1-1,i,11 (Christmas Island and Bikini Atoll)... . . . C. pallidum Fowler 11b. Coloration of alternating dark and pale streaks or dark wavy lines on sides. 12a. Sides of body with brown pigment specks, a few arranged in irregular lines on midsides; an oblique short dark brown streak on pectoral base; predorsal scales 13 to occiput... . . . . C. brocki (Schultz) 6 Grystes lunulatus Guichenot, Notes sur He dela Reunion. Faunae Ichthyologique, p. c-4, 1863 (Reunion Island). Pikea lunulata Steindachner, Sitzber. Akad. Wiss. Wien, vol. 69, p. 2, 1874; Denkschr. Akad. Wiss. Wien, vol. 47, pl. 6, fig. 2, 1883. Glaucosoma semilunifera Steindachner, in Bliss, Trans. Roy. Soc. Mauritius, new ser. vol. 13, p. 47, 1883 (Mauritius). 7 Liopropoma longilepis Garman, Mem. Mus. Comp. Zool., vol. 24, p. 45, 1899 (lat. 7°33’ N.; long. 78°34’20’” W.). 8 Pikea maculata Déderlein and Steindachner, Denkschr. Akad. Wiss. Wien, vol. 47, p. 234, pl. 6, figs. 1, la, 1b, 1883 (Japan). ° Liopropoma Gill, Proc. Acad. Nat. Sci. Philadelphia, p. 52, 1861 (type species, Perca aberrans Poey). Bathyanthics Giinther, Rep. Voyage Challenger, Zool., vol. 1, No. 6, p. 6, 1880 (type species Bathyanthias roseus Gunther). 10 Perca aberrans Poey, Memorias, vol. 2, p. 125, pl. 12, fig. 2, 1860 (Cuba). 1 Bathyanthias roseus Giinther, Rep. Voyage Challenger, Zool., vol. 1, No. 6, p. 6, pl. 1, fig. B, 1880 (Pernam- buco, Brazil). 12 Chorististium Gill, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, p. 15, 1862 (type species, Perca rubre Poey; misspelled Chorististum in heading). Ypsigramma Schultz, U. S. Nat. Mus. Bull. 202, vol. 1, p. 372, 1953 (type species, Ypsigramma lineata Schultz). 13 Chorististiwm pallidum Fowler, Monogr. Acad. Nat. Sci. Philadelphia, vol. 2, p. 199, fig. 20, 1938 (Christmas Island). 14 Ypsigramma brocki Schultz, U. 8. Nat. Mus. Bull. 202, vol. 1, p. 379, fig. 60, 1953 (Marshall and Gilbert Islands). NEW SERRANID FISHES—SCHULTZ 329 12b. Coloration not as above. 13a. Sides of body with 7 or 8 distinct, almost straight lengthwise dark brown streaks. 14a. Five brown streaks on each side of caudal peduncle; 3% or 4 scales between lateral line and last doral spine; predorsal scales 12 or 13 to occiput... . .... . . =. C.lineata (Schultz) 14b. Four brown streaks on each side of caudal peduncle; 3 scales between lateral line and second dorsal origin; predorsal scales 10 to occiput. . . . . . . . . C.susumi Jordan and Seale !4 13b. Four or 5 dark streaks on each side of body; 3 dark streaks on each side of caudal peduncle. 15a. Five dark streaks on each side ef body; pectoral rays ii,12; tips of posterior lobes of median fins with black blotches (Cuba; Bahama Islands)... . 2 ; . . CG. rubrum (Poey)!7 15b. Four dark streaks on ae side Gf body; pectoral rays ii,13 (Zanzibar) ........... . . C. africanum Smith '8 185 Ypsigramma lineata Schultz, U.S. Nat. Mus. Bull. 202, vol. 1, p. 355, fig. 59, 1953 (Marshall and Philippine Islands). Chorististium susumi (not Jordan and Seale) Smith, J. L. B., Ann. Mag. Nat. Hist., ser. 12, vol. 7, p. 862, pl. 27, fig. c, 1954 (Matemo Island, off East African Coast). 16 Chorististiwm susumi Jordan and Seale, Bull. U. 8. Bur. Fish., vol. 25 (1905), p. 256, fig. 48, 1906 (Apia, Samoa). 17 Liopropoma rubre Poey, Memorias, vol. 2, p. 418, 1861 (Cuba). 18 Chorististium africanum Smith, J. L. B., Ann. Mag. Nat. Hist., ser. 12, vol. 7, p. 866, fig. 1, pl. 27, fig. B, 1954 (Pinda Pembra, Zanzibar and Tekomazi Island). U. S. GOVERNMENT PRINTING OFFICE: 1958 AOKE CE REMORSE ANSE Mai © 6 Fan ae Bm 8 gh eR FA. copdulle dpe de) apttertaie dal drat. Beberdea cs. Ce iabataral to dete OR ell) ydlehCaveatdins.’ fe aia 3, silat... :dSneobi aie lene 0 Farce omasditeern se aiet ont SOfAde Lanse Pr iy -@ ta teenly dapl feta Ged ina. Lal adettecieemt “yr nts F Hi Mae Peer - 8 OE, aD Ghat? 5 JJudidowtaiiee a: (o> eer AB olde Oli oye Telia diese adele aver Oe, GT omy? Sipe nr! 4p) betas Pirkei? fies - TPF eae Yet roAL UPabiset Cu, ryt hy farsi Ses! eee er er “sale & xnoet AG ARE ror) At 1c. Create pict) WER eae te tab time Tease ae ee © e6s\0 Sed 8 - A aed) aiilulth fame a ei aot $.) Al = Aj;eje pte = “Fr eat! Ge" Tk PMT, ts ee eG ae - Re oe = ine it Se ah whe ns FE te 1B - CL SEs <= = a erprcaltnitt gmt fy ope a inane “eet Gi a a emi) IT chow pla Ue eg <1 1 Peg 2 — —. sade. et) o io ol MA WALLS wide Wn? 4 OME Flo me Palin + uy? BE gly 7 a DER Se ee, Oty a ee, ee ES st ORO a) as a = ae See Se ae ve a . - Oo oo > a o> oy - ff: 4 > — = (79 we a = ' ol vat - a » . =a “si ae > law i ieee - | a Se a a, SS aa = > a : > «= 4 ee ro > .; - = me ts, i isc, eo = ae a aa oo ae ee oe ~ G42 2 ear “ae = se Se ete : nS EC EO 7 re at) amt = | el a & Oe tak ae a, Ss PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1958 No. 3406 THE STATUS OF THE LIZARD CNEMIDOPHORUS PERPLEXUS BAIRD AND GIRARD (TEIIDAE) By T. Pau, Mastin,’ Ricwarp G. BEIDLEMAN,? and CHarLes H. Lowe, Jr.’ Several years ago Burger (1950), in a paper concerned with the systematics of Cnemidophorus, stripped the name Cnemidophorus perplezus of all specimens previously associated with it except the type. This action followed a period of some 50 years of taxonomic confusion and, as Burger himself pointed out, was a temporary measure until more material was available with definite locality data. Burger, with qualifications, followed Burt (1931) in his designation of the type and type locality. Burt in turn apparently based his designation of the type on indirect statements made by Cope in 1893 and again in 1900 and possibly from examinations of the catalog entries of the U.S. National Museum (USNM). But he based his designation of the type locality on the original description of Baird and Girard in 1852. His selection of the type locality has been accepted by most workers since then, but his action was unjustified. He designated one specimen as the lectotype of perplexus, and the habitat of different specimens as the type locality. This discrepancy and the recent description of two new striped species from the Southwest provided 1 University of Colorado Museum, Boulder, Colo. 2 Colorado College, Colorado Springs, Colo. 2 University of Arizona, Tucson, Ariz. 332 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 the original motivation for a reinvestigation of the status of the species, which poses the problem of answering three questions. First, what specimen actually constitutes the type; second, what is the type locality; and third, with what species in this area may the name perplerus be associated? ‘These questions while simple are not easily answered Their solutions have been greatly facilitated by the kind- ness of Dr. Remington Kellogg, director of the U.S. National Museum, in making available to us type material and other specimens; by the cooperation of Dr. Doris M. Cochran in her search of the USNM catalogs; and by the material assistance of the Council on Research and Creative Work of the University of Colorado, whose monetary aid has made possible the acquisition of comparative material. Tur Tyrer: Baird and Girard (1852) published descriptions of a series of lizards based largely on material collected by Dr. John H. Clark, under Col. J. D. Graham, head of the Scientific Corps, U. S. and Mexican Boundary Commission; but there were also included specimens collected by others. In this paper the extremely brief description of Cnemidophorus perplexus is based on an unstated number of specimens presumably collected by Clark in the valley of the Rio San Pedro of the Rio Grande del Norte (according to Smith and Taylor, 1950, p. 363, this is the Devils River, Val Verde County, Tex.) and specimens collected by Gen. Churchill on the Rio Grande west of San Antonio and by Dr. William Gambel, at no specific locality, on his last journey to California. No holotype is indicated, nor by eany clue from the description, title, or introduction does one speci- men seem to receive closer attention that any other. This is not too surprising in that the type concept was in its infancy at that time. As far as we are aware, the first indication that a type existed is Cope’s (1893, p. 34) remark that ‘The type specimen is the largest obtained, and is probably adult.”” He does not, however, indicate to which specimen he is referring; but this can be determined on the basis of size alone. Cope (1893, pl. 6, figs. 1, A-a; pl. 12, fig. B) also fig- ures the species. In his explanation of plates (p. 51) it is indicated that figure 1 of plate 6 is of “Specimen No. 3060 U.S. National Museum.” Cope used these figures again in a later work (Cope, 1900, p. 573, fig. 105), but here they lie above the legend ‘‘Cat. No. 3060, U.S. N. M.” This suggests, but does not specifically state, that the figures are of the type and that a lectotype, USNM 3060, had been designated. We can find no such designation in the literature and assume that Cope him- self selected a type sometime prior to 1893. Frequently Cope (1900) designated the types in the tables of specimens held by the Museum. But in this instance USNM 3060 is not even listed, nor does Yarrow (1882) list this specimen in his catalog. Furthermore, in his extended description of “the type’’ Cope’s (1900, p. 573) first sentence does not LIZARD CNEMIDOPHORUS PERPLEXUS—MASLIN, ET AL. 333 agree with the published figure above it. He states: “In the type specimen of this subspecies the interparietal plate is narrower than the parietals, and is twice as long as wide.’ In the figure it is much broader. Farther on he also states that there are 19 femoral pores on each side in the type; the excellent figures show 25. These discrepancies suggest that possibly the type is some specimen other than USNM 3060 or, as was frequently done in those days, more than one specimen was cataloged under that number. Through the kindness of Dr. Kellogg we have been able to examine all of the specimens upon which Baird and Girard based their description. The largest of these is now numbered 3060, and it is perfectly obvious that the figure, in spite of the legend beneath it, is not of this specimen. On the other hand Cope’s (1900, pp. 573-574) description of the type is a remarkably accurate description of this very same individual. Originally, however, there were, indeed, two specimens cataloged under this number. The smaller of the two was reidentified by Stejneger (Cochran, in lit.) as gularis and assigned a new number, USNM 30885. Actually the specimen—judging by its small size, seven light stripes, moderately enlarged hexagonal post-antebrachial scales, enlarged temporal scales, etc.—is a specimen of C. inornatus. It is of interest to note that Cope’s (1900, p. 588, fig. 112) figure of Onemidophorus tesselatus variolosus Cope is also of a specimen num- bered 3060. But this figure, too, is not of the specimen now bearing this number, nor is it a figure of USNM 30885. Cope obviously had been careless in assembling his manuscript, and his figures in this instance are valueless in determining the status of this species. His (1900) figure 105 is not a figure of a specimen of C. perplexus at all, but probably of some race of C. tigris. Before assuming that USNM 3060 is the type of perplexus as indi- cated but not specifically stated by Cope, it is essential to know if this specimen is a syntype. The original material upon which Baird and Girard based their description of this species is apparently intact, or nearly so, and in the U. S. National Museum; but the catalog entries of this material do not completely match Baird and Girard’s (1852, p. 128) locality notations. Dr. Doris M. Cochran, curator of herpetology at the U.S. National Museum, has kindly provided us with the catalog entries on this material. The specimens from the ‘Valley of the Rio San Pedro of the Rio Grande del Norte’’ are cataloged as having been collected along the ‘“‘Rio San Pedro to the Rio Grande, Texas’ by Col. Graham. However, Baird and Girard (1852) in their introduction specifically state the bulk of the material upon which their paper is based was “collected by John H. Clark, under Col. J. D. Graham.”’ There are now five specimens in this lot cataloged under USNM 3022; origin- 334 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 ally there were six. The specimens ‘collected’ by Gen. Churchill are two in number and are cataloged under USNM 3050, and desig- nated as having been collected in ‘‘Texas.’”? Gambel’s contribution is now entered as follows: ““Type/Cnemidophorus perplexus/‘Calif’??/ probably W. Texas/. Dr. W. Gambel/3060/1.”,. Dr. Cochran (in lit.) informs us that at Dr. L. Stejneger’s suggestion she added in her own hand the word “‘type,”’ the two question marks after ‘Calif’ and the notation “probably W. Texas.” These entries so closely match the notation of Baird and Girard in their original description of the species that there seems little doubt that they truly represent the specimens in hand when the original description was made, even though at that time the specimens were uncataloged and without numbers. The taxonomic fate of Clark’s and Churchill’s specimens will be mentioned later; it is Gambel’s large specimen that is of im- portance at the moment. The catalog entry of USNM 3060 was made on July 20, 1858, probably by Baird himself (Cochran, in lit.). Between that time and 1893 it had presumably been selected by Cope as the type of perplezus, for in 1900 he described a specimen which he designated in his de- scription as the type, and this specimen is unquestionably USNM 3060. It is not until Burt (1931) published his studies on the teiid lizards of the genus Cnemidophorus that a statement is finally made in the synonymy of Cnemidophorus sealineatus perplexus that the type is USNM 3060. This action of Burt’s constitutes a formal designation of a lectotype, properly selected from the syntypes of Baird and Girard. According to our interpretation of the proceedings of the 12th International Congress of Zoology, Paris, 1948, and the 14th International Congress of Zoology, Copenhagen, 1953, this action is final and the lectotype of Cnemidophorus perplexus Baird and Girard is what is now USNM 3060. THE TYPE LOCALITY: Examination of the Clark material shows that it is not Cnemidophorus perplexus but rather C. sacki gularis. Hence, the type locality of Rio Pedro of the Rio Grande del Norte is invalid for the former species. As will be seen, Gambel’s specimens, including the type, USNM 3060, could not have been collected in ‘‘W. Texas” as suggested by Dr. Stejneger inasmuch as Gambel never visited this part of the Southwest. Furthermore, the suggestion by several authors that Gambel made the collections on his second journey to California seems unlikely because of the northerly route followed and certain terminal events associated with that trip. Therefore it is necessary to reevaluate the actual type locality for Cnemidophorus perplexus, as evidence permits. William Gambel was a young Philadelphia protégé of the frontier botanist Thomas Nuttall. After making several eastern collecting LIZARD CNEMIDOPHORUS PERPLEXUS—MASLIN, ET AL. 335 excursions with this eminent scientist, Gambel, at 18, was encouraged to attempt a trip to the West, penetrating the only portion of this new country which Nuttall had not himself visited, namely the Southwest. Today, Gambel is most well known for making the first collections of plants from the Santa Fe, N. Mex., region and for his observations on and collections of birds from this frontier country of the 1840’s. However, he also acquired a small collection of reptiles, including the designated type of Cnemidophorus perplexus and the following: Holbrookia texrana (USNM 2787), Sceloporus graciosus (USNM 2861), Phrynosoma modestum (USNM 176), Crotaphytus wislizeni (USNM 2722), Heterodon nasicus nasicus (USNM 1277), Masticophis taeniatus (USNM 1979), and Thamnophis elegans vagrans (USNM 908). Because Gambel published on the birds and Nuttall published on Gambel’s plant collections for the Academy of Natural Sciences of Philadelphia, accurate information is available with respect to locali- ties and dates for these specimens. With respect to the reptiles, how- ever, no study of the material was ever published by Gambel or his immediate colleagues; consequently, collection details are vague. Baird and Girard (1852, p. 128) note that the specimens of Cnemi- dophorus perplerus were collected on Gambel’s “last journey to Cali- fornia.”” In Yarrow’s (1882) catalog, Gambel’s several species are for the most part designated as having been collected in “Calif.” The only specific locality is that given for Phrynosoma modestum. Yarrow (1882) cites this specimen (USNM 175) as coming from ‘‘Plaenis.”’ No such place can be located in the territories through which Gambel passed and it seems likely that Yarrow misread a label in making this notation. Cope (1900, p. 439) cites the same specimen as coming from the “plains,’’ presumably of eastern New Mexico. In order to reach some decision on the collection locality, especially of Cnemidophorus perplerus, it is necessary to scrutinize more closely the two western trips of Gambel. The first, in 1841, embarked upon undoubtedly at the behest of Thomas Nuttall, followed in general outline the Santa Fe Trail from Independence to Santa Fe and the Old Spanish Trail from that com- munity to California. The second expedition, which culminated in Gambel’s death, followed the eastern portion of the Oregon Trail from Independence to Wyoming and the Hudspeth Trail over Hastings Cut-Off through Nevada to California. This latter trip was in 1849. Despite suggestions to the contrary in the literature, especially Baird and Girard (1852), it is implausible that any of Gambel’s extant collections were from the second trip, either from California or en route. Indeed, it is most unlikely that any specimen material Gambel might have collected on this trip was ever returned to the East, due to the circumstances described below. 336 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 At the end of this second overland journey, Gambel died of typhoid fever in December 1849 in the northern Sierra Nevadas of California and was buried at Roses Bar above the Feather River. His grave was subsequently destroyed by placer miners. There were friends in Cali- fornia aware of Gambel’s death. D. B. Woods of Mountain House wrote an obituary of Gambel which later appeared in a Philadelphia newspaper. Also, on this trip Gambel had apparently been accom- panied by a man named Beesley from the Philadelphia area who returned to the East early in March of 1851 (Osborn, 1931, p. 259; letter, John Cassin to Baird, March 12, 1851, “. . . . Beesley who accompanied poor Gambel has returned within a day or two... ’’). Gambel, on this second trip, had made a journal which was returned to his wife after his death, most probably by Beesley. Later the journal was turned over to the Academy of Natural Sciences and subsequently lost. There is no evidence, however, that anything beyond this journal was returned from the young naturalist’s be- longings. Nonscientific friends on the scene of Gambel’s death would be most unlikely to take sufficient interest and effort to send or bring back from far away California any specimens, especially alcoholics, either by the torturous overland route or around Cape Horn by ship. Also, little among the known accessions of Gambel suggests a collec- tion point close to the more northerly route which apparently was followed in 1849. Therefore, the designation ‘second journey to California’’ seems invalid as a time and locality for any of Gambel’s collecting, particularly herpetological collecting. The route of Gambel’s first trip west, to Santa Fe and eventually to southern California, has been outlined by several authors but is herein changed somewhat to conform with newly discovered evidence. Gambel left Independence for Santa Fe with a party of 80 men, mostly merchants and their merchandise-loaded wagons, between May 8 and May 10, 1841. This yearly caravan to Santa Fe followed the Santa Fe Trail on to the Arkansas River in Kansas. It now appears evident that this particular caravan did not proceed to Fort Bent near the Colorado Rockies and then south over Raton Pass. Instead, it took the Cimmaron Cut-Off, leaving the Arkansas River in western Kansas, cutting across the panhandle of Oklahoma into northeastern New Mexico, probably encountering the mountains proper near Wagon Mound, passing on to Las Vegas and thence to Santa Fe. The basis for suggesting this change in route is a letter which was published in Niles’ National Register (vol. 61, p. 1575, 1841). This anonymous letter, dated July 1841, was written by a man who joined the annual Santa Fe caravan just before the crossing of the Arkansas River in Kansas. That he and Gambel were members of the same party from that time on is borne out by practically identical descrip- tions of two Indian encounters in letters of the two men to people in LIZARD CNEMIDOPHORUS PERPLEXUS—MASLIN, ET AL. 337 the East. Information in the Niles’ National Register letter makes it possible to designate the arrival date at Santa Fe as July 2, 1841. Gambel was in the vicinity of Santa Fe, along the Rio Grande and in the nearby mountains, for the period from July 2 to about Sep- tember 1. He made representative collections of plants, especially from the ‘sandy hills along the borders of the Rio del Norte, Santa Fe (Mexico).’”’ The Rio Grande River lies about 20 miles west of Santa Fe and today is generally inaccessible by highway in this area. Dur- ing Gambel’s visit, however, the main road from the east crossed Santa Fe Creek near the town and continued to the Rio Grande along the south side of this creek, thence turning south towards Albu- querque. It would seem justifiable to assume that Gambel gained access to the Rio Grande collecting localities by means of this road. Thus, specimens from “‘Rio del Norte, Santa Fe’’ presumably were col- lected south of the éntrance of Santa Fe Creek into the Rio Grande River, west and southwest of Santa Fe. None of the plant collections or bird observations warrant suspect- ing that Gambel went far south of Santa Fe; but one of the species of reptiles he collected suggests that he might have. This species, Holbrookia texana, extends up the Rio Grande Valley as far as Valencia County (4 miles north of Sabinal). It might even extend as far north as Albuquerque; but the likelihood of its occurring in Santa Fe is questionable. Although he makes comments about birds at Taos, it seems unlikely that Gambel went that far north. One of his com- ments concerns wintering juncos, and certainly he was not there during the winter. He probably learned about Taos birds from people in Santa Fe. On or within a day or so after September 1, Gambel departed from New Mexico, supposedly from Abiquiu, with John Workman’s party for California. This party is reported to have traveled northwest from New Mexico across southwestern Colorado, over the Colorado and Green Rivers in eastern Utah in September, into the mountains south of Salt Lake, then southwest across mountains and desert to southern California. There is some variance in arrival dates in Cali- fornia between the Workman party and Gambel, the former arriving in early November and the latter ‘‘the last of November being three month’s traveling over Rocky Mountains & barren deserts... .’’ The difference in arrival dates may simply mean that Gambel took time out for exploration on his own or with a small detached party some- where en route. In considering the most likely localities from which Gambel might have made his collections of reptiles on this first California expedition, the valley of the Rio Grande River southwest of Santa Fe seems the most probable choice, especially with respect to Cnemidophorus per- plexus. Inasmuch as Gambel’s type specimen was a female contain- 338 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 ing at least one large ovarian egg, the date it was collected probably fell somewhere between July 25 and not much later than August 7. During this period Gambel was collecting a number of plants near the river and, in the course of such collecting, had, like modern-day botanists with herpetological leanings, ample opportunity to collect lizards as well. Assuming that Gambel did collect C. perplezus on the Rio Grande River near Santa Fe, he may or may not have shipped it home before leaving for California. The fall express for Independence left Santa Fe shortly after July 25, on which date Gambel wrote a letter (Penn- sylvania Historical Society collection) home to his mother in Phila- delphia. Nuttall had urged that Gambel dispatch his plants to George Engelmann of St. Louis, who was to send them through a John H. Barnard on to Nuttall in Philadelphia. Nuttall (letter, Nuttall to Engelmann, Nov. 3, 1841; Missouri Botanical Gardens) received a letter from Gambel at Santa Fe stating that he mtended to send to Nuttall via Engelmann “a part of the collections he had made up to that place by another party returning to St. Louis in October.” There is nothing, however, among the Engelmann cor- respondence at the Missouri Botanical Gardens to suggest that Hn- gelmann ever received Gambel’s shipments from Santa Fe, either from the annual express or from an October party. If shipments were made, they must eventually have reached Philadelphia and the Academy of Natural Sciences but undoubtedly after Nuttall had de- parted for England at the end of 1841. Thus they probably would have remained in storage at the Academy. It seems logical that Gambel would ship east anything collected to this pomt, simce the next shipment point would have to be from California, over a thou- sand miles away across difficult country. But apparently no such shipment was made. Judging from the gravid condition of the type of Cnemidophorus perplerus the chances of its having been collected before the fall express left for Independence is very slight, It appears, then, as though Gambel carried all of his collections to California. How the specimens eventually reached the Smithsonian Institution is as obscure as where they were originally collected. In the Seventh Annual Report of the Smithsonian Institution (1853, p. 55) there is a notation under the section on new 1852 reptile accessions, ‘“‘A small number of specimens procured by Dr. Gambel, in the same country, has also come into the possession of the Institution.’ There is, un- fortunately, no further information on the accession. The accessions immediately preceding this entry were of specimens from California. It appears from this entry and Baird and Girard’s (1852) locality no- tations that Gambel must have shipped all of his herpetological col- lections from California and that the original recipient of these col- lections, whoever he may have been, simply assumed that it was there LIZARD CNEMIDOPHORUS PERPLEXUS—MASLIN, ET AL. 339 that they had been collected. The possible ways by which the Smithsonian Institution could have acquired them are many. They might have been donated through Gambel’s wife directly or through her brother, might have been sold to someone by Gambel’s wife, and subsequently donated, might have been received from the Academy of Natural Sciences, purchased by some Smithsonian agent such as John Kirk Townsend, a friend of Gambel’s, etc. How the other specimens of C. perplexus (so identified at that time) got into Baird’s hands at the Smithsonian is an easier problem. General Churchill was a close personal friend of the Bairds in Carlisle, Pa., and indeed became Baird’s father-in-law. Colonel Graham’s collections were actually made for him under orders by John H. Clark, and Clark had been a student of Baird’s at Carlisle College. Rowan Kennerly, who also collected one of the early, but not syntypic, USNM specimens of C. perplexus, had also been one of Baird’s students. In summary, the type of Cnemidophorus perplezus was undoubtedly not collected in “Calif.,”’? in ‘“W. Texas,” or during Gambel’s “last journey to California”; but was probably collected sometime during the last week of July 1841 during Gambel’s first journey to California. The type locality is probably the valley of the Rio Grande in Sandoval County, N. Mex., in the vicinity southwest of Santa Fe. Description: We are now faced with a paradox, namely the identification of a type. Cope’s (1900, p. 573-574) description of the specimen for the most part is accurate. The specimen is in fair condition but rather soft, and as Burt (1931, p. 122) points out the tail is now incomplete. Cope’s (loc. cit.) measurements, therefore, are probably as accurate as any which can now be made. Our own measurements agree essentially with his. USNM 3060, mature female, possessing nearly mature ovarian eggs. Total length, 260 mm. (according to Cope); length of head and body, 86 mm.; length of head to posterior edge of auditory meatus, 20.7 mm.; length of head to posterior face of jaw articulation, 20.3 mm.; length of forearm from axilla, 27 mm.; length of hind leg from inguen, 58 mm.; width of head, 11.6 mm. Interparietal twice as long as wide, narrower than parietals; a pair of frontoparietals; third and fourth supraoculars completely separated from fronto- parietals by a series of small scales; first and second supraoculars broadly in contact with frontal; five scales in both anterior and posterior occipital rows, anterior nasals 1-1; posterior nasals 1-1; loreals 1-1; preoculars 1-1; suboculars 4—3; frenoculars 0-0; supra- labials 6-6, counting first scale in contact with last subocular; infra- labials 7-6; chinshields 5-5, first pair in contact throughout their length; chinshields separated posteriorly from infralabials by 4-4 340 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 scales, these preceded by a few smaller granules. Scales of central intermandibular area slightly larger than adjacent scales, inter- mandibular scales gradually decreasing in size posteriorly; anterior gular scales about half diameter of lateral intermandibular scales; gular fold with medial, anteriorly directed open notch; fold bordered by small scales, medially small scales form a triangle, apex forward, anterior to notch, this triangle separating two patches of enlarged scales; these enlarged gular scales smaller than enlarged median intermandibular scales. Anterodorsal brachial scutes in four rows; anterodorsal antebrachial scutes in three rows; post-antebrachial scales small with central strip of very slightly enlarged scales; antero- dorsal femoral scutes in seven rows; anterodorsal tibial scutes in three rows. Three enlarged anal plates consisting of a pair separated from cloaca by four rows of small scales and preceded by an enlarged median scale. Femoral pores 19-19, median two separated by four scales. There are 178 vertebral scales counting from but not including enlarged occipitals to first row of enlarged scales at base of tail; 73 scale rows at region of greatest girth excluding ventral plates; 25 scales around base of tail. Color pattern. Median vertebral light stripe as distinct and as wide as paravertebral light stripes, commencing as a pair of light spots on occipital scales, extending posteriorly to first row of enlarged scales of tail, posterior three-fifths of line undulant or wavy; width of vertebral light stripe, paravertebral dark and light stripes about equal; paravertebral light stripes separated by 12 rows of scales (average of 10 counts) at midbody; paravertebral (first) dark stripes or fields darker than remaining ones; toward center of body vague indication of one or two light spots in these dark fields; second, third and fourth dark stripes successively wider, fourth stripe more than twice as wide as any light stripe which are all subequal in width. Second and third dark stripes each with a very vague indistinct median row of large light spots, somewhat confluent, difficult to count, approximately 12 or 13 spots in each field. Fourth dark stripes with one or two extremely vague, large, diffuse, light spots. Ventrum immaculate white. Comparisons: With this description in mind the type can now be compared with the species of Cnemidophorus that Gambel might have encountered from Missouri to California. These species are as fol- lows: tesselatus, tigris, sexlineatus, inornatus, neomexicanus, velox, sacki, and hyperythrus. The tesselated forms tesselatus and tigris may be dropped from the list immediately on the basis of color pattern alone. The species hyperythrus occasionally has a median light stripe but it has only a single frontoparietal scale and its distribution is such that it is the least likely to have been encountered by Gambel. LIZARD CNEMIDOPHORUS PERPLEXUS—MASLIN, ET AL. 34] Of the remaining forms sezlineatus can also be removed from the list on several counts. In this species there is never a median light line as bright or as narrow as the lateral light lines; the third and fourth supraocular scales are in contact with the frontoparietal; the posterior antebrachial scales are definitely though not abruptly enlarged; there are seldom more than 15-15 femoral pores; the intermandibular scales are larger and abruptly demarked by a sudden decrease in size from the small anterior gular scales; the gular fold is not notched, nor are the enlarged gular fold scales interrupted medially by a patch of smaller scales; the enlarged gular fold scales are larger than the median inter- mandibular scales. Furthermore, the species does not occur in the vicinity of Santa Fe, N. Mex. There is no question of doubt that —sexlineatus and perplerus are not the same. The remaining species all occur in the vicinity of Santa Fe, and it is these species which must be examined most critically. Of these four species perplerus resembles sacki the least. In this latter form there is no conspicuous median light stripe; the post-antebrachial scales are abruptly and greatly enlarged; the intermandibular scales are larger and abruptly demarked from the smaller anterior gular scales along a transverse line; the gular fold is unnotched and bordered by a row of enlarged, scutelike scales much larger than the median intermandibular scales and uninterrupted by a triangular patch of smaller scales. The paravertebral light stripes are separated by only about five scales; the second and, particularly, third dark bands are spotted by numerous light spots alternating from side to side of each band; the fourth and lowest band also contains numerous light spots. C. inornatus of New Mexico and Arizona differs from perplexus in that it lacks spots in the dark fields; has about eight scales between the paravertebral light stripes; is much smaller than the type; has larger post-antebrachial scales; the intermandibular scales are much larger, relatively few in number, and abruptly differentiated from the smaller anterior gular scales by one or two transverse rows of still smaller, granular, median scales; the gular fold is unnotched and the enlarged scales of the fold are larger than the median intermandibular scales; there is no patch of smaller scales interrupting the row of enlarged gular fold scales; temporal scales larger and fewer in number; femoral pores fewer. In spite of the conspicuous differences in scale counts and size, C. nornatus in the northwestern part of its range and C. velox resemble each more than they resemble any other species of Cnemidophorus. Lowe (1955) has discussed the relationship of these two species in a study demonstrating the validity of C. velox. While he did not em- phasize the fact that velox and inornatus are nowhere known to be sympatric, he pointed out that they exhibit a marked difference in 342 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 habitat and geographic distribution. Moreover, both species are relatively rare in those areas where their ranges approach one another. It is possible that C. velox is a direct derivative of C. inornatus. As far as we are aware, no definitely identified specimens of velox have been reported from Santa Fe, but the species has been collected in New Mexico at Taos, Taos County; 5 miles south of Youngsville, Rio Arriba County, a site some 50 miles northwest of Santa Fe; and 2 miles west of Sands, San Miguel County, some 30 miles southeast of Santa Fe. That the species will ultimately be collected near Santa Fe seems inevitable, for it is a member of similar pinyon- juniper communities not far distant and has been taken both to the north and south of this locality. C. velox differs from perplexus in that the median stripe if present and complete is indistinct, lighter in color and narrower than the paravertebral light stripes and never undulant; the paravertebral — light stripes are separated by about eight scales; there are no spots in the dark fields; the post-antebrachial scales are larger than those of perplezus but not as large as those of inornatus; the intermandibular scales are larger and abruptly demarked from the anterior gular scales along a transverse line (in an occasional specimen this is not marked) ; the largest scales on the gular fold are usually larger but often equal in size to the largest median intermandibular scales; the gular fold is unnotched and the enlarged scales along it are not interrupted medially by a patch of smaller scales; the temporal scales are slightly larger; the third supraoculars are in broad contact with the frontoparietals. C. neomexicanus has not yet been taken as near to Santa Fe as has velox. The nearest known locality for neomexicanus is 6 miles south of Bernalillo, Sandoval County, near the Rio Grande and about 50 miles southwest of Santa Fe. However, this locality more closely approaches the probable type locality for perplexus than does any in the presently known distribution of veloz. C. neomexicanus differs from perplexus in only a few characters, but these differences are for the most part concerned with variable characters and fall within the range of variation described by Lowe and Zweifel (1952). The chief and most important difference lies in the fact that in most specimens of neomexicanus the fourth, third, and often the second supraoculars are separated from the median head plates by small scales (circumorbitals). In the type of perplexus only the fourth supraocular and three-fourths of the third are so bordered. Another marked difference is in size. Of the 48 specimens available to Lowe and Zweifel (1952), all from Socorro County, N. Mex., the largest is a female measuring 76.4 mm. from snout to vent. The type of perplexus, also a female, measures 86 mm. In all other characters the two forms are strikingly similar. LIZARD CNEMIDOPHORUS PERPLEXUS—MASLIN, ET AL. 3483 The color patterns and the arrangement and size of the brachial, antebrachial, and post-autebrachial scales are identical; neomezxicanus usually has a peculiar, anteriorly directed open notch on the gular fold as does perplexus; the scales of the gular fold are only moderately and gradually enlarged anteriorly as they are in perplerus and their maximum size is less than or equal to the largest median intermandib- ular scales. The enlarged gular scales are occasionally interrupted medially, as in the type of neomezxicanus, by a patch of small scales. This condition closely approximates the condition of the type of perplexus. The intermandibular scales are larger than the anterior gular scales as they are in perplerus and usually are not sharply de- marked from them; however, on either side a few small scales, arranged in a short transverse row, often separate the two areas laterally in neomexicanus, but not medially. The arrangement and size of the temporal scales are as similar in the two forms as this type of scalation can be. The number of scales from the occipital region to the rump in the type of perplexus is 178, this is within six scales of the average given by Lowe and Zweifel (1952) for neomexicanus, namely 184.3+ 1.2; and the number of circumabdominal scales exclusive of the ventral plates is 73, two sceles less than the mean of neomezxicanus which is 74.9+0.62. In spite of the differences in body size and the extent of the anterior extension of the circumorbital semicircles we are convinced at this time that neomexicanus and perplerus are one and the same. It is appropriate to point out that this finding—the first proper allocation of the old name Cnemidophorus perplexus to a definitely known population—has been made possible through the detailed study of variation by Lowe and Zweifel (1952), where for the first time the body scale counts were determined and analyzed statistically in a study of variation in the genus Cnemidophorus. The remaining specimens upon which the description of perplexus was based have been examined and may be identified as follows. The five specimens (USN M 3022) collected by John H. Clark in the valley of the Rio San Pedro of the Rio Grande del Norte are all Cnemi- dophorus sacki gularis Baird and Girard. The two specimens of USNM 3050, ‘collected’ by General Churchill on the Rio Grande west of San Antonio, are both Cnemidophorus inornatus Baird. The second specimen, collected by Gambel and originally bearing the same num- ber as the type (USNM 3060), is now numbered USNM 30885. This is also a specimen of Cnemidophorus inornatus. For the most part descriptions of specimens of perplexus in the literature are so brief as to make identification virtually impossible. Furthermore, so many lined whiptails have been confused with each other that all of the earlier, more extended accounts of these forms, 344 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 when based on series, almost inevitably are based on more than one species. The synonomy given below is, then, tentative, and includes only those references which seem very likely to pertain to: Cnemidophorus perplexus Baird and Girard Cnemidophorus perplexus Baird and Girard, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 128, 1852; Appendix F, Reptiles, in Exploration of the Red River of Louisiana, in... 1852, p. 239, 1853 (reference to 1852 description).— Cope, U. S. Nat. Mus. Bull. 1, p. 46, 1875 (listed) —Gadow, Proc. Zool. Soe. London, p. 368, 1906 (Bernalillo County, N. Mex.; brief description).— Van Denburgh, Proc. California Acad. Sci., ser. 4, vol. 13, p. 211, 1924 (record from Santa Fe (San Ildefonso), N. Mex.; possible).—Burger, Chicago Acad. Sci., Nat. Hist. Mise., No. 65, p. 3, 1950 (diagnosis based on type). Cnemidophorus octolineatus, Yarrow, Report upon the collections of batrachians and reptiles, in Geographical and geological explorations and surveys west of the 100th meridian, vol. 5, chap. 4, p. 558, 1875 (San Ildefonso, - N. Mex.; possible). Cnemidophorus tesselatus perplerus, Cope, Amer. Nat., vol. 26, pl. 18, fig. B, 1892 (dorsal color pattern, possibly of type of perplexus); Trans. Amer. Philos. Soc., vol. 17, pp. 34-35, 1893 (redescribes type specimen; pl. 6, figs. a-@ not of perplexus; p. 12, fig. B, possibly of perplerus); in Ann. Rep. U. S. Nat. Mus. 1898, pp. 573-575, 1900 (figure 105 not of perplerus; redescription of type). Cnemidophorus sexlineatus perplexus, Burt, U. S. Nat. Mus. Bull. 154, pp. 122, 125, 1931 (designates lectotype, gives description). Cnemidophorus sexlineatus, Smith and Taylor, U. 8. Nat. Mus. Bull. 199, p. 185, 1950 (cites perplexus as synonym; specifies type, type locality unknown). Cnemidophorus perplexus perplexus, Smith and Taylor, Univ. Kansas Sci. Bull. 33, p. 363, 1950 (lists species as valid, identifies Rio San Pedro as Devils River). Cnemidophorus neomexicanus Lowe and Zweifel, Bull. Chicago Acad. Sci., vol. 9, pp. 230-247, fig. 1c, pl. 1, fig. a, 1952 (type, Mus. Vert. Zool. Univ. Cali- fornia No. 55807, McDonald Ranch Headquarters, 4,800 ft., 8.7 miles west and 22.8 miles south of New Bingham Post Office, Socorro County, N. Mex., Charles H. Lowe, Jr., collector; detailed description, comparisons, and ecology). Lectroryre: USNM 3060, adult female, type designation by Burt (1931, p. 122). Tyrer Locauity: Herewith restricted to valley of the Rio Grande in Sandoval County, N. Mex., in the vicinity southwest of Santa Fe. Type collected by Dr. William Gambel on his first journey to Cali- fornia, near the end of July 1841. Summary The type of Cnemidophorus perplexus was selected by Cope but not clearly designated as such. Burt (1931) followed the procedures recommended by the 13th and 14th International Congresses of Zoology and properly designated USNM 3060 as the lectotype of this species. The specimen was collected by William Gambel, probably on his first journey to California near the end of July 1841. The type LIZARD CNEMIDOPHORUS PERPLEXUS—MASLIN, ET AL. 345 locality is probably the valley of the Rio Grande in Sandoval County, N. Mex., in the vicinity southwest of Santa Fe. As this locality is geographically and ecologically apart from the area formerly recorded as being occupied by this species, extensive comparisons of C. perplexus with all the species living in the region of Santa Fe had not been made. Of the various species that are now known to occur here it is evident that one of them, C. neomexicanus Lowe and Zweifel, is conspecific with C. perplerus. This name, therefore, must be considered as a synonym of Cnemidophorus perplexus. Literature Cited Barrp, 8S. F., AND Grrarp, C. . 1852. Characteristics of some new reptiles in the museum of the Smithsonian Institution. Proc. Acad. Nat. Sci. Philadelphia, vol. 6, pp. 125- 129. Burcer, W. LEsLie 1950. New, revived, and reallocated names for North American whiptailed lizards, genus Cnemidophorus. Chicago Acad. Sci., Nat. Hist. Misc. No. 65, pp. 1-9. Burt, CHARLES E. 1931. A study of the teiid lizards of the genus Cnemidophorus with special reference to their phylogentic relationships. U. 8. Nat. Mus. Bull. 154, viii+286 pp. Cops, E. D. 1892. Parallel color-patterns in lizards. Amer. Nat., vol. 26, p. 522. 1893. A synopsis of the species of the teid genus Cnemidophorus. Trans. Amer. Philos. Soc., vol. 17, pp. 27-52. 1900. The Crocodilians, lizards, and snakes of North America. Ann. Rep. U.S. Nat. Mus. (1898), pp. 153-1270. Lowe, C. H., Jr. 1955. A new species of whiptailed lizard (genus Cnemidophorus) from the Colorado Plateau of Arizona, New Mexico, Colorado, and Utah. Mus. Comp. Zool., Breviora, vol. 47, pp. 1-7. Lowe, C. H., Jn., anD ZWEIFEL, R. G. 1952. A new species of whiptailed lizard (genus Cnemidophorus) from New Mexico. Bull. Chicago Acad. Sci., vol. 9, pp. 229-247. Osporn, H. F. 1931. Cope: Master naturalist. xvi+740 pp. SmiruH, H. M., anp Taytor, E. H. 1950. Type localities of Mexican reptiles and amphibians. Univ. Kansas Sci. Bull. 33, pp. 313-380. Yarrow, H. C. 1882, Check list North American Reptilia and Batrachia with catalogue of specimens in U. S. National Museum. U.S. Nat. Mus. Bull. 24, 249 pp. U. S. 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Rev iocy4 Arup PONT a wR ty HotaOMe re ee ae yee Se detable aR LGN iit “ate siahect tes 2M ernie Stay fro Oth" Ceara tbtied sits tions ‘by guil fasta: ahve We Midd, ge ee ‘Eohe atgeg gt hte Ok Eee geod's, Efe re 2 z SOM CMY t at ee pe an hehe ay 49 < PA freaks eh ¢ a m, esl OPN ee arr agree oR eT) sits daw ini non Le ae ori : ey vi ee. he ear ati ave wickom steel Sissi (GRR tel onl dee VBR I tenon aA EE bows 7d ORY Reg enemas, ONS: rhenl g hah a 4 ph une and dest ty Sime ene ‘soak Nat see Lore uy baa singed ‘eign ae Spor apart: ieee Pinto i) apa o a Ca WAR irene © Auer 98 ue aa at > _ i i jie eae hey Pr is ‘i iN a a a PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM issued 4 SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1959 No. 3407 SYNOPSIS OF THE SPECIES OF AGROMYZID LEAF MINERS DESCRIBED FROM NORTH AMERICA? By Kennetsa E. Frick’ Introduction A total of 206 described North American species are dealt with in this paper. All are separated by illustrated keys and briefly character- ized in short descriptions that include the known larval host plants and distributions. Although I am aware of some undescribed species in each of the large genera, I have attempted to define and illustrate each of the previously described species so that new species can be recognized as distinct. The objective of this paper is to lay a firm foundation upon which students of the agromyzid leaf miners may build. Originally only Nearctic species were to be considered. However, the types of the described West Indian and Central American species are in North American museums except for a few deposited in the British Museum (Natural History). Therefore, it was considered desirable to include the Neotropical area of North America not only because the types were available but because of the number of requests received by U. S. National Museum (USNM) personnel for identifica- tions of leaf miners from the southern extremities of North America. 1 Scientific Paper No. 1639, Washington Agricultural Experiment Stations, Pullman, Wash.; Project No 1260. Investigation supported by a grant-in-aid from the National Science Foundation. 2 Irrigation Experiment Station, Prosser, Wash. 347 348 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The North American species are distributed throughout 15 genera and 11 subgenera. Except for Xyracomyia and possibly Haplomyza, all are Holarctic. Several very small genera, mostly monotypic, are not found in North America. These are the Neotropical Triticomyza Blanchard and the Palaearctic Carinagromyza Sasakawa, Gymno- phytomyza Hendel, Ptochomyza Hering, Selachops Wahlberg, and Xeniomyza de Meijere. Each genus is redescribed and separated by the use of illustrated keys giving full consideration to the unusual species peculiar to North America. These unusual species are dis- cussed under their respective genera. The similarities and differences between the faunas of North America and the Palaearctic region, the two best collected regions in the world, are also taken up under each genus. It might be noted here, however, that the northern Neotropical region, poorly collected though it is, has contributed more than its proportionate number of extraordinary species. Among these are the reddish lunule and lower frontal vitta of Melanagromyza diadema, the white spot on the halter of Ophiomyia punctohalterata, the yellow scutellum of Phytobia (Phytobia) picta, the pictured wing of P. (P.) kallima, the yellow third antennal segment of P. (Calycomyza) meridiana, the distally narrowly yellow femora of P. (C.) cassiae, P. (C.) ipomoeae, and P. (C.) lantanae, the enlarged third antennal segment of males of Liriomyza commelinae, and the very large (3.5 to 4 mm. in wing length) South American L. braziliensis Frost and L. ecuadorensis Frost that have posterolateral setae medially on the midtibia and have larvae that mine the tubers and sometimes the stems of Solanum tuberosum Linnaeus. Two species groups that are all or nearly all Neotropical have representatives in the southern Palaearctic region also. The first of these is the viridula group in the genus Agromyza. There are four northern Neotropical species and one from eastern United States. There appear to be two Palaearctic species: Agromyza salicifolu Collin does not occur north of Israel, and A. quercus Sasakawa occurs in Japan. A similar situation is found in Melanagromyza. ‘There are five Neotropical representatives in the pulicaria group. The two Palaearctic species occur in the southern extremities of that region and one, M. inaequabilis Hendel, is not found as far north as Europe while the distribution of M. pulicaria Meigen includes Europe. The morphological characters used in the keys and descriptions were described by Frick (1952). The only change made in that system is the renumbering of the dorsocentral setae. Hendel (1931) numbered from the posterior of the mesonotum forward and Frick (1952) ac- cepted that arrangement. However, in 1953 Frick (1953b) rejected that system and counted from the anterior to the posterior or scutel- AGROMYZID LEAF MINERS—FRICK 349 lum. The mesonotum and its setae were later illustrated (Frick, 1956a). Because complete synonymies have been given previously (Hendel, 1931-1936; Frick, 1952, 1953a, 1956b, 1957a), only North American synonyms for genera, subgenera, and species are listed herein. The most recent paper (Frick, 1957a) gives all but two name changes not previously published so that the correct binomina would be avail- able for this synopsis. The two names not included are Liriomyza reverberata (Malloch) and L. sorosis (Williston); the changes are dis- cussed in full herein. The locations of those holotypes not mentioned in the original descriptions or in other papers are given herein. A number of types of Frost’s species had been retained in his personal collection and these he recently transferred to the U. S. National Museum. The references are limited to the original descriptions and to those papers that have illustrations or good redescriptions of previously poorly defined species. ‘The citations are not intended to be complete or to give all nomenclatural changes because those have been pre- viously published (Frick, 1952, 1953a, 1957a). Those papers cited in the text by author and date only are given in full in the list of references cited (p. 445). ACKNOWLEDGMENTS I am indebted to the following museum staff members for their kindness, cooperation, and helpfulness in locating types and literature and for providing study facilities during the course of this study: H. H. Ross, Illinois Natural History Survey, Urbana; R. H. Foote, Alan Stone, and C. W. Sabrosky, U. S. National Museum, Washing- ton, D. C.; C. H. Curran, American Museum of Natural History, New York; J. A. Wilcox and D. L. Collins, New York State Museum, Albany; P. J. Darlington, Jr., Museum of Comparative Zoology, Cambridge, Massachusetts; G. E. Shewell, Canadian National Col- lection, Ottawa; S. W. Frost, personal collection at Pennsylvania State University, University Park; R. E. Beer and W. E. LaBerge, Snow Entomological Museum, Lawrence, Kans.; and A. L. Melander, personal collection at Riverside, Calif. Without the cooperation of certain staff members of European museums, this study would be only partially complete. They lent type material or gave information on types, and I am deeply grateful to each of the following: E. M. Hering, Zoologisches Museum, Berlin; H. Oldroyd, British Museum (Natural History), London; René Malaise, Naturhistoriska Riksmuseum, Stockholm; S. L. Tuxen, Universitetets Zoologiske Museum, Copenhagen; A. Diakonoff, Rijksmuseum van Natuurlijke Historie, Leiden; Max Beier, Natur- 350 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 historsches Museum, Vienna; and Harald Schweiger, Niederéster- reichischen Landesmuseum, Vienna. So many individuals kindly lent specimens from various parts of the United States that space does not permit thanking each personally. I am thankful for reared and authentically identified European specimens for my collection from E. M. Hering, Arne Lundqvist, and K. A. Spencer. Marion Ownbey, Department of Botany, State College of Washington, kindly helped to bring the names of larval host plants up to date. Key to the genera of Agromyzidae found in North America 1. Subcosta developed throughout its length, coalescing with vein R; before reaching costa; vein R; somewhat broadened at union with the costa (fig. 1) (Agromyzinae) .. . hie > ; 2 Subcosta becoming a fold Gistally anal eneinen in nieoete a parately andl basad of vein R;; R; not broadened at its union with the costa (fig. 2) cape myzinae). . spaeaeeh Gi cuss 5 2. Halter usually all Blauk, roy ih a Sornite spot on neh! cereus : 3 Halter white or yellowish ..... BURST SEP Aged 3. Carina between antennae prominent, Gaaitennts or homnaphencen antennae somewhat divergent (fig. 45) ..... sfoat vat Carina absent, flattened between antennae even anowent bases made separat- ed; antennae anteriorly directed. . . . .... . . .2. Melanagromyza 4. Carina fusiform (fig. 45); orbital setulae erect or reclinate (fig. 47). 3. Ophiomyia Carina hemispherical; orbital setulae proclinate (fig. 55). . . .4. Tylomyza 5. Scutellum with four setae (fig. 99). . ......4... 6 Scutellum with two setae (fig. 95)... .. aff Ol cde nies Carcdanths 6. Orbital setulae erect or reclinate (fig. 133), aomcuimes weakly SSP er or absentee «2. Bhs, EY Oncaea ae 7 Orbital setulae eroclinnie! ell developed (fig. 141). Sais ihe hoon oa “12 7. Wing with costa reaching to end of vein M442, or (Phytobia) if only to Rass then ultimate section of Ma3+4 eas to the penultimate section (fig. Od) eae, a ehh ao eee Wing with costa reaching o ond of vein Byes: niltinaate eociian of Mas, at least twice as long as peer a section when crossvein m-m present (Git as IS): ) eee Sete Ue OE 8. Scutellum yellow, at leant centrally: vein Misa gaaing nearest wing tip (fig. 1118), hice Maer res Rib AS hed Scutellum usually dank aaa coucoloraus wale neahnonane if yellow vein Ry+5 ending nearest wing tip (fig. 60). .... . . .. . 5. Phytobia 9. Genovertical plates narrow, usually slightly but not abruptly raised above the plane of the frontal vitta; if abruptly raised then genovertical plates narrow except dorsad of antennae (fig. 104). . . . . ke ere eek O Genovertical plates broad, about one-third the width a the! frons and very abruptly raised above the plane of the frontal vittae (fig. 128). 8. Metopomyza 10. Mesonotum dull black, heavily gray pollinose; two sparse rows of acrostichal setae; crossvein m-m absent; only one upper-orbital seta, reclinate and outwardly directed (fig. 129). . ........... . 9. Haplomyza AGROMYZID LEAF MINERS—FRICK 351 Mesonotum shining black and yellow, or if gray pollinose then either four rows of acrostichals, or crossvein m-m a or two upper-orbital setae present ..... ... . .% Liriomyza 11. Crossvein m-m, when recent, never ‘basal ios crossvein r-m; basal section of Ms+4 present (fig. 134) ..... . ... . . 12. Phytagromyza Crossvein m-m basal to crossvein r-m and Appears to be the basal section of vein M344, which is absent (fig. 135). . . . . . . . 13. Pseudonapomyza 12. Costa reaching to the end of vein Mj+2; halter with a dark spot on knob. . 13 Costa reaching to the end of vein Ry+5; halter white or yellowish. . . . . 14 13. Wing with axillary lobe and calypter well developed; mesonotum with acrostichal setae present. ... . . .. . . 10. Phytoliriomyza Wing with axillary lobe very small nnd Galepter virtually absent; acrostichal setae absent. «26s sete eles la .... . LI. Xyraeomyia 14. Wing with crossvein m-m resent. oh sw ss wile eos » “he Napomyza Wing with crossveinm-mabsent .......... .. 15. Phytomyza Systematic Treatment 1. Genus Agromyza Fallén Agromyza Fallén, Specimen entomologicum novam Diptera disponendi methodum exhibens, p. 21 (No. 66), 1810 (Dissertat. Lund.). Domomyza Rondani, Dipt. Ital. Prodr., vol. 1, p. 121, 1856. This is a moderately large genus of 22 described species that has the subcosta complete and uniting with vein R, at the costa, which is ex- panded at that point (fig. 1), and the halteres white. The only species that I have seen that has the union of the subcosta and vein R, with the costa atypical in some specimens is Agromyza aristata. However, most of those examined have the typical costal union (fig. 4). The most diverse types have the subcosta about one-fourth incomplete and R, distinctly bent but not expanded near the costa. There are no diverse species that deviate from the Palaearctic pattern. The great majority of North American species have only two or three well-developed dorsocentrals with none to several small ones. Only four species have the usual arrangement of one strong presutural and three strong postsutural dorsocentrals. In the Palae- arctic region there are proportionately more species with the latter arrangement. As for the species groups, there are representatives in the rwbi group (couplets 3-5) but none in the cinerascens group, which contains three European species. The correlation between shining mesonota and grass mining larvae (ambigua group) holds true in North America as in the Palaearctic region. Of the six known North American species (couplet 14), four have larvae that mine the leaves of grasses. The alternative (couplets 7-13) contains eight species that have the mesonotum subshining to dull and larvae that mine in dicotyledons. The host plants are known for four. One of the more unusual features of the North American fauna is 302 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 the five species keying out at couplet 10 (viridula group). These are large dark species that may have a metallic sheen to the mesonotum and abdomen. Of the five, four are Neotropical, which seems to ac- count for the higher proportion in North America. There are two species in the Palaearctic region. A. salicifolii Collin is not found north of Israel and A. quercus Sasakawa occurs in Japan. Key to the described species of Agromyza 1. Mesonotum with two or three well-developed postsutural dorsocentrals plu, none to several small postsuturals; a presutural dorcocentral, when presents small and posterior to the presutural seta. ... beh: SPN OES ice Mesonotum with four strong well-developed dorsoceuirnlet one presutural and three postsutural, the a at least slightly anterior to the presutural seta. ... ace elO 2. Vein Myi4. at or nearest the wing ip inidtibia’ medially without postero- lateral setae ... . ood oho Vein R445 nearest the aa hans enrdutbia waediathe. wali one or 8 postero- lateral setae ... . 4 bedi gaa 3. Mesonotum dull, grayish pollinose a fesse res ele ievcloned postsutural dorsocentrals.... . ONE se see Mesonotum = shining black: tino _ well-developed oat dorsocen- trals 267, es" . . . varifrons 4, Femora, tibiae, and fori eens vein Miss coding onlay aletitly nearer the wing tip than R445 (figs. 4, 17) . Phases. 2 LAs tthe kone Femora, tibiae, and tarsi dark brown to black, pxccnt or foretibia being yellow distally; vein Mi+. ending at wingtip... . 3 a yes oe UD 5. Body with all setae yellowish; dorsal upper-orbital sciae Uren black ex- tending from vertex (fig. 14)..... . . . pallidiseta Body with all setae black; dorsal cpper orb ial arsine from yellow (fig. 5). aristata 6. Mesonotum and scutellum subshining to dull black, somewhat grayish pollinose, with or without a greenish sheen; larvae mine dicotyledons . . 7 Mesonotum and scutellum shining black; larvae mine grasses. .... 14 7. Thorax and abdomen yellow or light brown; femora mostly yellow. . . 8 Thorax and abdomen very dark brown or black, with or without a greenish sheen; femora jpredominately: dark, brown’ &.2acciu ese) © Pepe leneerp erred 8. Third antennal segment and maxillary palpus yellow. . ... . . diversa Third antennal segment and maxillary palpus brown . . .. . canadensis 9. ‘Tibiae and tarsi. dark browm: Or bach 3. os soe, ens: veh nons ene ee ance LO Tibiae and tarsi yellowish ..... .. . . reptans 10. Mesonotum, scutellum, and abdomen vith a erecsish eheen shea toepell Mesonotum, scutellum, and abdomen without a aaa sheen, subshining or dull brown or black... . 5 yin op oar ace fabian cael 11. Upper-orbital setae subequal in fenceh (fiz. 20) . Sih iste omgie . . . Viridula Upper-orbital setae unequal in length, the ventral about i 5 times as long asthe dorsal *(fipsO) 210.9 See? \ REPT Ss Gate, ¢eamaequatis 12. Calypter with margin and fringe white sit. dh ny s.wadalosrest hwo wanes Calypter with margin and fringe yellow. ....... . . centrosemae 13. Anepisternum entirely dark brown or black .......... . frosti Anepisternum with a narrow yellow dorsal margin. . . ... . . currani AGROMYZID LEAF MINERS—FRICK 353 14, Costa reaching to the end of vein Miz. . 2. 1. 2... 7 2 ee ee ee 15 Costa reaching to the end of vein Rays . . Sue elisha’ vs) Trappe 15. Calypter with fringe light brown; frons ioaeen tian wide . . . parvicornis Calypter with fringe white; frons wider than long. . . . . . . nigripes 16. Anterior half of mesonotum with acrostichal setae in six rather regular TOWS se ese Nera ne cae ee Anterior half of mieconotene witli RereeceHals in four fate regular rows. subnigripes 17. Gena in height midway between vibrissal and posterior angles about one- third the eye height (fig. 3)... .. aes Gena in height midway between ifriseal ad Sosteron seeks about one- seventh the eye height (fig. 6)... . . . . barberi 18. Third antennal segment dorsally atpened: Aictally wile a ee angle. niveipennis Third antennal segment rounded (fig. 3) ......... =... ambigua 19. Calypter with margin and fringe brown ..........2.2.+. +. 20 Calypter with margin and fringe white or yellow .......... 21 20. Midtibia medially with two osterolateral setae ....... . . isolate Midtibia medially without posterolateral setae ....... . . spiraeae 21. Tibiae and tarsi dark brown!) = =... 2). «es. =.» .% = . Eubiliceps Tibiae and tarsi yellowish ................ . albitarsis Agromyza albitarsis Meigen Agromyza albitarsis Meigen, Systematische Beschreibung der bekannten euro- paischen zweifliigeligen Insekten, vol. 6, p. 171, 1830.—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam.-59, p. 100, 1931 (figure of head). ee cinerascens (Macquart), Melander, Journ. New York Ent. Soc., vol. 21, p. 253, 1913 (3 from Washington: 107, 19, Monroe; 19, Port Gamble). This Holarctic species belongs with those species having four strong dorsocentral setae, one being presutural. The calypteral fringe is white and the tibiae and tarsi are yellowish. The larvae make blotch mines in the leaves of Populus spp. and Salix lasiandra Benth. I have studied reared specimens from Washington, California, and Pennsylvania. The species probably is widely distributed. Agromyza ambigua Fallén Agromyza ambigua Fallén, Diptera sueciae, vol. 2, No. 37 (Agromyzides), p. 4, 1823.—Hendel, in Lindner, Die Fliegen der Palaearktischen Region, fam. 59, p. 103, 1931 (figure of head and wing). Agromyza kincaidi Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 285, 1913 (figure of wing).—Shewell, Canadian Ent., vol. 85, p. 463, 1953. The holotype of A. kincaidi was compared with two specimens of A. ambigua in my collection and with four in the U. S. National Museum, all determined by E. M. Hering. This is a shining black species with three developed dorsocentrals, the costa reaching to the end of vein R4,;, a relatively wide gena, and a rounded third antennal segment (fig. 3). The larvae mine the leaves of some grasses. A Holarctic species, A. ambigua is at present known only from Alaska in North America. 354 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Agromyza aristata Malloch Agromyza aristata Malloch, Canadian Ent., vol. 47, p. 13, 1915. Agromyza ulmi Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 54, 1924.— Needham, Frost, and Tothill, Leaf-mining insects, p. 252, 1928 (figure of leaf mine). A. aristata belongs to the species group in which there are no medial setae on the midtibia and vein M,,, is at or nearest the wing tip (fig. 4). The head (fig. 5) has the arista and the setae somewhat shortened. The larvae mine in the leaves of Ulmus americana L. and probably Celtis occidentalis Mic. because I have seen a leaf with a mine very similar to that found on elm. Known from Iowa, Illinois, Indiana, Ohio, Pennsylvania, New York, and Virginia. Agromyza barberi Frick Agromyza abbreviata Malloch (not Fallén, 1823), Ann. Ent. Soc. Amer., vol. 6, p. 285, 1913 (fig. of wing). Agromyza barberi Frick, Univ. California Publ. Ent., vol. 8, p. 372, 1952. This is a shining species with three developed dorsocentrals, the costa ending at vein Ry,5, and a very narrow gena (fig. 6). The larvae probably mine the leaves of grasses. Known only from the holotype from New Mexico. Agromyza canadensis Malloch Agromyza canadensis Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 299, 1913 (figure of head)—Shewell, Canadian Ent., vol. 85, p. 462, 1953. This large species (3 mm. in wing length) has the mesonotum dull and bearing about five developed dorsocentrals. It is rather similar to A. diversa in that the mesonotum is light brown. However, this species has the third antennal segment and maxillary palpus brown. The head has a very narrow gena (fig. 7). There are only three specimens known, all from Ontario, Canada. Agromyza centrosemae Frost Agromyza centrosemae Frost, Ann. Ent. Soc. Amer., vol. 29, p. 301, 1936. The holotype male is in the U. S. National Museum (No. 62972). It is extremely teneral but the mesonotum lacks a metallic sheen and the calypteral fringe is yellowish. These characteristics separate this species from A. frosti and A. currani. Known only from the holotype which was reared from Centrosema pubescens Benth. in the Panama Canal Zone. Agromyza currani Frost Agromyza currani Frost, Ann. Ent. Soc. Amer., vol. 29, p. 305, 1936. The female holotype is in the American Museum of Natural History and there is a male paratype in the S. W. Frost collection. The head AGROMYZID LEAF MINERS—FRICK 355 is high and narrow and the arista is plumose (fig. 8). This species is rather distinctive in having the anepisternum and tergites bordered with yellow and the mesonotum heavily dull gray pollinose. The two known specimens are from Barro Colorado Island, Panama Canal Zone. Agromyza diversa Johnson Agromyza diversa Johnson, Occ. Pap. Boston Soc. Nat. Hist., vol. 5, p. 26, 1922.— Shewell, Canadian Ent., vol. 85, p. 462, 1953. This species belongs to Agromyza sens. str. even though the body, including the scutellum, is predominately yellow. A. diversa differs from the very similar A. ferruginosa van der Wulp of Europe in having the head mostly brown. I have seen specimens from Iowa, Illinois, Massachusetts, Vermont, and Ontario. Agromyza frosti Frick Agromyza schmidti Frost (not Aldrich, 1929), Ann. Ent. Soc. Amer., vol. 29, p. 302, 1936. Agromyza frosti Frick, Univ. California Publ. Ent., vol. 8, p. 373, 1952. The male holotype is in the U. S. National Museum (No. 62966). There are three paratypes in the S. W. Frost collection. The head has the antenna placed well below the middle of the head and the eye is ovoid (fig. 9). This is a species of moderate size (2 mm. in wing length) that has two developed dorsocentrals, the mesonotum subshining and without metallic sheen, and the anepisternum entirely dark. The type series is from La Caja, Costa Rica, and was reared from the stems of an unidentified plant. Agromyza inaequalis Malloch Agromyza inaequalis Malloch, Proc. Ent. Soc. Washington, vol. 16, p. 89, 1914 (figs. of head and wing). Agromyza iridescens Frost, Ann. Ent. Soc. Amer., vol. 29, p. 303, 1936. This species has the upper-orbital setae unequal in length, the ventral being the longer and stronger (fig. 10), and a bright green sheen to the mesonotum and abdomen. The larvae make blotch mines in the leaves of Vigna repens Baker, Phaseolus vulgaris L., and P. limensis Macfad. I have seen specimens from Puerto Rico, Cuba, Panama Canal Zone, and Venezuela. Agromyza isolata Malloch Agromyza isolata Malloch, Ann, Ent. Soc. Amer., vol. 6, p. 306, 1913. This species belongs with those species having four developed dorso- centrals, one being presutural. The calypter has the fringe brown and there are two medial setae on the midtibia. This last character and the high, narrow eye (fig. 11) serve to separate A. isolata from 356 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 A. spiraeae. In addition to the type female from California there is a male in the U. 8. National Museum that was collected in the District of Columbia. Agromyza nigripes Meigen Agromyza nigripes Meigen, Systematische Beschreibung der bekannten euro- piischen zweifliigeligen Insekten, vol. 6, p. 170, 1830.—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 137, 1931 (figures of head and wing). Agromyza dubitata Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 311, 1913. A relatively common Holarctic species, A. nigripes is rather dis- tinctive in having the frons wider than long and the third antennal segment subangulate (fig. 12). The mesonotum is shining black and bears three or four developed postsutural dorsocentrals. The larvae are grass miners and a series in the Illinois Natural History Survey collection were reared from Triticum aestivum L. at Wooster, Ohio. The species is known from Iowa, Illinois, Michigan, Ohio, New York, Massachusetts, Ontario, and Quebec. Agromyza niveipennis Zetterstedt Agromyza niveipennis Zetterstedt, Diptera Scandinaviae, vol. 7, p. 2741, 1848.— Hendel, in Lindner, Die Fleigen der palaearktischen Region, fam. 59, p. 139, 1931 (figures of head and wing). Agromyza cinerascens (Macquart), Melander, Journ. New York Ent. Soc., vol. 21, p. 253, 1913 (1 o&, Oroville, Wash.). This is a rather distinctive, shining black species with the third antennal segment angulate distally, the head in profile subtriangular, and six rows of acrostichals. It has not been reported previously from North America. I have reared it from leaf mines on the follow- ing grasses in central Washington: Secale cereale L., Hordeum jubatum L., Triticum aestivum L., and Agropyron repens L. There are other specimens from Utah and Kansas. Agromyza pallidiseta Malloch Agromyza pallidiseta Malloch, Canadian Ent., vol. 56, p. 192, 1924. This species is closely related to A. aristata, but has all body setae yellowish and the genovertical plates partially darkened (fig. 13). The frons is about as wide as long (fig. 14). Known only from the holotype female in the U. S. National Museum (64299), collected at Rock Creek Park, Washington, D. C. Agromyza parvicornis Loew Agromyza parvicornis Loew, Berliner Ent. Zeitschr., vol. 13, p. 49, 1869 (Centuria vil, No. 92).—Melander (part), Journ. New York Ent. Soc., vol. 21, p. 254, 1913 (excl. 19, Wisconsin).—Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 312, 1913 (figures of head and wing).—Phillips, Journ. Agr. Res., vol. 2, p. 15, 1915 (figures of leaf mines). AGROMYZID LEAF MINERS—FRICK 357 A. parvicornis has the mesonotum shining black and bearing two developed dorsocentrals. The costa reaches to the end of vein Mie and the calypteral fringe is brown. The larvae mine the leaves of Zea mays L. It is a widespread and well known species and I have seen specimens from most of the States in the United States and from Ontario. Agromyza reptans Fallén Agromyza reptans Fallén, Diptera sueciae, vol. 2, No. 37 (Agromyzides), p. 3, 1823.—Melander (part), Journ. New York Ent. Soc., vol. 21, p. 253, 1913 (1 o, Seattle, Wash.).—Hendel, in Lindner, Die Fleigen der palaearktischen Region, fam. 59, p. 144, 1931 (figures of head and wing). This is a large species (3 to 4 mm. in wing length) that has the meso- notum subshining black and bearing four postsutural dorsocentrals. The tibiae and tarsi are yellowish. A. reptans is a common species in Europe, but I have seen no other North American specimens than Melander’s and a long series that I reared from blotch mines in the leaves of Urtica californica Greene in California. Agromyza rubi Brischke Agromyza rubi Brischke, Schrift. Naturf. Ges. Danzig, vol. 5, p. 250, 1881.— Hering, Tijdschr. Ent., vol. 97, p. 118, 1954 (figures of larval characters). Agromyza sulfuriceps Strobl, Mitt. Naturw. Ver. Steiermark, vol. 34, p. 270, 1898.—Melander, Journ. New York Ent. Soc., vol. 21, p. 225, 1913,— Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 152, 1931 (figures of head and wing). This rather distinctive Holarctic species belongs to the group with vein M4. at or nearest the wing tip and with no medial midtibial setae. If differs from A. varifrons in having the mesonotum dull gray pollinose and from A. pallidiseta and A. aristata in having the femora, tibiae, and tarsi dark and vein Mj,4. ending at the wing tip. The larvae mine the leaves of species of Rubus, Potentilla, and Sanguisorba in Europe. I have seen swept specimens from Washington, Idaho, and California. Agromyza rutiliceps Melander Agromyza rutiliceps Melander, Journ. New York Ent. Soc., vol. 21, p. 261, 1913. The holotype male is in the collection of A. L. Melander. Its con- dition is poor so no illustrations were prepared from it. A. rutiliceps belongs with those species having four developed dorsocentrals, one being presutural. The calypteral margin and fringe are yellow and the tibiae and tarsi are dark. The type from Montana is the only specimen of this species that I have seen. 358 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Agromyza spiraeae Kaltenbach Agromyza spiraeae Kaltenbach, Verh. Naturh. Ver. Preuss. Rheinlande West- falens, vol. 24, p. 104, 1867—Malloch, Diptera of Patagonia and South Chile, pt. 6, fasc. 5, p. 477, 1934.—Hering, Tijdschr. Ent., vol. 97, p. 121, 1954 (figures of larval characters). Agromyza fragariae Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 307, 1913 (figure of wing).—Frost, Mem. Cornell Univ. Exp. Sta., vol. 78, p. 43, 1924 (figure of leaf mine on Rubus). Agromyza sanguisorbae Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 149, 1931 (figure of wing). Malloch’s holotype was compared with five specimens (one in the U. S. National Museum) sent by E. M. Hering. Hering restricted the concept of A. spiraeae to those specimens the larvae of which mine the leaves of the Rosoideae or tribe Roseae. This group in- cludes the genera Rubus, Fragaria, and Rosa, but not Spiraea. A. spiraeae belongs with those species having four developed dorso- centrals, one being presutural. The margin and fringe of the calypter are brown, and this species may be separated from or j6e soi, 5) paula Bom oSipaes Eoin Sok cle eens wel be ase Ceo ees moa sup cpe gra Femora distally yellow for one- third to one-half fhe fermoral length . muscina 4, Vein M344 with ultimate section about 1.5 times as long as penultimate; humerus and notopleural triangle dark. ... . LS, FARTS Vein M344 with ultimate section subequal to peauinmnte. mofoniearel triangle and. part of humerus yellow... °.-s.%)< sie.) = = . .. . . lateralis 5. Legs with all femora distally yellow. . ........ .. . . imconspicua Legs with.forefemur yellow, others:dark. .. 2... .. 4: . ss. incisa Phytobia (Poémyza) angulata (Loew) Agromyza angulata Loew, Berliner Ent. Zeitschr., vol. 13, p. 47, 1869 (Centuria vit, No. 87). Agromyza neptis Loew, Berliner Ent. Zeitschr., vol. 13, p. 50, 1869 (Centuria vir, No. 93). Dizygomyza semiposticata Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 49, 1931 (figures of head, wing, and puparial characters). Agromyza cinereifrons Frost, Canadian Ent., vol. 63, p. 276, 1931. AGROMYZID LEAF MINERS—FRICK 381 Phytobia angulata is close to P. subangulata because both have the prescutellar seta present. However, the former has the eye smoothly rounded anteriorly (fig. 69) as compared to the irregular curvature shown by P. subangulata (fig. 79). The lunule is not abruptly nar- rowed immediately above the antenna as it is in P. subangulata (fig. 78). All femora are narrowly yellow distally. The wing has cross- vein r-m at about the center of cell 1-M, and m-m is about 1.5 times its length from r-m. There are no North American specimens known to me that have been reared, but the larvae mine species of Carex in Europe. There are specimens from widely scattered States and it is probable that the species is present in every State of the United States. It is known from Ontario and Quebec in Canada. Phytobia (Poémyza) incisa (Meigen) Agromyza incisa Meigen, Systematische Beschreibung der bekannten europdischen zweifliigeligen Insekten, vol. 6, p. 182, 1830. Dizygomyza (Poémyza) incisa (Meigen), Hendel, Die Fliegen der palaearktischen Region, fam. 59, p. 38, 1931 (figures of head, wing, and puparial characters). This species is rather distinctive in having crossvein m-m approxi- mately beneath the junction of R,; in the costa. The prescutellar seta is not developed and the species is subshining black with only the forefemur yellow distally. Phytobia inconspicua is similar but has all femora distally yellow. The larvae form blotch mines in the leaves of many species of grass. There may be several larvae per mine. Frick (1952) listed four host plants; other North American grasses from which adults have been reared are Phleum pratense L., Triticum aestivum L., Setaria lutescens (Weigel) Hubb., and Phalaris arundinacea L. ‘There are specimens from many States in the north- ern half of the United States and from Ontario in Canada. Phytobia (Poémyza) inconspicua (Malloch) Agromyza inconspicua Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 310, 1913. This species is very similar to Phytobia incisa but is smaller (1.75 mm. in wing length), has the third antennal segment slightly angu- late, and has all femora distally yellow. The head of the type (fig. 70) is somewhat distorted because the specimen is teneral. This species is known from a single male reared from a mine on Agropyron sp. in Colorado. The puparium is on the same pin and illustrations of the posterior end and spiracles are included (figs. 71, 72, and 73). The spiracles are different from any others that I have seen. Phytobia (Poémyza) lateralis (Macquart) Agromyza lateralis Macquart, Histoire naturelle des Insectes, Diptera, vol. 2, p. 609, 1835. Dizygomyza (Poémyza) lateralis (Macquart), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 40, 1931 (figures of head and wing). 382 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Agromyza coquilletti Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 295, 1913 (figure of head). Agromyza marginata (Loew), Melander, Journ. New York Ent. Soc., vol. 21, p. 256, 1913. Paratype males of Phytobia coquilletti were compared with two males lent by E. M. Hering. The distinctive keel on the ninth tergite (figs. 74, 75), first described by Shewell (1953), is also present on the European males. Malloch’s figure of the head is somewhat inaccurate and I have redrawn it (fig. 76). This species is quite yellow with most of the head, most of the humerus, notopleural triangle, all femora distally, and all tergites posteriorly being yellowish. The larvae mine the leaves of many species of grass and there are North American records from the following: Avena sativa L., Elymus canadensis L., Hordeum jubatum L., Triticum aestivum L., and Zea mays L. I have seen specimens from many of the States in the northern half of the United States and from Manitoba in Canada. Phytobia (Poémyza) muscina (Meigen) Agromyza muscina Meigen, Systematische Beschreibung der bekannten euro- pdischen zweifliigeligen Insekten, vol. 6, p. 177, 1830. Dizygomyza (Poémyza) muscina (Meigen), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 44, 1931 (figures of head and wing). Agromyza marginata Loew, Berliner Ent. Zeitschr., vol. 13, p. 49, 1869 (Centuria vu, No. 91). Agromyza superciliosa (Zetterstedt), Melander, Journ. New York Ent. Soc., vol. 21, p. 256, 1913. Phytobia muscina is a distinctive species because the femora are distally yellow for about one-third of their length. The genovertical plates are yellowish on the dorsal half and usually are brownish for the full length, contrasting with the black frontal vitta. The larvae mine the leaves of several species of grass. The only reared specimens that I have seen from North America were from leaf mines on Agro- pyron repens (L.), Ehrharta erecta Lam., and Hordeum murinum L. I have seen specimens from California (including Los Angeles), Ore- gon, Washington, Idaho, Illinois, Indiana, Michigan, District of Columbia, Maryland, Pennsylvania, New York, and Massachusetts. The species also occurs in the North West Territories of Canada. Phytobia (Poémysa) subangulata (Malloch) Agromyza subangulata Malloch, Psyche, vol. 23, p. 51, 1916. This species is similar to Phytobia angulata in having the prescutellar seta developed. However, it differs in having only the forefemur distally yellow, the other femora distally reddish brown, crossvein r-m definitely distad of center of cell 1—-M; and slightly beyond the junction of R, in the costa (fig. 77). There is a narrow but rather AGROMYZID LEAF MINERS—FRICK 383 prominent keel between the antennae, and the lunule is abruptly narrowed immediately dorsad of the antenna (fig. 78). The eye is not as smoothly rounded anteriorly (fig. 79) as it is in P. angulata (fig. 69). I have seen no reared specimens but in Europe this group mine the leaves of species of Carex, Cyperus, and Scirpus. I have identified only a few specimens as belonging to this species, all from Illinois. Most of the specimens Malloch placed under Phytobia subangulata belong to P. angulata sens. str. df. Subgenus Dizygomyza Hendel Dizygomyza Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 83, 1951. Of the six described species of this subgenus two are Nearctic and four are Holarctic. The adults placed in this subgenus are char- acterized by a very large lunule. The lunule is in the plane of the frontal vitta and abruptly raised above the plane of the mesofacial plate (fig. 83). This is in contrast to subgenus Poémyza where the lunule is sunken somewhat below the frontal vitta and is in the plane of the mesofacial plate (fig. 78). The height of the lunule is usually less than one-half its width at the antennal bases. However, the lunule may be higher (fig. 83), as it is in Phytobia (Dizygomyza) thompsoni, but the lunule being in the plane of the frontal vitta sepa- rates the latter species from species of the subgenus Poémyza. Sub- genus Dizygomyza has the frontal triangle no larger than the ocellar triangle (fig. 83), in contrast to the elongate frontal triangle found in subgenus Jcteromyza which reaches to the lunule (fig. 85). The third antennal segment of the males is greatly enlarged. The larvae have three bulbs on each posterior spiracle. One or more of these is usually elongate, heavily sclerotized, acuminate, and variously curved. The posterior spiracles of some species are quite similar to some found in subgenus Poémyza. Key to the described species of Phytobia (Dizygomyza) 1. Anterior half of mesonotum with acrostichal setae numerous, in six to eight ’ PPrOOMIAL LOWS: 2 #) @ a8 .@ wood fey. he. Soups) Gee fe we reeks |S 2 Anterior half of mesonotum with acrostichals relatively sparse, in four to five WENO UALS POW Sag ca ey a ars oe cas war rige ace a oa d.da, Jar) Bi. ayles oe bur ae oLee: sel 4 2. Legs with all femora distally light yellow for a distance subequal to the femoral UIE CCR ees se, ss | Sy an) eee eae WES 8 tas em or eo te ee 3 Legs with forefemur distally light yellow, others light brown. ... . iridis 3. Head rounded in profile, genovertical plates at most barely extending beyond eye margin; tarsi dark brown; small species, two to 2.3 mm. in wing length. iraeos Head subtriangular in profile, genovertical plates strongly extending beyond eye margin (fig. 82); tarsi yellowish; large species, 2.75 to 3.0 mm. in wing Venp Calis seach ew colic as ule oc. 7d, car Loh eittel EQ od se, eghhW OA Go 3s) thompsoni 384 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 4. Legs with all femora distally light yellow for a distance subequal to the femoral diameter 0 4S) apres SHAAN Ty aie tisiaectwdtac fol msl peptie Benatisune mae Legs with forefemur distally light yellow, others black . . . . . . luctuosa 5. Abdomen with basal two or three tergites laterally yellow . . . . . morosa Abdomen with basal two or three tergites laterally black . . . magnicornis Phytobia (Dizygomyza) iraeos (Robineau-Desvoidy) Agromyza iraeos Robineau-Desvoidy, Rev. Mag. Zool., ser. 2, vol. 3, p. 393, 1851. Dizygomyza (Dizygomyza) iraeos (Robineau-Desvoidy), Hendel, Die Fliegen der palaearktischen Region, fam. 59, p. 86, 1931 (figures of larval spiracles). Agromyza taeniola (Coquillett), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 256, 1913 (19, Douglas, Alaska). This species is very similar to Phytobia (Dizygomyza) iridis, another Holarctic species, but may be separated by the three light yellow “Inees.”? The key characters of rounded head, dark tarsi, and small size serve to separate this species from P. (D.) thompsoni. The larvae mine in Jris spp. There are reared specimens from Indiana and Louisiana; other specimens that I have seen are from Michigan and Alaska. Phytobia (Dizygomyza) iridis (Hendel) Dizygomyza iridis Hendel, Zool. Anz., vol. 69, p. 253, 1927. This Holarctic species is very similar to Phytobia (Dizygomyza) iraeos but is distinguished by having only the forefemur light yellow distally, the others being light brown. The larvae mine in Jris spp. I have identified specimens from California and Michigan. Phytobia (Dizygomyza) luctuosa (Meigen) Agromyza luctuosa Meigen, Systematische Beschreibung der bekannten euro- paischen zweifliigeligen Insekten, vol. 6, p. 182, 1830. Dizygomyza (Dizygomyza) luctuosa (Meigen), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 88, 1931 (figures of head, wing, and puparial spiracles). Agromyza taeniola (Coquillett), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 256, 1913 (29 9, Portland, Oreg., and Monroe, Wash.). A rather distinctive species among the species having relatively sparse acrostichals in that the midfemora and hind femora are all black. No reared specimens are known from North America, but the larvae mine species of Carex and Juncus in Europe. At present this species is known from California, Oregon, Washington, and New York, in the United States, and from Nova Scotia in Canada. Phytobia (Dizygomyza) magnicornis (Loew) Agromyza magnicornis Loew, Berliner Ent. Zeitschr., vol. 13, p. 46, 1869 (Cen- turia vu, No. 86). This is a Nearctic species that may be separated from the closely related Phytobia (Dizygomyza) morosa in having the abdominal ter- AGROMYZID LEAF MINERS—FRICK 385 gites laterally black or brown; the basal two or three tergites of P. (D.) morosa are laterally yellow. The head (fig. 80) and wing (fig. 81) of the holotype male are illustrated in order to help identify this pre- viously misidentified species. The larval host plants are not known. I have seen specimens from Illinois, Indiana, Michigan, District of Columbia, Pennsylvania, and New Hampshire as well as from British Columbia. Phytobia (Dizygomyza) morosa (Meigen) Agromyza morosa Meigen, Systematische Beschreibung der bekannten euro- pdischen zweifliigeligen Insekten, vol. 6, p. 170, 1830. Dizygomyza (Dizygomyza) morosa (Meigen), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 90, 1931 (figures of head and puparial spiracles). Another Holarctic species that is little known in North America. It is distinctive in having the basal two or three tergites yellow later- ally. No larval host plants are known in North America but the larvae mine in species of Scirpus and Carex in Europe. Thus far known only from South Dakota, Illinois, Indiana, and Maryland. Phytobia (Dizygomyza) thompsoni Frick Agromyza magnicornis (Loew), Coquillett, in Thompson, Psyche, vol. 14, p. 74, 1907 (figure of leaf galls). Agromyza laterella (Zetterstedt), Malloch (part), Ann. Ent. Soc. Amer., vol. 6, - p. 300, 1913.—Claassen, Ann. Ent. Soc. Amer., vol. 11, p. 9, 1918 (figures of adult, puparium, larva, and leaf mines and galls). This is a large species (2.75 to 3 mm. in wing length) for a species of the subgenus Dizygomyza. The head is subtriangular in profile with the genovertical plates strongly extending beyond the eye margin (fig. 82). The lunule is high for this subgenus, extending halfway to the anterior ocellus from the antennal base (fig. 83). The male terminalia is characterized by the large yellow knob on the ninth tergite (fig. 84). The larvae mine the leaves of Iris versicolor L., Iris sp. (blue flag), and Typha spp. There are specimens from Illinois, Michigan, Penn- sylvania, New York, and Massachusetts. 5g. Subgenus Icteromyza Hendel Icteromyza Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 51, 1981. The adults are characterized by the bright yellow frontal vitta and lunule, the elongate frontal triangle reaching to the lunule, the very large lunule that is in the plane of the frontal vitta and abruptly raised above the mesofacial plate, and the widely spaced antennal bases (fig. 85). This subgenus may be separated from subgenus Poémyza by the bright yellow frontal vitta and the lunule in the plane of the frontal vitta and from subgenus Dizygomyza by the color of 386 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 the frontal vitta and the elongate frontal triangle. The third anten- nal segment of the males is not enlarged. The eye may be setulose, as it is in Phytobia (Icteromyza) pollinosa (fig. 87). The wing is rela- tively long and slender (fig. 86) as compared to the wings in the sub- genera Phytobia (fig. 60), Poémyza (fig. 77), and Dizygomyza (fig.81). The larva of Phytobia (Icteromyza) longipennis is the only one known from the subgenus. The anterior spiracle has two small bulbs and is very small. Each posterior spiracle of the third stage or full grown larva is long, ventrally directed, sclerotized, acuminate, and bears about 20 very small bulbs in a single row. Such a highly modified spiracle compares favorably with the unusual and diverse types found in the subgenera Poémyza and Dizygomyza and supports the close re- lationship shown by the adults of the three subgenera. Key to the described species of Phytobia (Icteromyza) 1. Maxillary palpus yellow; femora distally broadly yellow for a distance sub- equal to one-third the femoral length. . .........+++42+46-. 2 Maxillary palpus black; femora distally narrowly yellow for a distance sub- equal to the femoral diameter. . ... . Snag eS he. at capitata 2. Eye setulose; third antennal segment yellow... .......--. pollinosa Eye practically bare; third antennal segment mostly brown or black. longipennis Phytobia (Icteromyza) capitata (Zetterstedt) Agromyza capitata Zetterstedt, Diptera Scandinaviae, vol. 7, p. 2750, 1848. Dizygomyza (Icteromyza) capitata (Zetterstedt), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 52, 1931 (figures of head and wing).— de Meijere, Tijdschr. Ent., vol. 84, p. 15, 1941. Agromyza genualis Melander, Journ. New York Ent. Soc., vol. 21, p. 261, 1913. Agromyza coloradensis Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 297, 1913. This Holarctic species is large, with a wing length of 2.5 to 3.5 mm. The maxillary palpus is black and the femora are distally yellow for a distance equal to the femoral diameter. A front view of the head shows the form of the lunule and frontal triangle (fig. 85). De Meijere found an adult of this species on the stems of Juncus sp. and surmised that the larvae mined in Juncus spp. Dizygomyza (Icter- omyza) capitata is northern in distribution but extends down the Sierra Nevada mountain chain into southern California. It also occurs in Oregon, Washington, Idaho, Montana, Utah, Colorado, North Dakota, Michigan, Illinois, Maine, and Ontario. Phytobia (Icteromyza) longipennis (Loew) Agromyza longipennis Loew, Berliner Ent. Zeitschr., vol. 13, p. 48, 1869 (Centuria vit, No. 90).—Melander, Journ. New York Ent. Soc., vol. 21, p. 255, 1913.— Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 296, 1913. This species is smaller (2.5 mm. in wing length) than Phytobia (Icteromyza) capitata, has the palpus yellow, and all femora distally AGROMYZID LEAF MINERS—FRICK 387 broadly yellow for about one-third the length. From P. (J.) pollinosa it differs by having the third antennal segment brown or black and the eye bare. The wing is typical of those found in this subgenus in being relatively long and narrow (fig. 86). I have reared P. (J.) longipennis from leaf and stem mines in Juncus xiphioides Meyer in California. This is the first positive record of a host plant for species of the subgenus Jcteromyza. I have seen specimens from many States throughout the United States and from Manitoba and Quebec in Canada. Phytobia (Icteromyza) pollinosa (Melander) Agromyza pollinosa Melander, Journ. New York Ent. Soce., vol. 21, p. 263, 1913. The eyes are heavily pollinose (fig. 87), a character which separates this species from other North American species. It is closest to Phytobia (Icteromyza) longipennis in having the palpus yellow and the distal third of the femora yellow, but it has the third antennal segment yellow. There are only two known specimens, both from Sikta, Alaska. sh. Subgenus Calycomyza Hendel Calycomyza Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 65, 1931.—Frick, Ann. Ent. Soc. Amer., vol. 49, p. 284, 1956 (figures of larval characters and types of leaf mines of five species). This subgenus is placed among the more specialized subgenera on the basis of the usual lack of a presutural dorsocentral and the circular or irregular form of the posterior spiracles of two of the six described larvae. The species found north of Mexico were recently revised by Frick (1956b). Previously, species of the subgenus Calycomyza were distinguished by the three postsutural dorsocentral setae and the black legs. A number of North American species were described in which the forefemur is distally narrowly yellow and two—Phytobia (Calycomyza) majuscula and P. (C.) gigantea—that have four post- sutural and none to three presutural dorsocentrals. P. (C.) meridiana has the third antennal segment yellow, a unique character for this subgenus. The larvae of only six species are known. ‘There are usually three bulbs on each posterior spiracle but one Palaearctic species has from three to five while one Nearctic species has five to eight in a circle and one Holarctic species has about 10 irregularly arranged. The large number of species found in North America was sur- prising considering that only five are known in the well-collected Palaearctic region. Of the 17 species described from North America, three are Holarctic, three others are found in both Nearctic and Neotropical regions, eight are strictly Nearctic, and four are Neo- tropical only. 388 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The revision did not include species found south of the United States because of the poor condition of the type series of P. (C.) ipomoeae and P. (C.) cassiae. Complete descriptions are impossible and these species are placed in the key by using rather minor characters. Short descriptions are included for the three Neotropical species not included in the revision. The notes under the other species are limited to larval host plant records and distributions be- cause the revision included descriptions of all species considered therein. Key to the described species of Phytobia (Calycomyza) 1. Calypter with margin and fringe white or yellow. .......... 2 Calypter with margin and fringe dark brown orblack ......... 6 2. Third antennal segment rounded, subcircular; anterior half of mesonotum with about six rows of acrostichal setae... . : SAeERO Third antennal segment with an anterodorsal elon flattened dorcalls beyond base of arista; about five rows of acrostichals. ....... 5 3. Genovertical plates dark from vertex to at least the dorsal upper-orbital seta; forefemur entirely black. . .. . os HAA Genovertical plates dark not more than alias percent inner vertical seta and dorsal upper-orbital; forefemur distally narrowly yellow. lantanae 4. Second dorsocentral seta more than one-half the length of the third dor- socentral; genovertical plates dark from vertex to at least halfway be- tween the two upper-orbital setae... . . . premissa Second dorsocentral one-half or less the lene of he third; genovertical plates dark usually to dorsal ie Aart sometimes beyond, rarely to ventral upper-orbital. .... . . . jucunda 5. Mesofacial plate dark except for narrow wellcyy epistomale margin; calypter with margin and fringe white. ... . ~ + « . « /.(/)* humeralis Mesofacial plate yellow; margin and fringe pelle? - . .. . . .solidaginis 6. Mesofacial plate at least one-half dark. .... oh ar oVe SMP ohbeededs Mesofacial plate yellow, not more than antennal Races Hei RVs ee tere LO 7. Genovertical plates dark from vertex to at least ventral upper-orbital . . 8 Genovertical plates dark from vertex to dorsal upper-orbital . artemisiae 8. Genovertical plates dark, at least narrowly, from vertex to the ventral lower- orbital; second dorsocentral more than one-half the length of the third doreaceniea 3. Lapa Wicd eared O Genovertical plates dark ie Ganteal inperornrialt or Velorean logeronniale second dorsocentral less than one-half the third. . . . . . . cynoglossi 9. Mesonotum shining, six rows of acrostichals; humeral seta on black.majuscula Mesonotum dull, pollinose, four or five rows of acrostichals; humeral seta on yellow’. .. ~ - . « . gigantea 10. Genovertical plates eallow: rely ‘Gack from ferten to halfway to dorsal upper-orbital; third antennal segment ovoid, longer than wide. . . . 11 Genovertical plates dark at least to dorsal upper-orbital; third antennal segment subcircular. ... eth 4.2 °°. eee a ee 11. Inner postalar seta on yellow of meaometuiae Seay fit’ si\fethergeee | HAVENOLUI Inner postalar on black of mesonotum. .......... . . allecta AGROMYZID LEAF MINERS—FRICK 389 12. Third antennal segment blacks... . 6 6. 2 we ee sw ww ew ee 13 Third antennal segment yellow... . . . . meridiana 13. Genovertical plates dark from vertex to a feast fit eae upper-orbital . 14 Genovertical plates dark from vertex to dorsal upper-orbital. . . . . . 15 14. Arista subequal in length to the eye height; mesonotum with about 19 setu- lae in the intraalar row posterior to the transverse suture. . . . cassiae Arista not more than four-fifths of the eye height; about 16 setulae in the IMUAAIST TOW, <8 oe ay ch = Ss Se ve vee . . . . malvae 15. Forefemur brown or blacks at arost fichier brown distally. reg ae erat Ns, Forefemur distally yellow for a distance subequal to the femoral diameter. ipomoeae 16. Frontal vitta relatively wide, not more than 1.4 times as high as wide. ambrosiae Frontal vitta relatively narrow, not less than 1.7 times as high as wide. verbenae Phytobia (Calycomyza) allecta (Melander) Agromyza lateralis Williston, (not Macquart, 1835), Trans. Ent. Soc. London, 1896, pt. 3, p. 428, 1896 (figure of head). Phytobia (Calycomyza) allecta (Melander), Frick, Ann. Ent. Soc. Amer., vol. 49, p. 298, 1956. The larvae have elongate anal lobes and form large blotch mines in the leaves of Bidens frondosa L., Helianthus spp., and Rudbeckia spp. The species occurs east of the Rocky Mountains in the United States and Canada and on the islands of St. Vincent, Haiti, and Puerto Rico in the West Indies. Phytobia (Calycomyza) ambrosiae Frick Phytobia (Calycomyza) ambrosiae Frick, Ann. Ent. Soc. Amer., vol. 49, p. 299, 1956. This species has been reared from Ambrosia artemisiifolia L. and A. trifida L. Thus far it has been found in Indiana, Tennessee, and Florida. Phytobia (Calycomyza) artemisiae (Kaltenbach) Agromyza artemisiae Kaltenbach, Verh. Naturh. Ver. Preuss. Rheinlande West falens, vol. 13, p. 236, 1856. Dizygomyza (Calycomyza) artemistae (Kaltenbach), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 66, 1931 (figures of head, wing, and puparium). Phytobia (Calycomyza) artemisiae (Kaltenbach), Frick, Ann. Ent. Soc. Amer., vol. 49, p. 294, 1956 (figures of larval characters and leaf mines). The larvae of this Holarctic species make blotch mines in the leaves of Artemisia spp. and Rudbeckia spp. in North America. The species appears to be generally distributed over the United States and Canada. It is known from Guatemala. 390 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Phytobia (Calcomyza) cassiae (Frost) Agromyza sp., Frost, Sci. Month., vol. 30, p. 445, 1930 (figure of leaf mines). Agromyza cassiae Frost, Ann. Ent. Soc. Amer., vol. 29, p. 306, 1936. The holotype female is in the U. S. National Museum (No. 62965). Phytobia (Calycomyza) cassiae is very similar to P. (C.) malvae and, because the type series is in poor condition, separation of the two species is difficult. The very long arista, subequal in length to the eye height, is probably the best character to use at this time. The larvae make serpentine mines in the leaves of Cassia bacillaris L. The type series is from the Panama Canal Zone and I have identified two specimens from Puerto Rico as belonging to Phytobia (Calycomyza) cassiae. Phytobia (Calycomyza) cynoglossi Frick Phytobia (Calycomyza) cynoglossit Frick, Ann. Ent. Soc. Amer., vol. 49, p. 295, 1956. The larvae mine the leaves of Cynoglossum spp. This species is general throughout Canada but is not known west of Kansas in the United States. Phytobia (Calycomyza) flavinotum Frick Agromyza allecta (Melander), Frost (part), Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 38, 1924. Phytobia (Calycomyza) flavinotum Frick, Ann. Ent. Soc. Amer., vol. 49, p. 297, 1956 (figure of antenna). This is a distinctive species in that the inner postalar seta is on yellow. The larvae have elongate anal lobes and make large blotch mines in the leaves of Arctium lappa L., Eupatorium purpureum L., and Viburnum pubescens Pursh. Phytobia (Calycomyza) gigantea Frick Agromyza platyptera (Thomson), Malloch (part), Ann. Ent. Soc. Amer., vol. 6, p. 293, 1913. Phytobia (Calycomyza) gigantea Frick, Ann, Ent. Soc. Amer., vol. 49, p. 296, 1956. This is the largest species (1.75-2.5 mm. in wing length) known in the subgenus. The larval host plants are unknown. This species is recorded in the United States from Illinois, Virginia, Maryland, and in Canada from Ontario and from Quebec. Phytobia (Calycomyza) humeralis (von Roser) Agromyza humeralis von Roser, Korresp.-Blatt. Wuertt. Landw. Ver., vol. 8, p. 63, 1840. Phytobia (Calycomza) humeralis (von Roser), Frick, Ann. Ent. Soc. Amer., vol. 49, p. 290, 1956 (figures of antenna, larval characters, and leaf mines). The larvae are unique in having eight to ten bulbs in the posterior spiracles. Many larval host plants in the Compositae are known AGROMYZID LEAF MINERS—FRICK 391 both in Europe and the United States. Frick (1956b) reported the larvae mining Penstemon procerus Dougl., family Scrophulariaceae, in California. The species is widely distributed in the United States and Canada. Phytobia (Calycomyza) ipomoeae (Frost) Agromyza ipomaeae Frost, Ent. News, vol. 42, p. 74, 1931. The type series is teneral and therefore rather difficult to charac- terize fully. However, the forefemur is distally yellow for a distance subequal to its diameter and the midfemora and hindfemora are reddish distally. These characteristics separate Phytobia (Caly- comyza) ipomoeae from P. (C.) ambrosiae and P. (C.) verbenae. The - larvae mine the leaves of [pomoea batatas Poir. in Puerto Rico. Phytobia (Calycomyza) jucunda (van der Wulp) Agromyza jucunda van der Wulp, Tijdschr. Ent., vol. 10, p. 161, 1867. Phytobia (Calycomyza) jucunda (van der Wulp), Frick, Ann. Ent. Soc. Amer., vol. 49, p. 288, 1956 (figures of larval characters and leaf mines). Agromyza platyptera Thomson, Diptera, in Kongliga Svenska Fregatten Eugenies ..., vol. 6, pt. 2, p. 608, 1868. Agromyza coronata Loew, Berliner Ent. Zeitschr., vol. 18, p. 48, 1869 (Centuria vu, No. 89). This species is Nearctic in distribution and is commonly found throughout the United States and Canada. The larvae mine the leaves of a rather large number of plants in the family Compositae. Phytobia (Calycomyza) lantanae Frick Phytobia (Calycomyza) lantanae Frick, Ann, Ent. Soc. Amer., vol. 49, p. 287, 1956. The larvae mine the leaves of Lantana camara L., possibly other Lantana spp., and Lippia helleri Britt. This is a Neotropical species known from southern Texas, Mexico, Puerto Rico, and Trinidad. Phytobia (Calycomyza) majuscula Frick Phytobia (Calycomyza) majuscula Frick, Ann. Ent. Soc. Amer., vol. 49, p. 295, 1956 (figures of larval characters and leaf mines). This is a moderately large species, the larvae of which make blotch mines in the leaves of Senecio lugens var. exaltatus Gray in central California. Also known from Washington, Idaho, Saskatchewan, and Ontario. Phytobia (Calycomyza) malvae (Burgess) Oscinis malvae Burgess, U. S. Dep. Agr. Ann. Rep. (1879), p. 202, 1880. Phytobia (Calycomyza) malvae (Burgess), Frick, Ann. Ent. Soc. Amer., vol. 49, p. 298, 1956. Agromyza jucunda (Van der Wulp), Coquillett (part), U. 8S. Dep. Agr., Div. Ent. Bull. 10, p. 77, 1898. Agromyza cassiae Frost (part), Ann. Ent. Soc. Amer., vol. 29, p. 306, 1936. 392 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The larvae mine the leaves of the following malvaceous plants: Malva rotundifolia L., Abutilon theophrasti Medic., Althaea sp., Malvastrum coromandelianum L., and Sida spinosa L. The species has been collected in scattered locations throughout the United States and from the Panama Canal Zone. Phytobia (Calycomyza) meridiana (Hendel) Agromyza meridionalis Malloch (not Strobl, 1900), Trans. Amer. Ent. Soc., vol. 40, p. 35, 1914. Agromyza meridiana Hendel, Konowia, vol. 2, p. 145, 1923. This species is unique among species of the subgenus Calycomyza in having the third antennal segment yellow. No larval host plants are known. I have seen specimens from Costa Rica and Mexico. Phytobia (Calycomyza) promissa Frick Phytobia (Calycomyza) jucunda (van der Wulp), Frick (part), Univ. California Publ. Ent., vol. 8, p. 395, 1952. Phytobia (Calycomyza) promissa Frick, Ann. Ent. Soc. Amer., vol. 49, p. 287, 1956 (figures of antenna, larval characters, and leaf mines). This species was described from linear-blotch mines in the leaves of Aster chilensis Nees in central California. Several specimens from Manitoba and Ontario were placed under this name because the only observable difference was the slightly larger size. Phytobia (Calycomyza) solidaginis (Kaltenbach) Agromyza solidaginis Kaltenbach, Verh. Naturh. Ver. Preuss. Rheinlande Westfalens, vol. 26, p. 196, 1869. Phytobia (Calycomyza) solidaginis (Kaltenbach), Frick, Ann. Ent. Soc. Amer., vol. 49, p. 292, 1956. The larvae mine the leaves of Solidago spp. Found throughout the United States, eastern Canada, and Europe. Phytobia (Calycomyza) verbenae (Hering) Agromyza jucunda (van der Wulp), Coquillett (part), U.S. Dep. Agr., Div. Ent. Bull. 10, p. 77, 1898. Dizygomyza (Calycomyza) verbenae Hering, Notulae Ent., vol. 31, p. 42, 1951. Phytobia (Calycomyza) verbenae (Hering), Frick, Ann. Ent. Soc, Amer., vol. 49, p. 300, 1956. The larvae mine the leaves of Verbena spp. in the southwestern and eastern United States. The species appears to be common. Si. Subgenus Trilobomyza Hendel Trilobomyza Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 71, 1931. This is a small subgenus that contains three described Nearctic species. Itis characterized by thegenal height not exceeding one-third AGROMYZID LEAF MINERS—FRICK 393 that of the eye (fig. 90), two upper-orbital setae (fig. 89), the lunule low, and the genovertical plates not abruptly raised above the frontal vitta (fig. 88) as they are in species of the subgenus Praspedomyza (fig. 92). The frontal vitta is usually yellow but when it is dark the abdomen is also dark. A small pair of prescutellar setae may be present. The larvae have three short recurved bulbs on each posterior spiracle. The subgenus Trilobomyza is placed among the more spe- cialized subgenera because of the similarity of the adults to those of the subgenus Praspedomyza. One diverse species, Phytobia (Trilobomyza) pleuralis, is included here although it is an example of one that belongs strictly to no one subgenus. However, the adults have more characters in common with species of the subgenus Trilobomyza than with species in the other subgenera. Key to the described species of Phytobia (Trilobomyza) 1. Frontal vitta yellowish; femora yellow .....5......+2404604.8-. 2 Frontal vitta black; femora dark brown. . .......4.2.. calyptrata 2. Mesonotum with four dorsocentrals; anepisternum brown, dorsally about one=founthievellow a sage smh Mtew ee ne io Veg ei oes are an ee ei eae varia Mesonotum with three postsutural dorsocentrals; anepisternum yellow. pleuratis Phytobia (Trilobomyza) calyptrata (Hendel) Agromyza nigrisquama Malloch (not Malloch, 1914), Psyche, vol. 23, p. 53, 1916. Agromyza calyptrata Hendel, Konowia, vol. 2, p. 145, 1923. This species may be separated from others in the subgenus 7'rilo- bomyza by the over-all dark brown coloration, particularly that of the frontal vitta and the femora. The head is illustrated to show the subcircular eye (fig. 89). The holotype from Illinois is the only specimen that I have seen. Phytobia (Trilobomyza) pleuralis (Malloch) Agromyza pleuralis Malloch, Ent. News, vol. 25, p. 311, 1914. Agromyza clara (Melander), Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 41, 1924. Frick (1953a) discussed the subgeneric position of this atypical species, which has more characters in common with subgenus Trilo- bomyza than with any other subgenus. The head has the gena relatively wide (fig. 90). The larvae form serpentine-blotch mines in the leaves of Catalpa spp. At present the known distribution is limited to Illinois and Ohio. 394 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Phytobia (Trilobomyza) varia (Melander) Agromyza varia Melander, Journ. New York Ent. Soc., vol. 21, p. 264, 1913. This species is typical of those in the subgenus Trilobomyza in having yellow frons and femora. The head bears only one lower- orbital (fig. 91) and is of a much different shape than that of Phytobia (Trilobomyza) pleuralis (fig. 90). The dark area on the anepisternum and the four strong dorsocentral setae also serve to separate P. (T.) varia from P. (T.) pleuralis. The holotype and only known specimen is from Idaho. 5j- Subgenus Praspedomyza Hendel Praspedomyza Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 77, 1931. This subgenus is considered the most specialized because of the form of the posterior spiracles of the larvae. Each is circular and bears from 10 to 18 short bulbs. The adults are very similar to those in the subgenus Trilobomyza but may be separated by the broad prominent genovertical plates. These are distinctly above the plane of the frontal vitta and are abruptly raised above it (fig. 92). The adults are usually dark and may be distinguished from the dark Phytobia (Trilobomyza) calyptrata by the aforementioned genovertical plates. There are only two species in North America that belong to the subgenus Praspedomyza sens. str. One, Phytobia (Praspedomyza) morio, is Holarctic while Phytobia (Praspedomyza) subinfumata is Nearctic. Phytobia (Praspedomyza) clara is included here only because the posterior spiracles of the larvae each have 10 to 12 bulbs arranged in a circle. The adults are yellow and the gena is extremely wide, being nearly two-thirds as high as the eye (fig. 93). The Nearctic Phytobia clara and Palaearctic P. hilarella are very similar species that are not closely related to other Phytobia species. The larvae of both mine the fronds of Pteridium aquilinum (L.) Kuhn. Key to the described species of Phytobia (Praspedomy za) 1. Frontal vitta brown or black; femora primarily brown ......... 2 Frontal.vitta yellow; femora iyellowiisry. rid cya ie0e) * (AU OL I Bibs clara 2. Head with three or four lower-orbital setae; femora brown . . subinfumata Head with two lower-orbital setae; femora brown except for forefemur being narrowly yellow: distally oy oo... ic ep te eumelnaeion =) <8 ei ty ee morio Phytobia (Praspedomyza) clara (Melander) Agromyza clara Melander, Journ. New York Ent. Soc., vol. 21, p. 265, 1913. Agromyza citreifrons Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 290, 1913. This is a small (1.3 to 1.75 mm. in wing length) yellowish species that contrasts strongly with the predominately dark species typical AGROMYZID LEAF MINERS—FRICK 395 of the subgenus Praspedomyza. Phytobia hilarella, a very similar species, was placed into Praspedomyza by Hendel because of larval characters. Phytobia clara is retained here for the same reason although the demarcation between the frontal vitta and the geno- vertical plates is not as clearly defined as it is in most species of the subgenus Praspedomyza. I have illustrated the head (fig. 93) to show the contrast between Phytobia clara and P. hilarella, the latter figured by Hendel (1931). The larvae form mines in the fronds of Pteridium aquilinum (L.) Kuhn. I have seen specimens from Cali- fornia, Washington, Michigan, Tennessee, and Maine in the United States and from Ontario, Canada. Phytobia (Praspedomyza) morio (Brischke) Agromyza morio Brischke, Schrift. Naturf. Ges. Danzig, vol. 5, p. 258, 1881. Dizygomyza (Praspedomyza) morio (Brischke), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 80, 1931 (figures of head and wing). This is a small black species that has the third antennal segment and the forefemur distally yellow. Malloch had a Maryland specimen in the U. S. National Museum labeled ‘‘A. luteicornis Malloch MS.” I have reared the species from Galium trifidum L. and G. aparine L. in California. No other North American specimens are known. Phytobia (Praspedomyza) subinfumata (Malloch) Agromyza infumuta Malloch, (not Czerny and Strobl, 1909), Canadian Ent., vol. 47, p. 15, 1915. Agromyza subinfumata Malloch, Proc. U. 8S. Nat. Mus., vol. 49, p. 108, 1915. This species is dark brown and lacks the yellow third antennal segment and distally yellow forefemur of Phytobia morio. The head has the genovertical plates extending beyond the eye margin and bearing three or four lower-orbital setae (fig. 94). Known only from Illinois. 6. Genus Cerodontha Rondani Cerodontha Rondani, Dipt. Ital. Prodr., vol. 4, p. 10, 1861. This is a relatively small genus that is characterized by having only the distal pair of scutellar setae present (fig. 95). The third antennal segment is always angulate distally. Two widely divergent subgenera have been recognized. One, the subgenus Xenophytomyza, could be very properly placed as a synonym of Phytobia (Praspedo- myza) except for the two scutellar setae and the Cerodontha (Cero- dontha) type of male terminalia. 472314594 396 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Key to the subgenera of Cerodontha 1. Third antennal segment distally bearing a spine or strongly projecting into a very acute angle; thorax and abdomen partially yellow. 6a. Cerodontha Third antennal segment with a blunt angie distally; thorax and abdomen shining brown or black. . .......2. 2. =... .6b. Xenophytomyza 6a. Subgenus Cerodontha Rondani Cerodontha Rondani, Dipt. Ital. Prodr., vol. 4, p. 10, 1861. The adults placed in this subgenus are relatively slender and elongate and the body is always partially yellow. The third antennal segment either bears a spine or is produced distally into a very acute angle. Crossvein m-m is always present. There are eight species recognized in Europe but only one in North America. Cerodontha (Cerodontha) dorsalis (Loew) Odontocera dorsalis Loew, Berliner Ent. Zeitschr., vol. 7, p. 54, 1863 (Centuria 111, No. 98). Cerodontha dorsalis (Loew), Melander, Journ. New York Ent. Soc., vol. 21, p. 249, 1913 (figure of head) (east of Rocky Mountains).—Aldrich, Ann. Ent. Soc. Amer., vol. 11, p. 63, 1918. Cerodontha femoralis (Meigen), Melander, Journ. New York, Ent. Soc., vol. 21, p. 249, 1913 (Rocky Mountains, westward). The type is in the Museum of Comparative Zoology (No. 13433). This species varies greatly in the proportions of yellow and black. Aldrich (1918) showed that the dark western form came from the region of cool nights and the more yellowish eastern form is from the region of warm nights. Spring and fall collections, during seasons of cool nights, produced the dark form in the east. Frick (1952) showed that the male terminalia were the same for both color forms. The mesonotum of the holotype female is illustrated to show the color pattern (fig. 95). The larvae mine the leaves of many grasses, pupating in the sheath. Cerodontha dorsalis is one of the most com- mon and widespread species in the United States and Canada. 6b. Subgenus Xenophytomyza Frey Xenophytomyza Frey, Notulae Ent., vol. 26, p. 51, 1946.—Frick, Kansas Ent. Soc., vol. 25, p. 150, 1952 (figure of head). This subgenus contains three species, two Palaearctic and one Nearctic. The adults are robust and shining brown or black. The third antennal segment has a blunt angle distally in contrast to the very acute angle of the species of Cerodontha sens. str. Crossvein m-m is absent in one European species. AGROMYZID LEAF MINERS—FRICK 397 Phytobia (Xenophytomyza) illinoensis (Malloch) Agromyza illinoensis Malloch, Diptera of Patagonia and South Chile, pt. 6, fasc. 5, p. 488, 1934. Cerodontha (Xenophytomyza) illinoensis (Malloch), Frick, Journ. Kansas Ent. Soc., vol. 25, p. 151, 1952 (figure of head). A dark robust species having a blunt angle distally on the third antennal segment. It is most easily separated from the dark Phytobia species by the single pair of scutellar setae. It is known from swept specimens from Illinois and Virginia. 7. Genus Liriomyza Mik Liriomyza Mik, Wiener Ent. Zeit., vol. 18, p. 289, 1894. The genus Liriomyza is large, with 35 described North American species. Of these, 30 are Nearctic, four are Neotropical, and one is found in both regions. Six are Holarctic in distribution. This genus is characterized by the partially or totally yellow scutel- lum, vein M,,. being at or very near the wing tip (fig. 108), and the orbital setulae reclinate (fig. 107) or erect (fig. 110). There are usually numerous orbital setulae but they may be very sparse (fig. 112). Most of the species have some yellow on the head or body. There are a few that are as over-all dark as some species included in Meto- pomyza. The genovertical plates are narrower in Liriomyza species (the widest is shown in fig. 104). These plates are very broad through- out their full length in species of Metopomyza (fig. 128). There are no setae medially on the midtibia except for Liriomyza braziliensis (Frost) and ZL. ecuadorensis (Frost). Although Liriomyza and Haplomyza appear to be separated on rather superficial characters in the key, the separation is justified by the two widely divergent types of male terminalia and larvae found in the two genera. Certain European species of Liriomyza have as many as three of the Haplomyza characters; e. g., if the mes- onotum is heavily gray pollinose and there are two rows of acrostichals and only one upper-orbital, then crossvein m-m is present. In North America, two Haplomyza characters may occur together. As an example, Liriomyza assimilis has the mesonotum dull gray pollinose and has only two rows of acrostichals (fig. 99), but crossvein m-m is present and there are two upper-orbitals (fig. 98). Crossvein m-m is absent in L. chlamydata but the mesonotum is shining and bears four rows of acrostichals and there are two reclinate upper- orbitals. Several unusual characters might be briefly mentioned here. The sorosis group is unusual in having either two postsutural dorsocentrals (fig. 111) or three with the third anterior to the supraalar seta and 398 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 the third and fourth at least two times as far apart as the second and third (fig. 124). The usual arrangement is for the third dorso- central to be opposite or only slightly anterior to the supraalar with the third and fourth dorsocentrals about 1.5 times as far apart as the second and third (figs. 101, 120). JZ. commelinae is the only Liriomyza species known to me that has the third antennal segment of the male enlarged (fig. 102). This characteristic is found in all species of the subgenus Dizygomyza of Phytobia. Liriomyza fumicosta has the costal margin of the wing infuscated (fig. 108). Probably the most distinctive North American species that I have seen is L. deceptiva. The genovertical plates are very prominent and raised above the plane of the frontal vitta (fig. 104). Near the lunule these plates widen to equal about one-third of the width of the frons. The genovertical plates are proportionately broader for their full length in species of Metopomyza (fig. 128). Besides the aforementioned sorosis group, three others, all within the pusilla complex, may be designated. ‘These are based primarily on the colors of the vertex and genovertical plates (Frick, 1956a). The first may be called the trifoli: group (couplet 34). Both vertical setae are on yellow and there are two rows of acrostichals. The second is the eupatorit group (couplet 29). Here the verticals arise from black, the genovertical plates are yellow, and there are four rows of acrostichals. The last group (couplet 27) contains pictella and is named forit. The genovertical plates being infuscated separate this group from that of ewpatori1. The remaining species cannot be placed into groups because of wide differences between them. Several species are herein deleted from the North American list. These include L. pusilla (Meigen) and L. virgo (Zetterstedt), neither of which I have seen in my studies. The South American Agromyza zanthophora (Schiner) apparently belongs to Phytobia (Phytobia) and may be the same as P. (Phytobia) picta. No type specimens of A. zanthophora can be located. Key to the described species of Liriomyza 1. Mesonotum with a prescutellar yellow rectangular or triangular area between the dorsocentral setae extending anteriorly at least to the fourth dorsocentral (figsesO9, MUO)... itis PUPS ee a a aie bist i as dco a me ee 2 Mesonotum without such a prescutellar yellow area but sometimes with a wide yellow crossband, centrally not extending to the fourth dorsocentral (figs! TOS TO9)' SI Ob. SO as Ne Re AED ES RE A, 16 2. Third antennal segment all or partially brown or black (figs. 115, 126). . 3 Third. antennal segmentwyellow: 2.) ¢ 5) je) aya Ry ere @ Laden ety -Dimnoeee 6 3: ,Mesonotum subshining; not pollinoses, 5.42 ek aesslysi iss be ounce mea eeines ewes 4 Mesonotum dull, oray pollinose: gy (2 yc0 cn) vea bey episi ys! aucepee cn cians 3 10. 1: 12. 13. 14. 15. 16. 18. 19: AGROMYZID LEAF MINERS—FRICK 399 . Mesonotum with inner postalar seta about one-third as long as outer postalar (fig. 120); one or two lower-orbital setae .... . . . reyerberata Mesonotum with inner postalar subequal to outer; three lower-orbitals (fig. 2G) sees: ee 2 . . . vVariata . Wing with costal area light browit froth R, ‘eo Ree (Ge 108): sRottelinn without basal lateral black triangles (fig. 109) . ...... .. . . fumicosta Wing with costal area colorless; scutellum with large black triangles, basal scutellar seta on black... . . . . pacifica . Mesonotum with three or four devedcantyal Bean ‘he third ee anterior to the supraalar seta, third and fourth at least two times as far apart as the second and third (Giga. ja Ta Ps bY. a a Le Mesonotum with four dororeaateale| the third opposite or wont slightly anterior to the supraalar, third and fourth about 1.5 times as far apart as the second and third (figs. 101,113)... .. : ms seo . Mesonotum with dark area of two Galore. Biker light reddish Sentry with dark brown lateral stripes (fig. 111), or dark brown centrally and light reddish laterally .... SA oe ce Mesonotum with dark area Gniroimnly ‘dark ca or = black (Ge. 124) . SOrosis . Mesonotum with three dorsocentrals, central area reddish brown, lateral stripes dark brown (fig.111) a . . . marginalis Mesonotum with four dorsocentrals, central area bladk, lateral stripes light reddish. .... - 2... . .. . . . barrocoloradensis . Third antennal Seamierit fe indeah Sore Ay te eer eee | Third antennal segment angulate dorso anteriorly (fig. 96) . . angulicornis Scutellum without basal lateral dark triangles... ........=. 211 Scutellum with basal lateral dark triangles (fig. 101) ......... 12 Mesonotum dark brown; acrostichal setae in four rows, a small intraalar present, about twice as long as an acrostichal. . . . . . . melampyga Mesonotum light reddish brown; acrostichals in two rows; intraalar absent. lutea Anepisternum with a dark oval or triangular area (fig. 100) ...... 18 Anepisternum entirely yellow .-.. . .. . . sehmidti Humerus with a dark area; width of pene nOsenoEs nOt more than one-half the eye height (fig. 98) ..... 8 el a en Humerus entirely ern gena very mide! oniedions subequal to the eye height! Ji 2's 7% MA sheets © 8 Wim a2) Poe epeeeatae Maxillary palpus yellow: aes EN nc) eM an, A Es Ag oe Ott oe Maxillary palpus brown .. . . . . . tubifer Acrostichal setae about 13 in number, j in fours rows; ornenvellar yellow area subrectangular ... . ... . . flavonigra Acrostichals five or six in amber in two TOWS; yellow area triangular (fig. 99) ss ea: . . . assimilis Third antennal eeenint totally dank bone or blacks or if mgatly yellow then darkened distally (fig. 126) .... 5 nM, Sabatta at LO CNG al gi Third antennal segment entirely light yellow? ee al uF Sh) afk. mk 21 . Wing with m-m crossvein eine or if absent, then racerperon at least one- half yellow ... Rue bite LS Wing with m-m crossvein ny absent; aniesepiiiieror blacke. eee 2 ehlasiydats Third antennal segment yellowish, infuscated distally: . .'... ...°. . 19 Third antennal segment entirely black ..... af ee tates Sy 20 Mesepimeron at least one-half yellow; pteropleuron about cuetbalt yellow. langei Mesepimeron black; pteropleuron black or very dark grayish black . dianthi AOO PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 20. 21. 22. Ww on 26. 27. 28. 29. 30. dl. 32. 33. 34, Crossvein m-m about one-fourth of its length from r-m; distally femora brown. quadrisetosa Crossvein m-m about twice its length from r-m; femora brown; distally yellow for a distance subequal to the femoral diameter . . .. . . . baptisiae Anepisternum brown except for a narrow yellow dorsal band subequal in width to aisubure’: (vm eic .- ate oo 2e Anepisternum not more than Mires fouriis proaa Ge 116), dark area usually triangular ... . thas. Zo Genovertical plates yellow; inner C Actinal cn cH eam ellen (fig. 106). discalis Genovertical plates dark from vertex nearly to lunule; both vertical setae arising from black of occiput (fig. 104) . . ........ . deceptiva . Femora primarily yellow, ae marked with brown or black streaks or Spots ea. of De te his) eoprer-erypente thatpee: ee ee: Femora dark Browns Histall one- ‘enard sallow, siete vetye tyeichared., Arieiysthedlaveola {| Calypter, with, margin and fringe blackish) ....)).):.. (eyoelyst eerie sIamfier -§ 25 Calypter with margin and fringe white... . sl dnitioy lima . Head with both vertical setae arising from black of anne inner usually at edge of dark color... . iceventys Hoye? (hive ‘Sita newta- feo Head with both vertical setae on tallow LEE he carted Genovertical plates darkened between orbital mois amie eye aan pened Genovertical plates yellow, concolorous with frontal vitta . . . . . 29 Anepisternum at least one-half brown or black, dark area ee een . 28 Anepisternum less than one-half brown or black, dark area triangular. brassicae Anepisternum about three-fourths dark (fig. 116); first dorsocentral seta small, subequal to an acrostichal in length and about one-fourth the fourth dorsocentral in length. .... . . propepusilla Anepisternum about one-half dark; first dereccann larger, about one-third the length of the fourth Aoreocontral. ote. ee 8 . . . pictella Crossvein m-m not more than its length from r-m; aint section May. about three times as long as penultimate ... . Hreehisfee oo Crossvein m-m 1.5 to two times its length from r-m; ultimate section M344 about two times as long as the penultimate... . - . . eupatorii Inner postalar seta about one-half as long as the outer postalar: seven to nine setulae in the intraalar row posterior to the transverse suture . . munda Inner postalar seta about one-fourth to one-third as long as the outer; about five setulae in the intraalar row posterior to the transverse suture. verbenicola Mesonotum with four irregular rows of acrostichal setae. . ..... 32 Mesonotum with two irregular rows of acrostichal setae. . . ... . . 34 Humerus: without avdark‘spot: is tue SOL tee Ta ag tena, ees Humerus with a dark spot .. . . . . alliovora Head with black of the occiput roaclene oe eye margin (ae 107); anepister- num with a small dark ventral area. . . (de tee ey felti Head with black of the occiput not Posen fone eye rane aes 102); ane- pisternum entirely yellow ... . . . . comme\linze Head with black of the occiput not reaching t to eye margin; inner postalar seta on yellow... .- bie terete gop Head with black of the peciege fenchiae the} eye Sree inner postalar seta on blackijie jscorlie waniniactedn. (wollte Heslaue diel easiteyeytengteoln AGROMYZID LEAF MINERS—FRICK 401 35. Mesonotum dull, heavily gray ere basal scutellar seta arising from black of lateral triangle. . ... wwe, + vallia Mesonotum subshining, not nolrioges basal aeateline ae arising from yellow, at edge of black triangle. . ........ +... phaseolunata Liriomyza allia (Frost) Agromyza allia Frost, Journ. New York Ent. Soc., vol. 51, p. 257, 1943. Liriomyza allia (Frost), Frick, Journ. Kansas Ent. Soc., vol. 28, p. 90, 1955 (figure of head). The holotype male is in the U.S. National Museum (64300). This species varies from Liriomyza alliovora in having only two rows of acrostichals. JZ. allia has the mesonotum heavily gray pollinose in contrast to L. trifolii and L. phaseolunaia, which are subshining and not pollinose. No larval host plants are known. In addition to the holotype there is a paratype male; both are from Manhattan, Kans. Liriomyza alliovora Frick Liriomyza alliovora Frick, Journ. Kansas Ent. Soc., vol. 21, p. 88, 1955 (figure of head). This species differs from Liriomyza felti and L. commelinae in having a dark spot on the humerus and from ZL. allia in having four rows of acrostichals. The head of L. allivora does not have the black from the occiput reaching the eye margin, as does L. felti (fig. 107), and the anepisternum has a ventral black area, unlike L. commelinae. ‘The larvae mine the leaves of Alliwm sp. in Lowa. Liriomyza angulicornis (Malloch) Agromyza angulicornis Malloch, Canadian Ent., vol. 50, p. 79, 1918. Liriomyza angulicornis (Malloch), Hendel, zn Lindner, Die Fliegen der palaearkti- schen Region, fam. 59, p. 208, 1931 (figure of head and wing). This Holarctic species has the third antennal segment angulate dorsoanteriorly and the gena is about one-half as wide as the eye height (fig. 96). The anepisternum has a long slender ventral dark triangle (fig. 97). There are no known larval host plants. The holotype, from Illinois, is the only known North American specimen. Liriomyza assimilis (Mallech) Agromyza assimilis Malloch, Canadian Ent., vol. 50, p. 80, 1918. The head is entirely yellow in profile and both vertical setae are on yellow (fig. 98). The yellow prescutellar area is triangular in shape (fig. 99) and the dark ventral spot on the anepisternum is ovoid (fig. 100). Frost has a series of specimens reared from linear mines in the leaves of Helianthus sp. at Arendtsville, Pa. The holotype is from Hhinois. 402 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 108 Liriomyza baptisiae (Frost) Agromyza baptisiae Frost, Canadian Ent., vol. 63, p. 275, 1931. The holotype male is in the U. S. National Museum (No. 62962). The type series is in rather poor condition but this species belongs to those species that have the third antennal segment at least partially dark and lack the prescutellar yellow spot on the mesonotum. ‘This is a small species (1.25 to 1.5 mm. in wing length) that has the third antennal segment totally black, the femera distally yellow, and the wing with crossvein m-m about twice its length from r-m. The larvae form linear-blotch mines in the leaves of Baptisia tinctoria (L.) in Pennsylvania. Liriomyza barrocoloradensis (Frost) Agromyza barrocoloradensis Frost, Ann. Ent. Soc. Amer., vol. 29, p. 300, 1936. The holotype male is in the U. 8. National Museum (No. 62967). The head in profile is similar to that of Liriomyza sorosis (fig. 125). The mesonotum is shining and is marked much like that of L. mel- ampyga (fig. 113) except that the marginal dark stripes are light reddish. The anepisternum is entirely yellow. The larvae mine the leaves of an unidentified plant in the Panama Canal Zone. Liriomyza borealis (Malloch) Agromyza borealis Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 280, 1913 (figures of head and wing). The head of the holotype is partially collapsed but posteriorly the gena is as wide as the eye is high. The maxillary palpus is yellow. The mesonotum is marked as in the figure and the basal pair of scutellar setae arises from yellow (fig. 101). This species is known only from the holotype from British Columbia. Liriomyza brassicae (Riley) Agromyza brassicae Riley, U. S. Dep. Agr. Ann. Rep. (1884), p. 322, 1884. Liriomyza brassicae (Riley), Frick, Pan-Pacific Ent., vol. 33, p. 68, 1957. Agromyza diminuta (Walker), Coquillett (part), U. S. Dep. Agr., Div. Ent., Bull. 10, p. 78, 1898 (specimens from cabbage). Phytomyza mitis Curran, Canadian Ent., vol. 63, p. 97, 1931. Agromyza subpusilla Frost (part) (not Malloch, 1914), Journ. New York Ent. Soc., vol. 51, p. 255, 1943 (figure of mine of nasturtium leaf). Liriomyza brassicae belongs to the group having yellow antennae, both varietal setae on the dark color of the vertex, and the genovertical plates darkened between the orbital setae and the eye margin. Of the three species in this group, L. brassicae, L. propepusilla, and L. pictella, the latter two have the anepisternum at least one-half dark, the dark area being subrectangular. The anepisternum of L. brassicae is subtriangular. The larvae mine the leaves of a large number of AGROMYZID LEAF MINERS—FRICK 403 cruciferous plants and Tropaeolum sp. (nasturtium). I have seen specimens from many of the States throughout the United States and from Manitoba in Canada. Liriomyza brassicae is also a well known species in Europe. Liriomyza chlamydata (Melander) Antineura chlamydata Melander, Journ. New York Ent. Soc., vol. 21, p. 250, 1913. This species belongs to the group without a prescutellar yellow spot on the mesonotum and with the third antennal segment partially dark. This species may be separated from the other three with a darkened third antennal segment by the lack of crossveinm-m. How- ever, crossvein m-m is sometimes absent in Liriomyza langei, and L. chlamydata may be confused with those specimens of L. langer that lack the m-m crossvein. JL. chlamydata may be separated by the setulae of the third antennal segment being as long as the basal di- ameter of the arista, six or seven very strong orbital setulae that are about one-fourth as long as the orbital setae, the humeral seta on black, and the black mesepimeron. The holotype and only known specimen is from the State of Washington. Liriomyza commelinae (Frost) Agromyza commelinae Frost, Ent. News, vol. 42, pl 72, 1931. Liriomyza commelinae (Frost), Silva and Oliveira, Rev. Brasil. Biol., vol. 12, p. 293, 1952 (figures of adult and larval characters and leaf mines). The holotype male is in the U. 8S. National Museum (No. 62960). This species has the third antennal segment of the male greatly en- larged and the gena about one-third as wide as the eye height (fig. 102). The mesonotum is broadly yellow posteriorly but lacks a cen- tral rectangular or triangular area extending anteriorly of the fourth dorsocentral (fig. 103). The larvae form serpentine mines in the leaves of Commelina elegans H. B. K., C. longicaulis Jacq., and C. virginica L. I have seen specimens from St. Vincent and Trinidad in the British West Indies. This species is known from Brazil and apparently was redescribed by Blanchard (1954) as L. bahamondesi, from Commelina virginica L. in Argentina. Liriomyza deceptiva (Malloch) Agromyza deceptiva Malloch, Canadian Ent., vol. 50, p. 78, 1918. Liriomyza deceptiva is the most diverse North American species of Liriomyza known tome. The head is unique in having the genoverti- cal plates broadened immediately above the antennae (fig. 104). In profile the head has the genovertical plates strongly extending beyond the eye margin (fig. 105). The pleura and abdomen are all black. There are two females that represent this species, one from Illinois and one from Virginia. 404 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Liriomyza dianthi Frick Liriomyza dianthi Frick, Proc. Ent. Soc. Washington, vol. 60, No. 1, p. 1, 1958 (figures of head and pleura). This species is very black with relatively few yellow markings. It is similar to Liriomyza quadrisetosa and L. baptisiae, but differs in having the third antennal segment mostly yellowish and darkened distally. L. dianthi is very similar to L. langei but is darker over-all, having the mesepimeron and pteropleuron black or very dark grayish, the femora about three-fourths black, and the scutellum narrowly yellow with the distal scutellar setae being on the black triangles or not more than asetal baseremoved from them. L. dianthiis at present confined to the San Francisco Bay Area of California. The larvae make large white serpentine mines in the leaves of Dianthus caryophyllus L. Liriomyza discalis (Malloch) Agromyza discalis Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 277, 1913 (figure of head). This is a very dark species of Liriomyza that has the pleura black except for the sutures. The head has the inner vertical seta on yellow and has a few sparse reclinate orbital setulae (fig. 106). The margin and fringe of the calypter are white. The holotype is from Arizona and is the only known specimen. Liriomyze eupatorii (Kaltenbach) Argromyza eupatorii Kaltenbach, Die Pfanzenfeinde aus der Klasse der Insekten, p. 320, 1874. Liriomyza eupatorti (Kaltenbach), Frick, Canadian Ent., vol. 85, p. 72, 1953. This is a Holarctic species that is little known in North America. It is close to Liriomyza munda and L. verbenicola in having the genovertical plates yellow. The most obvious distinction of L. eupatorit is crossvein m-m being 1.5 to 2 times its length from r-m, m-m being about at its own length in the other two species. The larvae make serpentine mines, usually beginning as a spiral, in Solidago spp., including elongata Nutt. in Washington. In Europe also found mining species of Hupatorium, Lampsana, and Galeopsis. Thus far found only in Washington. Liriomyza felti (Malloch) Agromyza felti Malloch, Ent. News, vol. 25, p. 310, 1914.—Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 43, 1924 (figures of leaf mines). The holotype male is in the collection of the New York State Museum, Albany, N. Y. The head has the black of the occiput broadly reaching the eye margin and the eye is smoothly rounded anteriorly (fig. 107). This species differs from Liriomyza commelinae in having the gena centrally about one-fifth the eye height (about AGROMYZID LEAF MINERS—FRICK 405 one-third in commelinae) and in a small dark ventral spot on the anepisternum. The larvae make blotch mines in the leaves of the ferns Camptosorus rhizophyllus (L.) and Asplenium pinnatifidum Nutt. Known from Tllinois and New York. Liriomyza flaveola (Fallén) Argromyza flaveola Fallén, Diptera sueciae, vol. 2, No. 37 (Agromyzides), p. 6, 1823° Liriomyza flaveloa (Fallén), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 219, 1931 (figures of head and wing). Agromyza scutellata (Fallén), Malloch (part), Ann. Ent. Soc. Amer., vol. 6, p. 280, 1913 (1, Dunoon, Scotland). This species may be distinguished from the other species in the so-called pusilla complex by having the femora basally black for two-thirds of the length and yellow distally. The anepisternum is about one-half black with the dark area triangular. The abdomen is broadly yellow laterally. The larvae mine the leaves of grasses and I have reared it from Bromus carinatus H. & A., Hordeum murinum L., and Lolium multifiorum Lam. Thus far this Holarctic species is known in North America only from central California. Liriomyza flavonigra (Coquillett) Agromyza flavonigra Coquillett, Journ. New York Ent. Soc., vol. 10, p. 189, 1902— Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 281, 1913 (figure of head). Agromyza melampyga (Loew), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 258, 1913 (16, New Mexico). A rather large species (2.5 to 2.75 mm. in wing length) that has the genovertical plates extending beyond the eye margin. The mesono- tum is dull but not pollinose and bears four developed dorsocentral setae, all on yellow. The abdomen is mostly yellow. None of the three known specimens has been reared. All are from New Mexico and Melander’s is from Cloudcroft at an elevation of 9,000 feet. Liriomyza fumicosta (Malloch) Agromyza fumicosta Malloch, Ent. News, vol. 25, p. 310, 1914. Unique among Liriomyza species in having the costal area light brown (fig. 108). The mesonotum is slightly lighter brown centrally than the marginal stripes, there are two sparse rows of acrostichals, and the scutellum is entirely yellow (fig. 109). There are two known specimens, both from Illinois. Liriomyza langei Frick Agromyza orbona (Meigen), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 258, 1913 (29 9, Pullman, Wash., from peas). Liriomyza langet Frick, Pan-Pacific Ent., vol. 21, p. 81, 1951 (figure of anepister- num); Proc. Ent. Soc. Washington, vol. 60, No. 1, p. 1, 1958 (figures of head and pleura). 406 PROCEEDINGS OF THE NATIONAL MUSEUM YOL. 108 This species has the third antennal segment darkened distally and the femora marked with yellow streaks. Crossvein m-m is sometimes lacking. Liriomyza langei may be separated from L. chlamydata by having the mesepimeron at least one-half yellow, humeral seta mostly on yellow, and the setulae of the third antennal segment about one-half as long as the basal diameter of the arista. Of 1,000 specimens examined, 81.7 percent have m-m present in both wings, 5.7 percent with one m-m complete and one vestigial, 1.0 percent with one cross- vein complete and one absent, 6.8 percent with m-m vestigial in both wings, 2.9 percent with one vestigial and one absent, and 1.9 percent with crossvein m-m absent in both wings. Two similar species, L. quadrisetosa and L. baptisiae, have the third antennal segment black and the femora black except for being narrowly yellow distally. L. langei is very similar to L. dianthi. However, L. langei is over-all more yellow and has the pteropleuron and mesepimeron each about one-half yellow and the femora streaked with yellow. The larvae of L. langet have been found mining the leaves of a wide range of plants as follows: Allium cepa L. (Liliaceae), Beta vulgaris L. and Spinacia oleracea Li. (Chenopodiaceae), Brassica oleracea var. botrytis L. (Cruciferae), Pisum sativum L. (Leguminosae), Apium graveolens var. dulce Pers. (Umbelliferae), and Petunia sp. (Solanaceae), Aster spp., and Lactuca sativa L., cultivated varieties (Compositae). Thus far no uncultivated plants have been found as larval hosts. L. langei has been identified from Washington, Oregon, and California. Liriomyza lima (Melander) Agromyza lima Melander, Journ. New York Ent. Soc., vol. 21, p. 265, 1913. Agromyza holti Malloch, Canadian Ent., vol. 56, p. 191, 1924. This species is distinctive in the “pusilla complex” in having the margin and fringe of the calypter white. The head has a very wide gena and the eye subquadrate (fig. 110). Melander’s series is from Idaho while Malloch’s specimen is from South Dakota. Liriomyza lutea (Meigen) Agromyza lutea Meigen, Systematische Beschreibung der bekannten europaischen zweifliigeligen Insekten, vol. 6, p. 177, 1830. Liriomyza lutea (Meigen), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 230, 1931 (figures of head of wing).—Spencer, Ent. Gaz., vol. 5, p. 185, 1954. Liriomyza lutea is a very yellow species that has the scutellum entirely yellow. It is close to L. melampyga but differs in having only a few setulae on the mesonotum and the acrostichals in two rows. The dark color of the mesonotum is light reddish brown as contrasted with AGROMYZID LEAF MINERS—FRICK 407 the dark brown of LZ. melampyga. Spencer in England has recently reared this species for the first time from Asplenium rutamuraria L. I have identified one specimen from Alaska that was compared with two from Europe. Liriomyza marginalis (Malloch) Agromyza marginalis Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 283, 1913. This species is closely related to Liriomyza sorosis and L. barro- coloradensis but has only three dorsocentrals, one presutural and two postsutural (fig. 111). The mesonotum has the central area light reddish and the marginal stripes dark brown, whereas in L. barro- coloradensis the colors are reversed and in L. sorosis the dark area is uniformly dark brown. The head has the eye widest below the midline (fig. 112), while it is above themidline in the other two species (fig. 125). The larvae mine in Paspalum dilatatum Poir. I have seen specimens from Illinois, Texas, and South Carolina. Liriomyza melampyga (Loew) Agromyza melampyga Loew, Berliner Ent. Zeitschr., vol. 13, p. 48, 1869 (Centuria vi, No. 88)—Melander (part), Journ. New York Ent. Soc., vol. 21, p. 258, 1913 (19, Massachusetts). Agromyza flaviventris Johnson (not Strobl, 1898), Canadian Ent., vol. 34, p. 242, 1902. This species has the scutellum entirely yellow and the mesonotum bearing four developed dorsocentral setae (fig. 113). The mesonotal color pattern is like that of L. barrocoloradensis but is all dark brown. This species is close to L. lutea but the mesonotal dark area of L. lutea is light reddish brown and there are only two rows of acrostichals. The head used for the illustration was not that of the teneral type but of a female from Glen Echo, Md. (fig. 114). The larval host plants remain unknown. I have seen specimens from Michigan, District of Columbia, Maryland, New Jersey, New York, and Massachusetts. Liriomyza munda Frick Liriomyza munda Frick, Pan-Pacific Ent., vol. 33, p. 61, 1957. This species may be separated from Liriomyza eupatorit by having crossvein m-m at its own length from r-m and from L. verbenicola by a long inner postalar seta (about one-half the length of the outer) and with about twice as many setulae (seven to nine) in the intraalar row posterior to the transverse suture. The larvae make serpentine mines in the leaves of Lycopersicon esculantum Mill., rarely Solanum tuberosum L., and the native Datura meteloides DC. in California. 408 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Liriomyza pacifica (Melander) Agromyza pacifica Melander, Journ. New York Ent. Soc., vol. 21, p. 264, 1913. Agromyza longispinosa Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 276, 1913. This species is one of two having the mesonotum dull and pollinose and the third antennal segment black. The other, Liriomyza fumi- costa, varies in having the costal area tinged light brown and a com- pletely yellow scutellum. The head of L. pacifica is mostly yellow and the black of the occiput does not touch the eye margin (fig. 115). There is only one lower-orbital seta. Larval host plants are un- known. The species has been found in Washington, Illinois, Indiana, Michigan, Alaska, and from British Columbia and Ontario. Liriomyza phaseolunata (Frost) Agromyza phaseolunata Frost, Journ. New York Ent. Soc., vol. 51, p. 256, 1943. This species has two rows of acrostichals and is therefore closely related to Liriomyza trifolii and L. allia. L. phaseolunata is more yellow than L. trifolit and does not have the black of the occiput reaching the eye margin, and the inner postalar seta is on yellow. In contrast to LD. allia. the mesonotum is subshining and not polli- nose. The larvae mine the leaves of Phaseolus limensis Macfad. in New Jersey. Liriomyza pictella (Thomson) Agromyza pictella Thomson, Diptera, in Kongliga Svenska Fregatten Hugenies ..., vol. 6, pt. 2, p. 609, 1868. Liriomyza pictella (Thomson), Frick, Pan-Pacific Ent., vol. 33, p. 66, 1957. Liriomyza pictella is similar to L. propepusilla but differs in hav- ing the first dorsocentral developed and about one-third as long as the fourth dorsocentral. The anepisternum is about one-half yellow while that of L. propepusilla is about three-fourths dark (fig. 116). The larvae form serpentine mines in the leaves of about 16 species of native and cultivated plants throughout California. Frick (1957b) has tabulated all of the host plant and locality records. Since that paper was published I have seen a single specimen from Arizona, swept from alfalfa. Liriomyza propepusilla (Frost) Agromyza subpusilla Frost (not Malloch, 1914), Journ. New York Ent. Soc., vol. 51, p. 255, 1943. Phytomyza subpusilla Frost, Ent. News, vol. 65, p. 73, 1954. Liriomyza propepusilla Frost, Ent. News, vol. 65, p. 73, 1954.—Frick, Pan- Pacific Ent., vol. 33, p. 62, 1957. The name of this species erroneously has been associated with the common California leaf miner, Liriomyza pictella. L. propepusilla is darker over-all and has the episternum about three-fourths dark (fig. AGROMYZID LEAF MINERS—-FRICK 409 116). The first dorsocentral is very short and subequal to an acro- stichal seta in length. The head (fig. 117), wing (fig. 118), and anepisternum (fig. 116) are illustrated in order to help characterize this species, which is at present known from a single male from Kansas. Liriomyza quadrisetosa (Malloch) Agromyza quadrisetosa Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 332, 1913. The head of this rather diverse species is subtriangular in profile, and the genovertical plate is darkened from the vertex for about one- third of the length and bears one upper-orbital and four or five lower- orbitals (fig. 119). The third antennal segment is black, the femora are totally brown, and the wing has crossvein m-m about one-fourth of its length from r-m. There are specimens from Texas, Indiana, and New York. Liriomyza reverberata (Malloch), new combination Agromyza reverberata Malloch, 1924, Canadian Ent., vol. 56, p. 191, 1924.—Frick, Univ. California Publ. Ent., vol. 8, p. 375, 1952. Of the four specimens mentioned in Malloch’s description only one could be found; namely, a female paratype in the Canadian National Collection (No. 3398) labeled ‘““Glen Echo, Maryland, V—14—1922 (J. R. Malloch), Agromyza reverberata Paratype.” The other specimens were to remain in Malloch’s personal collection, or so he stated. The U. S. National Museum acquired Malloch’s collection but the types of Liriomyza reverberata were apparently missing. Because this species is quite distinct, a neotype designation is unnecessary. Left in Agromyza by Frick on the basis of the description, the species belongs in Liriomyza. This species is most closely related to ZL. variata but differs in the short inner postalar seta and in having one or two lower-orbitals. The mesonotum is illustrated to show the color pattern and the fully developed intraalar seta (fig. 120). The wing has vein Ro43 undulat- ing (fig. 121). The mesonotum is subshining but not pollinose. There are three specimens from Ontario in the Canadian National Collection together with the paratype from Maryland. Liriomyza schmidti (Aldrich) Agromyza schmidti Aldrich, Proc. Ent. Soc. Washington, vol. 31, p. 89, 1929. This species has the anepisternum entirely yellow, dark basal scutellar triangles present, and both vertical setae arising from the black of the vertex (fig. 122). The wing has the ultimate section of vein M,4. curved (fig. 123). The larvae make serpentine mines in the leaves of Gliricidia sepium (Jacq.) (=G. maculata) in Costa Rica. 410 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Liriomyza sorosis (Williston), new combination Agromyza sorosis Williston, Trans. Ent. Soc. London (1896), p. 429, 1896.— Melander (part), Journ. New York Ent. Soc., vol. 21, p. 258, 1913 (2 #0, Texas and Illinois, excluding 1 o@ from Bolivia). Agromyza melampyga (Loew), Coquillett (part), U. S. Dep. Agr., Div. Ent. Bull. 10, p. 78, 1898.—Malloch (part), Ann. Ent. Soc. Amer., vol. 6, p. 282, 1913 (specimens from Plantago spp.)—Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 47, 1924.—Frick, Univ. California Publ. Ent., vol. 8, p. 404, 1952. Lectotype male (British Museum, Natural History) from windward side of St. Vincent, Windward Islands, West Indies, 1907-66 (H. H. Smith). This specimen is labeled ““Cotype”’ while a headless male in the American Museum of Natural History (No. 20332) and a female in the Snow Entomological Museum are each labeled “‘Type.’”’ None of the British Museum syntypes, according to H. Oldroyd, bears the label “Mt. St. Andreas at Cavalries Forest, 1,200 feet. Oct. 16.” Such a specimen Williston considered as one of the typical specimens in a series that showed such variation that he mentioned the variation. Liriomyza sorosis includes the specimens of Agromyza melampyga (of authors) that had been reared from Plantago major L., P. media L., and possibly other species of Plantago. Liriomyza sorosis is close to L. marginalis and L. barrocoloradensis but has the mesonotum a uniform dark brown (fig. 124). There are usually four dorsocentrals, but frequently the second is greatly reduced or absent. The head is very similar to that of ZL. barrocolora- densis in having the eye widest above the midline (fig. 125). The type series is from St. Vincent, British West Indies, but specimens have been found in the United States from South Dakota, Illinois, Indiana, Michigan, Pennsylvania, Maryland, Virginia, Florida, and Texas. Liriomyza trifolii (Burgess) Oscinis trifolii Burgess, Ann. Rep. U. 8. Dep. Agr. (1879), p. 200, 1880. Liriomyza congesta (Becker), Mitt. Zool. Mus., vol. 2, p. 190, 1903. Agromyza pusilla (Meigen), Frost (part), Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 51, 1924 (figure of leaf mine on Trifolium repens). This species belongs to the group with only two rows of acrostichal setae. It is slightly darker than Liriomyza allia or L. phaseolunata, has the black of the occiput reaching the eye margin, and its mesono- tum has the inner postalar seta on black. The larvae form serpentine mines in the leaves of many species of lezumes in Europe and I have reared it from Trifolium hybridum L., T. repens L., Medicago sativa L., Melilotus alba Desr., M. indica All., and Vicia villosa Roth in Wash- ington. I have seen specimens from Washington, Oregon, California, Indiana, and Florida, but the species is probably present throughout the United States. | | | AGROMYZID LEAF MINERS——FRICK 411 Liriomyza tubifer Melander Liriomyza tubifer Melander, Journ. New York Ent. Soc., vol. 21, p. 266, 1913. This species is similar to Liriomyza borealis in coloration but differs in having the maxillary palpus brown and the gena narrow, posteriorly not more than one-fourth the eye height. The mesonotum has the yellow of the intraalar row reaching anteriorly about two-thirds of the distance to the transverse suture. There is a single female in Melander’s collection from Haiti. Liriomyza variata (Malloch) Agromyza variata Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 277, 1913 (figure of wing). The holotype female is in the Museum of Comparative Zoology (No. 27062). This species is close to Liriomyza reverberata in having the mesonotum subshining and not pollinose. However, both post- alars are subequal in length and the head has the maxillary palpus distally darkened, three lower-orbitals, and the third antennal segment subquadrate and distally darkened (fig. 126). The wing has vein Roy; undulating (fig. 127). Known from a single specimen from Maine. Liriomyza verbenicola Hering Liriomyza verbenicola Hering, Notulae Ent., vol. 31, p. 48, 1951. The holotype female is in the U. S. National Museum (64301). The type is in very poor condition and the head has turned black probably due to slow drying conditions following death. Therefore, the characters used to separate this species from Liriomyza munda are on the mesonotum. JL. verbenicola has a short inner postalar seta (about one-fourth to one-third the outer) and has very few (about five) setulae in the intraalar row posterior to the transverse suture. The larvae form serpentine mines in the leaves of Verbena sp. The type series is from New Mexico. Hering has a mined leaf from Salt Lake City, Utah, that he identified with this species. 8. Genus Metopomyza Enderlein Metopomyza Enderlein, Tierw. Mitteleuropas, vol. 6, No. 3, p. 180, 1936. This is a small genus closely related to Liriomyza. There are about seven Palaearctic species and one from North America. Metopomyza is characterized by the wide, prominent genovertical plates that are abruptly raised above the plane of the frontal vitta (fig. 128). Lirio- myza deceptiva has these plates prominent but they are proportionately narrower except immediately dorsad of the antenna (fig. 104). Species 472314—59——_5 412 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 included here are mostly black or dark brown and usually have some yellow markings. There are a few Liriomyza species that are as dark colored. Metopomyza interfrontalis (Melander) Agromyza interfrontalis Melander, Journ. New York Ent. Soc., vol. 21, p. 263, 1913. Thus far the only species of Metopomyza known in North America, it may be distinguished from Liriomyza species by the broad, prominent genovertical plates (fig. 128). This is a shining black species that has the frontal vitta dark yellow, scutellum medially yellow, calypter with margin and fringe yellow, some abdominal tergites posteriorly yellow, and all femora distally yellow for a distance subequal to the femoral diameter. No larval host plants are known. I have seen specimens from the northern half of California, Washington, Texas, Kansas, Illinois, Michigan, and Virginia. There are many specimens in the Canadian National Collection. 9. Genus Haplomyza Hendel Antineura Melander (not Osten Sacken, 1881), Journ. New York Ent. Soc., vol. 21, p. 219, 1913. Haplomyza Hendel, Ent. Mitt., vol. 3, p. 73, 1914. The characters used in the key to separate Haplomyza from Lirio- myza appear superficial. However, with the combination of four characters—(1) mesonotum dull, heavily gray pollinose, (2) two sparse rows of acrostichals, (3) crossvein m-m absent, and (4) only one upper-orbital (fig. 129)—go wide and clearcut differences in the male terminalia and posterior spiracles of the larvae. Certain species of Liriomyza in Europe have as many as three of the characters occurring together. Among North American Liriomyza not more than any two occur together. On the basis of this restricted concept of characters for Haplomyza species, Antineura chlamydata Melander belongs in Liriomyza because the mesonotum is shining and there are four rows of acrostichals and two upper-orbitals. Haplomyza is a small genus with three North American species. Of the Palaearctic species described by Hendel (1931) only one, Lirio- myza deficiens (Hendel), appears to belong here. Haplomyza lopesi Oliveira and Silva (1954) from Brazil seems to be properly placed. Key to the described species of Haplomyza 1. Inner postalar seta arising from black; scutellum with basal scuteliar seta arising from large lateral black triangle... . . 7a La OAt2 Inner postalar seta arising from yellow; scutellum broxelly allow basal scu- fellararisingefrom yellow 6; .< % a gras ale sled = ve el ae ee AGROMYZID LEAF MINERS—FRICK 413 2. Third antennal segment yellow; scutellum about one-half yellow, distal scutellar seta arising from yellow . . . . . . . togata Third antennal segment brow oe on distal One-thinds souteliurn about one- third yellow, distal scutellar arising from black ...... . . minuta Haplomyza minuta (Frost) Phytomyza minuta Frost (part), Mem. Cornell Univ. Agri. Exp. Sta., vol. 78, p. 86, 1924 (1 9, Fargo, N. Dak.). Haplomyza minuta (Frost), Frick, Canadian Ent., vol. 85, p. 73, 1953. This is the darkest of the three species. The third antennal seg- ment is distally brownish and the scutellum is not more than one- third yellow. The larvae make serpentine-blotch mines in the leaves of Chenopodium album L. in Washington. The lectotype female is from Fargo, N. Dak. Haplomyza palliata (Coquillett) Phytomyza palliata Coquillett, Journ. New York Ent. Soc., vol. 10, p. 191, 1902. The yellowest of the three species, Haplomyza palliata has the black of the occiput not reaching the eye margin, the inner postalar seta on yellow, and the basal scutellar seta on yellow. The larvae mine in Portulaca sp. in New Mexico. Haplomyza togata (Melander) Antineura togata Melander, Journ. New York Ent. Soc., vol. 21, p. 250, 1913. Phytomyza minuta Frost (part), Mem. Cornell Univ. Agri. Exp. Sta., vol. 78, p. 86, 1924 (3, New Mexico, Texas, and Bismarck, N. Dak.).—Frick, Canadian Ent., vol. 85, p. 73, 1953. Haplomyza togata is intermediate between the more yellow H. palliata and the darker H. minuta. The third antennal segment is yellow, the inner postalar is on black, and the scutellum has the basal seta arising from black and the distal from yellow. The head is illustrated to show the single reclinate upper-orbital (fig. 129). The larvae form serpentine-blotch mines in the leaves of Amaranthus hybridus L. and A. retroflecus L. The species is apparently wide- spread, with identified specimens from Washington, California, New Mexico, Texas, Kansas, and North Dakota. 10. Genus Phytoliriomyza Hendel Phytoliriomyza Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 203, 1931 (as subgenus). This genus is characterized by having the orbital setulae proclinate and the costa ending at vein Mi+.. Phytoliriomyza, as its name sug- gests, together with the closely related Xyraeomyia form a connecting link between the more generalized genera and the more specialized genera related to Phytomyza (Frick, 1952). The head shape is very 414 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 similar throughout the genus and is much like that of Xyraeomyia conjunctimontis (fig. 130). Phytolirriomyza may be separated from Xyraeomyia by the fully developed axillary lobe and calypter and the presence of acrostichal setae and crossvein m-m. Phytoliriomyza is a small but widely distributed genus. There are three species from North America, two from Europe, and one each from extreme southern South America, Formosa, and Hawaii. Frick (1953b) gave a key to the species of the world known at that time. Since then the correct status of P. arctica (Lundbeck), P. immaculata (Coquillett), and P. imperfecta (Malloch) has been determined. Key to the described species of Phytoliriomyza 1. Eye bare; anepisternum mostly yellow with a ventral dark area ..... 2 Eye setulose; anepisternum gray, dorsally narrowly yellow. . .. . arctica 2. Vein M344 with ultimate section about 1.5 times as long as penultimate section; abdominal tergites gray in lateral view . . . . . . immaculata Vein M344 with ultimate section about twice as dune: as 5 pemaltiniste section; abdominal tergites yellow in lateral view ....... . . . perpusilla Phytoliriomyza arctica (Lundbeck) Agromyza arctica Lundbeck, Vidensk. Medd. Naturh. For. Kjgbenhavn, vol. 5, p. 304, 1900 (figure of wing). Phytoliriomyza arctica (Lundbeck), Shewell, Canadian Ent., vol. 85, p. 469, 1953. This species is unique among North American species of Phytolirio- myza in having the eye setulose and the anepisternum mostly dark gray. P. montana Frick of Hawaii is the most similar species but has the maxillary palpi black while they are yellow in P. arctica. No larval host plants are known. Lundbeck notes that the species is found especially in willow thickets along the west coast of Greenland as far north as latitude 69°. The types are from Greenland and I have seen two specimens from Yale and Bear Lake, British Columbia. Phytoliriomyza immaculata (Coquillett) Odinia immaculata Coquillett, Journ. New York Ent. Soc., vol. 10, p. 185, 1902. Agromyza immaculata (Coquillett), Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 289, 1913 (figure of wing). Agromyza perpusilla (Meigen), Melander, Journ. New York Ent. Soc., vol. 21, p. 257, 1913. The holotype of Phytoliriomyza immaculata is teneral and therefore easily confused with the more yellowish P. perpusilla. The abdominal tergites of P. immaculata are dark when viewed laterally as contrasted with the laterally yellow tergites of P. perpusilla. Also, the ultimate section of M34, is shorter (1:1.5 vs. 1:2) in P. immaculata. This species is widespread and I have seen specimens from California, Washington, Idaho, New Mexico, South Dakota, Minnesota, Kansas, AGROMYZID LEAF MINERS—FRICK 415 Oklahoma, Missouri, Michigan, Indiana, Kentucky, Tennessee, Maryland, and New Hampshire as well as from British Columbia and Manitoba. Phytoliriomyza perpusilla (Meigen) Agromyza perpusilla Meigen, Systematische Beschreibung der bekannten euro- paischen zweifligeligen Insekten, vol. 6, p. 181, 1830. Phytoliriomyza perpusilla (Meigen), Hendel, in Lindner, Die Fliegen der palae- arktischen Region, fam. 59, p. 303, 1931 (figures of head and wing).—Frick, Univ. California Publ. Ent., vol. 8, p. 411, 1952 (figure of wing). This is the most yellow of the three species and the abdominal ter- gites are mostly yellow when viewed laterally. The wing has the ultimate section of M3., twice as long as the penultimate. Appar- _ ently rare, I have seen specimens from California, Washington, Montana, and New Mexico. ll. Genus Xyraeomyia Frick Xyraeomyia Frick, Univ. California Publ. Ent., vol. 8, p. 412, 1952 (figure of wing). This genus was erected for a single species that is very closely re- lated to Phytoliriomyza. Xyraeomyia and Phytoliriomyza form a con- necting link between the more generalized genera and the more spe- cialized genera related to Phytomyza (Frick, 1952). However, Ayracomyia is the more specialized of the two in having the axillary lobe and calypter greatly reduced, crossvein m—m absent, and no acrostichals. The head has the eye forwardly inclined, the orbital setulae proclinate, and a relatively long arista (fig. 130). Xyraeomyia conjunctimontis Frick Xyraeomyia conjunctimontis Frick, Univ. California Publ. Ent., vol. 8, p. 413, 1952 (figure of wing). This is a small species (1.15 to 1.3 mm. in wing length) that is yellow and brown, the latter being dull gray pollinose. The head has the eye forwardly inclined and an extremely long arista (fig. 130). The arista is slightly longer than the eye is high. The eye is setulose but the setulae are very short. There are no setulae on the mesonotum other than in the dorsocentral rows and the calypter is reduced to a narrow strip with brown margin and fringe. The type series, from central California, was swept. 12. Genus Phytagromyza Hendel Phytagromyza Hendel, Arch. Naturg., Abt. a, vol. 84, p. 145, 1920. This is a moderately large genus of species that vary widely from one another. They all have in common the orbital setulae either absent or reclinate (fig. 131), the costa ending at R,,;, crossvein m—m absent or, when present, distad of crossvein r—m (fig. 134). In fact, the 416 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 position of m—m and the presence or absence of the basal section of M344 are the only characters that separate all species of Phytogromyza from Pseudonapomyza. In Pseudonapomyza, m—m is basal to r-m and appears to be the basal section of M3,, (fig. 135). There are species in Phytagromyza that lack the orbital setulae and have only one upper-orbital, one to five lower-orbitals, the third antennal seg- ment as angulate as it is in Pseudonapomyza (fig. 136), two to seven dorsocentrals, and the intraalar and prescutellar setae present or ab- sent. ‘The larvae of the type species, Phytogromyza flavocingulata (Strobl), mine the leaves of grasses. The larvae of Pseudonapomyza species mine exclusively in species of Gramineae. Phytagromyza and Pseudonapomyza are the only genera in the Phy- tomyza group that have the orbital setulae erect or reclinate. The group of genera closely related to Phytomyza has the costa ending at R45, and M,4. is the weakest vein (figs. 134, 145). There are only five described species in North America, three of which are Nearctic in distribution and two are Holarctic. There are about 27 species described from the Palaearctic region. Key to the described species of Phytagromyza ts Crossvein ‘m—m' present’ (fig/ Toa) Si *2 a SF Eet SRN Me PONE ERS pe AD Crossvein m—m absent... . 4 2. Mouthparts not elongate, in Tengen eee ran thes eye eights (aoe 133); 1 meso- notum dull black, grayish pollinose. . . 3 Mouthparts elongate, in length subequal to choy eye Reieht igs AB1e 3 meso- notum shining dark brown .. . . . . Nitida 3. Head with two or three orbital ABtiiiae (ae 133); aotopieded tanele yellow. plagiata Head with six or seven orbital setulae; notopleural triangle brown. lonicerae 4. Veins Ry4s; and Mj42 with wing tip between them; head, pleura, and legs mostly dark brown. . . RUN Aiea Sorbitalis Vein Mj42 at wing tips heads lenne end jean vealog ... . . . populicola Phytagromyza lonicerae (Robineau-Desvoidy) Phytomyza lonicerae Robineau-Desvoidy, Rev. Mag. Zool., ser. 2, vol. 3, p. 396, 1851. Phytagromyza lonicerae (Robineau-Desvoidy), Hering, Notulae Ent., vol. 31, p. 36, 1951 (figures of larval and puparial characters). This dark Holarctic species has the mesonotum dull black and gray- ish pollinose and the notopleural triangle dark brown. There are about six developed dorsocentral setae, the head is dark except for the yellow dorsal half of the frontal vitta, and the eye is about four-fifths as long as high. The larvae form very wide serpentine mines in the leaves of Lonicera involucrata Banks and Symphoricarpos albus (L.) Blake. The larvae appear early in the spring and there is only one generation a year. I have seen no specimens other than my reared series from central California. AGROMYZID LEAF MINERS—FRICK 417 Phytagromyza nitida (Malloch) Agromyza nitida Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 288, 1913 (figures of head and wing). A diverse species having the mouthparts elongate, Phytagromyza nitida is distinct from other North American species. The head is partially yellow and has the eye much higher than long, only the dorsal upper-orbital reclinate, and the mouthparts elongate (fig. 131). There are only two developed dorsocentral setae. No larval host plants are known. I have seen specimens from Iowa, Illinois, Mary- land, Virginia, and North Carolina. Phytagromyza orbitalis (Melander) Phytomyza orbitalis Melander, Journ. New York Ent. Soc., vol. 21, p. 271, 1913. Phytagromyza orbitalis and P. populicola lack crossvein m-m, but P. orbitalis has the genovertical plates yellow in contrast to the darker frontal vitta, head, and antenna (fig. 132), dark brown mesonotum, pleura, and legs, and the wing tip between R4i; and M,.».._ In common with P. plagiata and P. lonicerae there are about six developed dorso- central setae. The larvae form blotch mines in the leaves of Symphor- icarpos albus (.), S. mollis Nutt., and S. rotundifolius Gray. I have seen mined leaves on Symphoricarpos at Kamiac Butte, Wash., the type locality. Thus far P. orbitalis is known from Washington, Idaho, and California. Phytagromyza plagiata (Melander) Napomyza plagiaia Melander, Journ. New York Ent. Soc., vol. 21, p. 273, 1913. Agromyza brevicostalis Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 283, 1918. This species, in common with Phytagromyza lonicerae, has crossvein m-m present, mouthparts not elongate (fig. 133), and mesonotum dull grayish pollinose with about six developed dorsocentral setae. It may be distinguished by the head having the gena mostly yellow and the two or three orbital setulae (fig. 133). The notopleural triangle is yellow. Larval host plants are unknown and the species is known from two specimens, one from Idaho and the other from Montana. Phytagromyza populicola (Walker) Phytomyza populicola Haliday (MS), Walker, Insecta Britannica, Diptera, vol. 2, p. 247, 1853.—Hendel, Arch. Naturg., Abt. a, vol. 84, p. 154, 1920. Phytomyza populicola Walker, Kaltenbach, Die Pflanzenfeinde aus der Insekten, p. 560, 1874.—Becker, in Becker, Bezzi, et al., Katalog palaéarktischen Dipteren, vol. 4, p. 258, 1905. Phytagromyza populicola (Haliday), Hendel, in Lindner, Die Fliegen der palaeark- tischen Region, fam. 59, p. 289, 1932 (figure of head). This species should be credited to Walker because Haliday did not publish a description. Phytagromyza populicola is a small yellow 418 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 species without crossvein m-m in the wing. It may be separated from P. orbitalis by the yellow head, antenna, pleura, and legs. The mesonotum is yellow with three light reddish brown vittae and vein M42 ends at the wing tip. E. M. Hering collected the greenish blotch mines on leaves of Populus deltoides Marsh. in Ontario and he identified adults in the Canadian National Collection from Ottawa, Ontario. G. E. Shewell kindly presented me with six adults and six larvae. He reports that this species suddenly appeared in large numbers for the first time in 1956. 13. Genus Pseudonapomyza Hendel Pseudonapomyza Hendel, Arch. Naturg., Abt. a., vol. 84, p. 115, 1920. Pseudonapomyza belongs to the Phytomyza group of genera but is most closely related to Phytagromyza in having the orbital setulae - reclinate (fig. 136). The species of Phytagromyza are so diverse that all of the Pseudonapomyza characters but one are to be found among one or more of the Phytagromyza species. Pseudonapomyza may be separated from Phytagromyza by having crossvein m-m basad of crossvein r-m and appearing to be the basal section of M34 (fig. 135). The wings of Phytagromyza species have m-m distal to r-m (fig. 134), or absent. There are three very similar species in the world. Pseudonapomyza atra is Holarctic, P. lacteipennis is Nearctic, and P. spicata (Malloch) is found on Formosa and throughout Micronesia and Hawaii. P. spicata may be separated from the other two by the black tarsi, shining mesonotum, and the presence of both the inner postalar and intraalar setae. All three species have one upper-orbital, three or four lower-orbitals, the third antennal segment strongly angulate (fig. 136), three dorsocentrals, and the wing as described and illustrated (fig. 135). The larvae of two species, P. atra and P. spicata, mine the leaves of grasses and have a row of elongate spines on each body segment. Key to the described species of Pseudonapomy za 1. Tarsi dark brown or black; mesonotum subshining, slightly gray pollinose, one intraalar seta and no inner postalar. . . . . a We Lip SA Tarsi with basal four segments yellowish, distal So ent, dark Grae n; meso- notum dull, bluish pollinose, no intraalar and one inner postalar. lacteipennis Pseudonapomyza atra (Meigen) Phytomyza atra Meigen, Systematische Beschreibung der bekannten europdischen zweifligeligen Insekten, vol. 6, p. 191, 1830. Pseudonapomyza atra (Meigen), Hendel, in Lindner, Die Fliegen der palaeark- tischen Region, fam. 59, p. 302, 1932 (figures of head and wing). Phytomyza acuticornis Loew, Wiener Ent. Monatsschr., vol. 2, p. 78, 1858. Phytomyza nitidula Malloch, Proc. U. 8. Nat. Mus., vol. 46, p. 151, 1913. AGROMYZID LEAF MINERS—FRICK 419 This is a subshining black species that has the tarsi dark brown or black. The mesonotum is only slightly gray pollinose and bears an intraalar seta but lacks the inner postalar. The head is very similar to that of Pseudonapomyza lacteipennis (fig. 136). The larvae mine the leaves of many grasses in Europe and I have reared it from Agro- pyron repens (L.) and Secale cereale L. in Washington. Pseudonapomyza lacteipennis (Malloch) Phytomyza lacteipennis Malloch, Proc. U. 8. Nat. Mus., vol. 46, p. 152, 1913. Phytomyza acuticornis (Loew), Melander, Journ. New York Ent. Soc., vol. 21, p. 269, 1913. This species is unusual in having the basal four tarsal segments yellowish. The mesonotum is dull black and heavily bluish pollinose and lacks an intraalar but bears the inner postalar seta. The head (fig. 136) is typical for a species of this genus in having the third antennal segment angulate, only one upper-orbital seta, and three or four lower-orbitals. No larval host plants are known but they are probably grasses. I have seen specimens from Washington, New Mexico, Kansas, North Dakota, and Michigan. This species is apparently common and widespread in North America. 14. Genus Napomyza Westwood Napomyza Westwood, An introduction to the modern classification of insects, vol. 2, p. 152, 1840. Napomyza is a small genus separated from Phytomyza by the pres- ence of crossvein m-m. Like Phytomyza, the orbital setulae are proclinate (fig. 137). There are about 20 Palaearctic species, one of which is found in North America. There are three in the Nearctic region, with N. lateralis being Holarctic in distribution. None is known from the Neotropical region. Napomyza anomala of Melander is deleted because Melander’s specimen is incorrectly identified. Key to the described species of Napomyza 1. Acrostichal setae in two irregularrows. ........,. 2% 2. se. 2 Acrostical setae in four or five irregular rows . . . . . . . . davisii 2. Wing with vein My42 ending at wing tip; maiany ae not broadened distally .... . . . lateralis Wing with tip Gace ne ane MG: iaheal ney daipne eseanaed distally to about one-half the diameter of the third antennal segment (fig. 138). parvicella Napomysa davisii (Walton) Agromyza davisii Walton, Ent. News, vol. 23, p. 468, 1912.—Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 284, 1918 (figure of wing). This is a valid species and may be separated from the other two Napomyza species by having four or five rows of acrostichal setae and 420 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 the genovertical plates not extending beyond the eye margin (fig. 137). Crossvein m-m is beyond r-m. The larvae mine the leaves of Ranunculus spp. I have seen specimens from Missouri, Indiana, Michigan, Wisconsin, and from Ontario. Napomyza lateralis (Fallén) Phytomyza lateralis Fallén, Diptera sueciae, vol. 2, No. 41 (Phytomyzides), p. 3, 1823. Napomyza lateralis (Fallén), Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 315, 1932 (figures of head and wing). Napomyza lateralis is a Holarctic species that has two rows of acrostichal setae which separate it from WN. davisii. From N. parvicella, N. lateralis may be distinguished by the maxillary palpus not being broadened distally and vein M,4. being at the wing tip. In Europe the larvae mine in the receptacles and the pith of stems of many annual plants. In North America, this species was reared from Achillea millefolium var. lanulosa Piper in California. I have seen specimens from California, Oregon, Colorado, and Alaska. Napomyza parvicella (Coquillett) Agromyza parvicella Coquillett, Journ. New York Ent. Soc., vol. 10, p. 189, 1902.— Malloch, Ann. Ent. Soc. Amer., vol. 6, p. 287, 1913 (figures of head and wing). Napomyza parvicella differs from N. davisii in having only two rows of acrostichals and the genovertical plates extending beyond the eye margin (fig. 138). The maxillary palpus is broadened and _ the wing tip is between veins Ry; and M4, characters which serve to separate N. parvicella from N. lateralis. The eye is setulose. Known from one female from Alaska. 15. Genus Phytomyza Fallén Phytomyza Fallén, Specimen entomologicum novam Diptera disponendi methodum exhibens, p. 21 (No. 67), 1810 (figure of wing). The most specialized North American genus, Phytomyza has the orbital setulae proclinate (fig. 139), crossvein m-m absent, and M4» the weakest vein (fig. 145). Napomyza is the only closely related genus and is separated by having crossvein m-m present. There are only 39 described Phytomyza species in North America as compared to the 201‘ listed for the Palaearctic region in 1936 by Hendel. Since: that’ time”*many more European species have been described. The North American species are rather evenly distributed throughout the genus as divided in Hendel’s key. The first species to be separated out by the use of that key to Palaearctic species is P. trivittata (couplet 7), while the last is P. Joewri, which keys out at the last couplet (No. 236), AGROMYZID LEAF MINERS—FRICK 421 The only closely related species that could be placed into a species group are those that are very similar to P. ilicicola (couplet 36). The larvae of these four species mine the leaves of woody shrubs and trees such as Symphoricarpos spp. (P. periclymeni), Lonicera spp. (P. gregaria), Prunus spp. (P. persicae), and Ilex spp. (P. alicicola). The following species names are not included in the present species list because I have found no specimens that agree with European specimens. These species are: P. affinis Fallén, P. analis Zetterstedt, P. hieracti Hendel, P. nigritella Zetterstedt, and P. obscurella Fallén. 6. 10. 12; 13. 14, . Frons yellow, light orange, or reddish yellow . Scutellum all or partially yellow. . Third antennal segment black or dark phate Te Key to the described species of Phytomyza Frons all or mostly brown or black, sometimes yellow eh Apnseily on Re Scutellum gray or black Bow on Third antennal segment yellow... . . . . major . Head with dorsal upper-orbital seta angetee fen he ventral upper orbital, or absent (fig. 147) ... . 5 Head with both upper-orbitals of renal length aad naan ee 168). trivittata . Vein Re+3 undulating, M142 straight or distally only Vee curved posteriorly ; one lower-orbital . . . . . . ranunculi Vein Rai3 APRS Mi+2 Gndulafing faedially: (Big. 148); tae lower-orbitals (fig. 147) . oo ; . . . . elemativora Femora Dateie or ee Salar a mace wit Shea streaks or spots. . 7 Femora black or gray, sometimes distally yellow for a distance subequal to the femoral diameter. . . . er . Antenna yellow, third segment pores ened aera 5 ead Antenna with third segment black, first and second yellow. . . . nervosa . Mesonotum heavily dull gray pollinose ; head with two upper-orbital setae. 9 Mesonotum shining black; only one upper-orbital. . . . . . . flavicornis . Anepisternum three-fourths dull gray, dorsal one-fourth and a very narrow posterior margin yellow. . . . .. . . rufipes Anepisternum yellow, except for a verittal eb aan panels of about one-half the height and width of the anepisternum . . . . genalis Third antennal segment yellow, sometimes Hankened ee ig, 152)2. oy UL Third antennal segment black. . . . ; : Ska aa Bete’, . Third antennal segment setulose (Gx. 144); tgaing row with two or three setulae anterior to and two or three eae to the transverse suture: . a): . . . auricornis Third antennal sou hent Dares ae 152); SniPaalar row mien about four or five anterior to and about six posterior to the transverse suture. flavinervis Mesonotum laterally yellow, including humerus and notopleural triangle. . 135 Mesonotum laterally brown or black. . . . ieee Oe wal Head with both upper-orbital setae of equal eet aoa Sennen: (fig. 164). .14 Head with dorsal upper-orbital seta shorter than the ventral, orabsent. . .15 Tarsi brown; mesonotum dull gray pollinose; gena snedually about two-fifths as high as the eye height (fig.149) . .... . . . . . delphinii Tarsi yellow; mesonotum subshining brown; gena ape einily about, one-eighth the eye height (fig. 164) .......4.4.4.2.4. 2... . . plumiseta 422 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 15. 16. lies 18. 19. 20. 21. 22. 23. 24. 26. 27. 28. 29. 30. Anepisternum dorsally one-third to one-half yellow; only one developed upper- orbital, seta: .)-.. 5: . . . . albiceps Anepisternum dark, donselly ned more ayia one-fourth pelle two developed upper-orbitals. . . . . . . sphondylii Head with dorsal upperorbial Mportar than the earal us ee (fig. 140), or absent . . . eo Ne we hae Head with both tip perlorbitale or 4 equal lengeli ind! streneh (ge. 166) EV Genovertical plates darkened (fig. 140); intraalar row with six to nine setulae anterior to and about five posterior to the transverse suture . . angelicella Genovertical plates yellow (fig. 141); intraalar row with 10 to 12 setulae an- terior to and about 13 to 15 posterior to the transverse suture . aquilegiana Acrostichal setae in two to five rows. . . . 4) pcos ees LO Acrostichal setae in six to eight rows (see also FeOuieG 33) whos ve, « - dhiecis Acrostichal setae in two rows or absent .............. ,..20 Acrostichal setae in four or five rows. . . . . . . aquilegiana Mesofacial plate, including subantennal peaouna pean or blade asco eb Mesofacial plate yellow, not more than the subantennal grooves dark . . . 23 Hive Daren = slat st °(% “Third Stab a Maate hia itwe babies boned Eye setulose ... . .. . Nigra Mesonotum without aneorucral seine or with ot more han one to four; one lower-orbital present (see also couplet 26). . . . . . . . atricornis Mesonotum with about 14 acrostichals in two rows; dene lower-orbitals present ..... S bk gay Sipe fe wien es aay i lee ke ETT S Forecoxa dark, distally. veitow a Be le hay ees en tee ROAEEe cette GLb rep on teak ae Oe: Forecoxa entirely dark. . .. . y fet Oe Antenna with first and second Eoementa yellowisH: sista not Deatty expanded (fig. IGG) a. sc. ee cure Antenna with first and ene Mee ienis peer or black: ‘grids greatly expanded), 2 12... ¢ . . . eYrassiseta . Head with two strong lower-orbital Betae) (fig. 166); inner postalar seta present... 2% . . . . subtenella Head with one strong Howerornical slebienes with a very small second; inner postalar absent ..... . . . . . plantaginis Mesonotum with at least eight AerostiGhal Bea! in big LOWS!) cic PRE eRe Mesonotum without acrostichal setae, or with not more than one to four (see also couplet 22)... .. . . . atricornis Third antennal segment with eouiee rot iducer than tie basal diameter of the arista (fig. 150)... . ee oS Third antennal segment a Beaune no +6 Tiree ines as ‘Tone as the basal diameter of the arista (fig. 156)... . . + MS to yet a WAetuUCcH Mesonotum with about 15 acrostichal setae; vein 1 WES ss gently undulating (fig. WAS) ee ee are ts . . . atripalpis Meonotum with abet nine aprostidtial setae: vein Mise straight (fig. NOW) et ay ek te vi aura Head with oth upper- Svoital seins of eee length ald ueatent (fig. 159) .30 Head with dorsal SSR shorter than the ventral upper-orbital (fig. 157), Or ADSCHGEy 3 cP as Pe ee ROO Antenna dark brown or : Pine tarsi and tibiae Blok, es or dark reddish brown 3.3" J). itt a atoll Antenna with fist, second, ‘and pasa portion of ae reddish yellow; tarsi and distal portion of tibiae yellow. . ......... =. . agromyzina AGROMYZID LEAF MINERS—FRICK 423 31. Acrostichal setae in two rows. ..........2.2.2.24.2.2.2.. .82 Acrostichal setae in four to eight rows. . . . eee OS 32. Third antennal segment subcircular (fig. 160); first dorseceritral seta about twice as far from the transverse suture asis the second .. . . melanella Third antennal segment subrectangular, longer than wide (fig. 159); first and second dorsocentral setae about equidistant from the transverse suture. marginalis 33. Acrostichal setae in four or five rows . . . Be fa: om, OE Acrostichal setae in six to eight rows (see Fits Eauplet 18) . wet 4 oo Ticis 34. Intraalar row with not more than 16 setulae anterior to and not more than five posterior to the transverse suture; abdomen mostly dark. . . . . 35 Intraalar row with about 24 setulae anterior to and about 12 posterior to the transverse suture; abdomen mostly yellow .. . . . . . . bicolor 35. Forefemur distally yellow to reddish brown; wing veins s basally yellow to light brown; wing length not exceeding 2.25mm... . ee eee OO Ravctemar entirely black; wing veins basally black; wing fength 2.5 to three mm . oe ee es.) 6cigrinervis 36. Recoeuchal setae extending BateKouty i. fe third dorsocentral . . . . 37 Acrostichal setae extending posteriorly to the fourth dorsocentral . . . . 38 37. Eye, viewed in profile, with length at most three-fourths the eye height; mesonotum dull brownish black... . . . gregaria Eye, viewed in profile, with length subequal a eye Gece maceoeatar dull bluish black ..... . . . periclymeni 38. Forefemur distally reddish Baa meenonin Binge dull grayish pollinose. persicae Forefemur distally yellow; mesonotum brown, dull bluish pollinose . ilicicola 39. Head with one or two upper-orbital aera one or two lower-orbitals; wing length upto2mm.... Ss eee 20) Head with one upper-orbital eetas thr ee Aoner orbitals wing easel at least 3mm 3 4 . . . nigripennis 40. Intraalar row ead ap feast three setulae DOseeviOr i fhe transverse suture; inner postalar about one-half as long as the outer. . ....... 41 Intraalar row with none to one setula posterior to the transverse suture; inner postalar small, up to one-third the length of the outer, or absent. minuscula 41. Forefemur distally brown; wing with second costal section about 1.5 times as long as the fourth (fig. 145) . SP a oak . . . centralis Forefemur distally yellow; second costal section ‘at least two times as long as the fourth (fig, 158) 95) 2 2 fe es se se doewia Phytomyza affinalis Frost Phytomyza affinalis Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 84, 1924. The holotype male is in the U. S. National Museum (No. 50025). This species has the frons yellow, mesofacial plate dark, third antennal segment, mesonotal margins, scutellum, and femora dark, and both upper-orbitals of equal length and strength. It differs from Phytomyza atricornis in having about 14 acrostichals in two rows and the head bearing three lower-orbitals and a wide gena (fig. 139). The type series is from Saskatchewan. 424 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Phytomyza agromyzina Meigen Phytomyza agromyzina Meigen, Systematische Beschreibung der bekannten europadischen zweifliigeligen Insekten, vol. 6, p. 191, 1830.—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 336, 1934 (figures of head and wing). This is a rather distinctive Holarctic species that has the frons dark and both upper-orbitals of equal length, the antenna mostly reddish yellow, and the tarsi and tibiae yellow. The larvae make serpentine mines in the leaves of Cornus spp. ‘Thus far known in North America only from California and Washington. Phytomyza albiceps Meigen Phytomyza albiceps Meigen, Systematische Beschreibung der bekannten europii- schen zweifliigeligen Insekten, vol. 6, p. 194, 1830.—Hendel, zn Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 337, 1934 (figures of head and wing). Phytomyza albiceps sens. str. is not well known in North America. It belongs to those species having a yellow frons, dark third antennal segment, scutellum, and femora, and the mesonotum bordered with yellow. The dorsal upper-orbital is absent, or, if present, not de- veloped and about the size of an orbital setula. The minute or absent dorsalmost orbital and the one-third to one-half yellow ane- pisternum separate this species from P. sphondylii. The larvae make serpentine mines in the leaves of Artemisia spp. that are rather dis- tinctive. Most mines are partially along the midrib and have the frass in two alternating lines of pellets. I have reared this Holarctic species from Artemisia vulgaris L. in California. Phytomyza angelicella Frost Phytomyza angelicella Frost, Ann. Ent. Soc. Amer., vol. 20, p. 218, 1927. This species and Phytomyza aquilegiana belong to the assemblage of species having a yellow frons, dark third antennal segment, mesonotal margins, scutellum, and femora, and the dorsal upper-orbital shorter than the ventral. This species is distinguished from P. aquilegiana by having the genovertical plates dark (fig. 140) and relatively few setulae in the intraalar row (six to nine anterior to the suture and about five posterior). ‘The larvae form linear mines in the leaves of Angelica atropurpurea in New York. Phytomyza aquilegiana Frost Phytomyza aquilegiae (Hardy), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 271, 1918 (1 9, Moscow Mountain, Idaho). Phytomyza bipunctata (Loew) Melander (part), Journ. New York Ent. Soc., vol. 21, p. 271, 1913 (1 9, Avon, Idaho). AGROMYZID LEAF MINERS—FRICK 425 Phytomyza plumiseta Frost (part), Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 87, 1924 (specimens from Aqutlegia sp.). Phytomyza aquilegiana Frost, Ann. Ent. Soc. Amer., vol. 23, p. 459, 1930 (figures of puparium and mined leaves). The holotype male is in the U. S. National Museum (No. 50023). This is a common species that is similar to Phytomyza angelicella in having a yellow frons, dark third antennal segment, mesonotal margins, scutellum, and femora, and the dorsal upper-orbital usually shorter than the ventral (fig. 141). The length of this seta is variable and it varies from as long as the ventral upper-orbital to three-fifths its length. There is usually a very small third lower-orbital and the _ genovertical plates are yellow (fig. 141). The mesonotum bears four to five rows of acrostichals, and the intraalar row has numerous setulae (10 to 12 anterior to the suture and 13 to 15 posterior to it). The larvae form blotch mines in the leaves of Aquilegia spp. and Thalictrum spp. I have seen specimens from California, Washington, Idaho, Pennsylvania, and New York. Phytomyza atricornis Meigen Phytomyza atricornis Meigen, Systematische Beschreibung der bekannten europdischen zweifltigeligen Insekten, vol. 7, p. 404, 1838.—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 358, 1934 (figures of head and wing). Phyltomyza chrysanthemi Kowarz, in Lintner, Report of the State Entomologist, in 44th Ann. Rep. New York State Mus., p. 243, 1892.—Smulyan, Massa- chusetts Agr. Exp. Sta. Bull. 157, p. 21, 1914 (figures of all instars and mined leaves). This species is probably the most widespread and omnivorous in the Agromyzidae. It has the frons yellow, third antennal segment, mesonotal margins, scutellum, and femora dark, and both upper- orbitals of equal length and strength. The mesofacial plate may be dark or it may be yellow with the subantennal grooves dark. There is only one lower-orbital. The acrostichals are frequently absent but some specimens have one to four. The larvae form serpentine mines in the leaves of many plants. There are 27 larval host plants listed in the table at the end of this paper. Nearly 300 more larval host plants are known in Europe (Hendel, 1934). I have seen reared specimens from California, Oregon, Washington, Massachusetts, and from British Columbia. Phytomyza atripalpis Aldrich Phytomyza atripalpis Aldrich, Proc. Ent. Soc. Washington, vol. 31, p. 89, 1929. This is a moderately large species (2.25 mm. in wing length) that has the frons yellow, the third antennal segment, mesonotal margins, scutellum, forecoxa, and femora dark. The two upper-orbitals are of equal length and strength (fig. 142) and the mesonotum has two 496 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 rows of acrostichals. From Phytomyza dura this species may be separated by the larger gena (medially two-fifths of the eye height) (fig. 142), about 15 acrostichals, and the wing with M,,, undulating (fig. 143). The larvae mine in the seeds of Anemone multifida Poir. Known only from British Columbia. Phytomyza auricornis Frost Phytomyza auricornis Frost, Ann. Ent. Soc. Amer., vol. 20, p. 217, 1927. The holotype female is in the U. S. National Museum (No. 50032). Phytomyza auricornis and P. flavinervis belong with those species having a yellow frons, dark scutellum and femora, and the third antennal mostly yellow. From P. flavinervis, P. auricornis can be distinguished by its smaller size (2.5 vs. 3 mm. in wing length), the third antennal segment setulose (fig. 144), and the intraalar row barer (2 to 3 vs. 4 to 6 setulae). Known only from the holotype from New York. Phytomyza bicolor Coquillett Phytomyza bicolor Coquillett, Journ. New York Ent. Soe., vol. 10, p. 191, 1902. A rather large species (3 mm. in wing length) that has a dark frons, antenna, and tarsi, two equal upper-orbitals, and four or five rows of acrostichals. Phytomyza bicolor differs from other closely related species by having the abdomen mostly yellow and the intraalar row of acrostichals very setulose, with about 24 anterior to the suture and 12 posterior to the suture. I have seen three specimens, all from New York. Phytomyza centralis Frost Phytomyza centralis Frost, Ann. Ent. Soc. Amer., vol. 29, p. 317, 1936. The holotype male is in the U. S. National Museum (No. 62978). This is a small dark species (about 1.5 mm. in wing length) that has the frons dark, the dorsal upper-orbital small, and with two lower- orbitals.. There are at least three setulae in the intraalar row posterior to the transverse suture, and the inner postalar is about one-half as long as the outer; these characters separate this species from Phyto- myza minuscula. From P. loewii, P. centralis can be distinguished by its fore-femur being distally brown and the wing having the second costal section about 1.5 times as long as the fourth (fig. 145). The larvae mine the leaves of Clematis sp. in Costa Rica. Phytomyza clemativora Coquillett Phytomyza clemativora Coquillett, Proc. Ent. Soc. Washington, vol. 12, p. 131, 1910. A relatively small yellow species with yellow frons and scutellum, dark third antennal segment, and with the mesonotum marked with AGROMYZID LEAF MINERS—FRICK 497 reddish brown (fig. 146). The head is rather denuded, but the dorsal upper-orbital is absent (fig. 147). The frons is wider than long. The mesonotum has the setae as shown, with only about seven acrostichals (fig. 146). The wing has R24; and Ry; straight and very close together and M,;, undulating medially (fig. 148). Known from two males reared from Clematis sp. in southern Texas. Phytomyza crassiseta Zetterstedt Phytomyza crassiseta Zetterstedt, Diptera Scandinaviae, vol. 14, p. 6469, 1860.— Melander (part), Journ. New York Ent. Soc., vol. 21, p. 271, 1913 (2 9 9, Mount Constitution, and 1 9, Chehalis, Wash.; 1 9, Collins, Idaho).— Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 387, 1935 (figures of head and wing). Phytomyza crassiseta is a Holarctic species that has the frons yellow, all three antennal segments, mesonotal margins, scutellum, and femora dark, and both upper-orbitals of equal length and strength. The mesofacial plate is dark and the forecoxa is distally yellow. Charac- ters that separate P. crassiseta from closely related species are the dark first and second antennal segments and the greatly expanded arista. The larvae make serpentine mines in the leaves of Veronica spp. I have seen specimens from Washington and Idaho. All are females because this species is parthenogenetic. Phytomyza delphiniae Frost Phytomyza delphiniae Frost, Canadian Ent., vol. 60, p. 77, 1928.—Griswold, Journ. Econ. Ent., vol. 21, p. 855, 1928. The holotype female is in the U. S. National Museum (No. 50024). This species belongs with those species having a yellow frons, dark third antennal segment, scutellum, and femora, and the mesonotum laterally bordered with yellow. Phytomyza delphinii, like P. plumiseta, has both upper-orbitals of equal length. From P. plumiseta, this species may be distinguished by the wide gena (medially about two- fifths the eye height) (fig. 149), the brown tarsi, and the mesonotum dull gray pollinose. The larvae form blotch mines in the leaves of Delphinium cultorum Voss. I have seen specimens from Ohio, Penn- sylvania, and New York. Phytomyza dura Curran Phytomyza dura Curran, 1931, Amer. Mus. Nov., No. 492, p. 10, 1931. Phytomyza dura has a yellow frons, dark third antennal segment, mesonotal margins, scutellum, forecoxa, and femora. The upper- orbitals are of equal length (fig. 150) and there are two rows of acrostichals. From P. atripalpis this species can be distinguished by its smaller gena (medially .30 of the eye height) (fig. 150) and by 472314596 428 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 having only about 9 acrostichals and vein My. straight (fig. 151). I have seen 16 swept specimens from Quebec. Phytomyza flavicornis Fallén Phytomyza flavicornis Fallén, Diptera sueciae, vol. 2, No. 41 (Phytomyzides), p. 4, 1823.—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 402, 1935 (figures of head and wing). This is a moderately large Holarctic species (2.3 to 2.6 mm. in wing length) that has yellow frons, dark scutellum, and femora mostly yellow. Phytomyza flavicornis is distinct from related species by having one upper-orbital and the mesonotum shining black. P. rufipes and P. genalis have the mesonotum gray pollinose and P. nervosa has the third antennal black. The species has not been reared in North America but the larvae mine the pith of stems of Urtica dioica L. in Europe. I have seen specimens from Indiana and Michigan. Phytomyza flavinervis Frost Phytomyza flavinervis Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 85, 1924. The holotype female is in the U. S. National Museum (No. 50022). This is a large species (3 mm. in wing length) that has the frons yellow, scutellum and femora dark, and the third antennal segment yellow. Phytomyza flavinervis may be separated from P. auricornis by the bare third antennal segment (fig. 152) and the more setulose (4 to 6 vs. 2 to 3) intraalar row posterior to the transverse suture. The two specimens of the type series are all that are known. They are from Texas. Phytomyza genalis Melander Phytomyza genalis Melander, Journ. New York Ent. Soc., vol. 21, p. 272, 1918. A large species (2.75 to 3.25 mm. in wing length) that has the frons yellow, scutellum dark, and the femora mostly yellow. The third antennal segment is yellow and the mesonotum is gray pollinose. The head has the gena extremely wide and the genovertical plates greatly extending beyond the eye margin (fig. 153). From Phyto- myza rufipes, P. genalis may be separated by its anepisternum being more than one-half yellow and with a ventral brown triangle. I have seen specimens only from Illinois. Phytomyza gregaria Frick Phytomyza nigritella (Zetterstedt), Melander, Journ. New York Ent. Soc., vol. 21, p. 270, 1913. Phytomyza obscurella (Fallén), Melander (part), Journ. New York Ent. Soe., vol. 21, p. 270, 1913 (excluding 19, Bellingham, Wash.). Phytomyza gregaria Frick, Ann. Ent. Soc. Amer., vol. 47, p. 371, 1954 (figure of leaf mines). AGROMYZID LEAF MINERS—FRICK 429 Phytomyza gregaria is one of a group of four species having the frons, antenna, and tarsi dark, the forefemur distally yellow to reddish brown, two equally strong upper-orbitals, and four or five rows of acrostichal setae. It may be separated from P. periclymeni, P. persicae, and P. ilicicola in having the eye ovoid, at most three-fourths the eye height, the mesonotum dull brownish black, and with the acrostichal setae extending posteriorly to the third dorsocentral. The larvae make serpentine mines in the leaves of Lonicera involu- crata Banks. As many larvae may be in a singie leaf, they frequently form a blotch. My specimens are from California, and Melander’s are from Washington and Idaho. Phytomysa ilicicola Loew Phytomyza ilicis Loew (not Curtis, 1846), Berliner Ent. Zeitschr., vol. 7, p. 54, 1863 (Centuria 111, No. 99). Phytomyza ilicicola Loew, Berliner Ent. Zeitschr., vol. 16, p. 114, 1872 (Centuria x, p. 290). Phytomyza ilicicola is one of a group of four species having the frons, antenna, and tarsi dark, the forefemur distally yellow to reddish brown, two equally strong upper-orbitals, and four or five rows of acrostichals. It may be separated from P. gregaria, P. periclymeni, and P. persicae in having the forefemur distally yellow, mesonotum brown, dull bluish pollinose, with the acrostichals extending posteriorly to the fourth dorsocentral. From P. ilicis this species may be dis- tinguished by the tan to brown frons, the narrow gena that is medially one-fourth to one-fifth the eye height (fig. 154), acrostichals in about four rows, and the second costal section 1.9 to 2.8 times as long as the fourth (fig. 155). The larvae form serpentine mines in the leaves of Ilex opaca Ait., I. glabra (.) Gray, J. decidua Walt., and J. vomitoria Ait. I have seen specimens from Texas, Alabama, South Carolina, Virginia, District of Columbia, West Virginia, New York, Khode Island, and Massachusetts. Phytomyza ilicis Curtis Phytomyza ilicis Curtis, Gardeners’ Chron., July 4, 1846, p. 444, 1846.—Downes and Andison, Journ. Econ. Ent., vol. 33, p. 948, 1941.—Hendel, zn Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 417, 1935 (figures of head and wing). The frons is yellow on the dorsal half and grayish black below. Therefore, this species could be considered as belonging to either part of couplet 1 in the key and is so treated in the key. One of the more distinctive characteristics of this species is the six to eight rows of acrostichals. Phytomyza ilicis may be separated from P. ilicicola by the following characters: frons about one-half yellow, gena medially about one-third the eye height, acrostichals in six to eight rows, and 430 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 the second costal section from 3 to 3.8 times as long as the fourth. This is an introduced European species whose larvae make serpentine- blotch mines in the leaves of lex aquifolium L. Now positively known from Oregon, Washington, and British Columbia. Phytomyza lactuca Frost Phytomyza lactuca Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 85, 1924 (figure of leaf mines); Canadian Ent., vol. 60, p. 77, 1928.—Needham, Frost, and Tothill, Leaf-mining insects, p. 263, 1928 (figure of leaf mine). This species has the frons yellow and the third antennal segment, mesonotal margins, scutellum, forecoxa, and femora dark. There are two equally strong upper-orbitals and two rows of acrostichals. Phytomyza lactuca may be distinguished by the very long setulae on the third antennal segment (fig. 156). The larvae make serpentine mines in the leaves of Lactuca scariola var. integrifolia (Bogenh.) G. Beck. I have seen specimens from Michigan, New York, and Pennsylvania. Phytomyza loewii Hendel Phytomyza clematidis Loew (not Kaltenbach, 1859), Berliner Ent. Zeitschr., vol. 7, p. 55, 1863 (Centuria 111, No. 100). Phytomyza loewti Hendel, Konowia, vol. 2, p. 145, 1923. Phytomyza nitida Melander (part), Journ. New York Ent. Soc., vol. 21, p. 271, 1913 (1 9, White Plains, N. Y.; 2 #0, Troy and Bovill, Idaho). This is a shining dark brown species with the frons dark, the dorsal upper-orbital very small, and the eye ovoid and about two-thirds as wide as high (fig. 157). It is distinguished from Phytomyza minuscula by the three setulae in the intraalar row posterior to the transverse suture and the inner postalar being at least one-half as long as the outer. P. loewii is close to P. centralis but is different in having the forefemur distally yellow and the second costal section of the wing at least twice as long as the fourth (fig. 158). The larvae make serpen- tine mines in the leaves of Clematis spp. and I have reared it from Clematis ligusticifolia Nutt. in Washington. I have seen specimens from Washington, Idaho, Indiana, District of Columbia, and Louis- jana. Phytomyza major Malloch Phytomyza major Malloch, Proce. U. S. Nat. Mus., vol. 46, p. 150, 1913. This is a large species (3.75 mm. in wing length) that is mostly yellow with the frons and scutellum yellow. The antennae are en- tirely yellow and the mesonotum has four brown vittae between the rows of setae, two between the dorsocentral rows. Both known specimens are teneral females from Labrador. AGROMYZID LEAF MINERS—FRICK 431 Phytomyza marginalis Frost Phytomyza marginalis Frost, Ann. Ent. Soc. Amer., vol. 20, p. 219, 1927. The holotype female is in the U.S. National Museum (No. 50030). This species and Phytomyza melanella are similar in having dark frons, antenna, and tarsi, two equal-sized upper-orbitals, and the acrostichals in two rows. From P. melanella this species can be distinguished by having the third antennal segment subquadrate, the genovertical plates moderately extending beyond the eye margin (fig. 159), and the first and second dorsocentrals equidistant from the transverse suture. Known from two specimens from New York. Phytomyza melanella Frost Phytomyza melanella Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 86, 1924 (figure of wing). The holotype male is in the U. S. National Museum (No. 50026). This species and Phytomyza marginalis are quite similar in having the frons, antenna, and tarsi dark, both upper-orbitals of equal length, and two rows of acrostichals. P. melanella has the third antennal segment rounded and the genovertical plates hardly extending beyond the eye margin (fig. 160). Also, the first dorsocentral is about twice as far from the transverse suture as the second. I hereby restrict this species to California. The specimen labeled as a paratype from Kentucky is in very poor condition and is of another species. Phytomyza minuscula Goureau Phytomyza minuscula Goureau, Ann. Soc. Ent. France, ser. 2, vol. 9, p. 154, 1851.—Frost, Ann. Ent. Soc. Amer., vol. 23, p. 457, 1930 (figures of puparium and leaf mines).—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 433, 1935 (figures of head and wing). Phytomyza nitida Melander, Journ. New York Ent. Soc., vol. 21, p. 271, 1913. Phytomyza aquilegia (Hardy), Coquillett (part), U. 8. Dep. Agr., Div. Ent. Bull. 10, p. 78, 1898 (specimens from Aquilegia sp.)—-Melander (part), Journ. New York Ent. Soc., vol. 21, p. 271, 1913 (2 # o, Ithaca, N. Y., and New Haven, Conn.).—Cory, Journ. Econ. Ent., vol. 9, p. 419, 1916 (figures of leaf mines).—Frost (part), Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 67, 1924 (figure of leaf mines). Phytomyza minuscula is a small dark Holarctic species (1.5 to 2 mm. in wing length) that has the frons dark. The dorsal upper- orbital is minute and the ventral lower-orbital is about one-half the size of the dorsal (fig. 161). This species is characterized by there not being more than one setula in the intraalar row posterior to the trans- verse suture and the inner postalar usually absent. The fourth costal section varies from 2 to 3.4 times as long as the fourth costal. The head has the eye relatively small and the antenna large (fig. 161). 432 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 The larvae form serpentine mines in the leaves of Aquilegia spp. and Thalictrum spp. I have reared it from Agquilegia, cultivated variety; A. truncata F. & M.; A. pauciflora Jepson; Thalictrum, cultivated variety; and 7. fendleri Engelm. I have studied specimens from California, Washington, Idaho, Tlinois, Indiana, District of Columbia, and Connecticut. Phytomyza nervosa Loew Phytomyza nervosa Loew, Berliner Ent. Zeitschr., vol. 13, p. 52, 1869 (Centuria vu, No. 99). The holotype female is in the Museum of Comparative Zoology (No. 13482). This species, together with several other species, has a yellow frons, dark scutellum, and femora primarily yellow. The third antennal segment is black, a character which separates Phy- tomyza nervosa from the other species. There are no acrostichals. The wing is relatively wide for its length and M,4. has a distinct curve outwardly (fig. 162). I have seen specimens from Iowa, Kansas, and District of Columbia. Phytomyza nigra Meigen Phyiomyza nigra Meigen, Systematische Beschreibung der bekannten europais- chen zweifliigeligen Insekten, vol. 6, p. 191, 1830.—Hendel, zn Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 436, 1935 (figures of head and wing). This Holarctic species is unusual in having the eye setulose. It also has a yellow frons and dark third antennal segment, mesonotal mar- gins, scutellum and femora. Both upper-orbital setae are of equal length and strength and the acrostichals are in two rows. The larvae make linear mines at the tips of leaves of Bromus tectorum L., Festuca arundinacea Schreb., F. rubra var. commutata Gaud., Hordeum juba- tum L., Lolium perenne L., and Triticum aestivum L. I have seen specimens from Oregon and Washington reared from the grasses listed here. Phytomyza nigrinervis Frost Phytomyza nigrinervis Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 87, 1924. The holotype male is in the U. S. National Museum (No. 50028). This is the darkest species among those species having a dark frons, antenna, and tarsi, two equally strong upper-orbitals, and the acrosti- chals in four or five rows. The head is distinctively shaped (fig. 163) and is totally brown or black. The femora are not lighter colored distally and the basal wing veins are black. The entire type series is from Colorado. AGROMYZID LEAF MINERS—FRICK 433 Phytomyza nigripennis Fallén Phytomyza nigripennis Fallén, Diptera sueciae, vol. 2, No. 41 (Phytomyzides), p. 2, 1823.—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 439, 1935 (figures of head and wing). Phytomyza clematidis (Loew), Melander, Journ. New York Ent. Soc., vol. 21, p. 269, 1913. This is a shining black species having the frontal vitta brown, the femora distally reddish with the forefemur more yellowish, the calypter with yellow margin and fringe, and the wing strongly tinged brown. Phytomyza nigripennis is a large species (wing length not less than 3 mm.) having one upper-orbital and three lower-orbitals. I have seen specimens of this Holarctic species from Wisconsin and Ontario. Phytomyza periclymeni de Meijere Phytomyza ilicicola (Loew), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 270, 1913 (1c, 19, Oroville, Wash.). Phytomyza nigra (Meigen), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 270, 1913 (1c, Oroville, Wash.). Phytomyza periclyment de Meijere, in Hendel, Wiener Ent. Zeitschr., vol. 39, p. 71, 1922.—Hendel, in Lindner, Die Fliegen der palearktischen Region, fam. 59, p. 452, 1935 (figures of head and wing).—Frick, Ann. Ent. Soc. Amer., vol. 47, p. 373, 1954. This Holarctic species is one of a group of four species having the frons, antenna, and tarsi dark, the forefemur distally yellow to red- dish brown, two equally strong upper-orbitals, and four or five rows of acrostichal setae. Phytomyza periclymeni may be separated from P. gregaria, P, persicae, and P. ilicicola in having the eye subcircular and the mesonotum dull bluish black with the acrostichal setae ex- tending posteriorly to the third dorsocentral. The larvae make blotch mines in the leaves of Symphoricarpos spp. and Lonicera spp. I have seen specimens from Washington and California. Phytomyza persicae Frick Phytomyza nigritella (Zetterstedt), Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 81, 1924 (figure of leaf mine). Phytomyza persicae Frick, Ann. Ent. Soe. Amer., vol. 47, p. 369, 1954. Phytomyza persicae is one of a group of four species having the frons, antenna, and tarsi dark, the forefemur distally yellow to reddish brown, two equally strong upper-orbitals, and four or five rows of acrostichal setae. It may be separated from P. gregaria, P. pericly- meni, and P. ilicicola in having the forefemur distally reddish brown, the mesonotum black, dull gray pollinose, and the acrostichals ex- tending posteriorly to the fourth dorsocentral. The larvae make serpentine mines in the leaves of Prunus persica L. I have not seen 434 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Frost’s specimens from Prunus serotina Ehrh. and Diervilla lonicera, so I restrict the species to peach at this time. I have seen specimens from Ohio, Virginia, Connecticut, and Ontario. Phytomyza plantaginis Robineau-Desvoidy Phytomyza plantaginis Robineau-Desvoidy, Rev. Mag. Zool., ser. 2, vol. 3, p. 404, 1851,—Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 82, 1924 (figure of leaf mine).—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 455, 1935 (figures of head and wing). Phytomyza genualis Loew, Berliner Ent. Zeitschr., vol. 13, p. 52, 1869 (Centuria vit, No. 100). Phytomyza crassiseta (Zetterstedt), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 271, 1913 (2 9 9, White Plains, N. Y., and Collins, Idaho.). This species has the frons and first and second antennal segments yellow, the third antennal segment, mesonotal margins, scutellum, and femora dark, and the forecoxa distally yellow. There are two equally strong upper-orbitals and two rows of acrostichals. It differs from Phytomyza subtenella in having only one strong lower-orbital and the lack of an inner postalar. The larvae make serpentine mines in the leaves of Plantago spp. I have seen this Holarctic species from many widely scattered States throughout the United States. Phytomyza plumiseta Frost Phytomyza plumiseta Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 87, 1924; Ann. Ent. Soc. Amer., vol. 23, p. 460, 1930. The holotype female is in the U. S. National Museum (No. 50027). Phytomyza plumiseta belongs to the assemblage of species that has a yellow frons, dark third antennal segment, scutellum, and femora, and the mesonotum bordered with yellow. Like P. delphinii, this species has both upper-orbitals of equal length and strength. P. plumiseta has a narrow gena (medially about one-eighth the eye height) (fig. 164), the tarsi yellow, and the mesonotum subshining. The wing is illustrated to show the undulating R23 and straight My42 (fig. 165). The larvae make blotch mines in the leaves of Thalictrum polygamum Muhl. and other Thalictrum spp. in Pennsylvania. Phytomyza ranunculi (Schrank) Musca ranunculi Schrank, Fauna Boiea, vol. 3, p. 140, 1803. Phytomyza ranunculi (Schrank), Hendel, in Lindner, Die Fliegen der palaearkti- schen Region, fam. 59, p. 463, 1935 (figures of head and wing).—Hering, Entomon, vol. 1, p. 207, 1949. This species is one in a group of species having a yellow frons and scutellum and dark third antennal segment. The dorsal upper- orbital is definitely shorter than the ventral. Hendel (1935) proposed several varieties in order to distinguish the seasonal color phases of AGROMYZID LEAF MINERS—FRICK 435 this Holarctic species, but he could find no morphological differences between the color phases, even in the male terminalia. Hering (1949) offered a key to subspecies based in part upon morphological differ- ences. Several of the color phases considered by Hendel are known in North America and a key for their separation is presented here (all three subspecies have the mesonotum grayish black with yellow vittae). 1. Mesonotum without a yellow prescutellar area. . . 2... .......2 Mesonotum with a yellow prescutellar area. . . . . . . ranunculi albipes 2. Femora mostly yellow; anepisternum about one-half yellow. ranunculi flavoscuiellata Femora black or brown; anepisternum about one-third yellow. ranunculi praecox The larvae form serpentine mines in the leaves of Ranunculus spp. and Anemone nemorosa L. in Europe. No host plants are known from North America. Phytomyza ranunculi albipes Meigen Phytomyza albipes Meigen, Systematische Beschreibung der bekannten euro- paischen zweifliigeligen Insekten, vol. 6, p. 195, 1830. Phytomyza flava (Fallén), Melander, Journ. New York Ent. Soc., vol. 21, p. 270, 1913. This subspecies is known from three specimens from Washington. Phytomyza ranunculi flavoscutellata Fallén Phytomyza flavoscutellata Fallén, Diptera sueciae, vol. 2, No. 41 (Phytomizides), p. 4, 1823.—Melander (part), Journ. New York Ent. Soc., vol. 21, p. 270, 1913 (40, Portland, Oreg., and Moscow Mountain, Idaho). Phytomyza zetterstedti (Schiner), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 270, 1913 (12, Moseow Mountain, Idaho). There are five specimens in Melander’s collection from Oregon and o> Idaho. Phytomyza tanunculi praecex Meigen Phytomyza rufipes Meigen, Systematische Beschreibung der bekannten euro- paischen zweifliigeligen Insekten, vol. 6, p. 194, 1830. Phytomyza flavoscutellata (Fallén), Melander (part), Journ. New York Ent. Soc., vol. 21, p. 270, 1913 (12, Moscow Mountain, Idaho). At present known from one specimen in Melander’s collection from Idaho. Phytomyza rufipes Meigen Phytomyza rufipes Meigen, Systematische Beschreibung der bekannten euro- pdischen zweifligeligen Insekten, vol. 6, p. 192, 1830.—Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 471, 1935 (figures of head and wing). Another rather large Holarctic species with yellow frons, dark scu- tellum, and mostly yellow femora. The third antennal segment is 436 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 yellow and the mesonotum dull gray pollinose. This species is most closely related to Phytomyza genalis but may be separated by P. rufipes having the anepisternum three-fourths dull gray. The larvae make serpentine mines in the leaves of various crucifers in Europe. The seven North American specimens that I have seen were from central Oregon. Phytomyza sphondylii Robineau-Desvoidy Phytomyza spondylii Robineau-Desvoidy,,Rev. Mag. Zool., ser. 2, vol. 3, p. 400, 1851. Phytomyza sphondylit Robineau-Desvoidy, Hendel, in Lindner, Die Fliegen der palaearktischen Region, fam. 59, p. 483, 1935 (figures of head and wing). Since this Holarctic species was named for the first known larval host plant, Heracleum sphondylium L., the proper spelling should be sphon- dylii and not spondylvi as originally proposed. Phytomyza sphondylii is very similar to P. albiceps in having a yellow frons, dark third an- tennal segment, scutellum, and femora, and the mesonotum bordered with yellow. From P. albiceps this species is distinguished by the two developed upper-orbital setae (although the dorsal is shorter than the ventral) and the anepisternum being three-fourths dark. I have reared this species from serpentine mines in the leaves of Heracleum lanatum Michx. in California. Phytomyza subtenella Frost Phytomyza subtenella Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 89, 1924. The holotype female is in the U. S. National Museum (No. 50021). This species has the frons yellow, the third antennal segment, meso- notal margins, scutellum, and femora dark, and the forecoxa distally yellow. The upper-orbitals are equally strong and the acrostichals are in two rows. From Phytomyza crassiseta this species can be sep- arated by its having the first and second antennals yellow, the arista not greatly expanded, and two strong lower-orbitals (fig. 166). From P. plantaginis this species differs in having two strong lower-orbitals and an inner postalar seta. The wing has M,,. undulating and the anal vein straight (fig. 167). At present known from two specimens from Washington and Wisconsin. Phytomyza trivittata Frost Phytomyza trivittata Frost, Mem. Cornell Univ. Agr. Exp. Sta., vol. 78, p. 89, 1924. This is a species of moderate size (1.75 mm. in wing length) with yellow frons and scutellum and a dark third antennal segment. The head is mostly yellow and has both upper-orbitals of equal length and strength (fig. 168). The wing has the radial sector strongly curved AGROMYZID LEAF MINERS—FRICK 437 forward (fig. 169). The mesonotum is marked with dull black and gray pollinose and is without setulae other than about four on either side anterior to the transverse suture (fig. 170). There are three known specimens: the type from Wells, Nev., and two specimens from the San Jacinto Mountains, Calif. Host Piants oF NortH AMERICAN AGROMYZID LARVAE (Species of Agromyzidae in italic. Plant names in parentheses are known larval hosts in Europe; to date no such host plant, or not more than one, has been reported in North America for each of the species of Agromyzidae so indicated.) Pteridophyta Polypodiaceae Asplenium pinnatifidum Nutt.: Liriomyza felti (Malloch) (Asplenium rutamuraria L.): Liriomyza lutea (Meigen) Camptosorus rhizophyllus (L.) Link: Liriomyza feltt (Malloch) Pteridum aquilinum (L.) Kuhn: Phytobia (Praspedomyza) clara (Melander) Spermatophyta Angiospermae Monocotyledoneae Typhaceae Typha spp.: Phytobia (Dizygomyza) thompsoni Frick (Typha latifolia L.): Phytobia (Dizygomyza) iraeos (Robineau-Desvoidy) Gramineae Festuceae Bromus carinatus Hook. & Arn.: Cerodontha (Cerodontha) dorsalis (Loew), Liriomyza flaveola (Fallén) Bromus tectorum L.: Phytobia (Poémyza) incisa (Meigen), Phytomyza nigra Meigen Dactylis glomerata L.: Cerodontha (Cerodontha) dorsalis (Loew) Eragrostis sp.: Cerodontha (Cerodontha) dorsalis (Loew) Festuca arundinacea Schreb.: Phytomyza nigra Meigen Festuca rubra var. commutata Gaud.: Phytomyza nigra Meigen Poa pratensis L.: Cerodontha (Cerodontha) dorsalis (Loew) Hordeae Agropyron repens (L.) Beauv.: Agromyza niveipennis Zetterstedt, Phytobia (Poémyza) incisa (Meigen), P. (P.) muscina (Meigen), Cerodontha (Cerodontha) dorsalis (Loew), Pseudonapomyza atra (Meigen) Agropyron sp.: Phytobia (Poémyza) inconspicua (Malloch) Elymus canadensis L.: Phytobia (Poémyza) lateralis (Macquart) E. glaucus Buckl.: Phytobia (Poémyza) incisa (Meigen), Cerodontha (Cerodontha) dorsalis (Loew) Hordeum jubatum L.: Agromyza niveipennis Zetterstedt, Phytobia (Poémyza) lateralis (Macquart), Cerodontha (Cerodontha) dorsalis (Loew), Pseudonapomyza atra (Meigen), Phytomyza nigra Meigen H. murinum L.: Phytobia (Poémyza) muscina (Meigen), Cerodontha (Cerodontha) dorsalis (Loew), Liriomyza flaveola (Fallén) H. vulgare L.: Phytobia (Poémyza) incisa (Meigen), Cerodontha (Cerodontha) dorsalis (Loew) 438 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 (Hordeum vulgare L.): Agromyza ambigua Fallén Lolium multiflorum Lam.: Phytobia (Poémyza) muscina (Meigen), Cerodontha (Cerodontha) dorsalis (Loew), Liriomyza flaveola (Fallén) L. perenne L.: Phytomyza nigra Meigen Lolium sp.: Liriomyza flaveola (Fallén) Secale cereale L.: Agromyza niveipennis Zetterstedt, Cerodontha (Cero- dontha) dorsalis (Loew), Pseudonapomyza atra (Meigen) Triticum aestivum L. (including T. sativum Lam. and T. vulgare Vill.): Agromyza nigripes Meigen, A. niveipennis Zetterstedt, Phytobia (Poémyza) incisa (Meigen), P. (P.) lateralis (Macquart), Cerodontha (Cerodontha) dorsalis (Loew), Phytomyza nigra Meigen Aveneae Avena sativa L.: Phytobia (Poémyza) lateralis (Macquart) Agrostidae Agrostis alba L.: Phytobia (Poémyza) incisa (Meigen) (Calamagrostis epigeios (L.) Roth): Agromyza ambigua Fallén Phleum pratense L.: Phytobia (Poémyza) incisa (Meigen), Cerodontha (Cerodontha) dorsalis (Loew) Chlorideae Hleusine indica (L.) Gaertn.: Cerodontha (Cerodontha) dorsalis (Loew) Phalaridae Ehrharta erecta Lam.: Phytobia (Poémyza) muscina (Meigen), Cero- dontha (Cerodontha) dorsalis (Loew) Phalaris arundinacea L.: Phytobia (Poémyza) incisa (Meigen) P. minor Retz.: Cerodontha (Cerodontha) dorsalis (Loew) Paniceae Digitaria sanguinalis (L.) Scop.: Cerodontha (Cerodontha) dorsalis (Loew) Echinochloa crusgalli (L.) Beauv.: Phytobia (Poémyza) muscina (Meigen), Cerodontha (Cerodontha) dorsalis (Loew) Panicum ecapillare L.: Cerodontha (Cerodontha) dorsalis (Loew) P. dichotomiflorum Michx.: Cerodontha (Cerodontha) dorsalis (Loew) P. miliacum L.: Cerodontha (Cerodontha) dorsalis (Loew) Paspalum dilatatum Poir.: Liriomyza marginalis (Malloch) Setaria lutescens (Weigel) Hubb.: Phytobia (Poémyza) incisa (Meigen) S. viridis (L.) Beauv.: Cerodontha (Cerodontha) dorsalis (Loew) Andropogoneae Sorghum vulgare Purs.: Cerodoniha (Cerodontha) dorsalis (Loew) Tripsaceae Zea mays L.: Agromyza parvicornis Loew, Phytobia (Poémyza) incisa (Meigen), P. (P.) lateralis (Macquart) Cyperaceae (Carex spp.): Phytobia (Poémyza) angulata (Loew), P. (Dizygomyza) luctuosa (Meigen), P. (D.) morosa (Meigen) (Scirpus maritimus L.): Phytobia (Dizygomyza) morosa (Meigen) Commelinaceae Commeline elegans H. B. K.: Liriomyza commelinae (¥rost) C. longicaulis Jacq.: Lirtomyza commelinae (Frost) C. virginica L.: Liriomyza commelinae (Frost) AGROMYZID LEAF MINERS—FRICK 439 Juncaceae (Juncus effusus L.): Phytobia (Dizygomyza) luctuosa (Meigen) Juncus xiphioides Meyer: Phyiobia (Icteromyza) longipennis (Loew) (Juncus sp.): Phytobia (Icteromyza) capitata (Zetterstedt) Liliceae Allium cepa L.: Lirtomyza alliovora Frick, L. lange Frick Asparagus officinalis L.: Melanagromyza simplex (Loew) Iridaceae Iris versicolor L.: Phytobia (Dizygomyza) thompsoni Frick Iris spp.: Phytobia (Dizygomyza) thompsoni Frick, P. (D.) traeos (Robineau-Desvoidy), P. (D.) zridis (Hendel) Dicotyledoneae Salicaceae Populus deltoides Marsh: Agromyza albitarsis Meigen, Phytagromyza populicola (Walker) P, nigra var. italica Muenchh.: Agromyza albitarsis Meigen P. trichocarpa T. & G.: Agromyza albitarsis Meigen Populus spp.: Agromyza albitarsis Meigen, Melanagromyza schineri (Giraud) Salix lasiandra Benth.: Agromyza albitarsis Meigen Salix spp.: Melanagromyza salicis (Malloch) (Salix spp.): Melanagromyza schineri (Giraud) Betulaceae Betula nigra L.: Phytobia (Phytobia) pruinosa (Coquillett) Fagaceae Quercus spp.: Agromyza viridula Coquillett Ulmaceae Celtis occidentalis Micocoulier: Agromyza aristata Malloch Ulmus americana L.: Agromyza aristata Malloch Urticaceae Urtica californica Greene: Agromyza reptans Fallén (Urtica dioica L.): Phytomyza flavicornis Fallén Chenopodiaceae Beta vulgaris L.: Liriomyza langei Frick Chenopodium album L.: Haplomyza minuta (Frost) (Chenopodium album L.): Phytobia (Amauromyza) abnormalis (Malloch) Spinacia oleracea L.: Liriomyza langei Frick Amaranthaceae Amaranthus hybridus L.: Haplomyza togata (Melander) A. retroflexus L.: Haplomyza togata (Melander) Amaranthus sp.: Phytobia (Amauromyza) abnormalis (Malloch) Caryophyllaceae Dianthus caryophyllus L.: Liriomyza dianthi Frick Portulacaceae Portulaca sp.: Haplomyza palliata (Coquillett) Ranunculaceae Anemone multifida Poir.: Phytomyza atripalpis Aldrich (Anemone nemorosa L.): Phytomyza ranunculi (Schrank) Aquilegia canadensis L.: Phytomyza aquilegiana Frost A. truncata F. & M.: Phytomyza aquilegiana Frost, P. minuscula Goureau PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 A. truncata var. pauciflora Jepson: Phytomyza aquilegiana Frost, P. minuscula Goureau A. vulgaris L.: Phytomyza aquilegiana Frost Aquilegia spp., cult. vars.: Phytomyza aquilegiana Frost, P. minus- cula Goureau Clematis ligusticifolia Nutt.: Phytomyza loewii Hendel Clematis spp.: Phytomyza centralis Frost, P. clemativora Coquillett, P. loewiit Hendel Delphinium cultorum Voss.: Phytomyza delphiniae Frost Ranunculus abortivus L.: Napomyza davisii (Walton) Ranunculus sp.: Napomyza davisii (Walton) (Ranunculus spp.): Phytomyza ranunculi (Schrank) Thalictrum fendleri Engelm.: Phytomyza aquilegiana Frost, P. minuscula Goureau T. polygamum Muhl.: Phytomyza aquilegiana Frost, P. plumiseta Frost Thalictrum spp., cult. vars.: Phytomyza minuscula Goureau Cruciferae (Armoracia lapathifolia Gilib.): Phytomyza rufipes Meigen Brassica arvensis (L.) B.S. P.: Liriomyza brassicae (Riley) B. campestris L.: Liriomyza brassicae (Riley) B. napus L.: Liriomyza brassicae (Riley) (Brassica napus L.): Phytomyza rufipes Meigen Brassica nigra (L.) Koch.: Liriomyza brassicae (Riley) (Brassica oleracea L.): Phytomyza rufipes Meigen Brassica oleracea var. botrytis L.: Liriomyza brassicae (Riley), L. langet Frick B. oleracea var. capitata L.: Liriomyza brassicae (Riley) Brassica rapa L.: Liriomyza brassicae (Riley) Descurainia (=Sophia) sp.: Ophiomyia texana (Malloch) (Diplotaxis tenuifolia L.): Phytomyza rufipes Meigen Erysimum inconspicuum (S. Wats.) MacM.: Liriomyza brassicae (Riley) Radicula palustris Moench.: Liriomyza brassicae (Riley) Raphanus sativus L.: Liriomyza brassicae (Riley) R. activus var. longipinnatus Bailey: Liriomyza brassicae (Riley) Rorippa sp.: Ophiomyia texana (Malloch) Sisymbrium altissimum L.: Lirtomyza brassicae (Riley) Rosaceae Amelanchier canadensis (L.) Medic.: Phytobia (Phytobia) amelan- chieris (Greene) Fragaria virginiana Duch.: Agromyza spiraeae Kaltenbach Fragaria spp., cult. vars.: Agromyza spiraeae Kaltenbach (Potentilla erecta L.): Agromyza rubi Brischke Prunus avium L.: Phytobia (Phytobia) pruni (Grossenbacher) P. domestica L.: Phytobia (Phytobia) pruni (Grossenbacher) P. persica (L.) Batsch.: Phytomyza persicae Frick ? P. serotina Ehrh.: Phytomyza persicae Frick Rubus idaeus L.: Agromyza spiraeae Kaltenbach R. occidentalis L.: Agromyza spiraeae Kaltenbach Rubus spp., cult. vars.: Agromyza spiraeae Kaltenbach (Rubus spp.): Agromyza rubi Brischke (Sanguisorba officinalis L.): Agromyza rubi Brischke AGROMYZID LEAF MINERS—FRICK 44] Leguminosae Baptisia tinctoria (L.) R. Br.: Liriomyza baptisiae (Frost) Cassia bacillaris L.: Phytobia (Calycomyza) cassiae (Frost) Centrosema pubescens Benth.: Agromyza centrosemae Frost Gliricidia sepium (Jacq.) Steud.: Liriomyza schmidti Aldrich Medicago lupulina L.: Liriomyza pictella (Thomson) M. sativa L.: Melanagromyza gibsoni (Malloch), Liriomyza pictella (Thomson), L. trifolii (Burgess) Melilotus alba Desr.: Liriomyza trifoliit (Burgess) M. indica All.: Liriomyza trifolit (Burgess), Phytomyza atricornis Meigen Phaseolus limensis Macfad.: Agromyza inaequalis Malloch, Lirtomyza phaseolunata (Frost), L. pictella (Thomson) P. vulgaris L.: Liriomyza pictella (Thomson) Pisum sativum L.: Liriomyza langet Frick, Phytomyza atricornis Meigen Trifolium hybridum L.: Liriomyza trifolii (Burgess) T. repens L.: Liriomyza trifolii (Burgess) Vicia gigantea Hook: Liriomyza pictella (Thomson) V. villosa Roth: Liriomyza trifoliz (Burgess) Vigna repens Baker: Agromyza inaequalis Malloch Tropaeolaceae Tropaeolum spp.: Liriomyza brassicae (Riley) Euphorbiaceae Croton billbergianus Muell.: Melanagromyza crotonis (Frost) ? Sauvia sp.: Phytobia (Calycomyza) allecta (Melander) Aquifoliaceae Ilex aquifolium L.: Phytomyza tlicis Curtis I. decidua Walt.: Phytomyza ilicicola Loew I. glabra (L.) Gray: Phytomyza ilicicola Loew I. opaca Ait.: Phytomyza ilicicola Loew I. vomitoria, Ait.: Phytomyza ilicicola Loew Aceraceae Acer rubrum L.: Phytobia (Phytobia) setosa (Loew) Tiliaceae Tilia americana L.: Melanagromyza tiliae (Couden) Malvaceae Abutilon theophrasti Medic.: Phytobia (Calycomyza) malvae (Burgess) Althaea rosea Cav.: Phytobia (Calycomyza) malvae (Burgess) Gossypium spp.: Liriomyza pictella (Thomson) Malva nicaeensis All. (=borealis): Liriomyza pictella (Thomson), Phytomyza atricornis Meigen M. rotundifolia L.: Phytobia (Calycomyza) malvae (Burgess) Malvastrum coromandelianum L.: Phytobia (Calycomyza) malvag (Burgess) Sida spinosa L.: Phytobia (Calycomyza) malvae (Burgess) Umbelliferae Angelica atropurpurea L.: Melanagromyza angelicae (Frost), Phy- tomyza angelicella Frost Apium graveolens var. dulce Pers.: Liriomyza langei Frick Heracleum lanatum Michx.: Phytomyza sphondylit Robineau- Desvoidy 442 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 Cornaceae Cornus californica C. A. Mey.: Phytomyza agromyzina Meigen C. stolonifera Michx.: Phytomyza agromyzina Meigen Convolvulaceae Ipomoea batatas Poir.: Melanagromyza caerulea (Malloch), Phytobia (Calycomyza) ipomoeae (Frost) I. laconosa L.: Melanagromyza caerulea (Malloch) J. sinuata Ort.: Melanagromyza caerulea (Malloch) Ipomoea spp.: Melanagromyza caerulea (Malloch) Boraginaceae Cynoglossum virginianum L.: Phytobia (Calycomyza) cynoglossi Frick Cynoglossum spp.: Phytobia (Calycomyza) cynoglossi Frick Verbenaceae Lantana camara L.: Ophiomyia lantanae (Froggatt), Phytobia (Calycomyza) lantanae Frick Lantana spp.: Ophiomyia lantanae (Froggatt), Phytobia (Calycomyza) lantanae Frick Lippia helleri Britt.: Phytobia (Calycomyza) lantanae Frick Verbena hybrida Voss.: Phytobia (Calycomyza) verbenae (Hering) V. noemexicana (Gray) Small: Phytobia (Calycomyza) verbenae (Hering) V. wrightii Gray: Phytobia (Calycomyza) verbenae (Hering) Verbena sp.: Liriomyza verbenicola Hering Labiatae Mentha sp.: Phytomyza atricornis Meigen Nepeta cataria L.: Ophiomyia proboscidea (Strobl) Stacbys bullata Benth.: Liriomyza pictella (Thomson), Phytomyza atricornis Meigen Solanaceae Datura meteloides DC.: Liriomyza munda Frick, L. pictella (Thomson) Lycopersicon esculentum Mill.: Liriomyza munda Frick Petura sp.: Liriomyza langet Frick Solanum tuberosum L.: Liriomyza munda Frick Scrophulariaceae Penstemon procerus Dougl.: Phytobia (Calycomyza) humeralis (von Roser) Veronica peregrina var. xalapensis (H. B. K.) Pennell: Phytomyza crassiseta Zetterstedt Bignoniaceae Catalpa bignonioides Walt.: Phytobia (Trilobomyza) pleuralis (Malloch) Acanthaceae Justicia (= Dianthera) americana (L.) Vahl.: Melanagromyza dian- thereae (Malloch) Plantaginaceae Plantago lanceolata L.: Phytomyza plantaginis Robineau-Desvoidy P. major L.: Liriomyza sorosis (Williston), Phytomyza plantaginis. Robineau-Desvoidy P. media L.: Liriomyza sorosis (Williston) Plantago sp.: Liriomyza sorosis (Williston) AGROMYZID LEAF MINERS—FRICK 443 Rubiaceae Galium aparine L.: Phytobia (Praspedomyza) morio (Brischke) G. trifidum L.: Phytobia (Praspedomyza) morio (Brischke) Caprifoliaceae Lonicera involucrata (Rich.) Banks: Phytagromyza lonicerae (Robi- neau-Desvoidy), Phytomyza gregaria Vrick Lonicera sp., cult. var.: Phytomyza periclymeni de Meijere Symphoricarpos albus (l.) Blake: Phytagromyza lonicerae (Robineau- Desvoidy), P. orbitalis (Melander), Phytomyza periclymeni de Meijere 5S. mollis Nutt.: Phytagromyza orbitalis (Melander) S. rotundifolius Gray: Phylagromyza orbitalis (Melander) Viburnum pubescens Pursh: Phytobia (Calycomyza) flavinotum Frick Cucurbitaceae Cucumis melo L., cult. vars.: Liriomyza pictella (Thomson) Compositae Cichorieae Lactuca sativa L.: Phylobia (Amauromyza) maculosa (Malloch), Liriomyza langet Frick Lactuca scariola var. integrifolia (Bogenh.) G. Beck: Phytomyza lactuca Frost Picris echioides L.: Phytomyza atricornis Meigen Sonchus asper (L.) Hill: Ophiomyia coniceps (Malloch), Phytomyza atricornis Meigen S. oleraceus L.: Phytomyza atricornis Meigen Taraxacum kok-sghyz Rodin: Phytomyza atricornis Meigen T. officinale Weber: Tylomyza nasuta (Melander) Eupatorieae Eupatorium odoratum L.: Melanagromyza mallochi (Hende!}) i. purpureum L.: Phytobia (Calycomyza) flavinotum Frick Astereae Aster chilensis Nees: Phytobia (Calycomyza) humeralis (von Roser), P. (C.) promissa Frick A. divaricatus L.: Ophiomyia maura (Meigen) A. leavis L.: Phytobia (Nemorimyza) posticata (Meigen) A. novae-angeliae L.: Ophiomyia maura (Meigen), Phytobia (Nemo- rimyza) posticata (Meigen) A. ramosissimus (T. & G.) Crong.: Ophiomyia maura (Meigen), Phytobia (Nemorimyza) posticata (Meigen) A. undulatus L.: Ophiomyia maura (Meigen), Phylobia (Nemorimyza) posticata (Meigen) Aster spp., cult. vars.: Phytobia (Amauromyza) maculosa (Malloch), P. (Calycomyza) humeralis (von Roser), Liriomyza langet Frick, L. pictella (Thomsen) Baccharis douglasii DC.: Phytobia (Amauromyza) maculosa (Malloch), P. (Calycomyza) humeralis (von Roser) Baccharis viminea DC.: Phytobia (Calycomyza) jucunda (Wulp) Erigeron canadensis L.: Phytobia (Amauromyza) maculosa (Malloch), P. (Calycomyza) humeralis (von Roser), P. (C.) jucunda (Wulp) Krigeron sp.: Phytobia (Calycomyza) jucunda (Wulp) Grindelia squarrosa (Pursh) Dunal: Phyiobia (Calycomyza) jucunda (Wulp) 472314——59—_7 444 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Heterotheca grandiflora Nutt.: Phytobia (Calycomyza) humeralis (von Roser), P. (C.) gucunda (Wulp) Solidago bicolor L.: Phytobia (Nemorimyza) posticata (Meigen) S. caesia L.: Ophiomyia maura (Meigen), Phytobia (Nemorimyza) posticata (Meigen), P. (Calycomyza) jucunda (Wulp) S. canadensis L.: Ophiomyia maura (Meigen), Phytobia (Nemorimyza) posticata (Meigen), P. (Calycomyza) jucunda (Wulp) S. elongata Nutt.: Phytobia (Nemorimyza) posticata (Meigen), Lirio- myza eupatortt (Kaltenbach) S. flexicaulis L.: Ophiomyia maura (Meigen), Phytobia (Nemorimyza) posticata (Meigen), P. (Calycomyza) jucunda (Wulp) S. juncea Ait.: Ophiomyia maura (Meigen), Phytobia (Nemorimyza) posticata (Meigen) S. leiophylla Fern.: Ophiomyia maura (Meigen), Phytobia (Nemori- myza) posticata (Meigen) 8S. macrophylla Pursh: Phytobia (Calycomyza) humeralis (von Roser) 8. nemoralis Ait.: Phytobia (Nemorimyza) posticata (Meigen) Solidago spp.: Phytobia (Calycomyza) solidaginis (Kaltenbach) Inuleae Antennaria plantaginifolia (L.) Hook: Phytomyza atricornis Meigen Gnaphalium leucocephalum Gray: Phytomyza atricornis Meigen Heliantheae Bidens frondosa L.: Phyltobia (Calycomyza) allecta (Melander), Phytomyza atricornis Meigen B. pilosa var. radiata Sch.: Phytobia (Amauromyza) maculosa (Malloch) Dahlia pinnata Cav.: Liriomyza pictella (Thompson) Encelia sp.: Melanagromyza viridis (Frost) Helianthus annuus L.: Phytobia (Amauromyza) maculosa (Malloch), P. (Calycomyza) humeralis (von Roser), P. (C.) jucunda (Wulp) Liriomyza pictella (Thomson) H. ealifornicus DC.: Phytomyza atricornis Meigen Helianthus spp.: Phytobia (Calycomyza) allecta (Melander) Parthenium argentatum Gray: Phytomyza atricornis Meigen Rudbeckia laciniata var. hortensis Bailey: Phytobia (Calycomyza) allecta (Melander), P. (C.) artemitsiae (Kaltenbach) Zinnia spp., cult. vars.: Melanagromyza viridis (Frost), Phytobia (Calycomyza) humeralis (von Roser), P. (C.) jucunda (Wulp), Liriomyza pictella (Thomson), Phytomyza atricornis Meigen Madieae Madia elegans Don.: Phytobia (Calycomyza) humeralis (von Roser) Ambrosieae Ambrosia artemisiifolia L.: Phytobia (Calycomyza) ambrosiae Frick, Phytomyza atricornis Meigen A. trifida L.: Phytobia (Calycomyza) ambrosiae Frick, P. (C.) jucunda (Wulp) Xanthium strumarium L.: Phytobia (Calycomyza) jucunda (Wulp) Anthemideae Achillea millefolium var. lanulosa Piper: Napomyza lateralis (Fallén) Artemisia douglasiana Bess.: Phyiobia (Calycomyza) artemisiae (Kaltenbach) AGROMYZID LEAF MINERS—FRICK 445 A. vulgaris L.: Phytobia (Amauromyza) maculosa (Malloch), P. (Calycomyza) artemisiae (Kaltenbach), P. (C.) humeralis (von Roser), Phytomyza albiceps Meigen, P. atricornis Meigen Chrysanthemum frutescens L.: Phyiomyza atricornis Meigen C. indicum L.: Phytobia (Amauromyza) maculosa (Malloch), Phyto- myza atricornis Meigen C. leucanthemum L.: Phytomyza atricornis Meigen C. morifclium Ramat.: Phytomyza atricornis Meigen Chrysanthemum spp., cult. vars.: Phytobia (Amauromyza) maculosa (Malloch), Phytomyza atricornis Meigen Matricaria sp., cult. var.: Phytomyza atricornis Meigen Senecioneae Petasites sp.: Phytomyza atricornis Meigen Senecio cruentus DC.: Phytomyza atricornis Meigen S. lugens var. exaltatus Gray: Phytobia (Calycomyza) majuscula Frick S. mikanioides Otto: Phytomyza atricornis Meigen Cynareae Arctium lappa L.: Phytobia (Amauromyza) maculosa (Malloch), P. (Calycomyza) flavinotum Frick Arctium spp.: Phytobia (Calycomyza) jucunda (Wulp) Carduus pyenocephalus L.: Phytomyza atricornis Meigen Cynara scolymus L.: Phytobia (Calycomyza) jucunda (Wulp), Phyto- myza atricornis Meigen Silvbum marianum Gaertn.: Phytomyza atricornis Meigen References AupRicH, J. M. 1918. Seasonal and climatic variations in Cerodonta. Ann. Ent. Soc. Amer., vol. 11, pp. 63-66. BLANCHARD, E. E. 1954. Sinopsis de los agromizidos Argentinos. Argentina Min. Agr. Gan., ser. A, vol. 10, No. 56, pp. 1-48. Prick, KH. 1952. A generic revision of the family Agromyzidae with a catalogue of New World species. Univ. Califonia Publ. Ent., vol. 8, No. 8, pp.339-452. 1953a. Some additions and corrections to the species list of North American Agromyzidae. Canadian Ent., vol. 85, No. 2, pp. 68-76. 1953b. Further studies on Hawaiian Agromyzidae with descriptions of four new species. Proc. Hawaiian Ent. Soc., vol. 15, No. 1, pp. 207-215. 1956a. Nearctic species in the Liriomyza pusilla complex No. 1. Introdue- tion. Pan-Pacific Ent., vol. 32, No. 1, pp. 11-19. 1956b. Revision of the North American Calycomyza species north of Mexico. Ann. Ent. Soc. Amer., vol. 49, No. 3, pp. 284-300. 1957a. Nomenclatural changes and type designations of some New World Agromyzidae. Ann. Ent. Soc. Amer., vol. 50, No. 2, pp. 198-205. 1957b. Nearctic species in the Liriomyza pusilla complex, No. 2, L. munda and two other species attacking crops in California. Pan-Pacific Ent., vol. 33, No. 2, pp. 59-70. GroscukKeE, F. 1954, Meszellen tiber Blattminen und Blattminierer. I. Deutsche Ent. Zeitschr., new ser., vol. 1, pts. 83-5, pp. 138-156. 446 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 HENDEL, F. 1931— Agromyzidae. Jn Lindner, Die Fliegen der palaearktischen Region, 1936 fam. 59, 570 pp. Herine, E. M. 1949. Biologische Unterarten bei Phytomyza ranunculi Schrk. Entomon, vol. 1, No. 1, pp. 207-210. Kaneas, E. 1955. Dendromyza cerasiferae n. sp. from England. Ann. Ent. Fennici, vol. 21, No. 4, pp. 165-170. Mautuocg, J. R. : 1913. A revision of the species in Agromyza Fallén, and Cerodontha Rondani. Ann. Ent. Soc. Amer., vol. 6, No. 3, pp. 269-336. Menanper, A. L. 1913. A synopsis of the dipterous groups Agromyzinae, Milichiinae, Ochthi- philinae and Geomyzinae. Journ. New York Ent. Soc., vol. 21, No. 3, pp. 219-278. De Ontverra, 8. J.. AND Da Sitva, G. A. 1954. Sdébre uma nova especie de Agromyzidae cujas larvas minam folhas de Solanum argenteum. Rev. Brasil. Ent., vol. 1, pp. 25-30. Prerr, C. V.,. AND), BEATITE: | R. 1K, 1914. Flora of southeastern Washington and adjacent Idaho. xi + 296 pp. Sasakawa, M. 1954. Neue Agromyzidae aus Japan. V. Trans. Shikoku Ent. Soc., vol. 4, No. 3, pp. 35-49. SHEWELL, G. E. 1953. Notes on the types of some American Agromyzidae. Canadian Ent., vol. 85, No. 12, pp. 462-470. AGROMYZID LEAF MINERS——FRICK 447 Figures 1-170 448 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ficures 1-20.—Agromyza and Napomyza species. The heads are shown in profile except where noted. ‘The solid line equals 1 mm. Fig. 1.—Section of wing base of Agromyza ambigua Fallén, showing the manner in which the subcosta and Ry unite at the costa, greatly enlarged. Fig. 2.—Section of wing base of Napomyza lateralis (Fallén), showing the manner in which the subcosta and R, end separately in the costa, greatly enlarged. Fig. 3—Head of Agromyza ambigua Fallén (holotype 9 of 4. kincaidi Malloch), dorsal upper-orbital and vertical setae missing. Fig. 4.—Wing of A. aristata Malloch (holotype 2 of A. ulmi Frost), one-half size. Fig. 5.—Head of holotype ? of A. artstata Malloch. Fig. 6.—Head of holotype o& of A. barberi Frick. Fig. 7—Head of holotype 2 of 4. canadensis Malloch. Fig. 8.—Head of holotype 9 of 4. currani Frost, one-half size. Fig. 9—Head of holotype @ of Agromyza frosti Frick, dorsal upper-orbital and vertical setae broken off. Fig. 10.—Head of Agromyza inaequalis Malloch (holotype co of Agromyza iridescens Frost), one-half size. Fig. 11—Head of holotype 9 of Agromyza isolata Malloch, dorsal upper-orbital and vertical setae missing. Fig. 12.—Head of Agromyza nigripes Meigen (holotype 2 of Agromyza dubitata Malloch). Fig. 13—Head of holotype 9 of Agromyza pallidiseta Malloch. Fig. 14.—Head of same, front view. Fig. 15.—Head of Agromyza spiracae Kaltenbach (holotype o of Agromyza fragariac Malloch). Fig. 16—Head of Agromyza subnigripes Malloch (holotype 9 of Agromyza aprilina Mal- loch). Fig. 17.—Wing of holotype 9 of Agromyza varifrons Coquillet, one-half size. Fig. 18.—Head of same, front view. Fig. 19.—Head of same, lateral view. Fig. 20.—Head of holotype 2 of Agromyza viridula Coquillett. AGROMYZID LEAF MINERS—FRICK 449 Ficures 1-20.—Agromyza species and Napomyza lateralis. Explanation on facing page. 45 Fic Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. 0 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 ures 21-38.—Melanagromyza species. Heads areshowninprofile. Solid line equals 1mm. 21.—Head of holotype @ of Melanagromyza aldrichi Frick. 22.—Head of holotype o@ of M. angelicae (Frost). 23.—Head of holotype @ of M. approximata (Frost). 24.—Portion of head of holotype 2 of M. burgessi (Malloch). 25.—Head of same, one-half size. 26.—Head of holotype 2 of M. caerulea (Malloch), one-half size. 27.—Antenna of allotype o& of M. dianthereae (Malloch), greatly enlarged. 28.—Antenna of holotype 9 of same, greatly enlarged. 29.—Head of same, one-half size. 30.—Head of paratype 9 of M. gibsont (Malloch). 31.—Head of holotype 9 of M. longiseta (Malloch). 32.—Head of holotype o of M. mallochi (Hendel). 33.—Head of holotype @ of M. minima (Malloch). 34.—Head of holotype o* of M. orbitalis (Frost), dorsal upper-orbital, ocellar, and inner vertical setae broken off. Fig Fig Fig Fig . 35.—Head of holotype Q of M. riparella (Hendel). . 36.—Head of holotype & of M. salicis (Malloch). . 37.—Head of holotype & of M. setifrons (Melander). . 38.—Head of holotype @ of M. similata (Malloch). AGROMYZID LEAF MINERS—FRICK 451 Ficures 21-38.—Melanagromyza species. Explanation on facing page. 472314—59—_8& 452 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ficures 39-56.—Melanagromyza, Ophiomyia, and Tylomyza species. Heads are shown i in eae except where noted. The solid line equals 1 mm. Fig. 39.—Head of Holowype 2 of Melanagromyza subvirens (Malloch). Fig. 40.—Head of lectotype 9 of M. tamia (Melander). Fig. 41.—Head of ‘paratype 2 of M. tiliae (Couden). Fig. 42.—Head of paratype 2 of M. virens (Loew). Fig. 43.—Head of holotype 2 of M. viridis (Frost). Fig. 44.—Head of holotype 9 of M. winnemanae (Malloch). Fig. 45.—Portion of head of Ophiomyia maura (Meigen) (o', Germany, ex Solidago virgaurea /L.), front view, greatly enlarged. Fig. 46.—Head of holotype @ of O. buscki (Frost). Fig. 47.—Head of holotype of of O. congregata (Malloch). Fig. 48.—Head of O. coniceps (Malloch) (@, Laurel, Santa Cruz County, Calif.). Fig. 49—Head of holotype @ of O. curvibrissata (Frost). Fig. 50.—Head of holotype 9 of O. insularis (Malloch). Fig. 51—Head of O. proboscidea (Strobl) (o', Germany, ex Hieracium dinbellaiue, L.). Fig. 52.—Head of holotype 9 of O. punctohalterata (Frost). Fig. 53.—Head of holotype co" of O. texana (Malloch), genovertical plates are not as in fig. 54 because the head could not be properly oriented, all setae missing. Fig. 54.—Head of paratype 9 of O. texana (Malloch). Fig. 55.—Head of Tylomyza nasuta (Melander) (holotype o of Agromyza youngi (Malloch)). Fig. 56.—Wing of same, one-half size. AGROMYZID LEAF MINERS—FRICK | 453 Ficures 39-56.—Melanagromyza, Ophiomyia, and Tylomyza species. Explanation on facing page. 454 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ficures 57-79.—Phytobia species, heads shown in profile except where noted. The solid line equals 1 mm. Fig. 57.—Head of holotype 9 of Phytobia (Phytobia) amelanchieris (Greene), one-half size, Fig. 58.—Mesofacial plate and lunule of same, full size. Fig. 59.—Head of holotype o of P. (P.) indecora (Malloch). Fig. 60.—Wing of holotype o& of P. (P.) kallima (Frost), one-half size. Fig. 61.—Mesonotum of holotype ? of P. (P.) picta (Coquillett), setulae on black not shown, ~ one-half size. Fig. 62.—Head of same, full size. Fig. 63.—Head of holotype @ of P. (P.) prutnosa (Coquillett). Fig. 64.—Head of P. (P.) setosa (Loew) (holotype 9 of Agromyza aceris Greene), one-half size. Fig. 65.—Mesofacial plate and lunule of same, full size. Fig. 66.—Head of P. (Cephalomyza) albidohalterata (Malloch) (co, White Heath, Ill.). Fig. 67.—Head of lectotype 9 of P. (C.) auriceps (Melander). Fig. 68.—Head of holotype 9 of P. (C.) indecisa (Malloch). Fig. 69.—Head of P. (Poémyza) angulata (Loew) (co, Savanna, Illinois). Fig. 70.—Head of holotype @ of P. (P.) inconspicua (Malloch). Fig. 71.—Posterior end of puparium of same, spiracles above, anal opening below, posterior view. Fig. 72.—Posterior spiracles of same, dorsal view. Fig. 73.—Posterior end of same, lateral view. Fig. 74.—Male terminalia of P. (P.) lateralis (Macquart) (paratype co of Agromyza coquil- letti Malloch), showing keel (K) of ninth tergite (9T). Fig. 75.—Male terminalia of same, lateral view. Fig. 76.—Head of P. (P.) lateralis (Macquart) (holotype 9 of Agromyza coquilletti Malloch). Fig. 77.—Wing of holotype @ of P. (P.) subangulata (Malloch), one-half size. Fig. 78.—Head of same, front view. Fig. 79.—Head of same, lateral view. AGROMYZID LEAF MINERS—FRICK 455 Figures 57-79.—Phytobia species. Explanation on facing page. 456 PROCEEDINGS. OF THE NATIONAL MUSEUM VOL. 108 Ficures 80-94.—Phytobia species. Heads are shown in profile except where noted. The solid line equals 1 mm. Fig. 80.—Head of holotype @ of Phytobia (Dizygomyza) magnicornis (Loew). Fig. 81.—Wing of same, one-half size. Fig. 82.—Head of lectotype of of P. (D.) thompsoni Frick. Fig. 83.—Head of same, front view. Fig. 84.—Posterior end of abdomen and male terminalia of same, lateral view. Fig. 85.—Head of P. (Icteromyza) capitata (Zetterstedt) (c’, Germany), front view. Fig. 86.—Wing of holotype 9? of P. (J.) longipennis (Loew), one-half size. Fig. 87.—Head of lectotype o of P. (I.) pollinosa (Melander). Fig. 88.—Head of P. (Trilobomyza) verbasci (Bouché) (o, Germany, ex Scrophularia nodosa L.), front view. Fig. 89.—Head of holotype 2 of P. (T.) calypirata (Hendel). Fig. 90.—Head of holotype @ of P. (T.) pleuralis (Malloch), third antennal segment missing. Fig. 91.—Head of holotype 9 of P. (T.) varia (Melander). Fig. 92.—Head of P. (Praspedomyza) approximata (Hendel) (9, Germany, ex Daphne mezereum \..), front view. Fig. 93.—Head of P. (P.) clara (Melander) (o', Mount Hermon, Santa Cruz County, Calif., ex Pteridium aquilinum (L.) Kuhn). Fig. 94.—Head of holotype @ of P. (P.) subinfumata (Malloch). AGROMYZID LEAF MINERS—FRICK 457 Ficures 80-94.—Phytobia species. Explanation on facing page. 458 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ficures 95-112.—Cerodontha and Liriomyza species. Heads are shown in profile except where noted. The solid line equals 1 mm. Fig. 95.—Mesonotum of holotype 2 of Cerodontha dorsalis (Loew). Fig. 96.—Head of holotype @ of Liriomyza angulicornis (Malloch). Fig. 97—Anepisternum of same, double size. Fig. 98.—Head of holotype o& of L. assimilis (Malloch). Fig. 99.—Mesonotum of same, right dorsocentrals omitted. Fig. 100.—Anepisternum of same, double size. Fig. 101—Mesonotum of holotype 9 of L. borealis (Malloch). Fig. 102.—Head of holotype & of L. commelinae (Frost), dorsal upper-orbital and both vertical setae missing. Fig. 103.—Posterior half of mesonotum and scutellum of same. Fig. 104.—Head of holotype 9 of L. deceptiva (Malloch), front view. Fig. 105.—Head of same, lateral view. Fig. 106.—Head of holotype 9 of LZ. discalis (Malloch). Fig. 107.—Head of holotype o” of L. felti (Malloch). Fig. 108.—Wing of holotype ? of L. fumicosta (Malloch), part of wing missing as shown. Fig. 109—Mesonotum of same, right dorsocentrals omitted. Fig. 110.—Head of lectotype @ of L. lima (Melander). Fig. 111—NMesonotum of holotype @ of L. marginalis (Malloch). Fig. 112.—Head of same. AGROMYZID LEAF MINERS—FRICK | 459 a a IY. 7 pe f 100 5* sf Figures 95-112.—Cerodontha and Liriomyza species. Explanation on facing page. bt 460 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Figures 113-130.—Liriomyza, Metopomyza, Haplomyza, and Xyraeomyia species. Heads Fig. are shown in profile except where noted. ‘The solid line equals 1 mm. 113.—Mesonotum of lectotype 9 of Liromyza melampyga (Loew), all setulae and setae on the right side omitted. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. 114.—Head of L. melampyga (Loew) (9 , Glen Echo, Md.). 115.—Head of lectotype @ of L. pacifica (Melander). 116.—Anepisternum of holotype o& of L. ssbropeniele (Frost), double size. 117.—Head of same. 118.—Wing of same. 119.—Head of holotype o of L. quadrisetosa (Malloch). 120.—Mesonotum of paratype 9 of L. reverberata (Malloch), left half only. 121.—Wing of same, one-half size. Fig. 122.—Head of holotype o& of L. schmidti (Aldrich), inner vertical seta missing. Fig. 123—Wing of same. Fig. 124.—Mesonotum of lectotype o& of L. sorosis (Williston), all setulae omitted. Fig. 125.—Head of same. Fig. 126.—Head of holotype 9 of L. variata (Malloch). Fig. 127.—Wing of same, one-half size. Fig. 128.—Head of Metopomyza interfrontalis (Melander) (? , Hope Valley, Alpine County, Calif.). Fig. 129.—Head of Haplomyza togata (Melander) (9, Bakersfield, Kern County, Calif., ex Amaranthus hybridus L.). Fig. 130.—Head of paratype 9 of Xyraeomyia conjunctimontis Frick. AGROMYZID LEAF MINERS—FRICK 461 129 -— 4H Ficures 113-130.—Liriomyza, Metopomyza, Haplomyza, and Xyraeomyia species. Explanation on facing page. 462 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ficures 131-149.—Phytagromyza, Pseudonapomyza, Napomyza, and Phytomyza species. Heads shown in profile. The solid line equals 1 mm. except where noted. Fig. 131.—Head of Phytagromyza nitida (Malloch) (co, White Heath, IIl.). Fig. 132.—Head of lectotype @ of P. orbitalis (Melander). Fig. 133—Head of holotype o&' of P. plagiata (Melander). Fig. 134.—Wing of P. flavocingulata (Strobl) (c&*, Germany), greatly enlarged, solid line equals 0.5 mm. Fig. 135.—Wing of Pseudonapomyza aira (Meigen) (o, Germany), greatly enlarged, solid line equals 0.5 mm. Fig. 136.—Head of holotype 2 of P. lactetpennis (Malloch). Fig. 137.—Head of holotype 9 of Napomyza davisti (Walton). Fig. 138.—Head of holotype 2 of N. parvicella (Coquillett). Fig. 139.—Head of paratype o& of Phytomza affinalis Frost. Fig. 140.—Head of holotype o& of P. angelicella Frost. Fig. 141.—Head of holotype o& of P. aquilegiana Frost. Fig. 142.—Head of holotype 9? of P. atripalpis Aldrich. Fig. 143.—Wing of same, one-half size. Fig. 144.—Head of holotype 9 of P. auricornis Frost. Fig. 145.—Wing of holotype of of P. centralis Frost. Fig. 146.—Mesonotum of holotype o& of P. clemativora Coquillett. Fig. 147.—Head of same, third antennal segment and both vertical setae missing. Fig. 148.—Wing of same. Fig. 149.—Head of holotype 9 of P. delphinii Frost. AGROMYZID LEAF MINERS—FRICK 463 Figures 131-149.—Phytagromyza, Pseudonapomyza, Napomyza, and Phytomyza species. Explanation on facing page. 464 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Figures 150-170.—Phytomyza species. Heads are shown in profile. The solid line equals 1 mm. 150.—Head of holotype 2 of Phytomyza dura Curran. 151.—Wing of same, one-half size. 152.—Head of holotype 2 of P. flavinervis Frost. 153.—Head of lectotype @ of P. genalis Melander. 154.—Head of lectotype 2 of P. ilicicola Loew. 155.—Wing of same, one-half size. 156.—Head of paratype @ of P. lactuca Frost. 157.—Head of lectotype 2 of P. loewit Hendel. 158.—Wing of same, one-half size. 159.—Head of holotype 2 of P. marginalis Frost. 160.—Head of holotype @ of P. melanella Frost. 161.—Head of P. minuscula Goureau (lectotype o of P. nitida Melander). 162.—Wing of holotype 2 of P. nervosa Loew, one-half size. 163.—Head of holotype &@ of P. nigrinervis Frost. 164.—Head of paratype 9 of P. plumiseta Frost. 165.—Wing of same, one-half size. 166.—Head of holotype 2 of P. subtenella Frost. 167.—Wing of same, one-half size. 168.—Head of holotype o@ of P. trivittata Frost. 169.—Wing of same, one-half size. 170.—Mesonotum of same. AGROMYZID LEAF MINERS—FRICK 465 a 169 Ficures 150-170.—Phytomyza species. Explanation on facing page. U.S. GOVERNMENT PRINTING OFFICE: 1959 PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM ; HOA Z issued W(4™\ SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 108 Washington : 1959 No. 3408 A REVISION OF THE BUTTERFLY GENERA THEOCHILA AND TATOCHILA (LEPIDOPTERA : PIERIDAE) By Jost Herrera aNd WitiraAM D. FIELD This is the second in a series of four papers treating the “Tatocheilae- Phulia” complex of genera. In this paper is offered a treatment of the 14 species included in the genera Theochila Field and Tatochila Butler. Because these genera have recently been defined by the junior author (Proc. U.S. Nat. Mus., vol. 108, No. 3396, pp. 103-131, 1958), they are not further described here. Both genera are Neotropical in distribution, Theochila being found in the southern nontropical parts of Brazil, in Paraguay, and in eastern Argentina and Tatochila being distributed from Colombia and Bolivia south through Tierra del Fuego in the tundra, temperate forests, and scrub areas. This study is based upon 620 specimens gathered from the collec- tions of the U. S. National Museum; Museo Nacional de Historia Natural, Santiago, Chile; Instituto Pedagégico, Universidad de Chile, at Santiago; Museu Nacional, Rio de Janeiro, Brazil; Fundacion Miguel Lillo, Tucuman, Argentina; Cornell University, Ithaca, N. Y.; American Museum of Natural History, New York City; Chicago Museum of Natural History; and British Museum (Natural History), London. Thanks for the loan of this material are due to the officials of all of these museums and most especially to Dr. Kenneth J. Hay- ward of the museum in Tucuman, Mr. D. 8. Fletcher of the British 472317—59—1 467 468 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Museum, and Dr. Emilo Ureta-R. of the Museo Nacional de Historia Natural, Santiago, Chile. All specific and subspecific descriptions were prepared to supplement the illustrations and should be studied with that in mind. The descriptions of the following new subspecies were prepared by William D. Field and are credited solely to him: Tatochila theodice staudingert and T. microdice fueguensis. Genus Theochila Field Theochila Field, Proc. U. 8. Nat. Mus., vol. 108, p. 106, 1958. Tyr: Pieris maenacte Boisduval= Theochila maenacte (Boisduval), by original designation. This genus with characters as given in the original description. Only one species is included and it is here rather provisionally con- sidered to consist of two subspecies. Its position until the present time has always been a matter of much speculation and doubt. It has, in a single decade, been placed in four different genera by four different authors. Theochila maenacte maenacte (Boisduval) Pieris maenacte Boisduval, Histoire naturelle des insectes, Species général des Lépidoptéres, vol. 1, p. 517, 1836.—Doubleday, in Doubleday and Westwood, The genera of diurnal Lepidoptera, vol. 1, p. 48, 1847. Pieris menacte Boisduval, Blanchard nec Boisduval (a misspelling), in Gay, Historia fisica y politica de Chile, Zoologia, vol. 7, pp. 10, 11, 1852.—Butler, Proce. Zool. Soc. London (1872), p. 41, 1872.—Burmeister, Description physique de la République Argentine . . . , vol. 5, pt. 1, p. 87, 1878; Atlas, sec. 5, pt. 2, p. 13, pl. 4, fig. 10, 1879-1880.—Herrich-Schaffer, Corresp.-Blatt Zool. mineral. Ver. Regensburg, vol. 21, No. 10, p. 126, 1867.—Gosse, Ento- mologist, vol. 13, p. 195, September 1880.— Mabilde, Guia practica para os principantes colleccionadores de insectos, p. 56, 1896.—Réber, in Seitz, Gross-Schmetterlinge der Erde, vol. 5, p. 58, pl. 19, figs. bl, b2, Sept. 26, 1908.—Giacomelli, Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 26, p. 403, Mar. 10, 1915.—J6érgensen, Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 28, p. 473, Nov. 10, 1916.—Giacomelli, Physis, vol. 3, No. 15, p. 379, 1917.— Kohler, Zeitschr. Wiss. Insekt.-Biol., vol. 18, pt. 12, suppl., p. 15, Dee. 15, 1923.—Drosihn, Ent. Rundschau, vol. 50, suppl., p. 81, pl. 20, fig. m, 1933.— Breyer, Verhandlungen VII Internationalen Kongress fiir Entomologie, vol. 1 (for 1938), pp. 34, February 1939.—Ureta, Rev. Chilena Hist. Nat., vol. 43 (for 1939), pp. 227, 228, 1940. Pieris automata Berg, Mabilde nec Berg (a misindentification), Guia practica para os principantes colleccionadores de insectos, p. 55, pl. 2, fig. 5, 1896. Tatochila menacte (Boisduval), Bartlett-Calvert nec Boisduval (a misspelling), Rev. Chilena Hist. Nat., vol. 2, No. 7, p. 98, July 1898.—Klots, Ent. Ameri- cana, new ser., vol. 12, No. 4, p. 218, December 1931. Pieris itaticayae Foetterle, Jorgensen nec Foetterle (a misidentification and mis- spelling), Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 28, p. 473, Nov. 10, 1916.—Rober, in Seitz, Gross-Schmetterlinge der Erde, vol. 5, p. 1016, Jan. 21, 1924. BUTTERFLIES—HERRERA AND FIELD 469 Ascia (Ganyra) menacte (Boisduval), Talbot nec Boisduval (a misspelling), in Strand, Lepidopterorum catalogus, pars 66, p. 635, 1935. Pieris phileta itaticayae Foetterle, Breyer nec Foetterle (a misidentification and mis- spelling), Verhandlungen VII Internationalen Kongress fir Entomologie, p. 34, February 1939. Synchloé maenacte (Boisduval), D’Almeida, Arg. Zool. Estado de Séo Paulo, vol. 2, p. 300, Jan. 3, 1941. Ascia maenacte (Boisduval), Talbot, in Strand, Lepidopterorum catalogus, pars 66, p. 211, 1935.—Hayward, Acta Zool. Lilloana Inst. ‘Miguel Lillo,” vol. 9, p. 97, Oct. 25, 1951. Theochila maenacte (Boisduval), Field, Proc. U. 8. Nat. Mus., vol. 108, p. 106, figs. 1, 9, 17, 25, 34, 1958. VenaTion: As illustrated (fig. 55). Lees: With paronychium, claw, and pulvillus as illustrated (fig. 69). Mate: Plate 1, figure 1. Head with an orange margin behind eyes and with outer face of pa)pus white to pale yellow. Wings white above, usually with apex of forewing, costal margin and distal portions of radial veins dark fuscous in color. Extreme bases of both wings above irrorated with dark fuscous scales, especially along base of cubitus in the hindwing. Hindwing and apex of forewing below white with a faint yellowish cast, remainder of forewing white. There is a faint orange spot in extreme base of hindwing below and many specimens have the veins on this surface faintly outlined with fuscous, especially along the base of cubitus. Length of forewing, 23-28 mm. (average 25.5 mm.). Male genitalia as illustrated (figs. 27, 41). No differences were found between the genitalia of typical 7. maenacte and its subspecies T’. maenacte itatiayae. The genitalia of four males were studied. Frmaue: Plate 1, figure la. Head as in the male. Wings above white with veins heavily outlined with fuscous and with dark apical area of forewing more extensive than in the male. Wings below as in the male, with ground color more yellow and with fuscous along the veins more distinct. Length of forewing, 23-28 mm. (average, 25.5 mm.). Female genitalia as illustrated (fig. 83), with anterior lobe of inner genital plate large, subtriangular, and having its ventral margin triundulate. As in the male, no differences were observed between the two subspecies. The genitalia of six females were studied. Typr Locatity: ‘Paraguay, Republique Argentina’ = Paraguay. LocaTIon oF TyPE: British Museum (Natural History). _ ADDITIONAL TYPE DATA: Described from a number of male speci- mens (number not specifically stated) in the Boisduval collection. Of these, only a single specimen is in the British Museum (Natural His- tory), and we consider this to be the holotype. 470 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 M®sTHOD OF IDENTIFICATION: By means of characters mentioned in the original description that are quite adequate for the identification of this name. SYNONYMICAL NoTES: As shown in the references above, a number of workers have misspelled the name by dropping the first ‘a.’ MIsIDENTIFICATIONS: J6rgensen (1916), Réber (1924), and Breyer (1939) have all confused some specimens of 7. maenacte with T. maenacte itatiayae and, in addition, have misspelled the latter. Mabilde (1896) has mistakenly identified this species as Pieris automata Berg. DistripuTion: Province of Buenos Aires in Argentina north into Paraguay and northwest into the states of Santa Catarina and Parandé in Brazil. ARGENTINA: Province of Buenos Aires, Buenos Aires. Paraguay: no specific record. Braztu: State of Santa Catarina, Joinville; State of Parana, Castro. MATERIAL STUDIED: 7 males, 12 females. Theochila maenacte itatiayae (Foetterie) Pieris itatiayae Foetterle, Rev. Mus. Paulista, vol. 5, pp. 624-627, pl. 16, fig. 5, 1902.—Jérgensen, Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 28, p. 473, Nov. 10, 1916.—Giacomelli, Physis, vol. 3, No. 15, p. 379, 1917.—Zik4n, Ent. Rundschau, vol. 45, p. 7, Jan. 15, 1928. Pieris itaticayae Foetterle, Réber nec Foetterle (a misspelling), in Seitz, Gross- Schmetterlinge der Erde, vol. 5, p. 60, pl. 19, fig. d1, 1908; ibid, vol. 5, p. 1016, Jan. 21, 1924. Synchloé maenacte itatiayae (Foetterle), D’Almeida, Arq. Zool. Estado de Sao Paulo, vol. 2, p. 300, pl. 1, figs. 1, 5, Jan. 3, 1941. Leptophobia itaticayae Foetterle, Talbot nec Foetterle (a misspelling), in Strand, Lepidopterorum catalogus, pars 66, p. 52, 1935. VenatTion: Not illustrated, not different from typical maenacte. Lees: With paronychium, claw, and pulvillus not illustrated, not different from the typical subspecies. Maus: Plate 1, figure 2. This subspecies is quite similar to typical 7’. maenacte, differing chiefly in having the fuscous on apex of forewing covering a considerably larger area and usually with dark scaling along veins on hindwing below slightly more distinct. Length of forewing, 22-28 mm. (average, 25 mm.). Male genitalia not illustrated, not different from the genitalia of typical 7. maenacte. The genitalia of four males were studied. Frmauu: Plate 1, figure 2a. Palpi as in the male but all five females available for study lacked the orange scaling behind the eyes (in four of these the heads were in poor shape). Wings similar to females of T. maenacte maenacte with fuscous on apical area more extensive and forming a more solid band. Wings below not distin- guishable from the typical subspecies. BUTTERFLIES—HERRERA AND FIELD A471 Length of forewing, 23-27 mm. (average 25 mm.). Female genitalia not illustrated and not different from the typical subspecies. The genitalia of three females were studied. VARIATION: One male (of a series of four) and one female (of a series of seven) labeled “Castro, State of Parand, Brazil’ are like T. maenacte itatiayae, while the remaining specimens from this locality are clearly T. maenacte maenacie. If these two specimens are correctly labeled (which we have reason to doubt) we will have to accept the fact that either the two subspecies overlap in distribution or that they are found a very short distance from each other although perhaps under different ecological conditions. For the time being this pair is classified as ttatiayae, but Castro is not included as a locality for that subspecies. TypE Locatity: ‘Itatiaya, a partir de 1700 metros para cima,” Brazil. The Itatiaya Mountains in the Serra do Mar chain, located in the region where the three states of Minas Gerais, Rio de Janeiro, and Sado Paulo join. Location or tyres: In the collection of José G. Foetterle, which according to Hern and Kahle (Ent. Beih. Berlin-Dahlem, vol. 2, p. 78, 1935) is in the Natural History Museum in Vienna, Austria. ADDITIONAL TYPE DATA: Described from four males and one female, of which Foetterle considered one male and ene female as “types” (=cotypes). MertTHoD OF IDENTIFICATION: Identified by comparison of topo- typical material with the original description and with the excellent original figure (in color). SYNONYMICAL NoTEs: There are no synonyms; however both Rober and Talbot, as shown in the references, have misspelled the name by adding a ‘‘c’’ between the second ‘%”’ and the second “‘a.”’ Distrisution: Only known from in and around the Itatiaya Mountains in the Serra do Mar chain of eastern Brazil. Braziu: State of Minas Gerais, Passa Quatro (1,600 m., March, April, Novem- ber), Virginia (1,500 m., January), Caldas (July); State of Rio de Janeiro, Itatiaya (February—April, August, October). MATERIAL STUDIED: 14 males, 6 females. Genus Tatochila Butler Tatochila Butler, Cist. Ent., vol. 1, pp. 38, 51-52, 56, Sept. 12, 1870; Proc. Zool. Soc. London (1872), p. 67, Jan. 16, 1872.—Reed, Anal. Univ. Chile, vol. 49, p. 665, September 1877; Una Monografia de los Mariposas Chilenas, p. 20, 1877.—Butler, Trans. Ent. Soc. London (1881), pp. 472-474, December 1881.—Bartlett-Calvert, Anal. Univ. Chile, vol. 69, p. 314, March 1886.— Schatz, in Staudinger and Schatz, Exotische Schmetterlinge, Theil 2, p. 61, 1892.—Berg, Anal. Mus. Nac. Buenos Aires, vol. 4, pp. 217-255, figs. 1-5, Nov. 18, 1895.—Bartlett-Calvert, Rev. Chilena Hist. Nat., vol. 2, p. 98, 472 PROCEEDINGS OF THE NATIONAL MUSEUM YOU. 108 1898.—Elwes, Trans. Ent. Soc. London (1903), pp. 292-293, October 1903.— Réber, in Seitz, Die Gross-Schmetterlinge der Erde, vol. 5, pp. 55-57, pl. 18, 1908.—Dixey, Proc. Ent. Soc. London (1909), p. exxxv, Mar. 30, 1910.— Giacomelli, Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 26, pp. 403-415, Mar. 10, 1915; Rev. Chilena Hist. Nat., vol. 20, pp. 41-57, 3 pls., June 30, 1916.—Jérgensen, Anal. Mus. Nac. Hist. Buenos Aires, vol. 28, pp. 430, 433-470, figs. 3-7, Nov. 10, 1916.—Giacomelli, Physis, vol. 3, pp. 371-378, 1917.—Rober, in Seitz, Die Gross-Schmetterlinge der Erde, vol. 5, pp. 1015-1016, Jan. 21, 1924.—Klots, Ent. Americana, new ser., vol. 12, No. 3, pp. 154, 161, December 1931; No. 4, pp. 217-218, 230, pl. 12, fig. 87, March 1932.—Talbot, in Strand, Lepidopterorum catalogus, pars 53, pp. 57-59, Oct. 24, 1932; pars 66, p. 626, Aug. 30, 1935.—Ureta, Bol. Mus. Nac. Hist. Nat., vol. 16, p. 123, 1937; Rev. Chilena Hist. Nat., vol. 41, pp. 239-283, pls. 11, 13, figs. 33-39, 1938.—Breyer, Verhandlungen VII Internationalen Kongress fiir Entomologie, vol. 1 (for 1938), pp. 28-33, 1938.—Apolinar, Rev. Acad. Colombiana Cienc. Exact. Fis. Nat., vol. 4, p. 349, 1941.— D’ Almeida, Pap. Avulsos Dep. Zool., Sec. Agr. Brazil, vol. 3, p. 103, June — 4, 1943.—Bryk, Ark. f. Zool., vol. 36a, pp. 5-7, pl. 1, figs. 2, 9, Nov. 6, 1944.— Hayward, Acta Zool. Lilloana, vol. 9, pp. 91-94, Oct. 25, 1951.—Field, Proc. U. S. Nat. Mus., vol. 108, pp. 108-110, 1958. Tatocheila Butler, Scudder nec Butler (a misspelling), Proc. Amer. Acad. Arts Sci., vol. 10, p. 276, 1875.—D’Almeida, Pap. Avulsos Dep. Zool., Sec Agr. Brazil, vol. 3, p. 102, June 4, 1943. Synchloé Hiibner, D’ Almeida nec Hiibner (falsely states that Synchloé was orig- inally monotypic and makes a pseudotype of S. autodice Hiibner), Arq. Zool. Estado de Sado Paulo, vol. 2, p. 299, Jan. 3, 1941; Pap. Avulsos Dep. Zool., Sec. Agr. Brazil, vol. 3, p. 102, June 4, 1943. Tyrer: Synchloé autodice Hiibner=Tatochila autodice (Hibner). Type by reason of original designation and also by being sole included species. Since this genus has just recently been redefined (Field, Proc. U.S. Nat. Mus., vol. 108, pp. 108-110, 1958), there is no need for discussion of its characters here. The 13 species included in this genus are divided into five species groups on the basis of the type of aedeagus and differences in the wing patterns. It was not found necessary to include genitalia characters in the key. Key to the species and subspecies of Tatochila 1. Hindwing below with a distinct white spot covering the lower discocellular vein and a considerable area of white along both sides of this vein (Groups Band ©). 2 e098 28 Ly Hindwing below jacinta. chi mates et ned with ees er discoceltulae vein outlined with black; or if white spot is present, it is restricted to the vein only (Groups A, D, 'B). Se ae A EMM oak, BO 2. Hindwing below with interspace Se-+R, Contains a small oval or irregularly shaped spot or streak in the middle (Group B). . . ae Hindwing below with interspace Sc+R, lacking this Sas (creas ©) ie iS 10. i 12. 13. 14. 15. BUTTERFLIES—-HERRERA AND FIELD 473 INVIGICShe ence uisennesde resect cee PULA CMD eh cl ss or chrar ae 14 Females. .... Lae ve 4a, 16 Hindwing above laplante distinc: Gincie. merenel: Ww aiciees inigcd markings on ends of veins and lacking the distnes sagittate, submarginal, and intervenal black spots... . . ... . . T. autodice (p. 481) Hindwing above with distinct, bisole naoiant wedge-shaped markings on ends of veins and with distinct sagittate, submarginal and intervenal black spots .... ... .. T.blanchardii. . . 5 Hindwing below with prea coer very pale yellow or white washed with yellow... . . . . TU. blanchardii blanchardii (p. 484) Hindwing below with pron color distinctly yellow. T. blanchardii ernestae (p. 486) Hindwing above with ground color white, sometimes slightly tinged with yellow; apex of forewing with wedge-shaped marks extending inward only to submarginal black spots. . .. . . .. . . T. auteodice (p. 481) Hindwing above with ground color Aeneas yellow; apex of forewing with wedge-shaped marks extending inward to the cell. T. blanchardii. . . 7 Apex of forewing and hindwing below, light yellow (central Chile, Province of Coquimbo south to Temuco). T. blanchardii blanchardii (p. 484) Apex of forewing and hindwing below, dark yellow (northern Chile (Prov- ince of Tarapaca), southern Peru, and Bolivia (Department of Potosf)). T. blanchardii ernestae (p. 486) IVIGCS: Mega ceca ine ar aah ave te Cad deh tel, altar WE weclen a 0 wt cele ek ee OO Females .... 2. di be WS ei pain 1? oe Rend, Wess Be Ke a ae A Hindwing above ential snare wee Seat 2 a a ee lO Hindwing above with considerable black acai on or near ilie ends of the veins ... Sie Cave. Suse. glo Seales behind ae ee aie helo ees es T. vanvolxemi (p. 495) Seales behind eyes white to pale yellow; hindwing below mainly yellow . 11 Marginal wedge-shaped spots of forewing distinctly divided in middle by white scaling on veins. . .. ... . . T. mercedes (p. 497) Marginal wedge-shaped spots of ee SOUGe SS) ao. ee ee Forewing averaging about 22 mm. in length; with daa Tabane and discocellular bar on forewing reduced and with white spot over lower discocellular vein on hindwing underneath small. T. microdice fueguensis (p. 490) Forewing averaging about 24 mm. in length; with dark markings and discocellular bar on forewing more extensive and with white spot over lower discocellular vein on hindwing underneath larger. T. microdice microdice (p. 488) Hindwing below with a series of distinct, submarginal, sagittate markings. T. microdice macrodice (p. 492) Hindwing below lacking this series of sagittate markings. T. miecrodice arctodice (p. 494) Scales behind eyes distinctly orange. ..........+24.2... 17 Scales behind eyes white to pale yellow. ........2.2.2... «15 Ground color of wings above distinctly yellow. T. microdice fueguensis (p. 490) Ground color of wings above white, sometimes with a faint yellow cast . 16 474. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 16. Hindwing above with dark scaling along veins restricted to the outer margin. T. mercedes (p. 497) Hindwing above with dark scaling along entire length of veins. T. microdice microdice (p. 488) 17. Dark sealing along either side of veins on hindwing not covering the veins. T. vanvolxemi (p. 495) Dark scaling covering the veins. . . Marts A) Mls 18. Hindwing below with a distinct series i fae couR subaiageeall sagittate markings; with white striping on veins narrow, usually covering only the: -veins® . 2 : . . . . . VT. mierodice macrodice (p. 492) Hindwing below with Pos oons: sagittate markings obsolete, sometimes entirely replaced with faint red spots; with white striping on veins broader, usually covering slightly more than the veins. TF. microdice arctodice (p. 494) 19. Ground color of hindwing white washed with yellow; submarginal sagittate markings on both pair of wings ee extending to at least below vein Cug . 2 1./% SESE . theedice and its subspecies (p. 475) Ground color of hndyane ath et distinctly yellow; submarginal sagittate markings usually absent from both pairs of wings underneath or, if present, very faint and not extending below vein Cuz (Groups D, EP isegess f oc 5), tere OY 515 Ai. 20 20. Palpi with miter surface OLANSEM ETOYS ae T. orthodice: (p. 504) Palpi with outer ‘surface ‘yellow or white 2-005. [C102 2724 2. 21 Zist iscales4behine: evesOrange... «is lesz keeles ve Biss, See Seal eh 2 Scales behind eyes white... .. . 24 22. Outer edge of forewing entirely black at eneceoaeil wedze-ahaped aedcines solidiy colored; hindwing below with distinct orange streaks in base of wing below cell and on outer half of wing between veins. . .... 25 Outer edge of forewing white except for the wedge-shaped markings, at least some of which are divided in the middle by white along the veins; hindwing below with orange restricted to coastal margin. T. inversa (p. 501) 23. Hindwing below with large orange streak covering upper half of cell; fore- wing above with a submarginal series of connected sagittate markings. T. xanthedice (p. 511) Hindwing below with this area yellow; forewing above lacking sagittate markings 2! (57 .. . . . T.homoecdice (p. 503) 24. Hindwing and apex lee forewinie below, with long orange streaks between the veins extending from near the cell to the outer margin; veins on hindwing narrowly covered with dull white... . eee 25 Hindwing and apex of forewing below lacking biiese erates streniess veins on hindwing broadly covered with silvery white . . T. distincta (p. 513) 25. Outer margin of hindwing above and below bordered by a distinct but narrow black line .. . ... . . T. sagittata (p. 506) Outer margin of hindwing Peleg: Ww a thie black line lacking or represented only by short indistinct bars at the ends of the veins. T. stigmadice (p. 507) Grour A. THe THEODICE GRouP In this group the distal end of the aedeagus is deeply incised on its ventral side only, forming a large triangular-shaped lobe on its right side. BUTTERFLIES—HERRERA AND FIELD AT5 Hindwings above with solid-colored wedge-shaped markings along the outer margins and usually with a submarginal series of sagittate markings. Hindwings below with white ground color; with a sub- marginal series of sagittate markings; with narrow orange streaks through the middle of the interspaces from the outer margin to the cell and lacking a white spot that in some other groups covers the lower discocellular vein. A single species that divides into three subspecies comprises this group. Tatechila theodice theedice (Boisduval) Pieris theodice Boisduval, Voyage de découvertes de l Astrolabe, pt. 1, Lépidop- téres, p. 51, 1832; Histoire naturelle des insectes, Species général des Lépi- doptéres, vol. 1, pp. 540, 541, 1836.—Doubleday, 7x Doubleday and West- wood, The genera of diurnal Lepidoptera, vol. 1, p. 51, 1847.—Blanchard, in Gay, Historia Fisica y Politica de Chile, Zoologia, vol. 7, p. 12 (description of female theodice only, which actually is male theodice), 1852.—Lucas, Rev. Mag. Zool., ser. 2, vol. 4, p. 337, July 1852.—Herrich-Schaffer, Corresp. Blatt Zool.-Min. Ver. Regensburg, vol. 21, No. 10, p. 125, 1867.— Kirby, A synonymic catalogue of diurnal Lepidoptera, p. 450, 1871.— Berg, Actas Acad. Nac. Cienc., Cérdoba, Argentine Rep., vol. 1, p. 66, 1875. Pieris demodice Blanchard, in Gay, Historia de fisica y politica de Chile, Zoologia, vol. 7, pp. 138-14, 1852.—Kirby, A synonymic catalogue of diurnal Lepi- doptera, p. 451, 1871.—Berg, Bull. Soc. Imp. Nat. Moscou, 1875, No. 4, pp. 195, 196, 1876.—Burmeister, Description physique de la République Argentine . . . vol. 5, pt. 1, p. 89, 1878.—Staudinger, in Staudinger and Schatz, Exotische Schmetterlinge, vol. 1, p. 31, vol. 2, pl. 18, fig. al, 1888. Tatochila theodice Boisduval, Butler, Proc. Zool. Soc. London, 1872, p. 67, Jan. 16, 1872.—Reed, Anal. Univ. Chile, vol. 51, pp. 668-669, 1877.—Dixey, Proc. Ent. Soc. London, 1909, p. exxxiv, Mar. 30, 1910.—Klots, Ent. Ameri- cana, new ser., vol. 12, No. 4, p. 218, March 1932.—Ureta, Rev. Chilena Hist. Nat., vol. 38, p. 78, 1934.— Breyer, Verhandlungen VII Internationalen Kongress fiir Entomologie, p. 30, February 1939.—Hayward, Act. Zool. Lilloana Inst. ‘Miguel Lillo,” vol. 9, p. 92, Oct. 25, 1951. Tatochila demodice (Blanchard), Butler, Proc. Zool. Soc. London (1872), p. 67, Jan. 16, 1872.—Reed, Anal. Univ. Chile, vol. 51, pp. 666, 667, 1877.— Butler, Trans. Ent. Soc. London (1881), p. 478, December 1881.—Bartlett- Calvert, Anal. Univ. Chile, vol. 69, p. 314, 1886.—Berg, Anal. Mus. Nac. Buenos Aires, vol. 4, pp. 240-245, Nov. 18, 1895.—Bartlett-Calvert, Rev. Chilena Hist. Nat., vol. 2, p. 98, July 1898.—Porter, Rev. Chilena Hist, Nat., vol. 3, p. 36, April 1899.—Elwes, Trans. Ent. Soc. London, 1903, pt. 3, pp. 292-293, pl. 12, figs. 7, 8, October 1903.—Rober, in Seitz, Die Gross- Schmetterlinge der Erde, vol. 5, pp. 56, 57, May 25, 1908—Paravicini, Mitteil. Schweiz. Ent. Ges., vol. 12, pt. 1, p. 22, May 1910.—Giacomelli, Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 26, pp. 406, 408, 414, 415, Mar. 10, 1915; Rev. Chilena Hist. Nat., vol. 20, pp. 44, 46, 57, 51, June 30, 1916.—Jérgensen, Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 28, pp. 443, 462-464, figs. 3, 4, Nov. 10, 1916.—Giacomelli, Physis, vol. 3, p. 376, 1917.—Rober, in Seitz, Die Gross-Schmetterlinge der Erde, vol. 5, p. 1015, pl. 194, figs. a2, a3, Jan. 21, 1924—Talbot, in Strand, Lepidopterorum catalogus, pars 53, Pieridae I, p. 58, Oct. 24, 1982.—Breyer, Rev. Soc. Ent. 476 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Argentina, vol. 8, p. 63, 1936; Verhandlungen VII Internationalen Kongress fir Entomologie, vol. 1 (1938), p. 30, February 1939.—Hayward, Act. Zool. Lilloana Inst. ‘‘Miguel Lillo,”’ vol. 9, p. 938, Oct. 25, 1951. Pieris xanthodice Lucas, Mabille nec Lucas (a misidentification, figure only), Mission scientifique du Cap Horn, vol. 6, Zoologie, Insectes, pl. 1, fig. 1, 1888. Tatochila theodice theodice (Boisduval), Ureta, Rev. Chilena Hist. Nat., vol. 41 (1937), pp. 265-270, fig. 37, Nos. 1-7; pl. 11, figs. 9, 10, 1938; Bol. Mus. Nac. Hist. Nat., vol. 16, p. 1238, 1938. Tatochila theodice flammivolans Bryk, Ark. f. Zool., vol. 36a, pp. 6, 7, Nov. 6, 1944, Tatochila theodice flammivolans f. antiquincune Bryk, Ark. f. Zool., vol. 36a, p: 7, pl. 1, fig. 9, Nov.'6;,1944: Tatochila theodice autodice (Boisduval), Forster nec Boisduval (in error for T. theodice theodice), Veroffentl. Zool. Staatss. Mimchen, vol. 3, p. 134, Dec. 1, 1955. Venation: As illustrated (fig. 56). Leas: With paronychium, claw, and pulvillus as illustrated (fig. wo) Mate: Plate 1, figures 3, 4. Head with scales behind eyes and with outer faces of palpi white to pale yellowish white. Wings above white with dark markings as illustrated, having the dark scaling along veins more heavy on outer margin, forming inwardly directed wedge- shaped marks. Both wings with a submarginal series of sagittate markings. Forewing with a large discal spot. Wings below with apex of forewing and ground color of hindwing pale yellow to white. Veins white and outlined on both sides with fuscous. Both pair of wings with a submarginal and intervenal row of sagittate markings. Hindwing with a fuscous streak through cell, forked near end of cell, with costal margin ornage-yellow, with a long orange-yellow streak below cell and with long narrow intervenal streaks of the same color on outer margin and directed inward through the sagittate markings, continuing toward the cell. Length of forewing, 25-30 mm. (average 27.1 mm.). Male genitalia as illustrated (figs. 29, 42), with characters as described for the group. No differences were observed between the three subspecies. The genitalia of seven males of typical 7. theodice were studied. Femae: Plate 1, figures 3a, 4a. Head as in the male. Wings above very similar to those of the male, differing in having ground color somewhat yellowish or cream colored and with all dark mark- ings more distinct than in the male. Wings below with ground color as in the male with yellow streaks usually darker and more distinct. Length of forewing, 25-29 mm. (average, 26.8 mm.). Female genitalia as illustrated (fig. 86), with posterior lobe of inner genital plate uncinate and with a small basal fingerlike process. Anterior lobe of inner genital plate subtriangular, with posterior and BUTTERFLIES—-HERRERA AND FIELD 477 anterior free angles greatly produced and with ventral margin un- dulate. Ductus bursae with ribbonlike plate large, fused to anterior subtriangular plate and with a pair or more of small irregularly shaped plates anterior to this. As in the male, no differences are displayed between the three subspecies. The genitalia of six females were studied. SEASONAL VARIATION: In the spring and sometimes in the fall, this Chilean species produces a form that is smaller (o, 19-25 mm., average 22.6 mm.; 9, 22-24 mm., average 22.8 mm.) with all markings correspondingly reduced in size (pl. 1, figs. 4, 4a). Type Locauity: Herewith restricted to central Chile. A mixup over the type locality is the reason for the restriction. Boisduval, in his original description, cited the type locality as “Bourou” (Buru, an island in Indonesia). Later, he (1836) corrected this to Payta, Peru. However, this locality is also in error as no species found in Peru fits either the original description or his subsequent description, while the species here discussed from central Chile and southward does agree with his descriptions. ADDITIONAL TYPE DATA AND LOCATION OF TyPES: J’. theodice was originally described from the female sex, the number of specimens not being given. The type should have gone to the British Museum from the Oberthtir collection. Mr. D. S. Fletcher of that museum writes that the type is not in the collection there. He states further that there is a Boisduval specimen labeled ‘‘type’”’ by some subse- quent worker but that it is certainly not the type, ‘for even if Bois- duval were to mistake his sexes’”’ (the specimen in question is a male) he would never mistake pure white for ‘‘white, a little yellowish.” Mr. Jean Bourgogne of the Paris Museum searched the collections there and reported that the type of theodice is not present. The type is apparently lost. MeETrFOD OF IDENTIFICATION: Based upon the original and subse- quent descriptions of Boisduval which positively fix the name theodice to the species here described. )": oe ey mF S i Xt _ VANSS Ei Sy JSF } ¢ Sate Vv: a Se}. san FNS - - Gor Sa ee ee em Seescasd Pare x ty So Oe eet oie ene Tes wm eh FL Se ag re Nes 563 Figure 6.—a, Paraplanocera fritillata, sagittal view of copulatory complex, male above. b, c, Asolenia deilogyna: b, entire animal; c, sagittal view of copulatory complex, male above. (Explanation on page 597.) 564 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 prostatic vesicle and the ejaculatory duct of the seminal vesicle meet and unite in the proximal part of the penis papilla and the common duct so formed extends along the center of the penis papilla to its tip. The penis papilla is an elongated structure with a thick muscular wall covered with an epithelium that is thickened at the penis tip and proximally becomes continuous with the lining epithelium of the rather spacious male antrum, which exits ventrally by a narrow tubu- lar passage. Under the epithelium of the male antrum is found a considerable muscular investment. The female apparatus is very curious, much reduced, and springing from the roof of the male antrum, something quite unusual in poly- clads. This condition could hardly result from contraction on pres- ervation. The female antrum as it leaves the male antrum is a nar- row tube encircled by a thick muscular sphincter. The parts of the female tract are unfortunately poorly preserved in the sections. The antrum widens to an ascending vagina that then descends as a tube with a wall of cuboidal cells. This seems to make a curve and ascend but preservation was poor here. There next comes an oval glandular part of the vagina, definitely seen, entered by cement glands. A narrowed tube leaves this and descends, then widening to a sac that seems to receive two ducts; the connection with these ducts could not be followed although very probably it is as shown in the figure. These two ducts descend behind the male antrum and then curve anteriorly. They seem to be the uteri and could be followed, although not too clearly, to the rear ends of the egg-filled uteri shown in figure 6b. Itis possible that the two ducts are the double Lang’s ves- icle characteristic of the genus Callioplana, but in that case the uteri would enter the vagina beyond the glandular region and no indication of ducts here could be found. Hototrypr: USNM 28686, one set of sagittal sections (one slide). Remarks: Since the female tract was unclear, it was difficult to make a decision about this worm. If the two ducts discussed above are Lang’s vesicles rather than uteri, the specimen could be fitted into the genus Callioplana, with which the male system is in accord al- though the lack of tentacles and the characters of the female system disagree with the two known species of that genus: C. marginata (Stimpson) (1857) and evelinae (Marcus) (1954). The matter cannot be decided until better material is obtained. Qn o> or POLYCLAD FLATWORMS—HYMAN Section EMPROSTHOMMATA Family CESTOPLANIDAE Juvenile Cestoplanid Remarks: There was present in the material from Ifaluk Atoll one very juvenile cestoplanid, collected Oct. 1, 1953, from the reefridge at the north end of Falarik. A drawing was not made. It may be recalled that a juvenile cestoplanid was previously reported from the same general area (Hyman, 1955b), but in view of the immaturity of both specimens a specific comparison is impossible. The specimen is de- posited in the U. S. National Museum. Suborder CoTyLEa Family PsEUDOCERIDAE Lang, 1884 Genus Pseudoceros Lang, 1884 Remarks: The genus Pseudoceros is one of the largest polyclad genera, comprising at present over 100 valid species. The genus is easily recognized by the combination of smooth dorsal surface, ten- tacles as upfolds of the anterior margin, and ruffled pharynx. The species, however, are distinguishable mainly on shape and color pattern and the one is distorted and the other often totally lost in preserved specimens. Hence the identification of preserved specimens usually offers formidable difficulty. Whether the male apparatus is single or paired is a useful character and details of the male copulatory apparatus may be of value in specific diagnoses. It now appears that the shape of the pharynx may be decisive. In most species the pharynx has a compact outline but several species are now known in which the pharynx takes what I have termed the butterfly shape, putting out lateral lobulations that increase in length in the anteroposterior di- rection as in figure 10, c. In 1950 Marcus published a useful list of the described species of the genus. In the thought of increasing the usefulness of this list I here add some old species overlooked by Marcus and the species described since that date: fulminatus (Stimpson) (1855), guttatomar- ginatus (Stimpson) (1855), interruptus (Stimpson) (1855), albicornis (Stimpson) (1857), coccineus (Stimpson) (1857), gaponicus (Stimpson) (1857), niger (Stimpson) (1857), affinis (Kelaart) (1858), atraviridis (Kelaart) (1858), dulcis (Kelaart) (1858), fuscus (Kelaart) (1858), pur- pureus (Kelaart) (1858), luteus (Plehn) (1897), czwensis Kato (1944), nipponicus Kato (1944), evelinae Marcus (1950), mopsus Marcus (1952), bajae Hyman (1953a), canadensis Hyman (1953a), meaicanus Hyman (1953a), montereyensis Hyman (1953a), coralliferus Hyman (1954), micronesianus Hyman (19538b), and texanus Hyman (1955c). 566 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 I am of the opinion that Pseudoceros gratus Kato (1937, also 1943) is identical with Hurylepta striata Schmarda (1859). To be sure, the former is described as white with black stripes and the latter as buff with dark brown stripes but the identity of size, shape, and color pattern outweigh such slight color differences which can be caused by ingested food. As the name striata is preoccupied by Eurylepta striata Kelaart (1858), it appears that Kato’s name gratus is valid. Therefore the proposal of strigosus by Marcus (1950) to cover the Kelaart-Schmarda homonymy is unnecessary. The genus is characteristic of tropical and subtropical waters and appears to center in the Indo-West Pacific area. Pseudoceros perviolaceus, new name Eurylepta violacea Schmarda, 1859, p. 27. Pseudoceros violaceus, Stummer-Traunfels, 1933, p. 3544. Not Planaria violacea Kelaart, 1858, p. 135. Remarks: A specimen referred to this species that was collected in the Palau Islands, July 22, 1954, was sent to the American Museum of Natural History in a lot of miscellaneous material. No other data are available. The specimen conforms satisfactorily to the original description and figure and with Stummer-Traunfels’ statements made from Schmarda’s specimen. The original specimen from Ceylon measured 60 by 40 mm., the present one 25 by 13, but presumably Schmarda’s measurements are from life. The present specimen re- tains the shape depicted by Schmarda, that is, is broad across the anterior end and narrows regularly toward the posterior end. There is but a single male apparatus as discovered by Stummer-Traunfels. The violet or purple color had dissolved badly in the alcohol which was stained a reddish purple, leaving the animal of a uniform medium brown color. It was necessary to create a new specific name for Schmarda’s species as the name violacea is preoccupied by Planaria violacea Kelaart, which is a Pseudoceros but definitely not identical with Schmarda’s species for it has a broadly oval shape and a yellow margin and middorsal stripe. It appears improbable that Schmarda’s species can be a variant of Pseudoceros velutinus as supposed by Lang (1884, p. 540), for the shape of this is quite different although it, too, has but a single male apparatus. The specimen is retained in the invertebrate section of the American Museum of Natural History. Pseudoceros bedfordi Laidlaw, 1903 Remarks: This large and handsome pseudocerid is easily recog- nized by the distinctive color pattern, accurately depicted by Bock POLYCLAD FLATWORMS—HYMAN 567 (1913) and Kato (1943). The present specimen (USNM 28650)? was found swimming near the reef at Ifaluk Atoll, Aug. 12, 1953. Alive it was stated to be at least 100 mm. long, hence is the largest on record; preserved it measured 35 by 36 mm., having contracted strongly to a rounded shape whereas the natural shape is an elongated oval. This species was reported by Kato (1943) from the Palau Islands where it was stated to be common although surprisingly not recovered in the present material from these islands. It is also re- corded from Singapore, the Philippines, and Heron Island in the Great Barrier Reef and is evidently widely spread in the western tropical and subtropical Pacific. Pseudoceros izuensis Kato, 1944 FIGURE 7a-c Remarks: One specimen was taken on a reef in the Palau Islands by the Stanford team, Aug. 8, 1955, Sta. 69, lagoon margin of reef southeast of Malakal Pass. As the original description is brief, an expanded account of the present specimen appears desirable, espe- cially as there is some slight doubt of its identity with Kato’s species. The preserved specimen (USNM 28651) measures 22 by 16 mm. and is of broadly oval shape and thin consistency (fig. 7a). Kato gave the length, preserved, as 20 by 13 mm., but 60 by 28 in life. The tentacles as sketched by Bayer in life appear in figure 76; their appearance in the preserved animal is shown in figure 7c. The ruffed pharynx had ruptured through the ventral surface. The color in life as described by F. M. Bayer, who also took a color photograph, was pale green above mottled with white and dotted with black dots, with a median ridge of sepia brown laced with white. The tentacles in life (fig. 7b) are sepia brown with white tips and white spots. The body is encircled by a marginal band of olive green composed of radial streaks. Preserved, the animal appears light gray dotted with black dots with a dark margin and a dark middorsal band laced with white. The radial streaks composing the marginal band are still detectable in the preserved specimen. Figure 7a attempts to depict the color pattern. The colors agree well with Kato’s description except for the tentacles which he described as purplish and the absence of black at the extreme outer margin. Behind the tentacles is seen the bilobed cluster of cerebral eyes (fig. 7c). This disagrees with Kato’s statement that the eyes form a single cluster although his figure indicates a slight bilobulation. Kato figured the tentacular eyes, not well discernible in the present specimen. 2 A photograph, in color, of this specimen was published in National Geographic Magazine, vol. 109, No. 4, p. 557, April 1956. 568 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ficure 7.—Pseudoceros izuensis: a, entire animal; b, tentacles from life; c, tentacles of preserved specimen. (Explanation on page 597.) The specimen is sexually mature but because of the rupture of the pharynx, the number of male pores could not be ascertained, although two are probably present. Kato stated a paired condition of the male apparatus. At the posterior end of the pharynx the female gonopore with accompanying cement glands is obvious (fig. 7a). The specimen has two suckers, one behind the other, as shown in figure 7a, but this is presumably an anomaly. A central position of the sucker is indicated by Kato. POLYCLAD FLATWORMS—HYMAN 569 The small differences mentioned above do not seem adequate grounds for separating the present specimen from Kato’s species. Pseudoceros izuensis shows considerable resemblance in color pattern to P. viridis (Kelaart) (1858), Ceylon, which according to the descrip- tion and colored figure in Collingwood (1876) also is green with brown tentacles, a brown middorsal stripe, and a brown streaky margin; but the dorsal surface is splotched with brown rather than dotted with black. Contrary to Lang (1884, p. 567), I think viridis (Schmarda) (1859) is not identical with viridis (Kelaart) as the color patterns of the two species appear somewhat different. It therefore is necessary to rename Schmarda’s species, and I propose virescens. Pseudoceros habroptilus, new species FIGurRE 8 MateEriAL: One specimen taken by the Templeton Crocker Expe- dition, No. 1014, June 5, 1933, at Vanikoro Island, in the Solomons. GENERAL CHARACTERS: The worm was badly crumpled and the posterior part could not be straightened. After flattening as much as possible the worm appeared as in figure 8. It measures 10 by 6 mm. and is probably of elongated oval form. At the anterior end appear the usual tentacular upfolds of which one is well preserved while the other is contracted and distorted. The tentacular eyes were poorly discernible; the single oval cerebral cluster is shown in the figure. The black-and-white color pattern is pretty and distinctive. The worm is white with a narrow, very black border that also edges the tentacular folds. Paralleling the margin is a wider band of grayish black hue and centrally there is a paired band of similar width and color. The two median bands converge anteriorly and fuse just behind the cerebral eyes. The black bands could not be followed posteriorly because of damage and crumpling here but the black margin obvi- ously encircles the entire worm. The pharynx had probably been shed as rupture was evident ventrally at the appropriate place. The sucker was not clearly evident and is shown conjecturally in figure 8. The worm is juvenile without any traces of the reproductive system. DIFFERENTIAL DIAGNOSIS: The color pattern is sufficiently distinc- tive. Houotyrpre: USNM 28652, the worm mounted whole. Pseudoceros caeruleocinctus, new species FIGURE 9a Materia: The single specimen was taken by the Stanford team at the Palau Islands, Sta. 77, Aug. 10, 1955, crawling on clean sand in a shallow bay on the south shore of Auluptagel Island. 570 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 : Figure 8.—Pseudoceros habroptilus, entire animal. (Explanation on page 597.) GENERAL CHARACTERS: The preserved worm is of oblong shape, 20 mm. long by 12 mm. wide. The tentacular foldings are prominent (fig. 9a), but because of the black color the eyes could not be seen in the cleared worm. The color is described by the collector as velvety black with a narrow, brilliant blue border. Some trace of the blue border remains in the preserved worm. The pharynx was not detectable. There is a single male apparatus with the male pore located 5 mm. from the anterior end. The female pore occurs close behind the male pore. The sucker is located at about the body middle, 10 mm. from the anterior margin. POLYCLAD FLATWORMS—HYMAN 571 DIFFERENTIAL DIAGNOSIS: The blue border distinguishes this species from other black species with a colored border. This pattern, black with a narrow border of a bright contrasting color, is common among species of Pseudoceros, but usually the border is red, orange, or yellow, and I know of no previously described species with a blue border. Ho.totyrze: USNM 28653, in alcohol. Pseudoceros ferrugineus, new species Fiaure 9b,c Martsriau: The single specimen was taken by the Stanford team at Sta. 2364, in Iwayama Bay, Palau Islands, Oct. 20, 1955, crawling in about a meter of water on the rocky shelf of the east end of Koror Island, in Oyster Pass. GENERAL CHARACTERS: The worm is of elongated oval shape (fig. 9b), measuring 18 mm. in length by 11 mm. in width, preserved. The tentacular foldings appear slightly developed, both in the pre- served worm (fig. 9c), and on the color photograph of the live worm. The brillant coloring in life is preserved on the photograph taken by F. M. Bayer, but, alas, the preserved worm is a dull grayish brown. In life the general color is a bright rusty red blending into a narrow margin of brilliant orange. The dorsal surface is liberally flecked with white on the rusty background and here and there these flecks fuse to form nebulous patches of clear white or tinged with pink in places. The white flecks diminish towards the margin where the rusty red background is much more in evidence. The bright orange marginal line is free from spots. There is some indication of a narrow middorsal line of pale rust that fades away posteriorly. In the cleared worm (fig. 96) there could be seen the slightly bilobed cluster of cerebral eyes (fig. 9c), the ruffled pharynx having the but- terfly shape that I have noticed in some species of Pseudoceros, the pair of male pores in the concavity of the last lobulations of the pharynx, the female pore close behind the male pores, and the sucker slightly behind the middle, about 10 mm. from the anterior end. DIFFERENTIAL DIAGNOSIS: The color pattern is distinctive. Hotorypre: USNM 28654, in alcohol. Pseudoceros ater, new species FIGURE 9d MatTerRIAL: The single specimen was taken by the Stanford team on Raeldil reef, Palau Islands, Sta. 254, Oct. 27, 1955. It was col- lected at night with a light on the outer reef flat among branching corals. GENERAL CHARACTERS: This is a small worm, measuring 9 by 7 mm. preserved, although evidently somewhat contracted, hence longer in 572 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 Ficure 9.—a, Pseudoceros caeruleocinctus, entire animal. 6, c, P. ferrugineus: b, entire animal; c, tentacles enlarged. d, P. ater. (Explanation on page 597.) POLYCLAD FLATWORMS—HYMAN 573 life. The form is an elongated oval (fig. 9d). The tentacular upfold- ings are well preserved. Behind them is seen the small, compact ruffed pharynx of a few folds, and directly behind this occurs the single male gonopore, followed by the female pore. The sucker is situated somewhat posterior to the middle of the worm. Despite its small size the worm was fully mature and laid a small globular ege mass during the night in the jar in which it had been placed. The color is a uniform very dark grayish black, or practically black, without any markings. Because of the black color, eyes could not be seen. DIFFERENTIAL DIAGNOSIS: There does not seem to be any previously described species of Pseudoceros that is small and uniformly of a dull black coloration. Pseudoceros velutinus (Blanchard) (1847) is uni- formly black but the black is of a velvety texture and tinged with blue or violet; this species is further quite large, being still not fully mature at a length of 50 mm. Pseudoceros bajae Hyman (1953a) is also quite large, with a pair of male apparatuses. Houotryre: USNM 28655, in alcohol. Pseudoceros fulvogriseus, new species Ficure 10a,b MatTertrAL: One specimen was taken by the Stanford team in Geruherugairu Pass, Iwayama Bay, Palau Islands, Sta. 85, Aug. 12, 1955. GENERAL CHARACTERS: The worm is a large, elongated species, 50 mm. long, preserved, 20 mm. across the widest part. The exten- sive ruffling of the margins indicates a much greater length when crawling, extended. The worm appears widest across the middle and tapers from this to the rounded anterior end, with typical tentac- ular foldings, and posteriorly to the pointed tail (fig. 10a@). The shape is unusually elongated for the genus. An enlarged view of the tentac- ular foldings is shown in figure 10, which also gives the eye distribu- tion. Shortly behind the tentacles is found the group of cerebral eyes, forming a single large rounded cluster. Behind the cerebral eye cluster is seen the compact ruffled pharynx, surprisingly small for the size of the worm. Posterior to the pharynx are the two male pores, unusually close together, located 10 mm. from the anterior end. They are followed by the female pore, 13 mm. from the anterior end. The sucker is located at about the middle, 25 mm. from the anterior end. The worm appeared in breeding condition for the branched, anastomosing uterus (shown on one side only in figure 10a) was very evident in the cleared specimen. 574 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 aay By “ EEO ERT pag i s th hen, aw A Qe Sas mY ® we forts * oa enter ® oo SFeRXSO Ficure 10.—a, b, Pseudoceros fulvogriseus: a, entire animal; b, tentacles enlarged. c, d, P. fuscogriseus: c, entire animal; d, tentacles enlarged. (Explanation on page 597.) The preserved worm is of a uniform medium gray color, marked with dark reticulations, which probably actually are the uterine anasto- moses. A color photograph taken by F. M. Bayer shows that in life POLYCLAD FLATWORMS—HYMAN 5d the color is light gray flushed with yellowish brown. Along the center of the median ridge there runs a narrow light line flanked on each side by yellowish brown. The yellowish brown color deepens near the margin which is edged by a very narrow light line. The tentacles appear brown edged with a light margin. There is some suspicion in my mind that the tawny brown color may be caused, at least in part, by the presence of ripe eggs in the interior. DIFFERENTIAL DIAGNOSIS: The worm is characterized by the large size, elongated shape, small compact pharynx, and color pattern. Houoryre: USNM 28656, in alcohol. Pseudoceros fuscogriseus, new species Fiaure 10c,d MATERIAL: Two specimens were collected by the Stanford team in eel grass in the channel between Peliliu and Ngedebus, Palau Islands, Sta. 37, July 25, 1955. GENERAL CHARACTERS: The worms are of moderate size and general oval form, tapering posteriorly to a pointed tail (fig. 10c). The larger specimen measured, preserved, 23 by 10 mm., the smaller one, 16 by 8mm. Both worms were bent in the pharyngeal region and ruptured there when straightened out, hence it did not appear profitable to section the copulatory region of one of them as this would probably be involved in the rupture. The anterior end of the larger specimen while cleared is shown in figure 10d. The tentacles, much distorted by preservation, are provided with numerous eyes and well behind them is seen the group of cerebral eyes, consisting of two oval clusters. The relatively large ruffled pharynx has the butterfly shape already mentioned in connection with P. ferrugineus. The single male pore is embraced by the posterior pharyngeal folds and shortly behind it is seen the female gonopore. Thesucker of the larger worm is located at about 12 mm. from the anterior end. The usual middorsal ridge seen in pseudocerids is indistinct here. The main intestine could be seen in the posterior part of the worm. Unfortunately, no information is available about the color in life but probably it was the same as in the preserved worms. They are dusky grayish brown in color with the anterior central part of a somewhat lighter shade. The extent of the light region is indicated by the dotted line in figure 10c. The line passes along the edge of the pharynx, slightly behind the sucker, and diverges at the anterior margin. The boundary between the lighter and darker shades of grayish brown is quite sharp. DIFFERENTIAL DIAGNOSIS: This species lacks very definite charac- teristics but the combination of paired cerebral eye clusters, large 472590—59 8 576 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 ruffed pharynx of the butterfly type, single male gonopore, and coloration should afford recognition. Hototyre: The larger specimen, USNM 28657. Pseudoceros tristriatus, new species FiagurRE lla Pseudoceros concinnus, Stummer-Traunfels, 1933, p. 3565, fig. 9 on color pl. Not Proceros concinnus Collingwood, 1876, p. 90, fig. 4, pl. 17. MareriAu: A single specimen was taken from beneath rocks on the seaward shore of Ella Islet, Ifaluk Atoll, western Carolines, Sept. 20, 1953, by F. M. Bayer under the auspices of the Pacific Science Board Atoll Research Program. GENERAL CHARACTERS: The preserved worm (fig. lla) is of oval form, measuring 12 by 5.5 mm., but a color photograph taken by F. M. Bayer shows that it is more elongated in life, about 4 times as long as broad. The preserved worm is entirely black but in life it is light blue with three longitudinal orange stripes. These stripes appear faintly bordered with black and extend from shortly behind the tentacles almost to the posterior end. The two lateral stripes are confluent posteriorly behind the median stripe. In the preserved worm the tentacular folds are fairly well retained and a few eyes can beseen upon them. Behind their bases is a rounded cluster of cerebral eyes but the eye arrangement could not be satisfactorily ascertained because of the dense black color of the preserved worm. The pharynx is of the butterfly type, that is, with pronounced lateral lobes increasing in length posteriorly where they slant backwards. Behind the pharynx the main intestine 1s conspicuous in the cleared worm, giving off numerous side branches that enter a dense black network of intestinal branches spread throughout the body. The sucker is located about 5 mm. from the anterior end in the preserved specimen. The worm is juvenile, being devoid of any traces of the reproductive system. Hototrypr: USNM 28659, one whole mount. Remarks: Despite the immaturity of the specimen, the very dis- tinctive color pattern justifies giving it a name. I believe this specimen is identical with the one figured in Stummer-Traunfels (1933, fig. 9 on col. pl. following p. 3596), which he identified as Pseudoceros concinnus (Collingwood). I believe this identification is erroneous, for Collingwood’s colored figure gives an entirely different color pattern: cream with a blue border and blue middorsal stripe. I previously called attention to Stummer-Traunfels’ error when I described as P. concinnus specimens from New Guinea (Hyman, 1954). Stummer-Traunfels’ colored figure that I regard as identical with the 577 POLYCLAD FLATWORMS—HYMAN \. \ wn) mae PLUS) JARI TIN II Ficure 1l.—a, Pseudoceros tristriatus. b, c, Nymphozoon bayeri: b, female copulatory apparatus, sagittal view; c, sagittal view of male copulatory apparatus. (Explanation on page 597.) present species differs from it only in that the lateral orange stripes do not reach as far posteriorly, hence are not confluent behind the median stripe. This appears an insignificant difference. Stummer- Traunfels’ specimen was part of the Semper material collected in the Philippines and the Palau Islands and not yet published; the colored figure was made from life by Mrs. Semper. 578 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 Nymphozoon, new genus Dertnition: Pseudoceridae with multiple female apparatuses, arranged in a midventral longitudinal row; sucker wanting; otherwise as in Pseudoceros. Typr spuctus: Nymphozoon bayert, new species. Nymphozoon bayeri, new species Figures 116,c; 12a,b MateEriAL: Two specimens were taken by the Stanford team, one on a reef flat at Iwayama Bay, Palau Islands, Sta. 133, Aug. 28, 1955; the other on shallow coral sand and eel grass in the same region, Sta. 85a, Oct. 29, 1955. GENERAL CHARACTERS: This is a very large, handsome, black-and- white pseudocerid of delicate consistency. The larger specimen is 70 mm. long by 55 mm. wide, preserved, the smaller one 50 mm. by 30 mm. From the ruffling of the margins one may surmise that a considerably greater length may be attained in life. A photograph of the smaller specimen in life shows it to have been about 75 mm. long. The shape is broadly oval tapering to a somewhat pointed posterior end (fig. 12a); anteriorly there are present the usual tentacular folds. Figure 12a attempts to depict the striking color pattern. There is a narrow, very black, sharply delimited band along the margin that also edges the tentacular folds. Medially there is a moderately broad black longitudinal band that tapers to a point behind the tentacular folds and narrows posteriorly, but it could not be followed completely here because of damage. Between the median band and the margin there is present on each side a broad lateral band of grayish black hue that also could not be followed to the posterior end. The remainder of the animal is pure white. The large ruffled pharynx with central mouth is drawn in figure 126. The smaller specimen is definitely devoid of any indication of the reproductive system but the larger specimen is fully mature. As shown in figure 126, there is a pair of male pores behind the pharynx and this is succeeded by a midventral longitudinal row of eight pores, somewhat unevenly spaced. The nature of these pores could not be ascertained without sections and these showed that they are female gonopores. 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