I ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME 37 • 1952 • NUMBERS 1 TO 21 PUBLISHED BY THE SOCIETY The ZOOLOGICAL PARK, New York Contents Part 1. June 30, 1952 Page 1 . A New Dwarf Toad from Southeastern Brazil. By George S. Myers & Antenor Leitao de Carvalho 1 2. Sexual Broods and the Production of Young Queens in Two Species of Army Ants. By T. C. Schneirla & Robert Zanes Brown. Plates I-III; Text-figures 1&2 5 3. Spectral Effects on the Growth Rate and Endocrine Histology of the Teieost, Astyanax mexicanus. By Phyllis H. Cahn. Text- figures 1-4 33 4. Fungus Associated with a Granuloma in a Turkish Fish, Aphanius chantrei Gaillard. By Recai Ermin. Plates I & II; Text-figures 1-11. 43 5. Deep-sea Fishes of the Bermuda Oceanographic Expeditions. Fam- ilies Cetomimidae and Rondeletiidae. By Robert R. Harry. Plate I; Text-figures 1-4 55 Part 2. September 19, 1952 6. The Effects of Prolonged Treatment with Acriflavine on the Killifish, Fundulus heteroclitus (Linnaeus). By Grace E. Pickford 73 7. Effects of ACTH and Cortisone on the Pituitary, Thyroid and Gonads of the Teieost Astyanax mexicanus. By Priscilla Rasquin & Ethel Hafter Atz. Plates I & II; Text-figures 1 & 2 77 8. The Effects of Holothurin on Fish, and Mice with Sarcoma 180. By Ross F. Nigrelli 89 9. Sex Determination in Xiphophorus (Platypoecilus) maculatus. III. Differ- entiation of Gonads in Platyfish from Broods Having a Sex Ratio of Three Females to One Male. By Myron Gordon. Plates I & II; Text-figure 1 . . . . 91 10. Four New Species of Geometridae (Moths) from Rancho Grande, North- central Venezuela. By D. S. Fletcher. Plate I; Text-figures 1-7 101 Part 3. October 31, 1952 Page 11. The Mutillidae (Wasps) of British Guiana. By Clarence E. Mickel. . 105 12. A Revision of the Fishes of the Subfamily Alfarinae in the Family Poeciliidae. By Donn Eric Rosen. Text-figures 1-10 151 Part 4. December 31, 1952 13. Introduction to the Ecology of the Arima Valley, Trinidad, B.W.I. By William Beebe. Plates I-V; Text-figures 1-5 157 14. Spontaneous Neoplasms in Fishes. VI. Thyroid Tumors in Marine Fishes. By Ross F. Nigrelli. Plates I-IX 185 15. Further Comparisons of Length and Voltage in the Electric Eel, Electro- phorus electricus (Linnaeus). By M. Vertner Brown & C. W. Coates. Text -figures 1-7 191 16. A Contribution to the Life History of Colobura ( Gynaecia auct.) dirce dirce (Linnaeus). (Butterfly). By William Beebe. Plates I & II 199 17. The Saturnioidea (Moths) of Rancho Grande, North-central Venezuela. By Henry Fleming 203 18. The Functional Morphology of the Egg-eating Adaptations in the Snake Genus Dasypeltis. By Carl Cans. Plates I-IV; Text-figures 1-15 209 19. An Annotated List of the Mantids (Orthoptera, Mantoidea) of Trinidad, B.W.I. By William Beebe, Jocelyn Crane & Sally Hughes-Schrader. Plates I-VIII; Text-figures 1 & 2 245 20. A Comparative Study of Innate Defensive Behavior in Trinidad Mantids (Orthoptera, Mantoidea). By Jocelyn Crane. Plates I-X; Text-figures 1 & 2 259 21. An Experimental Study of the Color Vision of the Giant Tortoise. By John V. Quaranta. Plates I & II; Text-figures 1&2 295 ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME 37 • PART I • JUNE 30, 1952 • NUMBERS 1 TO 5 PUBLISHED BY THE SOCIETY The ZOOLOGICAL PARK, New York Contents Page 1 . A New Dwarf Toad from Southeastern Brazil. By George S. Myers & Antenor Leitao de Carvalho 1 2. Sexual Broods and the Production of Young Queens in Two Species of Army Ants. By T. C. Schneirla & Robert Zanes Brown. Plates I-III; Text-figures 1&2 5 3. Spectral Effects on the Growth Rate and Endocrine Histology of the Teleost, Astyanax mexicanus. By Phyllis H. Cahn. Text- figures 1-4 33 4. Fungus Associated with a Granuloma in a Turkish Fish, Aphanius chantrei Gaillard. By Recai Ermin. Plates I & II; Text-figures 1-11. 43 5. Deep-sea Fishes of the Bermuda Oceanographic Expeditions. Fam- ilies Cetomimidae and Rondeletiidae. By Robert R. Harry. Plate I; Text-figures 1-4 55 1 A New Dwarf Toad from Southeastern Brazil George S. Myers & Antenor Leitao de Carvalho Stanford University and Museu Nacional, Rio de Janeiro THE authors have examined a large series of the comon South American toad, Bufo granulosus Spix (called also B. globulosus Spix), from the lower Amazon (Manaus, San- tarem), the Rio Araguaya, Ceara, Minas Gerais, and the upper Rio Paraguay (Sao Luiz de Ca- ceres). These show clearly that this widespread species varies geographically with considerable consistency, and that several subspecies will probably be recognized eventually. The large series from the Araguaya (Rio Tapirape) shows a flattened muzzle which easily separates this lot from the toads we collected at Pirapora in Minas Gerais, and, indeed, from the Manaus and Santarem examples. The Sao Luis de Caceres individual exhibits a peculiar shortened physiognomy and strong infranarial crest different from all the others. In some ways this specimen approaches two toads from Asun- cion, Paraguay, which we were at first inclined to call Bufo d’orbignyi Dumeril & Bibron. How- ever, we have not seen typical examples of the latter (from Uruguay) and Boulenger’s 1882 figure of the head of B. d’orbignyi (Montevideo specimen), which shows no serrations or warts on the cranial crests, leads us to suspect that these Asuncion toads may lend weight to Park- er’s (1936) suggestion that the flat-headed B. granulosus and the deep-headed B. d’orbignyi intergrade in the Paraguay Basin. Strikingly different from all the toads of this type that we have examined is a fine series col- lected by the junior author in the restinga (scrub beach vegetation) of the eastern part of the State of Rio de Janeiro. These represent a form close to typical B. granulosus but sharply dis- tinct in its minute size, striking coloration and different cranial morphology. While there is a possibility that this strange little toad (perhaps the smallest South American Bufo) will even- tually be found to intergrade with B. granulosus, we are not sure of this, and, on the basis of our material, venture to predict the existence, in the littoral district of the State of Rio and perhaps a wider area, of a quite distinct, pigmy species of Bufo belonging to the granulosus complex and agreeing fairly well with the following diagnosis. Bufo pygmaeus, new species Holotype — An adult male of average size, 37.5 mm. in snout-to-vent measurement, taken while calling by A. L. de Carvalho and L. de Castro Faria at Sao Joao da Barra, at the mouth of the Rio Parahyba (right bank). State of Rio de Janeiro, Brazil, in December, 1948. To be deposited in the Museu Nacional, Rio de Janeiro. Paratypes — Eighty-five examples, taken with the holotype while in breeding congress, and at the same time and place. Most of the paratypes are in the Museu Nacional, Rio de Janeiro. Others are in the Natural History Museum of Stanford University (nos. 10720-10723). Two have been sent to the U. S. National Museum. The specimens are all adult, even the smallest males possessing a vocal sac and a nuptial pad on the first finger of each hand. Other material— Three examples in the Museu Nacional from the Restinga de Marambaia, to the westward of the city of Rio de Janeiro, indi- cating a general distribution in the littoral region of the State of Rio de Janeiro. Diagnosis— Similar in most respects to Bufo granulosus Spix but differing especially in the very small size (largest female 42 mm. in head and body length; largest male 41 mm.); in the much bolder light-and-dark mottling and mar- bling of the dorsum; in the much stronger de- velopment of a longer, curved, supratympanal crest; in the unserrated subocular crest; and, above all, in the development of a strong parietal crest. Discussion.— This small toad is a vicarious (allopatric) form of the granulosus complex but, insofar as our material shows, it is a distinct species. In B. granulosus, most of the bony crests 1 JOL 10 1952 Table 1. Measurements of Bufo pygmaeus in Millimeters 2 Zoologica: New York Zoological Society [37:1 Of CN in CN 00 in ■ CN NO 00 00 NO in in o Ov CN Of o ON in NO m N- in O in m d 1-H NO r-H r”' oo cn CN in cn CN on »-h r— 1 00 rH ON CN in oo on on 'O • 03 • .5 ’ OX) G *0 G G C M no CL, £ G H 03 O hJ d> - o o CD b on <2 o- ° I p a - > a x 2 CQ 'So •o "b <1> g Q-t £ CU »~ a o ~ e > 03 O O 03 g 3 •£ X « -r-t .5 "d T3 »-i o r ) (D Ui g-* 05 p 1) ^ 00 C/5 03 X « Q o . ON fej w-i s 5 Q 3 s o u ° o CO Jj ' d> o3 OO o ; gs - c : ojD C3 .E d <-W cr o > a JJ o p* cS — -C 6fl 3 « ° -w « ' — 1 G « « 13 ■ S3 ~ g- g>2 8 - 5 0) oo Q 0 1 .a <3 03 .u&Pgp* CO C 03 O ‘-d c 3 d> ■e * ‘C 2 05 „ ’^s <*> Q .s> c o £ hi 03 a Q a b 05 § 8 ^ c 2 o 2§w® lO . . .-b c 2-2 « 3 e| 13 .22 > *d *- a «Q s ° qj -C "5 rt ** ’b -S £ /2 30.0 10.0 * Standard errors not calculated when less than three fish left in group. green filters, whereas radiant energy under the yellow filter was considerably greater, although still inferior to the controls. Results General behavior. — No behavior derange- ments could be attributed to the colored filters. As far as could be determined, feeding was nor- mal and regular in all of the groups. Astyanax mexicanus is known to exhibit frequent “ner- vous” behavior patterns. In the course of one or several days these fish sometimes alternate Zoologica: New York Zoological Society [37: 3 36 CONTROL RED YELLOW GREEN BLUE Text-fig. 3. Monthly mean lengths and depths of the fish are indicated by small circles. Monthly extreme values of length and depth are indicated by vertical lines. from a relatively quiescent state to an extremely active hostile condition, which occasionally re- sults in a group annihilation (Breder, 1943). Survival.— There were no differences in the ability to survive under the colored filters. Dur- ing the course of the experiment one fish from each group except the control died from un- known causes. It appeared, however, that death was not caused by the experimental environ- ments. Growth.— The filtered light did not inhibit the body growth of the fishes, as seen from the data in Table 1, where the monthly mean lengths and depths and their standard errors are listed. The growth curves (Text-fig. 3), prepared from the mean lengths and depths plotted against time, have similar slopes, except in the last few months of the experiment. The fluctuations seen at this time were caused by a reduced number of fish as they were killed for histological study. Comparisons of the monthly size ranges of the fish, represented by the vertical lines in Text- fig. 3, indicate that the most extreme size dif- ferences were shown by the fish under the red and blue filters. The greatest percentage of growth in mean length and depth at the end of five months oc- curred in the group under the yellow filter, and in the controls, whereas under the red filter this value was smallest. From the percentage growth at monthly intervals it was found that growth during the first month under the red filter was small, but by the end of the second month in- creased to the greatest value found in any of the other groups (Table 2). This probably reflects a slower adjustment to the red filter. Statistical examination of the data in this table indicates that these growth differences were not statisti- cally significant. Thyroid.— In the thyroid glands of Astyanax under one year old, maintained under normal laboratory conditions, the follicular epithelium varies from squamous to low cuboidal, and oc- casionally high cuboidal. The location of the follicles, which are diffusely scattered as in most other teleosts, and their normal histological structure, were described by Rasquin (1949). No extreme deviations in thyroid histology were produced by the colored filters. Monthly variations in the height (Table 3) and activity of the follicular epithelial cells and differences in the amount and staining reaction of the col- loid were seen in all of the groups. The thyroid hyperplasia observed by Rasquin (1949) in Astyanax maintained in complete darkness was not produced in these fish. Small differences in thyroid activity revealed by careful analysis of the data are of sufficient interest to be further examined. It can be seen from the graphic representation of the mean 1952] Calm: Spectral Effects on the Teleost, Astyanax mexicanus 37 Table 2. Percentage Increase in Mean Length and Depth at Monthly Intervals and After a Five-Month Interval Months Number Percentage Increase Under Colored Filters of Fish Control Red Yellow Green Blue Mean Length 1 6 41.2 27.1 40.0 35.6 43.5 2 6 24.8 42.9 38.0 23.0* 29.7 3 6 15.7 10.5 5.3 23.7 5.1 4 5 7.8 13.5 8.7 10.4 9.4 5 4 10.5 6.0 8.0 7.4 12.3 1-5 — 73.9 64.3 78.9 65.9 67.5 Mean Depth 1 6 30.8 24.1 38.9 30.3 37.8 2 6 25.6 55.2 46.2 36.4* 8.1 3 6 7.7 deer. deer. 00.0 13.5 4 5 15.4 deer. deer. 9.1 2.7 5 4 20.5 20.7 17.9 24.2 37.8 1-5 — 47.0 31.8 46.6 41.3 44.6 * Average of five fish. thyroid epithelial cell heights (Text-fig. 4) that the control group showed the smallest variations in height, whereas all of the colored filters pro- duced more atypical curves when compared with the controls, especially the blue and yellow filters. The values for the difference between the extremes of the mean thyroid epithelial cell heights were respectively: blue, 1.01; yellow, .70; red, .59; green, .56; control, .29 (all in microns) . Over 50 percent, of these variations in the mean thyroid cell height associated with the colored filters were found to be statistically sig- nificant, as seen in the lower part of Table 3. The mean cell heights were significantly de- creased by the yellow filter, increased by the green and red filters, and both decreased and increased, in different fishes, by the blue filter. Other changes in thyroid histology accom- panied the variations in cell height. When cell height was high, the follicles were small, the epithelial cells contained vacuolated cytoplasm, and the stored colloid which had decreased in amount and changed in staining reaction, con- tained chromophobic vacuoles. When cell height was low, the follicles were enlarged and were filled with deeply staining dense colloid. Pituitary.— The examination of serial sections revealed no changes in the histology of this gland that could be referred to the colored filters. The relative proportions and the size, distribution, cytoplasmic granulation and staining reactions of the acidophiles, basophiles and chromo- phobes of the transitional lobe— normally asso- ciated with cyclic changes in secretory activity —corresponded to the description given by Ras- quin (1949) for the normal six- to twelve- month-old Astyanax pituitary. Vacuolated baso- philes, normally found in this lobe, were more numerous in the five- to eight-month-old fish grown under the blue, green and yellow filters, than in the controls of the same age, whereas in all of the nine-month-old fish many vacuolated basophiles were present. Ovary.— Fully mature yolk-filled ova were found in nine of the thirteen ovaries examined. Under each colored filter, however, there was one fish with only a few mature eggs. The data were insufficient to evaluate this finding because under similar normal conditions it is known that some fish mature earlier and develop faster than others. Some resorbing eggs, probably caused by a failure to ovulate, were found in all ovaries. This resorption process was best ob- served in the extremely large, fully mature ovaries from the seven-month-old control fish, and the six-month-old fish under the yellow fil- ter. In the middle and posterior regions of these ovaries, all of the mature eggs appeared to be breaking down. They had become confluent into one large mass which contained eosinophilic yolk, fat-like vacuoles, reticular-like tissue, cho- 38 Zoologica: New York Zoological Society [37: 3 Table 3. Individual and Group Mean Thyroid Epithelial Cell Heights in Microns of Fishes under Colored Filters Filter Age in Months Sex Size in mm. Length by Depth Mean Cell Height Standard Error Group Mean Cell Height Standard Error Control 5 M 34x11 2.585 .0368 6 F 34x11 2.725 .0356 7 F 38x13 2.510 .0399 2.55 .0444 8 F 40x12 2.500 .0400 9 F 32x11 2.435 .0358 Red 5 F 32x10 2.600 .0339 6 M 34x11 3.010 .0464 7 M 35x10 2.663 .0429 2.65 .2761 8 F 40x13 2.420 .0417 9 M 34x9 2.575 .0378 Yellow 5 F 35x12 2.670 .0443 6 F 33x11 2.625 .0335 7 F 35x12 2.300 .0367 2.36 .3623 8 M 35x10 1.970 .0468 9 M 32x9 2.280 .0285 Green 5 M 31x9 2.800 .0452 6 F 31x11 2.935 .0375 7 F 36x12 2.380 .0395 2.62 .3052 8 M 34x9 2.425 .0389 9 M 36x11 2.545 .0368 Blue 5 M 30x10 3.255 .0631 6 F 34x11 2.655 .0379 7 M 39x15 2.250 .0391 2.70 .1536 8 M 35x11 2.470 .0528 9 F 35x11 2.855 .0334 Comparisons made for statistical significance of individual mean thyroid cell heights Fishes Compared Significance 5 mos. control with 5 mos. green 3.6 5 mos. control with 5 mos. blue 9.2 6 mos. control with 6 mos. red 4.9 6 mos. control with 6 mos. green 4.1 7 mos. control with 7 mos. red 2.5 prob. sig. 7 mos. control with 7 mos. yellow 3.8 7 mos. control with 7 mos. blue 4.6 8 mos. control with 8 mos. yellow 8.6 9 mos. control with 9 mos. red 2.7 9 mos. control with 9 mos. yellow 3.5 9 mos. control with 9 mos. blue 8.5 rionic membrane remains and hyperplastic fol- licular epithelium. In three of the four fish with ovaries that con- tained few mature eggs, many ova were resorb- ing prematurely. It is of interest that these fish were seven to nine months old (from the groups under the blue, green and yellow filters). In the five-month-old fish from the group under the red filter that was not yet fully mature, resorb- ing eggs were not found. Testis.— The testis is organized into lobules which contain cysts of spermatogenic cells. As the cells mature they are shed into the lobular cavities where they are stored prior to passage into the sperm duct. Rasquin & Hafter (1951) described the normal structure of the testicular 1952] Calm: Spectral Effects on the Teleost, Astyanax mexicanus 39 Text-fig. 4. Mean thyroid epithelial cell heights, from three to seven months after the start of the experiment, for all of the fish. Numbers 1, 2, 3, 4 and 5 represent respectively the controls and the groups under the green, red, yellow and blue filters. lobules and Rasquin (1951) demonstrated the presence of osmicated granules in the sperm duct epithelium which are indicative of a prob- able secretory function. Only one of the twelve males examined was from the control group. This made difficult the determination of specific effects of the filtered light. It was observed, however, from compari- sons of serial sections through corresponding regions, that all of the testes contained mature sperm in the lumina of the lobules and in the sperm duct. In five of the seven- to nine-month-old fish from the experimental groups (two from yellow, one each from the red, green and blue groups) the testes contained very little spermatogenic tissue. This was also observed in some of a series of control males of approximately the same age, examined for another experiment, which indi- cates that this lack was probably not caused by the filtered light. Discussion The reduction of luminous intensity and ra- diant energy by the red, green and blue filters, compared with the controls, could account for the decreased total growth and increased thyroid activity observed in these groups. Similar but vastly greater changes were found when Asty- anax were maintained in total darkness (Ras- quin, 1949). Yung (1881) reported a spectral effect on the growth rate of salmon and tadpoles, but he used filters that differed in the transmis- sion of radiant energy. Rugh (1935) analyzed the properties of the filters used by Yung and found that the transmission of the most radiant energy resulted in the fastest growth. Rugh also found, in his own experiments, that when ra- diant energy was equalized, differences in wave length and luminous intensity produced no dif- ferential effects on the growth rate of tadpoles. This relationship between radiant energy and growth rate was also found in these experiments on Astyanax. The red filter transmitted more light than the blue and green filters, but about the same amount of radiant energy. The per- centage increase in length was approximately the same under all three filters. The absence of most of the shorter wave lengths under the red filter may also have contributed to the small length increase. The presence of both the shorter and longer wave lengths of visible light were found to be essential for the normal growth of young chicks (Sheard & Higgins, 1928; Sheard, Higgins & Foster, 1930). Constant housing of these chicks behind an amber glass filter that transmitted only the longer visible wave lengths of sunlight retarded growth and development. 40 Zoologica: New York Zoological Society [37: 3 Crawford (1930) noted that blue light re- sulted in the slowest rate of growth of young salmon, and also the heaviest mortality; red light produced growth effects similar to those pro- duced by total darkness; and green and yellow light were optimal for growth. He did not pub- lish any data, however, on the properties of the filters he used. Red and blue light were found to accelerate the growth of young rats (Ludwig & von Ries, 1931), but this was not confirmed by Allardyce, et al. (1942), who found no con- sistent differences in growth rate following main- tenance under colored, colorless and black filters. The variability in the growth of young fishes often encountered under normal conditions may be partially responsible for the small differences observed in the groups under the colored filters. Consideration should also be given to the dif- ference in sex distribution in the different groups, since the fish were sexually immature at the start of the experiment. In Astyanax, females usually attain a larger size than males, so that the predominance of females in the control and yellow groups may therefore, in part, explain the greater total growth that oc- curred in these groups. Modlinger (1941) found that red light and darkness increased the epithelial cell height and decreased the iodine content of the colloid of the dove thyroid, and these changes coincided with those that took place during the annual cycle. Pighini (1941) also reported that mono- chromatic red, yellow, green and blue light all produced increased thyroid activity in guinea pigs. The effects were not wave-length specific, and may have been caused by the decreased light and radiant energy transmitted by the col- ored lights. In Astyanax, the yellow filter sig- nificantly decreased thyroid cell height, and this appeared to be a wave-length effect, since the light and radiant energy transmitted did not differ substantially from the control. The blue filter significantly increased thyroid cell height in two fish, but significantly decreased cell height in another fish. The increased cell height was explained by a decreased transmission of light and radiant energy, but it is impossible to explain the opposite effect without additional data. It is possible that these results, although statistically significant, are not biologically significant be- cause of the nature of the material. Other studies also indicated that decreased light or darkness results in increased thyroid activity (Bergfeld, 1931, in rats; Puntriano & Meites, 1951, in mice; and Rasquin, 1949, in the teleost, Ameiurus nebulosus) , although Ma- yerson & Branch, 1934; Mayerson, 1935; and Kenyon, 1935, in rats; and Stein & Carpenter, 1943, in the salamander, Triturus viridescens, did not confirm this response. That all of the colored filters altered the thy- roid activity of Astyanax indicates the role of this gland in the general response to a changed environment. An altered metabolic rate may have been involved in this response, although it is not definitely known that the teleost thyroid gland is a regulator of metabolism (Etkin, Root & Mofshin, 1940, and Matthews & Smith, 1947). Some indication of the thyroid’s meta- bolic function is seen by the reports that total darkness increased both the oxygen consump- tion (Schlagel & Breder, 1947) and the thyroid activity (Rasquin, 1949) of Astyanax. Although no prominent changes in the pitui- tary of Astyanax were produced by the colored filters, Pighini (1941) found that red, yellow, green and blue light all produced the early ap- pearance of acidophiles in the anterior lobes of young guinea pigs. This was attributed more to the lower luminous intensity and radiant energy transmitted by the colored lights than to specific wave-length effects. In addition, darkness re- duced the relative number and activity of pitui- tary basophiles in Astyanax (Rasquin, 1949) and in frogs (Florentin & Stutinski, 1936; Stu- tinski, 1936; and Woitkewitsch, 1944). The many resorbing ova found in all of the ovaries of Astyanax did not appear to be effects of specific wave lengths. The breakdown of ma- ture ova, in the absence of spawning, and their subsequent resorption, has been described as a normal occurrence in other teleost ovaries (Brock, 1878; Barfurth, 1886; Cunningham, 1897; and Wallace, 1904). These degenerating ova undergo fatty changes, accompanied by the immigration of much cell debris into the sub- stance of the egg. It is still uncertain whether this cell debris comes from a growth and pro- liferation of the follicular epithelium (as in trout ova, Barfurth; and in Zoarces viviparus, Wallace), from a proliferation of the cells from the connective tissue in the wall of the follicle (as in Trigla hirunda, Cunningham), or from both of these locations (as in the bitterling, Rhodeus amarus, Bretschneider & de Wit, 1941). It has been demonstrated by the present work that despite all of the spectral, light intensity and radiant energy reductions provided by the colored filters, the fish were able to maintain the essential integrity of their endocrine systems. Evidently, for this stability, the presence of some visible radiation as such is of greater significance than the exact spectral quality of that radiation, for which there are many corroborating studies 1952] Calm: Spectral Effects on the Teleost, Astyanax mexicanus 41 in the literature. There is need for further ex- periments which will utilize filters of more re- fined properties and which will operate at in- tensities between those used in the present study and the zero intensity used by Rasquin (1949). The minimal light intensity that will support normal endocrine structure and function, as well as the possible importance of specific wave lengths at such a lowered intensity of light, are problems yet to be studied. Summary and Conclusions 1. Two-month-old Astyanax mexicanus were grown under daylight filtered by red, yellow, green and blue cellophane sheets, for three to eight months, after which the histology of the thyroid, pituitary and gonads was compared with that of controls maintained without any filter. The cellophane sheets were analyzed for the transmission of the spectrum and for lumi- nous intensity. 2. At the end of five months, the fish under the yellow filter showed about the same percent- age of increase of growth in body length and depth as the controls. A smaller percentage of increase was attained under the red, green and blue filters, because of the decreased light trans- mitted, but these lower figures were not signifi- cantly different from the controls. 3. Thyroid activity in some of the fish under the red, green and blue filters was significantly increased over the controls, and was probably' produced by the decreased transmission of these filters. Some spectral influence of the yellow filter was presumed to be responsible for the significantly decreased thyroid activity observed in some of these fish. Additional data are needed to explain the significantly decreased thyroid cell height in one fish grown under the blue filter. The biological significance, if any, of these alterations in thyroid activity is yet to be as- certained. 4. The structure of the transitional lobe of the pituitary and the testes showed no significant changes induced by the colored filters. 5. Some ovaries that were not fully mature were found in fish raised under the colored fil- ters, with prematurely resorbing ova, especially under the yellow, green and blue filters. The significance of this item is unknown. 6. It was demonstrated that despite all of the spectral, light intensity and radiant energy re- ductions provided by the colored filters, the fish were able to maintain the essential integrity of their endocrine systems. 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Beitrage zur anatomie und histologie der geschlectsorgane der Knochenfische. Morph. Jahrb., 4: 505-569. Crawford, D. R. 1930. Some considerations in the study of the effects of heat and light on fishes. Copeia, 173: 89-92. Cunningham, J. T. 1898. On the histology of the ovary and of the ovarian ova in certain marine fishes. Quart. Jour. Micr. Science, 40: 101-164. Etkin, W., R. W. Root & B. P. Mofshin 1940. The effect of thyroid feeding on the oxygen consumption of the goldfish. Physiol. Zool., 13: 415-429. Florentin, P. & F. Stutinski 1936. Modifications cytologiques de la gland pituitaire des grenouilles maintenues a l’obscurite. Compt. Rend. Soc. Biol., Paris, 122: 674-676. Judd, D. B. 1933. The 1931 I. C. I. Standard Observer and coordinate system for colorimetry. Jour, of Optical Soc. America, 23: 359-374. 42 Zoologica: New York Zoological Society [37: 3: 1952] Kawamoto, N. & M. Takeda 1950. Studies on the phototaxis of fish. I. 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Modificazioni di Chiandola endocrine in animali allevati a luci monochromatiche. Rivista di Biologie, 31: 186-208. PUNTRIANO, G. & J. MEITES 1951. The effects of continuous light or dark- ness on thyroid function in mice. Endo- crin., 48: 217-224. Rasquin, P. 1949. The influence of light and darkness on thyroid and pituitary activity of the chara- cin Astyanax mexicanus and its cave de- rivatives. Bull. Amer. Mus. Nat. Hist., 93: 501-531. 1951. Effects of carp pituitary and mammalian ACTH on the endocrine and lymphoid systems of the teleost Astyanax mexicanus. Jour. Exp. Zook, 117: 317-358. Rasquin, P. & E. Hafter 1951. Age changes in the testis of the teleost Astyanax mexicanus. Jour. Morph., 89: 397-407. Rawson, R. W. & P. Starr 1938. Direct measurement of the height of thyroid epithelium. Arch. Intern. Med., 38: 726-738. Rugh, R. 1935. The spectral effect on the growth rate of tadpoles. Physiol. Zool., 8: 186-195. Schlagel, S. R. & C. M. Breder, Jr. 1947. A study of the oxygen consumption of blind and eyed cave characins in light and darkness. Zoologica, 32: 17-27. Sheard, C. & G. M. Higgins 1928. The effects of selective solar irradiation on the growth and development of chicks. Amer. Jour. Physiol., 85: 290-298. Sheard, C., G. Higgins & W. I. Foster 1930. The growth and development of chicks as influenced by solar irradiation of long visible and ultraviolet wave lengths, re- spectively with and without supplementary irradiation of various types. Amer. Jour. Physiol., 94: 84-90. Stein, K. & E. Carpenter 1943. The effect of increased and decreased light on the thyroid gland of Triturus viridescens. Jour. Morph., 72: 491-513. Stutinski, F. 1936. Effets de L’Eclairment Continu sur la Structure de la Gland Pituitaire de la Grenouille. Compt. Rend. Soc. Biol., Paris, 123: 421-423. Wallace, W. 1904. Observations on the ovarian ova and follicles in certain teleostean and elasmo- branch fishes. Quart. Jour. Micr. Sci., 47: 161-214. Warner, L. H. 1931. The problem of color vision in fishes. Quart. Rev. Biol., 6: 329-348. Walls, G. 1942. The vertebrate eye and its adaptive radia- tion. Cranbrook Institute of Science, Bull, no. 19: 462-490. W OITKEWITSCH, A. A. 1944. Influence of light upon thyrotropic activity of anterior lobe of pituitary body. Compt. Rend. Acad. Sci. U. R. S. S., 45: 357-360. Yung, E. 1881. De l’lnfluence des Lumieres Colorees sur le Developpement des Animaux. Mitt. Aus. Der. Zool. Stat. zu Neapel., 2: 233. 4 Fungus Associated with a Granuloma in a Turkish Fish, Aphanius chantrei Gaillard Recai Ermin University of Istanbul and the New York Zoological Society 1 (Plates I & II; Text-figures 1-11) Introduction RELATIVELY few fungus infections that produce tumors in fishes have been stud- ied in detail. Montpellier & Dieugeide (1933) briefly described a “pseudo-tumeur my- celienne” in a North African Cyprinodon fas- ciatus, and recently Walker (1951) reported a case in Salmo sp. The present paper describes a granuloma caused by an undetermined fungus in Aphanius chantrei, a small, freshwater cy- prinodont fish indigenous to Turkey. Since the blood cells play an important role in the development of granulomas, it is fortunate that the papers by Duthie (1939) and Catton (1951) on fish blood elements are now avail- able. This makes possible a more complete analysis of the piscine granuloma than has pre- viously been presented. The present study may have special signifi- cance in view of the recent investigations of Diller & Fisher (1950), who demonstrated the presence of inter- and intracellular fungal struc- ture including mycelia (Fungi Imperfecti related to Blastomycetes) from transplanted, induced and spontaneous tumors of the mouse and of man. Materials and Methods A number of Aphanius chantrei were col- lected near Samsun, a town in northern Anatolia on the Black Sea. They were kept for about a year in the aquarium of the Zoological Institute of the University of Istanbul at Baltalimani on 1 From the Zoological Institute of the University of Istanbul, Turkey, and from the Genetics Laboratory of the Aquarium, New York Zoological Society, at the American Museum of Natural History, New York 24, N. Y. Supported in part by a grant from the Anna Fuller Fund and facilities made possible by a grant to the New York Zoological Society from the National Cancer Institute, U. S. Public Health Service. the Bosporus. One of these specimens, a female 45 mm. in standard length, developed a tumor on the dorsal surface over the operculum and pectoral fins (PI. I, Fig. 1). The swelling did not interfere with the normal behavior of the fish. The fish was fixed in a mixture of 76 parts of saturated aqueous mercuric chloride, 20 parts of formalin and 4 parts of glacial acetic acid. The anterior half of the animal was decalcified in nitric acid, dehydrated in alcohol, cleared in xylol, embedded in paraffin and cross-sectioned at 10 p. Sections were stained either with May- er’s hematoxylin (hemalum) and eosin, Fleiden- hain’s iron hematoxylin and eosin, Mallory’s azan stain or van Gieson’s collagen stain, or by Gram’s method. Gross Description of the Tumor The swelling was smooth, soft, and projected dorsad from the right side of the body (PI. I, Figs. 1-3; Text-figs. 1, 2). It measured 7 mm. in length, 5.5 mm. in depth and 4 mm. in height. The entire mass was covered by a layer of skin and was the same color as the body of the fish. The scales over the swelling were separated and no longer imbricated. Scales were missing on the right dorsal surface and in some other areas (Text-figs. 1, 2). Examination of the sections showed that the growth was located in the dorsal musculature extending from the region just back of the head (PI. I, Figs. 1,2) to that near the gonads. In some regions the skin in contact with the tumor was hyperplastic (PI. I, Fig. 3; Text-fig. 1). In other regions the skin was normal and there was a space between the boundary of the tumor and the dermis. At the most anterior portion of the growth there was no external swelling but the skin was hyperplastic and the scale pockets were larger 43 44 Zoologica: New York Zoological Society [37: 4 Text-fig. 1. A cross-section of the granuloma over the gill region. The new growth arises cone-like above the dorsal surface of the body. Note the hyperplastic epi- dermis on the right and the nodules in the granuloma. Ep =: epidermis, Gr = granuloma, I.t. = infiltrating tissue, M = musculature, Me — melanophore, Sc = scale. Hemalum-eosin. than those over the unaffected regions of the body. Here the growth, which contained some melanophores, had infiltrated the muscles be- tween the skin and the skull. A little further caudad the growth enlarged and extended above the body surface. The center of the growth contained nodule-like formations. On the right side of the mass, the epidermis and dermis of the skin were hyperplastic and ex- tended into the growth. The scales and the scale pockets were wanting (Text-fig. 1). The left side was normal. Still further caudad, in the gill region, the growth was larger and rounder. The overlying skin was thin and contained scales. The growth had infiltrated the dorsal trunk musculature of the right side of the body. Here the growth was composed primarily of whorl-like nodules which were surrounded by a fibrillar, capsule-like tissue. The largest part of the growth developed in the region above the liver (Text-fig. 2). The entire mass was round and was composed of two large and four small foci, each of which was encapsulated. The centers of some of the foci were necrotic. At this level the infiltration of the muscles was seen only on the right side, but the pressure of the growth had forced the central muscle mass to the left. Posterior to this region, the tumor as a whole was smaller. Microscopic examination of the region just beyond the ex- ternally visible tumor revealed granulation tissue cells. Histological and Cytological Observations The granuloma is composed of a reticular connective tissue stroma and of proliferated lymph cells, leucocytes, mast cells, giant cells, fibroblasts and capillaries. No bacteria were found in sections stained by Gram’s method. The mycelia of an undetermined species of fungus are surrounded by lymphocytes, leuco- cytes and fibroblasts (Text-figs. 3-5). Most of the hyphae show septa and branches (Text-fig. 3; PI. II, Fig. 4). The hyphae may be intra- cellular or intercellular. They lie either in granu- lation tissue as individual filaments or in an irregular network of mycelia (Text-figs. 3, 4). In hemalum-eosin-stained sections they were yellowish-pink in color; stained by Mallory’s they are blue. All hyphae are Gram negative. They show a hyalin structure and homogeneous plasma. The width of the hyphae varies from 1.2 to 2.5 [x while their length varies approxi- mately from 15 to 110 ju,. 1952] Ermin: Fungus and Granuloma in Aphanius chantrei 45 Text-fig. 2. A cross-section in region of the liver show- ing different foci and necrotic area (N.a.) in the granu- loma. Ep — epidermis, Gr = granuloma, M = muscu- lature, Me = melanophore, Sc = scale. Hemalum-eosin. The fibers of the connective tissue stroma, in the center of the growth, are reticular and run approximately parallel around the whole granuloma (Text-figs. 6, 7). The mesenchyme cells or reticulum cells are 9 to 13 p, spindle- shaped, and have elongated nuclei (3.5 to 13 p) which contain granular chromatin. These cells are attached to the fibers. Lymphocytes of different shapes and sizes, many of which show mitosis, are scattered be- tween the reticular network. The majority of them are round, except those around the nodules which are elongate. These lymphocytes have a relatively thin plasma mass around their nuclei, which contain either granular chromatin or a chromatin network. They measure 6 to 9 p. Their nuclei are 3.5 to 5.5 p. These cells are characteristically found around mycelia (Text- figs. 6-8). More finely granulated lymphocytes, called “fine granulocytes” by Duthie (1939) and Cat- ton (1951), are found in the reticulum of the granuloma. These cells possess fine eosinophilic granules and measure 4.5 to 8.5 p. Their nuclei, which measure 3 to 4.5 p, contain a granular network in which the nucleoli can be seen in some instances. These fine granulocytes may be 46 Zoologica: New York Zoological Society [37: 4 Text-fig. 4. Part of the granuloma showing a giant cell (G.c.) with dividing nuclei (D.n.). One of two small nodules (N) contains a hypha (Hy), seen in cross-section, and the other contains a degenerating cell. Hemalum-eosin. Text-fig. 5. Part of the granuloma showing three different giant cells (G.c.), each of which contains a hypha (Hy). Hemalum-eosin. 1952] Ermin: Fungus and Granuloma in Aphanius chantrei 47 10/i. Text-fig. 6. Reticular stroma of the granuloma showing reticular fibers (R.f.), reticulum cells (It.c.), lympho- cytes (Ly), fine granulocytes (F.g.) and small lympho- cytes (S.I.). found either in capillaries or in the reticulum (Text-figs. 6-8). The fine granulocytes appar- ently pass through the blood vessels as shown in Text-fig. 8. Occasionally they may assume an elongated form, forming oval nuclei. Some very small, round lymphocytes, having a diameter of 2.5 to 3.5 p, are also seen, but rarely. These “small lymphocytes” have small amounts of cytoplasm around their nuclei. The nuclei, which are 1.5 to 2.5 p in size, contain fine and extensive granular chromatin (Text-fig. 6, S.I.). These cells are similar to those called “lymphoid hemoblasts” by Duthie (1939) and “small lymphoid hemoblasts” by Catton (1951). Another type of granuloma cell is represented by coarse granulocytes or mast cells. These are found mostly in the fibrillar, infiltrating, capsule- like zone around the granuloma (Text-fig. 9) and in the hyperplastic epidermis between the basal cells (Text-fig. 10). These mast cells or coarse granulocytes (10 to 20 p) are spindle- shaped, round or oval and contain large eosino- philic granules. The nuclei are relatively small (2.5 to 3.5 /(.)• They are mostly eccentric and contain granular chromatin. These cells are found in the tissue of the new growth, but not in the blood vessels, nor are they found in the blood of a normal Aphanius. The fibroblasts, which are elongated cells with oval or spindle-shaped nuclei, are scattered in the granuloma. They measure 13 to 23 p and their nuclei, which very often contain a visible nucleolus, are 9.5 to 14.5 p in size (Text-figs. 7-9). The fibroblasts take part in the infiltration of muscle and the formation of nodules in the granuloma. Macrophages are found in necrotic areas. They are smaller than fibroblasts, similar to the lymphocytes and difficult to distinguish from the latter. Giant cells are distributed in the granulation tissue, particularly where the mycclia cf fungus are scattered (Text-figs. 4, 5). They are poly- morphic and contain round or oval nuclei which have nucleoli and loose chromatin nets. The nuclei are very often at the periphery of the cells and often form “pearls.” The number of nuclei varies, many cells containing 10 to 27. The cells measure 19 to 40 p and their nuclei, 3.2 to 8 p. Text-fig. 5 shows three different types of giant cells embedded in granulation tissue, each of which has phagocytized one hypha. Whether giant cells are derived from fibro- blasts or lymphocytes or endothelial cells has not been decided. Nuclear division was found in one of the giant cells. As shown in Text-fig. 4, the nuclear divisions are most peculiar. They probably represent a kind of amitotic division which may produce the pearl-like distribution of nuclei. Plehn (1906) found a similar type of nuclear division in a sarcoma in the trunk muscles of Phoxinus laevis. According to her, after such division multinuclear giant cells may appear. Some of the giant cells contain pyknotic nuclei. The cells including lymphocytes and fibro- blasts, which accumulate around the mycelia, form the nodules of granulation tissue (PI. II, Figs. 5-6). In the formation of nodules lympho- cytes are especially active (Text-figs. 7, 8). The nodules are round, vary in size, and are encap- sulated by a reticular stroma, as shown in Text- fig. 7. Some necrotic areas are found in the center of the granuloma. These areas contain cell debris, hyphae and macrophages (PI. II, Fig. 5). Capillaries are found either in or around the granuloma (Text.-fig. 8). They are hyperemic and their endothelial cells are sometimes hyper- trophic. Some erythrocytes lie free in the granu- lation tissue. The epidermis over the granuloma is partly normal and partly hyperplastic. The normal epidermis of Aphanius is 20 to 30 p thick and is composed of about six layers of squamous epi- thelium cells, oval or round mucous cells and basal cells, and some melanophores in the basal part of the epidermis. The hyperplastic epi- dermis over the granuloma is approximately 130 p thick and has 20 to 25 cell layers (Text- fig. 10). Besides the squamous epithelial cells, mucous cells and basal cells, some coarse granu- locytes are found in the proximal part of the hyperplastic epidermis. Since the coarse granu- locytes in normal fish are only in connective tissue, they probably come from granulation 48 Zoologica : New York Zoological Society [37: 4 I OJX Text-fig. 7. A nodule (N) in the granuloma, surrounded by fibrillar tissue. D.ly = dividing lymphocyte, F.g. = fine granulocyte, Fb = fibroblast, Hy == hypha, Ly == lymphocyte, R.c. = reticulum cell. Hemalum-eosin. tO/L Text-fig. 8. Part of the granulation tissue containing blood capillaries and dividing lymphocytes (D.ly). En.c. = endothelium cell, F.g. = fine granulocyte, Fb = fibroblasts, Ly = lymphocytes. Hemalum-eosin. 1952] Ermin: Fungus and Granuloma in Aphanius chantrei 49 S* I. Text-fig. 9. Part of the fibrillar tissue which surrounds the granuloma. C.g. = coarse granulocyte (mast cell), F.g. — fine granulocyte, Fb fibroblast, Ly = lymphocyte, R.c. = reticulum cell, S.l. — small lymphocyte. Hemalum-eosin. tissue which in some regions is adjacent to the epidermis. The part lying between the squamous and basal cells of the hyperplastic epidermis is looser than in the normal epidermis. In some regions, the hyperplastic epidermis resembles granulation tissue in character and contains lymphocytes, fine granulocytes and hyphae of fungus besides epidermis cells. As the granu- loma grew upwards, the overlying epidermis apparently became stretched. The epidermis thus grew parallel to the new growth. The scales are disarranged in some regions because of the epidermis growing under them, elevating them away from the dermis. The fibrillar, capsule-like tissue which sur- rounds the granuloma infiltrates the subjacent musculature (Text-fig. 11; PI. II, Fig. 7). At its deepest penetration, the invading granuloma cells appear fibrosarcoma-like and malignant. Discussion In the granuloma described above, two phenomena must be distinguished. One is the cellular reaction of the body to the fungus in- fection. This results in the formation of granu- lation tissue in the infected area. Some of the cellular elements of granulation tissue, such as lymph cells, leucocytes and fibroblasts, form a nodule around either the mycelia or a single hyphae. The second phenomenon is the forma- tion of a fibrosarcoma-like growth. The granu- lation tissue appears to lose its protective func- tion and becomes a malignant growth. This tumor in Aphanius may be regarded as an “in- fective granuloma,” as defined by Hamperl (1944) and Ogilvie (1947) in human pathology. Human granulomas are essentially similar in cellular elements to those of fishes (Ash & Spitz, 1945; Ewing, 1940). 50 Zoologica: New York Zoological Society [37: 4 Text-fig. 10. Hyperplastic epidermis overlying the granuloma. Ba.c. = basal cells, C.g. = coarse granu- locyte, De.c. = dividing epithelial cell, Mu.c. = mucous cell, S.e. = squamous epithelium. Hemalum-eosin. The species of infectious fungus could not be identified. As Ash & Spitz (1945) have point- ed out, “It is usually not possible to identify a fungus in a histologic section. . . . The spores and mycelia of different species are too much alike to permit classification solely on morphol- ogy.” Since the hyphae found in Aphanius con- tain septa, it can be said that this fungus does not belong to the Saprolegniaceae (Plehn, 1912). The following reactions to the fungus were seen in Aphanius : a hyperplasia of lymph cells and leucocytes and the formation of giant cells. Similar general responses were found by Mont- pellier & Dieugeide (1933) in Cyprinodon, and by Walker (1951) in Salmo. Some specific inflammatory responses and the formation of granulation tissue in fishes are mentioned by Plehn & Mulsow (1911), Taka- hashi (1929), Thomas (1931), Plehn (1938), Drew (1910-13), Wurmbach (1951), Breider & Schmidt (1951), etc., but the authors give few, if any cytological and histological details. Plehn (1910) and Schaperclaus (1941) report that in a disease called “Drehkrankheit,” a spo- rozoon, Lentospora cerebralis, causes a granu- loma in the auditory organs of trout, destroying this organ. With regard to the origin of coarse granulo- cytes or mast cells, which are seen only rarely in the blood of fishes, several authors indicate that they come from the connective tissue. Michels (1923), studying a whitefish ( Leuciscus sp.) and the carp ( Cyprinus carpio), suggested this origin. Duthie (1939) and Catton (1951) have frequently observed these cells in connective tis- sue and rarely in the blood of a variety of teleost fishes. From my observations, the views of Michels seem valid. Summary 1. The histology and cytology of a granuloma associated with an undetermined fungus infec- tion are described in a Turkish freshwater cy- prinodont, Aphanius chantrei. 2. The granuloma mass, covered by a layer of skin, is found on the dorsal surface between the posterior margin of the operculum and the region of the pectoral fins. 3. The granuloma is composed of reticular connective tissue stroma and of proliferated lymph cells, leucocytes, mast calls, phagocytic giant cells, fibroblasts and macrophages. This mass is highly vascularized. 4. The mycelia of a fungus are scattered in the granulation tissue and are surrounded by lymph- ocytes and fibroblasts. These form definitive nodules in the granuloma. 5. The overlying epidermis, in some regions, is hyperplastic and shows partly the charac- teristics of granulation tissue. 1952] Ennin: Fungus and Granuloma in Aphanius chantrei 51 Text-fig. 11. Muscle infiltrating and fibrosarcoma-like tissue (I.t.). In.m. = infiltrated musculature, M = musculature. Hemalum-eosin. 6. The granuloma mass is surrounded by a fibrillar tissue which infiltrates the subjacent musculature. This infiltrating tissue is fibrosar- coma-like and appears to be malignant. Acknowledgments It is a pleasure to make the following acknowl- edgements: to Dr. Sadao Otani, Mt. Sinai Hos- pital, New York, and to Drs. Uveys Maskar and Kemal Akgiider of the Faculty of Medicine of the University of Istanbul, for their aid in diagnosis; to Dr. Uveys Maskar and Cafer Turkmen for the photomicrographs; to Drs. Ross F. Nigrelli, Leonard J. Goss, Myron Gor- don, Olga Aronowitz and Messrs. James W. Atz and Donn E. Rosen for aid in preparing the manuscript; to the New York Zoological Society and the American Museum of Natural History for laboratory facilities. Bibliography Aschoff, L. 1936. Pathologische Anatomie. 8th Ed. Gustav Fischer, Jena. 52 Zoologica: New York Zoological Society [37: 4 Ash, J. E., & Spitz, S. P. 1945. Pathology of tropical diseases. 1st Ed., W. B. Saunders Co., Philadelphia, Penna., and London, England. Breider, H., & Schmidt, E. 1951. Melanosarkome durch Artkreuzung und Spontantumoren bei Fischen. Strahlen- therapie, 84: 498-523: Brumpt, E. 1949. Precis de Parasitologie. 6th Ed., Masson, Paris, France. Catton, W. T. 1951. Blood cell formation in certain teleost fishes. Blood, Jour. Hematol., 6: 39-60. Diller, I. C., & Fisher, M. 1950. Isolation of fungi from transplanted, chemically induced and spontaneous tumors. I. General considerations. Cancer Research, 10: 595-603. Drew, G. H. 1910-13. Some cases of new growth in fish. Jour. Marine Biol. Assoc., 9: 281-287. Duthie, E. S. 1939. The origin, development and function of the blood cells in certain marine teleosts. I. Morphology. Jour. Anat., 73: 396-411. Ewing, J. 1940. Neoplastic diseases. 4th Ed., W. B. Saunders Co., Philadelphia, Penna., and London, England. Hamperl, H. 1950. Lehrbuch der allgemeinen Pathologie und der pathologischen Anatomie. 18th Ed., Springer Verlag, Heidelberg. Hofer, B. 1906. Handbuch der Fischkranheiten. 1st Ed., E. Schwizerbart’sche Verlagsbuchhand- lung. Stuttgart. Kahls, O. 1930. tjber das Vorkommen von Algen und Pilzen bei Fischen. Z. f. Fischerei, 28: 253-262. Mac Callum, W. G. 1940. A textbook of pathology. 7th Ed., W. B. Saunders Co., Philadelphia, Penna., and London, England. Michels, N. A. 1923. The mast cell in the lower vertebrates. Cellule, 33: 339-462. Montpellier, J., & Dieugeide, R. 1933. Pseudo-tumeur mycelienne chez un pois- son ( Cyprinodon fasciatus Val.). Bull, des Trav. publies par la Stat. d’Agric. et de Peche de Castiglione (Algere), 1. fasc. Ogilvie, R. F. 1947. Pathological histology. 3d Ed., E. and S. Livingstone, Ltd., Edinburgh, Scotland. Plehn, M. ’ 1906. fiber Geschwiilste bei Kaltbliitlern. Z. f. Krebsf., 4: 525-564. 1909. fiber einige bei Fischen beobachteten Geschwiilste und geschwulstartige Bil- dungen. Berichte d. K. Bayer. Biol. Ver- suchsstation in Miinchen, 2: 55-76. 1910. Die pathogene Bedeutung der Myxo- boliden fur die Fische. Sitz. Berichte d. Ges. f. Morph, und Physiol, in Miinchen, 26: 20-27. 1912. Eine neue Karpfenkrankheit und ihr Erreger Branchiomyces sanguinis. Zentb. f. Bakterilogie, Parasitologie, Parasiten- kunde und Infektionskr. I. Abt. Originale, 62: 129-134. 1924. Praktikum der Fischkrankheiten. 1st Ed., E. Schweizerbart’sche Verlagsbuchhand- lung, Stuttgart. 1938. Pankreas-Fettnekrose bei karpfenartigen Fischen (Cyprinoiden). Virch. Arch. f. Pathol. Anat. und Physiol., 302: 9-38. Plehn, M., & Mulsow, K. 1911. Der Erreger der “Taumelkrankheit” der Salmoniden. Centralbl. f. Bakter., Para- sitk. und Infektionskr. I Abt. Originale, 59: 63-68. Schaperclaus, W. 1941. Fischkrankheiten. 2d Ed.. Gustav Wenzel und Sohn, Braunschweig. Takahashi, K. 1929. Studien fiber die Fischgeschwfilste. Z. f. Krebsf., 29: 1-73. Thomas, L. 1931. Les tumeurs des poissons. Bull, de l’Assoc. Frang. pour l’etude du cancer. 20: 703-760. Walker, R. 1951. Mycetoma in a landlocked salmon. Anat. Rec., Ill: 115 (Abstract). 1952] Ermin: Fungus and Granuloma in Aphanius chantrei 53 Willis, R. A. 1948. Pathology of Tumours. 1st Ed., Butter- worth and Co., London, and C. V. Mosby, St. Louis, Missouri. WURMBACH, H. 1951. Geschlechtsumkehr bei Weibchen von Lebistes reticulatus bei Befall mit Ich- thyophonus Hoferi Plehn-Mulsow. Roux Arch. f. Entw.-mech., 145: 109-124. 54 Zoologica: New York Zoological Society [37: 4: 1952] EXPLANATION OF THE PLATES (All photomicrographs were made from hemalum-eosin stained sections). Plate I Fig. 1. Granuloma on the dorsal surface in the region Fig. between the operculum and the pectoral fins of a female cyprinodont, Aphanius cliantrei. pj„ About 1.4 X. Fig. 2. Cross-section through the gill and operculum region showing the granuloma as a round, pin. cone-like mass arising over the dorsal surface. 8.5 X. Fig. 3. Another, more posterior, cross-section show- ing several foci of granuloma, one of which contains a large necrotic area. 8.5 X. Plate II 4. Section showing two hyphae of a fungus in the granulation tissue. 1600 X. 5. Section through one part of the granuloma showing many nodules. A necrotic area (gray) may be seen in lower left portion. 65 X. 6. Section showing a part of the granuloma. Two large and some small nodules in the granula- t on tissue are shown at the lower right. Fi- brillar and capsule-like tissue are at the lower left. These tissues are composed of fibroblasts, lymph cells and leucocytes, which infiltrate the musculature (extreme left). 375 X. Fig. 7. The muscle infiltrated by fibrillar, fibrosar- coma-like tissue. Some muscle is not yet com- pletely destroyed. PLATE I. FIG. 1. FIG. 3. FUNGUS ASSOCIATED WITH A GRANULOMA IN A TURKISH FISH, APHANIUS CHANTREI GAILLARD. i :rmin, PLATE II. FIG. 6. 7 * H V V §b T i jf* iuf 1 i': *- -<$ F ? si N ? f -4 flJ \ ■ •# f FUNGUS ASSOCIATED WITH A GRANULOMA IN A TURKISH FISH. APHANIUS CHANTREI GAILLARD. 5 Deep-sea Fishes of the Bermuda Oceanographic Expeditions. Families Cetomimidae and Rondeletiidae Robert R. Harry Natural History Museum, Stanford University, California, and California Academy of Sciences, San Francisco ( Plate I; Text-figures 1-4) Contents Introduction 55 Acknowledgments 56 Family Cetomimidae 58 Genus Ditropichthys Parr 58 Ditropichthys storeri (Goode & Bean) 58 Genus Cetomimus Goode & Bean 59 Key to the species 59 Cetomimus gillii Goode & Bean 60 Cetomimus teevani, new species 61 Cetomimus craneae, new species 64 Genus Cetostoma Zugmayer 68 Cetostoma regani Zugmayer 68 Genus Gyrinomimus Parr 68 Gyrinomimus simplex Parr 68 Family Rondeletiidae 69 Rondeletia bicolor Goode & Bean 69 Summary 69 Literature Cited 70 Introduction IN connection with the author’s systematic and morphological studies on the order Iniomi, Dr. William Beebe and Mr. John Tee-Van kindly made available the strange whale-fishes of the families Cetomimidae and Rondeletiidae obtairied by the Bermuda Ocean- ographic Expeditions of the New York Zoologi- cal Society. While this collection comprises only six species and 16 specimens of the family Ceto- mimidae, it is the largest collection yet studied of one of the rarest families of deep-sea fishes. Apparently only 14 specimens and seven species have previously been recorded in the literature. Of the monotypic family Rondeletiidae, 14 ex- amples have been mentioned in the literature. The Bermuda Expeditions add two more to this number. The Bermuda material includes all previously known genera of both families and provides the second records and second known specimens of Cetomimus gillii Goode & Bean and of Gyri- nomimus simplex Parr; the third record of Rondeletia bicolor Goode & Bean; the fourth record of Ditropichthys storeri (Goode & Bean) ; the fifth record for Cetostoma regani Zugmayer; and two new species of the genus Cetomimus (C. craneae and C. teevani). The most impor- tant results of the present investigation are the discovery for the first time of bioluminescent organs in most species of the family Cetomimi- dae and a unique development of the optic nerve in Ditropichthys storeri. In this form the eye is absent and the optic nerve passes to the surface and branches in the region where the eye should be. Also described is a new subgenus, Psapharocetus, of the genus Cetomimus, for the unique C. kerdops Parr. The Bermuda ma- terial has been deposited in the fish collection of Stanford University and a series has been retained by the New York Zoological Society.1 Practically the entire published work on the families Cetomimidae and Rondeletiidae has been done by three authors. Goode & Bean (1895) discovered both families and described two species of Cetomimus and one of Ronde- letia. The other important reviews of the fami- lies are by Parr, in a series of six papers which appeared between 1928 and 1946. His main systematic review (1934) recognized four ge- nera of Cetomimidae, two of which he described as new, and five species, two of which were new. Myers (1946) and Parr (1946b) have an inter- esting exchange of arguments on the relation- 1 It is of interest to note that the remaining unworked deep-sea fishes collected by the Bermuda Oceanographic Expeditions have recently been transferred to Stanford University and are now deposited at that institution. The work on this report was done in the Natural History Museum of Stanford University. 55 56 Zoologica: New York Zoological Society [37: 5 ships of the families Cetomimidae, Rondeletiidae and Barbourisiidae. The only osteological studies of the families Cetomimidae and Rondeletiidae are by Parr (1929), describing the osteology of Cetostoma regani and Rondeletia bicolor. The single other anatomical study is by Brauer (1908) on the histology of the eye of “Ceto- mimus gillii" (identification doubtful). The his- tory of these families can be obtained from Parr’s 1928 and 1934 papers. While considerably more research has been completed on the families Cetomimidae and Rondeletiidae than is included here, it has been deemed advisable to publish at this time only a relatively brief record of the Bermuda material and to save the remainder of the results for a general monographic review to be published in the Carlsberg Foundation reports. Dr. A. Vedel Taning of the Carlsberg Foundation has sent me the important world-wide collections made by the “Dana,” and Dr. Carl L. Hubbs of the Scripps Institution of Oceanography is lending material obtained in the northeastern Pacific. For data in regard to nets, localities, depths, etc., concerning the capture of cetomimids and rondeletiids treated in this report, the reader may refer to the articles by Dr. Beebe in Zoologica (1931a, 1931b, 1932, 1936). The numbers before the net data are the catalogue numbers of the New York Zoological Society Collection. Part of the cetomimid and ronde- letiid material was listed by Beebe (1937). Acknowledgements To the New York Zoological Society, Dr. William Beebe and Mr. John Tee-Van, I am much indebted for the opportunity to examine the remarkable Bermuda material and for their enthusiastic cooperation during the preparation of this report. In addition they kindly supplied all field data and illustrations previously pre- pared. Also I wish particularly to thank Dr. Daniel Merriman and the Bingham Oceano- graphic Collection for the loan of the types of Cetomimus kerdops Parr, Gyrinomimus myersi Parr and G. simplex Parr; this allowed the study of all known species except Cetomimus picklei (Gilchrist) while preparing this report. Dr. Taning and the Carlsberg Foundation have been most gracious in providing their extensive ma- terial of whale-fishes. Dr. George S. Myers aided considerably by intervening on my behalf to borrow the Bingham Oceanographic Collection types and to examine the collections of ceto- mimids and rondeletiids of the Carlsberg Foun- dation, and by helping in many other ways. The figures of Cetomimus teevani and C. craneae were ably prepared by Miss Florence Sprague. The morphological drawings are by the author. Systematic Position of the Cetomimidae, Barbourisiidae and Rondeletiidae The family Cetomimidae is a very little known group of deep-sea fishes, members of which have seldom been taken, only a few authors being able to examine specimens. The relationships of the group have been little understood, but it was placed in the order Iniomi by Goode & Bean (1895) and has remained there ever since. Jordan (1923) recognized the Cetomimidae and Rondeletiidae in a distinct suborder (Cetun- culi). In general form and a great number of characters the Cetomimidae are most similar to the families Rondeletiidae and Barbourisiidae, and there are many reasons for considering these families to be naturally related. In fact, Myers (1946) goes so far as to place the Bar- bourisiidae as a subfamily of the Cetomimidae. The situation is paradoxical, however, for the Rondeletiidae are currently placed in the order Berycomorphi and the other two families in the order Iniomi. The rondeletiids used to be recognized as iniomous fishes, but Parr (1929) transferred them to the order Berycomorphi on the basis of osteological characters. I am not yet prepared to enter wholeheartedly the argument on the systematic position of either the Cetomimidae or the Rondeletiidae, because I have not yet done enough morphological in- vestigation, but perhaps some preliminary com- ments are in order. Parr’s statement (1929, p. 38) that “an examination of Rondeletia bi- color Goode and Bean 1895a revealed this species to be a typical representative of the order Xenoberyces introduced by Regan 1911” does not seem justified at this time, nor does his state- ment “one may . . . also call attention to such a great number of other osteological characters in which Rondeletia shows a perfect concord- ance with the true Berycomorphi . . .” Actually very little has yet been published on the osteol- ogy of the group Xenoberyces so that a com- parison could apparently be only very limited. Also Rondeletia does not seem to show such a perfect agreement with the order Berycomorphi as Parr concludes, for it possesses a great num- ber of structural peculiarities unknown in any other group of fishes. In fact, there is no bery- comorph that looks like a rondeletiid, so far as general appearance goes. While Parr may be perfectly right in placing the Rondeletiidae in the group Xenoberyces of the order Bery- comorphi, considerably more knowledge is needed of this order before such statements can be fully justified. One of the questions which comes to mind is that, despite the similarity in general form, and the placing of these families together in pre- 1952] Harry: Deep-sea Fishes : Cetomimidae and Rondeletiidae 57 vious classifications, it remains unclear why Parr did not compare the osteology of Ronde- letia with that of the cetomimid ( Cetostoma regani) he examined. A possible answer became apparent while examining the cetomimids col- lected by the Carlsberg Foundation. This ma- terial contains some fine large Cetostoma regani from many widely separated regions, and it was clearly evident that the osteology of this form is probably the most aberrant in the family, and would be the most difficult from which to trace relationships. One of the most interesting recent finds of new deep-sea fishes is Parr’s (1945) discovery of Barbourisia rufa, which he placed in a dis- tinct family of the order Iniomi and compared to the Cetomimidae. Myers (1946) immediately disputed Parr’s recognition of the Barbourisiidae and suggested that the genus should be recog- nized as a subfamily of the Cetomimidae. Parr (1946b) then published a rebuttal to Myers’ arguments, and continues to recognize the family Barbourisiidae. Briefly, the main points of their argument are as follows. Parr (1946b, p. 262) separated these two families primarily on the basis of three characters: “Barbourisia differs from all cetomimids, 1 ) by the presence of normal pelvic fins as compared with their complete absence; 2) by a densely spinulose skin as compared with a completely smooth and scaleless skin; 3) by a red color in life and com- plete absence of melanic elements in the color of the skin, as compared with a melanic pig- mentation.” Myers (1946) minimizes the im- portance of these differences, particularly in regard to separating families by the presence or absence of pelvic fins. What is more revealing is to compare the Barbourisiidae to the Rondeletiidae, which neither Parr nor Myers did. Both groups have almost exactly the same general appearance except that the eye is larger in the latter; the pelvic fins are present in the same position in both groups, but are absent in the Cetomimidae. The distribution and formation of the head pores are the same, differing quite strikingly from the type of pore structure of the cetomimids. There are four gills and a moderate slit behind the fourth in both. The gillrakers are long and slender in both, while they are club-shaped tubercles or flat granular plates in the ceto- mimids. There are no teeth on the palatines in both, while teeth are always present on the palate in the Cetomimidae. Barbourisia differs from both the Cetomimidae and the Ronde- letiidae, taken together, in only one significant character: the densely spinulose skin. In regard to Parr’s character concerning the red color in life, it is now clear that all three families are red in life. Also, a species of Gyrinomimus from the Indo-Pacific lacks most of the melanic pig- mentation normally characteristic of the Ceto- mimidae and is probably quite similar to Bar- bourisia in this respect. The significance of this comparison between the Rondeletiidae and Barbourisiidae is that, on the basis of Parr’s figure and description of Barbourisia, the two groups are closely related and separated by relatively few characters. Thus it now appears to be a question of whether this genus warrants separation from the Ronde- letiidae, and not from the Cetomimidae. Carry- ing this idea farther— because Barbourisia shows several revealing intermediate characters in re- gard to lateral-line, size of eye, and palate den- tition-future investigation might show that all three families should be considered as one. Such a possibility also points out the doubtful pro- priety of placing these families in separate orders. This is further complicated by the fact that neither family closely resembles any other members of either of the two orders (Iniomi, and Berycomorphi) in which they have been placed. There is some reason to suspect relationships to the order Lyomeri. It is of significance that the type of luminous tissue found in the Ceto- mimidae is apparently similar to that found in only one other family of fishes— the Sacco- pharyngidae of the order Lyomeri. Sacco- pharynx harrisoni Beebe (1932b, p. 64) pos- sesses a unique luminous caudal organ that appears from the illustration to be structurally similar to the cavernous luminous tissue around the dorsal and anal fin of the Cetomimidae. There are other superficial similarities, too. The size of the gape is approximately the same in the whale-fishes and this genus. The texture and structure of the skin seems to be the same. The granular dentition is the same in both groups. These families should be compared further. Despite a lack of knowledge of their sys- tematic position, the Cetomimidae, Barbourisii- dae and Rondeletiidae are so similar as to sug- gest that they form a natural group widely dis- tinct from all other families of fishes, and de- serve at least subordinal designation, no matter where placed in the classification of fishes. Jordan (1923, p. 156) recognized them in the suborder Cetunculi of the Iniomi, and the name would be Cetomimina, using the subordinal terminology of Stenzel (1950, p. 94) that was sanctioned by the Committee on Fish Classi- fication of the American Society of Ichthyolo- gists and Herpetologists (see Committee on Fish Classification, 1950, and Schultz, 1951). 58 Zoologica: New York Zoological Society [37: 5 Family Cetomimidae The Cetomimidae comprise at the present time four genera and nine species from the At- lantic and Indian Oceans. The Indian material is inadequately known from collections of the Deutschen Tiefsee-Expedition of the “Valdivia,” reported by Brauer (1906), and of the John Mur- ray Expedition. Following is the classification of the family as presently understood. Subfamily Ditropichthyinae Parr (1934) Genus Ditropichthys Parr (1934) Ditropichthys storeri (Goode & Bean, 1895) Subfamily Cetomiminae Goode & Bean (1895) Genus Cetomimus Goode & Bean (1895) Cetomimus gillii Goode & Bean (1895) Cetomimus teevani, new species Cetomimus craneae, new species Cetomimus picklei (Gilchrist, 1922) Cetomimus kerdops Parr (1934) Genus Cetostoma Zugmayer (1914) Cetostoma regani Zugmayer (1914) Genus Gyrinomimus Parr (1934) Gyrinomimus myersi Parr ( 1934) Gyrinomimus simplex Parr (1946) Dr. Carl L. Hubbs has obtained large ex- amples of the genera Cetomimus and Gyrinomi- mus from the Pacific, but no specific determi- nations have yet been made. Most of the Carls- berg Foundation’s material is from the Indo- Pacific and the group is now known to be circumtropical in distribution at great depths. Diagnosis.— Body moderately short, somewhat compressed, scaleless. Head large, also scaleless. Mouth enormous, generally extending to or moderately behind a vertical from posterior end of cranium. Eyes very small, rudimentary or absent, situated distinctly nearer to upper jaw than to profile of top of head, in middle third of the distance between tip of snout and posterior tip of upper jaw. Teeth granular or cardiform; present on jaws, vomer and palatines; teeth in numerous rows on both jaws. Similar teeth on fused copula and upper pharyngeals. Gillrakers club-shaped or reduced to flat bony plates cov- ered with granular teeth. Gill openings large. Gill membrane free from isthmus. Gills three or four. Dorsal and anal fins far back on body, opposite each other, with a similar number of rays. Adipose and pelvic fins absent. Pectoral fins short, placed rather low. Caudal fin short, moderately forked, truncate, or gently rounded. Lateral-line and lateral-line openings on head and body usually very large. Lateral-line on body a broad hollow tube pierced by large pores. Cavernous, intricately ridged, luminous organs often present around the anus and dorsal and anal fin bases. Luminosity presumed to be red. No photophores present. Body and head cov- ered by a smooth, loose, dark skin. General color when alive brown or orange with brilliant orange or red jaws and fins. Genus Ditropichthys Parr The genus Ditropichthys is superficially char- acterized by ( 1 ) the posterior tips of the lower jaw situated approximately midway between tip of snout and opercular margin, (2) eyes absent, optic nerve extending out to opaque cornea, (3) granular teeth on the jaws and gillrakers, (4) copula (lingual bone) small, short, the sides parallel and the ends rounded, (5) the presence of spinous, club-shaped gillrakers, (6) four gills, (7) the presence of cavernous luminous tissue only in a small area at the anterior margin of the anus, (8) less than 20 rays in either the dorsal or anal fin, (9) a pair of thin dermal ridges along the ventral edge of the abdomen and a scalloped fold on each side of the anal fin base. The genus Ditropichthys presently includes only one species, D. storeri (Goode & Bean). Ditropichthys storeri (Goode & Bean) Specimens taken by the Bermuda Oceano- graphic Expeditions.— Two specimens 45.7 and 94.0 mm. in standard length; original standard lengths taken in the field before shrinkage 50 and 103 mm. respectively; from a cylinder of water eight miles in diameter (five to thirteen miles south of Nonsuch Island, Bermuda), the center of which is at 32° 12' N. Lat., 64° 36' W. Long. No. 18,531; net 882; 700 F.; Sept. 13, 1930; 45.7 mm. No. 22,030; net 1148; 700 F.; Aug. 7, 1931; 94 mm. Specimens previously recorded.— Three speci- mens from the Atlantic and Indian Oceans. For synonymy and discussion see Parr ( 1 928, p. 177; 1934, p. 22) and Norman (1939, p. 32). The specimens Parr and Murray & Hjort (1912, pp. 613, 681, fig. 497) record and figure do not appear to be this species. Description of Bermuda material. — These specimens agree well with the original descrip- tion, but the smaller example is dried out. Of particular interest is the discovery that there is no eye in this form. Instead, the optic nerve appears to extend out to the opaque cornea and branch in the region where the eye should be. The implication is that the optic nerve is light sensitive. I know of no similar condition in vertebrates and plan to investigate in greater 1952] Harry: Deep-sea Fishes: Cetomimidae and Rondeletiidae 59 detail. The cavernous luminous tissue is little developed, and is confined to a small half-moon shaped area bordering the anterior margin of the anus. Field color notes prepared by Beebe.— No. 18,531, net 882, 45.7 mm. in standard length. The entire fish is a solid rich dark sepia brown. No. 22,030, net 1148, 94.0 mm. in standard length. Interior of mouth and gills Carnelian Red2. Skin blackish brown. The dorsal and anal bases, and the mid-dorsal line from behind the fin almost to the caudal fin base pigmented with the same shade of red as the mouth. The bases of the caudal and pectoral fins are faintly tinged with a somewhat lighter shade of red. Finrays Raw Umber. Genus Cetomimus Goode & Bean The genus Cetomimus is superficially charac- terized by (1) the posterior tips of the lower jaw reaching or almost reaching the opercular opening, (2) the presence of relatively well de- veloped eyes, (3) granular teeth on the jaws and gillrakers, (4) copula (lingual bone) fairly broad, slightly indented laterally or dumb-bell shaped, (5) the reduction of the gillrakers to flat bony plates, (6) three gills, (7) usually the presence of cavernous luminous tissue around the anus, along each side of the first anal fin rays and often around the dorsal fin origin, (8) less than 20 rays in either the dorsal or anal fin, (9) the absence of ridges on the belly or folds along each side of the anal fin. The genus Cetomimus presently includes five species, of which C. ker- dops Parr and C. picklei (Gilchrist) are not in- cluded in the Bermuda material. The latter species was not available for study. Since it was discovered to belong in the genus Cetomimus subsequent to Parr’s review (1934), the original description by Gilchrist (1922) and redescrip- tion by Smith (1935) should be consulted. Generic type of Cetomimus by subsequent desig- nation Cetomimus gillii Goode & Bean. The five species of Cetomimus clearly fall into two subgenera, because C. kerdops is so strikingly different from the other species in many characters. In addition the other three species examined, C. gillii, C. teevani and C. craneae, are closely related to each other, form- ing a distinct group of their own. It appears from the available descriptions that C. picklei belongs with the latter group. Many of the differences of C. kerdops from the other species are pre- sented in the relationships of C. craneae and C. teevani, and in the key to the species of Cetomimus. 2 The colors that were identified by comparison to the plates in Ridgway (1912) are capitalized. Subgenus Cetomimus Goode & Bean.— This group contains four species: C. gillii, C. craneae, C. teevani and C. picklei. Subgeneric type Ce- tomimus gillii Good & Bean. Diagnosis: Head moderately compressed or depressed, consid- erably wider than body. Posterior tips of lower jaw well separated, not extending posterior to opercular border. Eyes very small, but fully formed. Outline of upper jaw straight, slightly convex or slightly concave. Lateral-line pores 23 or less. Each jaw with five or more longi- tudinal rows of teeth. Subgenus Psapharocetus, new.— This group contains only one species, and therefore the subgeneric type is Cetomimus kerdops Parr. Diagnosis: Head strongly compressed, as wide as body. Posterior tips of lower jaw touching, extending posterior to opercular border. Eye comparatively the largest in the family except for the genus Cetostoma; eye fully formed. Outline of upper jaw sigmoid. Lateral-line pores on body 25. Each jaw with three irregular longi- tudinal rows of teeth. Psapharo (delicate) ; cetus (whale) . Key to the Species of the Genus Cetomimus la. Posterior tips of lower jaw widely separated; head rounded when viewed dorsally; head distinctly broader than body. (Subgenus Cetomimus Goode & Bean) . 2a. Approximately 10 large pores in the lat- eral-line between gill opening and caudal fin base. . .Cetomimus picklei (Gilchrist) 2b. Seventeen or more large pores in lateral- line between gill opening and caudal fin base. 3a. Posterior lateral-line pores without dis- tinct lappets; pectoral fin rays approxi- mately 16; no cavernous luminous tis- sue around dorsal fin origin Cetomimus gillii Goode & Bean 3b. Posterior lateral-line pores with large lappets; pectoral fin rays 20-23; caver- nous luminous tissue present around dorsal fin origin. 4a. Outline of upper jaw straight pos- teriorly, slightly convex anteriorly; cavernous dorsal luminous tissue confined to a tiny circular patch around the base of the first dorsal ray; posterior lateral-line pores small, less than half the width of the lateral- line tube, their lappets twice the di- ameter of the pores .... Cetomimus teevani, new species 4b. Outline of upper jaw distinctly curved (concave); dorsal luminous 60 Zoologica: New York Zoological Society [37: 5 Text-fig. 1. Main types of premaxillary dentition of whale-fishes. A, side view, and B, cross section, of premaxillary of Cetomimus gillii Goode & Bean, NYZS no. 22,677, illustrating the granular dentition characteristic of Rondeletia, of all known cetomimids except the genus Gyrinomimus, and presumably of Bourbourisia. C, side view, and D, cross section, of premaxillary of Gyrinomimus simplex Parr, NYZS no. 17,198, illustrating the unique dentition among whale-fishes of the genus Gyrinomimus. Since the more primitive cetomimids have the granular teeth, it is likely that the dentition of Gyrinomimus is a specialization from an original granular condition. tissue extensively developed on each side along the base of the first four dorsal rays; posterior lateral-line pores large, equal to the width of the lateral-line tube, their lappets less than the diameter of the pores . . . Cetomimus craneae, new species lb. Posterior tips of lower jaw meeting pos- teriorly; head sharply pointed when viewed dorsally; head as broad as body. (Subgenus Psapharocetus, new) Cetomimus kerdops Parr Cetomimus gillii Goode & Bean Text-figs. 1 & 2 Specimens taken by the Bermuda Oceano- graphic Expeditions.— Eight specimens 34.8 to 59.8 mm. in standard length; August 9, 1929, to August 21, 1931, at 700 to 1,000 fathoms; from a cylinder of water eight miles in diameter (five to thirteen miles south of Nonsuch Island, Ber- muda) , the center of which is at 32° 12' N. Lat., 64° 36’ W. Long. No. 12,138; net 359; 1,000 F.; Aug. 9, 1929; 37.4 mm. No. 13,136; net 428; 1,000 F.; Sept. 5, 1929; 38.2 mm. No. 13,473; net 467; 900 F.; Sept. 12, 1929; 47.7 mm. No. 17,197; net 780; 900 F.; July 5, 1930; 45.4 mm. 1952] Harry: Deep-sea Fishes: Cetomimidae and Rondeletiidae 61 Text-fig. 2. Spongy cavernous luminous tissue around the anus and anterior rays of the anal fin of Cetomimus gillii Goode & Bean. Side view facing left of NYZS no. 22,677. This figure is typical of the kind of luminous tissue found around the dorsal and anal fin origins of most cetomimids, although it is often not so well developed. It is believed that luminosity is accomplished by the exudation of a red substance, forming a luminous mucus layer over large portions of the head and body. The luminous tissue is most persistently developed around the anus and first anal fin rays, since it is seldom absent from this region. No. 18,105; net 862; 800 F.; Sept. 8, 1930; 42.8 mm. No. 18,628; net 892; 800 F.; Sept. 15, 1930; 59.8 mm. No. 22,677; net 1203; 900 F.; Aug. 19, 1931; 51.6 mm. No. 22,778; net 1216; 1,000 F.; Aug. 21, 1931; 34.8 mm. Specimens previously recorded. — Definitely one specimen from the Atlantic and doubtfully a second from the Indian Ocean. For synonymy and discussion see Parr (1928, p. 176; 1934, p.24). Description of Bermuda material. — These specimens agree well with the original descrip- tion, but as few essential characters were con- sidered in that account, little is actually known of this species. The fact that the present material has approximately 16 pectoral rays, significantly agreeing with the original description in this respect, leads me to believe that this determina- tion is correct. Cavernous luminous tissue is ex- tensively developed around the anus and on each side of the bases of the first five anal rays; com- pletely absent dorsally. Standard lengths were taken by Beebe in the field on two specimens. No. 18,628 was 65 mm. and no. 22,677 was 52 mm. Field color notes by Beebe— No. 22,677 , net 1203, 52 mm. in standard length. General color blackish brown with a delicate outer membran- ous skin of bright Orange Chrome. The orange is especially evident on the ventral region. Inside of mouth Violet-Gray. Buccal valves bright Orange Chrome. Cetomimus teevani, new species Text-fig. 3 Specimens taken by the Bermuda Oceano- graphic Expeditions.— Holotype 98.5 mm. in standard length; original standard length taken in field before shrinkage 105 mm.; original num- ber 19,519; Stanford University number 17101; net 961 ; 700 fathoms; September 29, 1930; from a cylinder of water eight miles in diameter (five to thirteen miles south of Nonsuch Island, Ber- muda) , the center of which is at 32° 12' N. Lat., 64° 36' W. Long. Description of Bermuda material.— The only known specimen of this species is in excellent condition and well preserved. Even most of the fin rays are still complete. Body oval in cross section immediately behind head. Greatest depth in region of pectoral fins. Caudal peduncle as long as deep. Anus imme- diately before anal fin; anus surrounded by ex- tensive cavernous luminous tissue. Flead large, slightly compressed; head width slightly less than greatest head depth. Profile of snout interrupted by moderate bony projections before and above eye. Nostrils not easily dis- tinguished, appearing exactly the same as large head pores. Nostrils without raised rims, close to upper jaw, nearer to tip of snout than to eye. Anterior nostril round, its diameter slightly less than that of the eye. Posterior nostril crescent shaped, twice as large as other nostril, close to but obliquely superior to anterior nostril. Nasal organ oblong, situated in center of anterior nos- tril, slightly ridged, without development of olfactory laminae. Internasal area equal to di- 62 Zoologica: New York Zoological Society [37: 5 'ext-fig. 3. A, Side view of holotype of Cetomimus teevani, new species. B, Top view of holotype of ’ etomimus teevani, new species. 1952] Harry: Deep-sea Fishes: Cetomimidae and Rondeletiidae 63 ameter of anterior nostril. A line between eye and tip of snout touches lower border of anterior nostril. Eye very small, slightly nearer upper jaw than profile of head. Interorbital very broad, strongly convex, with a lateral and a median projection. Central part of occiput moderately rounded. Mouth enormous, moderately oblique. Dentigerous margin of upper jaw straight pos- teriorly, slightly convex anteriorly; that of lower jaw slightly concave. Rictus approximately 3.5 eye diameters before posterior tip of premaxil- lary. Jaws extend posteriorly almost to opercular margin. Mandible with a short lateral process behind tip of upper jaw and the tip of the angu- lar produced into a short blunt spine. Distance between ricti about 2.2 into upper jaw length. Teeth on jaws granular, in many series on both jaws. The individual teeth are tiny, always pointed, and either conical or incisiform. Denti- gerous edge of jaws rounded in cross section and teeth extending from outer to inner face of both jaws. Teeth in upper jaw in five (rarely six) irregular series along entire length of pre- maxillary. Teeth in lower jaw similar to those in upper, with approximately 120 teeth in the middle series along the mandible, extending posteriorly well beyond angle of gape. Teeth on vomer and palatines similar to those on jaws. Teeth on vomer incisiform, arranged in a cir- cular dome-shaped patch; posteriorly they are in concentric rings; approximately 10 teeth in width and 1 1 teeth in length across center. Pala- tine series divided into two patches on each side. The teeth in the anterior section are arranged in about five concentric series; there are about 40 teeth in the longest row. The posterior tip of the anterior patch is slightly over two eye diameters before a vertical from eye. Posterior palatine dentition in a long narrow patch, ex- tending far posterior to rictus; approximately 95 teeth in a median longitudinal series; gene- rally five (sometimes six) series of teeth in width. Teeth on copula in a dumb-bell shaped patch with moderately pointed ends. Length of copuiar patch about 1.8 in snout length; with 34 rows of irregular teeth along its length; at widest point 14 series of teeth; width of copula one sixth its length. Anteriorly and posteriorly the teeth are formed in distinct wedge-shaped rows somewhat separated from the other copu- iar teeth. The individual teeth on the copula are similar to those on the jaws. Three gill arches present, no slit behind the last. Holobranchs well developed, their length approximately twice the width of the bony gill arch. Gill-teeth granu- lar, similar to those on jaws but smaller. Teeth on hypobranchial of first arch in a long club- shaped patch with five series of teeth at greatest width and 35 series of teeth in the central row along its length. Teeth on ceratobranchial of first arch in a single long patch with 95 teeth in a longitudinal row and five (rarely six) across. Teeth above angle on epibranchial of first arch in two patches. Near angle there is a tiny circular patch of 15 teeth; other patch long and thin, with about 40 teeth in a row along length and four teeth in width. Pharyngobranchial teeth in two elongate dome-shaped patches on each side. Individual teeth similar to those on jaws. Posterior pharyngeal patch long; 30 teeth in length, nine teeth in width. Anterior patch 24 teeth in length and 10 teeth in width. Lateral-line a broad tube, pierced by 19 pores on body from upper edge of gill opening to base of caudal fin, the last seven pores with well developed flaps on their anterior margins; pos- terior pores small, much narrower than lateral- line tube; posterior flaps as long as or longer than pores. Lateral-line membrane between pores generally has one or two median mucus- tubes anteriorly and two mucus-tubes (one above the other) posteriorly. Longitudinal keel on membranes between pores only slightly de- veloped. There are eight or nine scattered mucus-tubes on the caudal fin behind the last pore. A few scattered mucus-tubes present on back near head. Numerous large pores and a few mucus-tubes scattered over head. No cres- cent shaped rows of mucus-tubes above eyes. Skin loose and flabby over head and body; skin smooth, completely lacking scales or spicules. Dorsal cavernous luminous tissue confined to a small patch around the base of the first dorsal ray. Anal luminous tissue moderately well de- veloped around anus and on each side of the bases of the first four anal rays. Otherwise no luminous organs noted. Dorsal and anal fins far back on body. Dorsal fin origin exactly opposite anal fin origin. Dorsal fin with seven short simple rays followed by eight branched rays and one simple ray. Last two rays separate at base and counted indi- vidually. End of dorsal base exactly opposite end of anal base, measured from where the last ray contacts the profile of the body. Posterior rays of dorsal and anal fins much longer than anterior rays. Anal fin with seven short simple rays followed by eight divided rays and two simple rays. Last rays separate at base and counted individually. Pectoral fin short, low, directed obliquely upward, outline evenly rounded; pectoral rays 20 on each side, counted by dissecting the base and staining with alizarin red S. Pelvic fins absent. Caudal fin short, trun- cate; principal caudal rays 7 + 8. Coloration.— Uniform solid dark brown in 64 Zoologica: New York Zoological Society [37: 5 alcohol. Tips of fins light. Interior of mouth lightly pigmented with brown. Buccal valves dark brown. Inside of operculum more lightly pigmented than interior of mouth. (There are no notes on life-colors accompanying this specimen). Measurements in percent, of standard length. —Body depth measured at a vertical from region of pectoral fins, 21.5. Caudal peduncle length, which is the oblique distance between the end of the anal base and mid-base of caudal fin, 7.81. Least depth of caudal peduncle, 7.81. Head length measured from the anterior tip of the snout (upper jaw) to the most distant point of the opercular margin including mem- branous flaps, 37.8. Snout length measured from the tip of the snout (upper jaw) to the anterior margin of the fleshy orbit, 14.1. Upper jaw length, which is the distance between the tip of the snout and posteriormost point of the pre- maxillary, 30.1. Eye diameter, which is the greatest visible distance, without dissection, across the orbit, and corresponding to the di- ameter of the opaque cornea, 1.32. Interorbital width, which is the least fleshy width, 12.1. Greatest head width measured between the sides of the premaxillaries, 17.9. Distance between tip of snout and visible base without dissection of first dorsal fin ray, 77.6. Dorsal fin base length from origin of luminous tissue to base of last ray, 17.9. Distance between tip of snout and anus, 76.5. Length of anal fin base from anus to base of last ray, 18.0. Distance of dorsal luminous tissue behind a vertical from anus, 1 .32. Length of pectoral fin measured from base of longest middle ray (found by dissection) to its tip, 9.55. Length of caudal fin from mid-base (measured from the fold when the fin is flexed from side to side) to a vertical from tip of longest ray, 12.7. Distance between origin of dorsal luminous tissue and mid-base of caudal fin, 26.7. Relationships— This new species belongs in the genus Cetomimus, agreeing completely with all characters used to distinguish the genus. In general body proportions and many characters it is intermediate between Cetomimus gillii and C. craneae, but shows only a distant relation- ship to C. kerdops. The comparisons with C. gillii in drawing relationships have been taken from the Bermuda specimens of this species. Cetomimus teevani differs from the other species of the genus in (1) the formation of the dorsal cavernous luminous tissue (absent in C. gillii, well developed along the sides of the first dorsal rays in C. craneae and C. kerdops, confined to a small patch around the first dorsal ray in C. teevani ) ; and (2) the size of the lateral- line pores (in the other species the body pores are the width of the lateral-line, in C. teevani most of the posterior pores are much smaller than the lateral-line width). This new species differs from C. kerdops and C. gillii but agrees with C. craneae in the possession of well de- veloped lappets on the posterior lateral-line pores. Cetomimus teevani further differs from C. gillii in the following characters: (1) pectoral rays 20 in the former, 16 in the latter; (2) lower jaw outline when viewed from below distinctly pointed versus evenly rounded; (3) pores bor- dering upper jaw between nostrils and eye dis- tinctly smaller than eye versus considerably larger; (4) teeth in jaws considerably smaller and less distinct in C. teevani, about 160 teeth in the middle row of the premaxillary versus approximately 100. Cetomimus teevani further differs from C. craneae in the following characters: (1) pec- toral rays 20 versus 23; (2) head compressed versus as broad as deep, head width 17.9% of standard length as compared to 23.8%; (3) most of the lappets of the posterior lateral-line pores much longer than diameter of pores versus always less than diameter of pores; (4) region above tip of snout lacking mucus-tubes versus covered with over 50 tubes; (5) ends of dorsal and anal fin bases exactly opposite versus dorsal ending distinctly before a vertical from end of anal base. Cetomimus teevani further differs from C. kerdops in the following characters: (1) pos- terior tips of lower jaw over half the snout length apart versus practically meeting in C. kerdops; (2) posterior process of lower jaw not reaching opercular border versus extending well beyond; (3) 19 lateral-line pores on body versus 25; (4) lower jaw inferior versus extending slightly before upper jaw; (5) opercular mem- branes at isthmus join well behind a vertical from eye versus joining below a vertical from eye. Name— This species is named in honor of Mr. John Tee-Van in appreciation of his im- portant part in the Bermuda Oceanographic Expeditions. Cetomimus craneae, new species Text-fig. 4 Specimens taken by the Bermuda Oceano- graphic Expeditions.— Holotype 82.4 mm. in standard length; original number 11,370; Stan- ford University number 17102; net 269; 800 fathoms; July 8, 1929; from a cylinder of water eight miles in diameter (five to thirteen miles 1952J Harry: Deep-sea Fishes: Cetomimidae and Rondeletiidae 65 south of Nonsuch Island, Bermuda), the center of which is at 32° 12' N. Lat., 64° 36' W. Long. Description of Bermuda material— The only known specimen of this species is in excep- tionally fine condition and not damaged in any way. Most of the fin rays are still complete. Body moderately oval in cross section imme- diately behind head. Greatest depth immedi- ately behind head. Length of caudal peduncle slightly greater than its least depth. Anus imme- diately before anal fin, surrounded by extensive cavernous luminous tissue. Head large, its width equal to its depth. Profile of snout evenly rounded onto occiput. Nostrils not easily distinguished, appearing exactly like the head pores. Nostrils round, of equal size, close to premaxillary; both large, without raised rims except that the anterior nostril has a slight swollen lobe at its anterior margin. Nasal organ tear-drop shape, without any trace of distinct olfactory laminae, situated internally between the two nostrils. Internasal membrane same size as diameter of nostrils. A line between eye and tip of snout passes through anterior nostril; pos- terior nostril above this line. Eye very small, much nearer upper jaw than profile of head. Interorbital very broad, strongly convex, with two median longitudinal humps. Central part of occiput flat. Mouth enormous, moderately oblique. Dentigerous margin of both jaws slight- ly concave. Rictus approximately four eye di- ameters before posterior tip of premaxillary, laws extend posteriorly to opercular margin. Mandible with a short lateral process behind end of upper jaw and the tip of the angular is produced into a short blunt spine. Distance between ricti about 1.3 into upper jaw length. Teeth on jaws granular, in many series on both jaws. The individual teeth are tiny and either conical with their lengths equal to their basal diameter or else incisiform with a flat outer margin. Edge of jaws rounded in cross section and teeth extending from outer to inner face of both jaws. Teeth in upper jaw in six irregular series along entire length of premaxillary; ante- riorly the innermost series is separated from the remaining teeth by a narrow space. There are approximately 1 40 teeth in the middle row along the premaxillary. Teeth in lower jaw similar to those in upper, with approximately 120 teeth in the middle series along the mandible, extend- ing posteriorly behind angle of gape. Teeth on vomer and palatines similar to those on jaws. Teeth on vomer incisiform, in an oval dome- shaped patch; anteriorly the teeth are in con- centric rings. There are approximately 12 teeth across and 14 teeth along the length of the vomer. Palatine series divided into two patches on each side. The teeth in the anterior section are arranged in about twelve concentric series; there are about 40 teeth in longest row. The posterior tip of the anterior patch is an eye diameter anterior to a vertical from the eye. Posterior palatine patch long, fairly broad, ex- tending well posterior to rictus; there are slightly over a hundred teeth in a longitudinal series, 10 series of teeth in width. Teeth on copula in a dumb-bell shaped patch; copula with a moder- ately pointed anterior end and rounded posterior margin. Length of copular patch about 1.5 in snout length, with 28 rows of irregular teeth along its length; at widest point 17 series of teeth; width of copula one-third its length. An- teriorly the copular teeth are arranged in dis- tinct wedge-shaped rows, posteriorly these merge into a closely packed mass. The individual teeth on the copula are round at the base and incisi- form at the tip. Three gill arches present, no slit behind last. Holobranchs well developed, their lengths approximately twice the width of the bony gill arch. Gill-teeth granular, similar to those on jaws but more feebly developed. Teeth on hypobranchial of first arch in a single flat club-shaped patch; six series of teeth at greatest width and 30 teeth in middle row along its length; the individual teeth are in irregular longitudinal series. Teeth on ceratobranchial of first arch in a single long bony patch of irregular outline; the teeth are in irregular patches, some in longitudinal series and others in oblique series; there are approximately 90 teeth in longitudinal series and 10 across the bone. Teeth above the angle on epibranchial of first arch in two patches; near angle there is a small crescent-shaped patch of about 10 teeth in width; other patch long and thin with about 28 teeth along its length and six teeth across its width. Pharyngobranchial teeth developed in two cone-shaped bony patches on each side; individual teeth similar to those on jaws. Pos- terior pharyngobranchial patch oval, 24 teeth in length, 15 teeth in width. Anterior patch 23 teeth in length and 12 teeth in width. Lateral-line a broad tube pierced by 22 large pores on body, the last nine pores with well developed flaps on their anterior margins; lateral-line pores same size as tube, their lappets always shorter than diameter of pores. In addi- tion, the last sections have a longitudinal keel on the membranes between the pores. Lateral- line membranes between pores generally have one median mucus-tube anteriorly and two mucus-tubes (one above the other) posteriorly. There are eight mucus tubes in groups of two on the caudal fin behind the last lateral-line pore. Scattered mucus-tubes present along mid- 66 Zoologica: New York Zoological Society [37: 5 1952] Harry: Deep-sea Fishes: Cetomimidae and Rondeletiidae 67 line of back. Lateral-line pores continued on head forward onto snout. Numerous large cavernous pores and many mucus-tubes scat- tered over head. No crescent-shaped rows of mucus-tubes above eyes, but a curved row is present across interorbital on top of head. Skin loose and flabby over head and body, completely lacking scales or spicules. Cavernous luminous tissue well developed around origins of dorsal and anal fins and on each side of the bases of the first four rays of these fins; also extensively formed around anus. Otherwise no luminous organs noted. Dorsal and anal fins far back on body. Dorsal fin origin slightly anterior to anal fin origin. Dorsal and anal fin with seven short simple rays followed by nine branched and one simple ray. Last two rays separate at base and counted in- dividually. End of dorsal base slightly in advance of anal fin base. Posterior rays of dorsal and anal fins much longer than anterior rays. Pec- toral fin short, fairly low, directed obliquely upward, outline evenly rounded. Pectoral rays 23 on each side; counted by dissecting pectoral fin base. Pelvic fins absent. Caudal fin short, slightly emarginate; principal rays 6 + 8. Coloration.— Uniform solid dark brown in alcohol. Tips of fins light. Roof of mouth lightly pigmented brown; floor of mouth darker. Buccal valves dark brown. Inside of operculum lightly pigmented brown. Field color notes prepared by Beebe.— The fish at first was a dark brown tinged with Terra Cotta. Before being put in preservative the epidermis became loose. After a night in 3% formalin, the entire epidermis was blown away with a pipette. The fish then became a dark Seal Brown. (It is probable that the “epidermis” referred to was a coating of coagulated mucus, for the fish still possesses its epidermis). Color notes from field drawings.— Two color illustrations were prepared of the holotype by Helen Tee-Van (B127, no. 284). One of them shows the specimen as being a solid orange over head, body and fins and must have been pre- pared before preservation. The other illustra- tion is in much greater detail and corresponds to Beebe’s notes. The general color is a dark brown. The jaws, fins and pores are tinged with orange, particularly the cavernous luminous tissue and vertical fins. Measurements in percent, of standard length. —Body depth measured at a vertical from pos- terior tip of head, 20.9. Caudal peduncle length, which is the oblique distance between the end of the anal fin base and mid-base of caudal fin, 10.6. Least depth of caudal peduncle, 5.70. Head length measured from the anterior tip of the snout (upper jaw) to the most distant point on the opercular margin including membranous flaps, 36.7. Snout length measured from the anterior tip of the snout (upper jaw) to the anterior margin of the fleshy orbit, 12.5. Upper jaw length, which is the distance between the tip of the snout and posteriormost point of the premaxillary, 28.9. Eye diameter, which is the greatest visible distance without dissection across the orbit, .85. Interorbital width, which is the least fleshy width, 14.9. Greatest head width measured between posterior tips of lower jaw while head was freely suspended, 23.8. Dis- tance between tip of snout and visible base without dissection of first dorsal fin ray, 76.7. Dorsal fin base length from origin of luminous tissue to base of last ray, 18.6. Distance between tip of snout and anus, 70.8. Length of anal fin base from anus to base of last ray, 18.4. Distance of dorsal luminous tissue before a vertical from the anus, 2.31. Distance between end of dorsal fin base posteriorly to a vertical from end of anal fin base, .85. Length of pectoral fin mea- sured from base of longest middle ray (found by dissection) to its tip, 8.86. Length of caudal fin from mid-base (measured from the fold when the fin is flexed from side to side) to tip of longest ray, 10.2. Distance between origin of dorsal luminous tissue and mid-base of caudal fin, 29.4. Relationships.— This new species clearly be- longs in the genus Cetomimus, agreeing com- pletely with all characters used to distinguish the genus. In general body proportions and many characters this species is most closely re- lated to C. teevani and shows the closest rela- tionships to the genus Gyrinomimus of any species outside that genus. The head is broad, and the outlines of the jaws distinctly concave as in Gyrinomimus. This new species is much more closely related to C. gillii than to C. kerdops. Cetomimus craneae differs from the other species of the genus in (1) the more extensive development of the cavernous tissue than any other known species in the family; (2) having the head as broad as deep versus distinctly com- pressed in the other species of Cetomimus; (3) having the highest number of pectoral fin rays in the genus, 23 versus 16-20; (4) the presence of great numbers of mucus-tubes over the head and back which are only sparsely present in the other species; this condition is most closely ap- proached by C. gillii, but there are considerably fewer in this form. Cetomimus craneae further differs from C. gillii in the following characters : ( 1 ) outline of jaws distinctly concave versus straight in C. 68 Zoologica: New York Zoological Society [37: 5 gillii; (2) posterior lateral-line pores possessing well developed lappets versus lappets vestigial or absent; (3) dorsal cavernous luminous tissue present versus absent; (4) pores bordering upper jaw between nostrils and eye distinctly smaller than eye versus considerably larger than eye; (5) sides of copular patch strongly indented versus practically straight. Cetomimus craneae further differs from C. kerdops in the following characters: (1) Pos- terior tips of lower jaw over three-fourths of the snout length apart versus practically meeting in C. kerdops; (2) opercular membranes at isthmus join well behind a vertical from eye versus joining below a vertical from eye; (3) posterior lateral-line pores possessing well de- veloped lappets versus lappets absent; (4) sides of copular patch strongly indented versus prac- tically straight; (5) head considerably wider than body versus as wide as body. Further comparisons to the other species of Cetomimus are given in the section on rela- ships of C. teevani. Name— This unusual fish is named in honor of Jocelyn Crane in recognition of her work on deep-sea fishes. Genus Cetostoma Zugmayer The genus Cetostoma is superficially charac- terized by (1) having the posterior tips of the lower jaw extend well posterior to the opercular opening, (2) the presence of relatively com- pletely developed eyes, (3) having granular teeth on the jaws and gillrakers, (4) having the copula (lingual bone) very narrow, long and slender, somewhat expanded at anterior tip, (5) the reduction of the gillrakers to flat bony plates, (6) having three gills, (7) the presence of a tiny area of modified cavernous luminous tissue on each side of anus, but none elsewhere, (8) having approximately 30 rays in both dorsal and anal fins, and (9) the presence of a low ventral median fold on belly from below pec- toral fins to anus, but lack of folds on each side of anal fin. The genus Cetostoma includes a single remarkable species, C. regani Zugmayer, which shows many unique differences from all other genera. Its closest relationships lie with Cetomimus kerdops. It should be noted that this determination of genus and species is made on the basis of Parr’s findings, because Zug- mayer’s original description is useless. Cetostoma regani Zugmayer Specimens taken by the Bermuda Oceano- graphic Expeditions.— Two specimens 41.8 and 43.7 mm. in standard length; original number 17,196; net 778 at 700 fathoms, July 5, 1930; from a cylinder of water eight miles in diameter (five to thirteen miles south of Nonsuch Island, Bermuda), the center of which is at 32° 12' N. Lat., 64° 36' W. Long. Specimens previously recorded.— Three speci- mens from the Atlantic. See Parr (1934), and Murray & Hjort (1912, pp. 613, 682, fig. 498). Description of Bermuda material.— Both spec- imens are in rather poor condition, having par- tially dried out at one time, but there appears to be no question of their identity. The standard lengths were probably 10-15 mm. longer before preservation. Normal cavernous luminous or- gans are not present in this form, but there ap- pears to be modified luminous tissue of this basic type in a small narrow ridge on each side of the anus. Genus Gyrinomimus Parr The genus Gyrinomimus is superficially char- acterized by ( 1 ) the posterior tips of the lower jaw reaching the opercular openings, (2) the presence of relatively well developed eyes, (3) having elongate cardiform teeth on the jaws and granular teeth on the gill arches, (4) having the copula (lingual bone) broad, shield-shaped, moderately pointed anteriorly and expanded laterally posteriorly, (5) the reduction of the gillrakers to flat bony plates, (6) having three gills, (7) the complete absence of cavernous luminous tissue, or if such is present, by having it only faintly formed around base of first dorsal ray and around anus, (8) having less than 20 rays in both the dorsal and anal fins, and (9) the absence of ridges on the belly or folds along each side of the anal fin. The genus Gyrinomi- mus includes two species of which G. myersi is not included in the Bermuda material. Generic type by original designation, Gyrinomimus myersi Parr. This genus is most closely related to Cetomimus, particularly through C. craneae, to which it shows the greatest similarities. Gyrinomimus simplex Parr Text-fig. 1 Specimens taken by the Bermuda Oceano- graphic Expeditions.— One specimen 48.3 mm. in standard length: original standard length before shrinkage 54 mm.; original number 17,198, net 732 at 900 fathoms, "June 27, 1930; from a cylinder of water eight miles in diameter (five to thirteen miles south of Nonsuch Island, Ber- muda) , the center of which is at 32° 12' N. Lat., 64° 36' W .Long. Specimens previously recorded— One speci- men from the Atlantic. See Parr ( 1946). Description of Bermuda material.— This speci- men agrees well with the holotype. The posterior 1952] Harry: Deep-sea Fishes: Cetomimidae and Rondeletiidae 69 lateral-line pores are round, completely lacking lappets. No cavernous luminous tissue. Family Rondeletiidae The family Rondeletiidae has been reported upon only three times. The original description (later copied by various authors) was made by Goode & Bean (1895) from a single specimen, which was placed in the order Iniomi. Parr (1928) recorded and briefly described 13 speci- mens, and in 1929 he considered the osteology in considerable detail, placing the family in the group Xenoberyces of the order Berycomorphi. As previously stated, Myers (1946) and Parr (1946b) have interesting arguments about the systematic position of this family. Apparently only two illustrations giving the aspect of the unique species have been prepared. The first figure appeared with the original description by Goode & Bean and has been used by several sub- sequent authors. The second illustration is a colored plate prepared under the direction of Dr. William Beebe that has appeared in the National Geographic Magazine (Beebe, 1931). Since this latter plate is of considerable impor- tance in establishing the coloration of this group and is more accurate than the original figure, it has been reproduced with this paper. The Rondeletiidae include only one genus and species, Rondeletia bicolor Goode & Bean. Rondeletia bicolor Goode & Bean Plate I Specimens taken by the Bermuda Oceano- graphic Expeditions— Two specimens 24.7 and 86.7 mm. in standard length; original stand- ard lengths taken in field before shrinkage 25.7 and 97 mm. respectively; from a cylinder of water eight miles in diameter (five to thirteen miles south of Nonsuch Island, Bermuda), the center of which is at 32° 12' N. Lat., 64° 36’ W. Long. No. 13,434; net 463; 1000 F.; Sept. 11, 1929; 86.7 mm. No. 23,339; net 1291; 600 F.; Sept. 12, 1931; 24.7 mm. Specimens previously recorded. — Fourteen specimens from the Atlantic. See Parr (1928, p. 179; 1929, p. 39). Description of Bermuda material.— The two Bermuda examples agree quite closely with the previous descriptions. No luminous structures are found, but it is believed that the orange areas in life are luminous. It is of particular signifi- cance that its coloration in life is very similar to that known for most members of the Cetomi- midae. Field color notes by Beebe. — The colored plate of this species has obviously been prepared from no. 13,434, but the original color notes are not available. From the color plate it can be seen that the fish is dark brown with bright red- dish-orange jaws, and that the fin bases are also reddish-orange. No. 23,339, net 1291, 24.7 mm. in standard length. General color blackish-brown. The pel- vic fins are strikingly webbed with scarlet. Re- mainder of color notes not complete enough to be included. Summary The families Cetomimidae and Rondeletiidae are remarkable groups of rare luminous whale- like fishes, about two to six inches long, which have been sporadically collected in the Atlantic and Indian Oceans. The systematic position of these families has aroused some argument, but appears to be in the neighborhood of the orders Iniomi, Berycomorphi, and perhaps the Ly- omeri. The families Cetomimidae, Barbourisii- dae and Rondeletiidae seem to form a naturally related group with the second named group in- termediate in position, but considerably closer to the Rondeletiidae. Previous specimens of the family Cetomimi- dae had been placed in four genera and seven species. In the course of the oceanographic work of the Department of Tropical Research of the New York Zoological Society, during nine years’ investigation within an eight-mile circle off Bermuda, sixteen cetomimids and two rondeletiids were obtained. This collection, while small in number, has added considerably to our knowledge of these excessively rare fishes. This material again shows the great value of concentrated research to find out just what part of the total bathypelagic fish fauna could be taken in a restricted area. In regard to the Ce- tomimidae and Rondeletiidae, all known genera were taken and all but three of the previously known species. In addition two new species of the genus Cetomimus were collected (C. craneae, C. teevani), these being of particular value in tracing the relationships of the genera Cetomi- mus and Gyrinomimus. All the Bermuda ma- terial was taken between the depths of 700 and 1 ,000 fathoms. Dr. Beebe’s field notes give the first life color information ever obtained on these strange fishes. These notes reveal the “orange-mouths,” or “giant-mouth orange-skins” as Beebe termed them, to be usually of a solid orange or brown in life with bright orange or reddish-orange mouths and fins. Bioluminescence has not been previously dis- 70 Zoologica: New York Zoological Society [37: 5 cussed in the family Cetomimidae, although these fishes are apparently luminous. Most species of this family have peculiar webbed, spongy, cavernous tissue around the anus and the first dorsal and anal fin rays; these tissues appear to represent luminous organs. This type of luminous structure appears to be similar to that found in only one other species of fish, Saccopharynx harrisoni Beebe, a gulper eel of the order Lyomeri. Since the cavernous tissue of the cetomimids is red or reddish-orange in life, it is presumed that the luminescence may also be of these colors. The peculiar luminous tissue of S. harrisoni glows scarlet, apparently showing a similarity to the cetomimids in this respect. A remarkable discovery is the complete ab- sence of eyes in the cetomimid, Ditropichthys storeri. Instead, the optic nerve extends out to a heavily pigmented area in the opaque cornea and branches near the surface of the skin. The implication is that the optic nerve is light-sensi- tive but further study is needed before more can be said of this condition, which is unique among vertebrates. Literature Cited Beebe, William 193 la. Bermuda Oceanographic Expeditions 1929-30. No. 1— Introduction. Zoologica, vol. 13, no. 1, pp. 1-14, figs. 1-7. 1931b. Bermuda Oceanographic Expeditions 1929-30. No. 2— List of nets and data. Zoologica, vol. 13, no. 2, pp. 15-36, 20 tables. 1931c. A round trip to Davy Jones’s Locker. Peering into mysteries a quarter mile down in the open sea, by means of the bathy- sphere. National Geogr. Mag., vol. 59, no. 6, pp. 653-678, 15 figs., 8 pis. 1932a. Bermuda Oceanographic Expeditions 1931. Individual nets and data. Zoologica, vol. 13, no. 3, pp. 37-45, 8 tables. 1932b. Nineteen new species and four post-larval deep sea fish. Zoologica, vol. 13, no. 4, pp. 47-107, figs. 8-31. 1936. Bermuda Oceanographic Expeditions. Individual nets and data, 1932-1935. Zoologica, vol. 21, no. 3, pp. 69-73, 5 tables. 1937. Preliminary list of Bermuda deep-sea fish. Based on the collections from fifteen hundred metre-net hauls, made in an eight-mile circle south of Nonsuch Island, Bermuda. Zoologica, vol. 22, no. 14, pp. 197-208. Brauer, August 1906. Die Tiefsee-Fische. L, Systematischer Teil. Wiss. Ergeb. Deutschen Tiefsee-Exped. “Valdivia” 1898-1899, vol. 15, pp. 1-432, figs. 1-176, pis. 1-18. 1908. Die Tiefsee-Fische. II., AnatomischerTeil. Wiss. Ergeb. Deutschen Tiefsee-Exped. “Valdivia” 1898-1899, vol. 15, pp. 1-266, figs. 1-11, pis. 19-44. Committee on Fish Classification 1950. Report of the Committee on Fish Classi- fication. Copeia, 1950, no. 4, p. 326. Gilchrist, John Dow Fisher 1922. Deep-sea fishes procured by the S.S. “Pickle” (Part I). Rep. Fish. Mar. Biol. Surv. Union S. Africa, vol. 2, pp. 41-79, pis. 7-12. Goode, George Brown, & Tarleton Hoffman Bean 1895. On Cetomimidae and Rondeletiidae, two new families of bathybial fishes from the northwestern Atlantic. Proc. U. S. Nat. Mus., vol. 17, pp. 451-454, pi. 17. Jordan, David Starr 1923. A classification of fishes including families and genera as far as known. Biol. Sci. Stanford Univ., vol. 3, no. 2, pp. 79-243, i-x. Murray, John, & Johan Hjort 1912. The depths of the ocean. A general ac- count of the modern science of oceanogra- phy based largely on the researches of the Norwegian steamer Michael Sars in the North Atlantic. MacMillan, London, xx + 82 1 pp., 575 text figs., 1 1 pis. Myers, George Sprague 1946. On a recently proposed new family of deep-sea fishes (Barbourisiidae, Parr, 1945). Copeia, 1946, no. 1, pp. 41-42. Norman, John Roxbrough 1939. Fishes. John Murray Expedition 1933-34. Sci. Rep., vol. 7, no. 1, pp. 1-116, text-figs. 1-41. Parr, Albert Eide 1928. Deepsea fishes of the order Iniomi from the waters around the Bahama and Ber- muda Islands. Bull. Bingham Ocean. Coll., vol. 3, no. 3, pp. 1-193, figs. 1-43. 1929. A contribution to the osteology and classi- fication of the orders Iniomi and Xeno- beryces; with description of a new genus and species of the family Scopelarchidae, from the western coast of Mexico; and some notes on the visceral anatomy of Rondeletia. Occ. Pap. Bingham Ocean. Coll., no. 2, pp. 1-45, figs. 1-18. 1952] Harry: Deep-sea Fishes: Cetomimidae and Rondeletiidae 71 1934. Report on experimental use of a triangular trawl for bathypelagic collecting with an account of the fishes obtained and a re- vision of the family Cetomimidae. Bull. Bingham Ocean. Coil., vol. 4, art. 6, pp. 1-59, figs. 1-21. 1945. Barbourisidae, a new family of deep sea fishes. Copeia, 1945, no. 3, pp. 127-129, pi. 1. 1946a. A new species of Gyrinomimus from the Gulf of Mexico. Copeia, 1946, no. 3, pp. 116-117, pi. 1. 1946b. On taxonomic questions related to the classification of Barboitrisia, the Cetomi- midae and the Iniomi. Copeia, 1946, no. 4, pp. 260-262. Ridgway, Robert 1912. Color standards and color nomenclature. Washington, D. C., 43 pp., 53 pis. Schultz, Leonard Peter 1951. Results of a preliminary survey of group endings in zoological classification above the category of genus. Science, vol. 113, pp. 655-657. Smith, J. L. B. 1935. New and little known fishes from South Africa. Rec. Albany Mus., vol. 7, pp. 169- 235, pis. 18-23. Stenzel, Henry Bronislaw 1950. Proposed uniform endings for names of higher categories in zoological systematics. Science, vol. 112, p. 94. Zugmayer, Erich 1914. Diagnoses de quelques poissons nouveaux provenant des campagnes du yacht Hiron- delle II (1911-1913). Bull. Inst. Ocean. Monaco, no. 288, pp. 1-4. 72 Zoologica: New York Zoological Society [37: 5: 1952] EXPLANATION OF THE PLATE. Plate I. Adults of the rare whale-fish, Rondeletia bicolor, feeding on very small crustaceans. The coloration was noted before preservation, establishing that these unusual fishes are red in life to a considerable extent. (From a painting by E. Bostelmann). HARRY. PLATE I. DEEP-SEA FISHES OF THE BERMUDA OCEANOGRAPHIC EXPEDITIONS. FAMILIES CETOMIMIDAE AND RONDELETIIDAE. NEW YORK ZOOLOGICAL SOCIETY GENERAL OFFICE 30 East Fortieth Street, New York 16, N. Y. PUBLICATION OFFICE The Zoological Park, New York 60, N. Y. OFFICERS PRESIDENT Fairfield Osborn SECRETARY Harold J. O’Connell SCIENTIFIC STAFF GENERAL John Tee-Van Executive Secretary Sam Dunton Photographer ZOOLOGICAL PARK Lee S. Crandall General Curator James A. Oliver Curator of Reptiles Leonard J. Goss Veterinarian Charles P. Gandal. . .Assistant Veterinarian VICE-PRESIDENTS Alfred Ely Laurance S. Rockefeller Donald T. Carlisle TREASURER Cornelius R. Agnew William Bridges Editor and Curator of Publications Robert M. McClung. . Assistant Curator of Mammals and Birds Grace Davall Assistant to General Curator AQUARIUM Christopher W. Coates . Curator and Aquarist James W. Atz Assistant Curator Ross F. Nigrelli Pathologist Myron Gordon Geneticist C. M. Breder, Jr Research Associate in Ichthyology Homer W. Smith Research Associate in Physiology DEPARTMENT OF TROPICAL RESEARCH William Beebe Director William K. Gregory Associate Jocelyn Crane Research Zoologist JohnTee-Van Associate Henry Fleming Entomologist SCIENTIFIC ADVISORY COUNCIL A. Raymond Dochez Alfred E. Emerson W. A. Hagan Caryl P. Haskins K. S. Lashley John S. Nicholas EDITORIAL COMMITTEE Fairfield Osborn, Chairman James W. Atz William Beebe William Bridges Christopher W. Coates Lee S. Crandall Leonard J. Goss James A. Oliver John Tee- Van ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME 37 • PART 2 • SEPTEMBER 19, 1952 • NUMBERS 6 TO 10 PUBLISHED BY THE SOCIETY The ZOOLOGICAL PARK, New York Contents Page 6. The Effects of Prolonged Treatment with Acriflavine on the Killifish, Fundulus heteroclitus (Linnaeus). By Grace E. Pickford 73 7. Effects of ACTH and Cortisone on the Pituitary, Thyroid and Gonads of the Teleost Astyanax mexicanus. By Priscilla Rasquin & Ethel Hafter Atz. Plates I & II; Text-figures 1 & 2 77 8. The Effects of Holothurin on Fish, and Mice with Sarcoma 180. By Ross F. Nigrelli 89 9. Sex Determination in Xiphophorus (Platypoecilus) maculatus. III. Differ- entiation of Gonads in Platyfish from Broods Having a Sex Ratio of Three Females to One Male. By Myron Gordon. Plates I & II; Text-figure 1 . . . . 91 10. Four New Species of Geometridae (Moths) from Rancho Grande, North- central Venezuela. By D. S. Fletcher. Plate I; Text-figures 1-7 101 6 The Effects of Prolonged Treatment with Acriflavine on the Killifish, Fundulus heteroclitus (Linnaeus) Grace E. Pickford Bingham Oceanographic Laboratory, Yale University THE experiments reported below were un- dertaken to determine whether prolonged treatment with acriflavine, used in an at- tempt to control an infectious epidemic similar in its symptoms to that described by Wells & Zobell (1934) for Fundulus parvipennis but not curable by the heat treatment recommended by these authors, was responsible for the develop- ment of cirrhosis of the liver due to the accumu- lation of large amounts of a ceroid-like material. The bacteriocidal and trypanocidal acri- flavine dyes (acriflavine, trypaflavine, euflavine) are commonly used in the treatment of proto- zoan infections of fish. Schaperclaus (1941), for example, recommends a bath in solutions containing 1 gm in 50 to 100 liters of water for ten hours, but the dye may be allowed to remain in the water until the color “wears off” (Nigrelli, personal communication). The latter procedure had been used in the treatment of infected fish which, at autopsy, showed the liver condition mentioned above. It was thought that the acri- flavine might have a direct action or, alterna- tively, that it might act indirectly through the elimination or partial elimination of the normal intestinal bacterial flora which, as is well known in higher vertebrates, may play an important role in supplementing the natural source of vita- mins derived from the food. The experiment was designed to test both these hypotheses. My best thanks are due to the staff of the Bingham Oceanographic Laboratory, especially to my former assistants, Miss Harriet A. Chambers and Mrs. Shirley M. Conover, and to Mr. A. Charles Laws, all of whom participated in the preparation of the tissues, and to Mrs. Joyce Lownsbery who assisted with the sta- tistical calculations. I am also indebted to Dr. Ross F. Nigrelli, Dr. Donald F. Poulson, and Dr. Daniel Merriman for many valuable sug- gestions. Materials and Methods Four tanks each containing 52 liters of sea water were arranged as follows: Tanks 9 and 10 had plain sea water, Tanks 11 and 12 each re- ceived in addition 10.4 cc of 1% neutral acri- flavine so that the approximate strength of the dye was 2 gm per 1,000 liters. On Oct. 26, 1950, eight two-year-old specimens of Fundulus heteroclitus were placed in each tank. All were supposedly males but, at this time of year, the sexes are difficult to separate and at autopsy one fish in Tank 9 was discovered to be a female. Each fish was marked with colored beads sewn on with tantalum wire, and its weight and length were recorded. The fish were in complete sexual regression with at most a mere trace of nuptial coloration. During the course of the experi- ment one fish died in Tank 11, from unknown causes. The female in Tank 9 has been ex- cluded from the analysis. The color of the water in the tanks containing acriflavine faded slowly with time and 10 cc of 1 % stock solution were added on the following dates: November 14, January 8, February 11 and March 4. The fish were fed daily, with rare exceptions. All tanks received a regular diet of cooked liver- pablum (Gordon, 1943). Tanks 10 and 12, one without and one with acriflavine, received nothing but this diet during the whole course of the experiment. In accordance with the plan outlined above. Tanks 9 and 11 received a dietary supplement intended to provide an ad- ditional source of vitamins. This supplement was offered about thrice weekly and consisted at first of chopped raw beef liver but, beginning on February 11, this was replaced with frozen, raw daphnia in generous amounts. Also, after Febru- ary 11a trace of iodine was added to the liver- pablum, since results of previous experiments, which became available at this time, showed that 73 74 Zoologica: New York Zoological Society [37: 6: Table 1. Summary of the Experimental Results No acriflavine Acriflavine Daphnia No daphnia Daphnia No daphnia Tank 9 10 11 12 No. of fish 7 8 7 8 Weight % incr. 40. 0± 10.7 44.2±13.3 25.1±4.8 28.0± 1 1 .9 Length % incr. 14.4±6.5 10.3±2.4 6.9±1.4 9.3±4.9 Testis GSI1 3.9+0.9 4.2±2.4 3.5±1.2 1.4±0.4 Nuptial colors Strong Strong Strong Slight Liver HSI2 3.5±0.6 3.8±0.7 3.4±0.5 3.9±0.7 Liver color Pinkish-brown Pinkish, reddish, or grayish-brown Pinkish, reddish, or grayish-brown Ochraceous Liver glycogen Abundant Abundant Abundant Abundant Liver fat Abundant Abundant Abundant Abundant Liver ceroid Trace Trace Trace Trace Thyroid epith. ht.3 5.4±0.7 5.0±0.6 4.6±0.6 5.2±0.5 Thyroid active colloid4 4 fish 3 fish 2 fish 1 fish 4GSI = gonosomatic index. 2HSI = hepatosomatic index. 3Thyroid epith. ht., standard deviation calculated for the differences between the means for each fish. 4Thyroid active colloid: predominently blue or particolored with Azan. fish fed exclusively on this diet tended to develop a low to moderate grade of goiter. On April 2-3, 159-160 days after the start of the experiment, the fish were killed and sub- jected to autopsy. Weight, length and nuptial coloration were again recorded. The liver was weighed and its size expressed in terms of the hepatosomatic index. Liver color was noted against the Ridgway color chart, and slices were fixed in alcohol-formalin (for glycogen) and calcium formol (for fat). Glycogen was esti- mated subjectively on sections stained by the periodic-Schiff method, with malt diastase con- trols. Fat was similarly estimated on frozen sec- tions stained with scarlet red and haematoxylin, control sections being treated with scarlet red dissolved in 100% alcohol which removes most if not all of the soluble fat in one hour. Ceroid was stained red by the periodic-Schiff method on the glycogen controls and remained as un- dissolved material stained by scarlet red on the fat controls. The development of nuptial coloration is a visible indication of the maturation of the male gonad. In addition, the testes were weighed and the relative size expressed in terms of the go- nosomatic index. The hypobranchial region was fixed in Bouin’s fluid and the thyroid was studied on serial sections, alternate slides being stained with haematoxylin-eosin and with Azan. The height of a single follicle cell was measured from each follicle on every twentieth section and the average cell height calculated from 100 cells for each fish. Results The overall results are summarized in Table I. Weight.— Fish in the two acriflavine tanks showed a retardation of growth as measured by percentage of weight increase. The dietary sup- plement had no effect on untreated fish and conferred no beneficial effect in the presence of acriflavine. Length— Effects on growth in length are more difficult to interpret. The best growth was ob- tained in Tank 9 (daphnia, no acriflavine) but there is no significant difference between Tanks 9 and 10, although growth in the latter was somewhat less. Therefore, in the absence of acriflavine the dietary supplement had little or no effect. Similarly when Tanks 11 and 12 are compared there is no significant difference. However, the fish in Tank 11 (daphnia and acriflavine) showed a very marked retardation of growth in length, which is significantly dif- ferent from that in either Tanks 9 or 10 (no acriflavine). Contrary to expectation, and in disagreement with the results based on weight increase, the length increase in Tank 12 is not statistically less than that in Tanks 9 and 10. The less striking retardation of growth in Tank 12 may be correlated with the inhibition of sexual development discussed below. The values of P in the comparison of length increase are given in Table 2. Analysis of vari- ance supports the hypothesis that the four tanks are not parts of a homogenous population (F = 5.40, df — 3 and 26) but, in view of the 1952] Pickford: Effects of Acriflavine on Killifish 75 small numbers, a more detailed study was not attempted. It is felt that a larger series would show that growth in length, like growth in weight, is retarded by the presence of acrifla- vine. Sexual development— It was evident by in- spection that the fish in Tank 12 were in a state of partial sexual regression, as manifested by the weak development of nuptial coloration. This was fully confirmed by the low values of the gonosmatic index. The small differences be- tween Tanks 9, 10 and 11 are not statistically significant. These results indicate that sexual re- tardation due to treatment with acriflavine may be abolished by a diet enriched with raw daph- nia. The effects on the testes are therefore in- direct or moderated by a more efficient metab- olism of the dye. Table 2. Values of P in Comparisons of Growth in Length Tank 9 Tank 10 Tank 1 1 Tank 10 0. 1-0.2 Tank 11 0.01 0.001-0.01 — Tank 12 0.1 0.6-0. 7 0.2-0. 3 Liver.— It had been anticipated at the start of the experiment that the liver would show marked differences correlated with the acrifla- vine treatment, hence the careful study of fat and glycogen. The results do not confirm this expectation. In all groups the hepatosomatic indices were in the normal range for spring males. There is a slight tendency for the fish in Tanks 10 and 12, which did not receive a diet- ary supplement, to have larger livers, but the differences are not statistically significant. Simi- larly the study of sections revealed no appre- ciable differences in fat or glycogen content. All livers were heavily laden with reserves, showing the fish to be in a well-nourished con- dition. It is probable that if the experiment had been continued through the advancement of the breeding season the fish with well-developed gonads would have developed a depletion of liver reserves similar to that which is known to occur in the wild population (Chambers, 1951; Pickford, 1952). No fish showed cirrho- sis of the liver and the small amounts of ceroid which were present in all cases are such as occur normally in wild fish. The data regarding liver weight, and glycogen and fat content, do not appear to tell the whole story. The livers of fish from Tank 12 were all more or less strongly ochraceous in color and after fixation a yellow, non-fluorescent, water soluble coloring matter dissolved out into the fixing fluid. The livers of fish in the three remaining tanks showed the normal range of pinkish or reddish-brown, with an occasional grayish-brown, and did not yield this bright yel- low solution after fixation. It appears that in the absence of a suitable dietary supplement the livers are unable to metabolize the acrifla- vine in a normal manner. Thyroid— In contrast to earlier experiments in which no iodine had been added to the liver- pablum mixture, no fish developed goiter. The condition of the thyroids is essentially the same in all tanks. The slightly lower epithelial height noted in the fishes of Tank 11 is just barely statistically significant, at the 5% level, when this group is compared either with Tank 9 or with Tank 12, but it is not significantly different from Tank 10. Analysis of variance does not support the hypothesis that the Tank 1 1 group is significantly different from the pop- ulation as a whole (F = 2.09, df = 3 and 26). No correlation was observed between the av- erage epithelial height and the condition of the colloid as seen after staining with Azan. In most cases the thyroid follicles were predominantly inactive, with homogenous, red-staining colloid. In each tank some individuals showed a more active condition, with predominantly blue-stain- ing, granular, or parti-colored colloid. The inci- dence of active colloid was certainly higher in those groups whose growth was not retarded by acriflavine. Discussion In a study of the antibacterial action of acri- flavine, Mcllwain (1942) has shown that inhibi- tion of growth caused by this dye may be re- versed by nucleic acid and to a lesser extent by degradation products of nucleic acid. Acrifla- vine forms complex salts with nucleic acid, lacking the fluoresence of the free substance. A second type of reversal could be obtained with marmite and with phenylalanine, the effect be- ing augmented by riboflavin and other hydrogen carriers such as methylene blue. Mcllwain con- cludes that the inhibited system lacks both substrate and normal H-transport mechanisms and that acriflavine probably combines with enzyme systems involving nucleotides. As Mcllwain points out, the inhibitory action of acriflavine on cell respiration is well known; the work of Quastel & Wheatley (1931) on bac- teria is cited, also the work of Scheff & Hassko (1936) on trypanosomes. The latter authors conclude that the effect is on a cyanide-sensi- tive hydrogen carrier. More recently Ephrussi and co-workers (Ephrussi, 1950, and references 76 Zoologica: New York Zoological Society [37: 6: 1952] therein) have shown that acriflavine induces an inheritable cytoplasmic mutation in yeast. The mutants lack cytochrome oxidase and succinic dehydrogenase, and are unable to utilize glyco- gen except by anaerobic metabolism. A biochemical study was beyond the scope of the present investigation, although an exami- nation of the respiratory enzyme systems of the liver in acriflavine-treated fish would doubtless be most illuminating. The inhibitory action on the growth of the fish may be a direct effect, since it cannot be offset by a dietary supple- ment of raw daphnia. The pronounced retarda- tion of the maturation of the testes is in a different category since it may be prevented by a suitable enrichment of the diet. It is suspected that this may be due to a vitamin deficiency but it is also possible that raw daphnia provides factors similar to those noted by Mcllwain in marmite which restore the growth of acriflavine- inhibited bacteria. The retardation of the gonads supports a pop- ular belief, to which I can find no scientific ref- erence, that treatment with acriflavine causes temporary sterility in fish1. However, it is known from the work of Hertwig (1924) and Dalcq ( 1931 ) that spermatozoa may be inactivated by acriflavine, although the mechanism of this in- activation has not been elucidated. In respect to the development of hepatic cirr- hosis, towards which the investigation was pri- marily directed, the results are entirely negative. This disease is not caused by the presence of acriflavine in the water, nor by the prolonged use of a cooked liver-pablum diet without sup- plementary sources of vitamin. The problem of the development of ceroid in the liver of Fun- dulus is still under investigation. Summary 1. Prolonged treatment with acriflavine retard- ed the growth of Fundulus heteroclitus on a standard cooked liver-pablum diet. This ef- fect was not offset by a dietary supplement of raw, frozen daphnia. 2. Acriflavine caused a marked retardation of the testes, prevented by the addition of raw daphnia to the standard diet. 3. Acriflavine had no effect on liver weight, nor on the glycogen and fat reserves of the liver. It did not induce hepatic cirrhosis resulting from the accumulation of ceroid in the liver. However, in the absence of raw daphnia, a liver disfunction was indicated by the ac- 1In the popular aquarium literature, Earle E. Patterson (Aquarium Journal [San Francisco], Vol. 21, p. 36, 1950) has reported a loss in reproductive capacity in fishes treated with acriflavine. cumulation of a yellow, non-fluorescent, water soluble coloring matter. 4. Acriflavine had little or no effect on the function of the thyroid gland as reflected in average follicle cell height and colloid con- dition. References Chambers, H. A. 1951. The effect of thiourea on male Fundulus heteroclitus. Anat. Rec., 109 (2): 106. Dalcq, A. 1931. Contribution a l’analyse des fonctions nu- cleates dans l’ontogenese de la grenouille. 1. Etudes cytologiques des oeufs fecon- des par du sperme trypaflavine. Arch. d. Biol., 41: 143-220. Ephrussi, B. 1950. Induction par l’acriflavine d’une mutation specifique chez la levure. Pubbl. Staz. Zool. Napoli, 22 Suppl.: 1-16. Hertwig, G. 1924. T rypaflavin als Radiumsalz zur gewinnung haploidkerniger Froschlarven. Anat. Anz., 58 Suppl.: 223-227. Gordon, M. 1943. Feeding platyfishes and swordtails. Aquar- ium, 12: 86-88. McIlwain, H. 1942. A nutritional investigation of the anti- bacterial action of acriflavine. Biochem. Journ., 35: 1311-1319. Nigrelli, Ross F. 1951. (Personal communication). Quastel, J. H. & A. H. M. Wheatley 1931. The action of dyestuffs on enzymes. 1. Dyestuffs and oxidations. Biochem. Journ., 25: 629-638. PlCKFORD, G. E. 1952. Hormonal regulation of liver size in fish. Anat. Rec., 112 (2): 127-128. SCHAPERCLAUS, W. 1941. Fischkrankheiten. Gustav Wenzel & Sohn, Braunschweig. Scheff, G. & A. Hassko 1936. Einfluss einiger Chemotherapeutika auf den Stoffwechsel der Trypanosomen, mit Riicksicht auf das Interferenzphanome- non. Zbl. Bakt., Abt. 1, 136: 420-424. Wells, N. A. & C. E. Zobell 1934. Achromobacter ichthyodermis, n. sp., the etiological agent of an infectious derma- titis of certain marine fishes. Proc. Nat. Acad. Sci., 20: 123-126. 7 Effects of ACTH and Cortisone on the Pituitary, Thyroid and Gonads of the Teleost Astyanax mexicanus" Priscilla Rasquin & Ethel Hafter Atz The American Museum of Natural History (Plates I & II; Text-figures 1 & 2) A LTHOUGH considerable work has been /\ done on the effects of ACTH and corti- JL JLsone on mammals, comparatively little attention has been given cold-blooded verte- brates. Much of the mammalian histological work has been limited to a study of one or two different organs under identical experimental procedures. The use of a small teleost, Astyanax mexicanus (Filippi), affords an opportunity to study the action of administered hormones on several affected organs simultaneously because it is practicable to section the entire fish serially. Rasquin ( 1951) and Rasquin & Hafter (1951a), in experiments dealing with adrenal cortical function in normal and tumorous tele- osts, found evidence tending to show a depres- sion of thyroid activity and some inconclusive evidence concerning pituitary function after ad- ministration of ACTH. The present experiment was designed to clarify these results and to pro- vide information on the effects of continuous injection of ACTH and cortisone on both these glands as well as the gonads, and to discover whether the effects were reversible. Materials and Methods Ninety-six sexually mature fish, Astyanax mexicanus (Filippi), of the same spawning, 1 An abstract of this report may be found in Anat. Rec., suppl. Ill (3) : 41. 1951. The work was supported in part by grants from the National Cancer Institute, U. S. Public Health Service, and from the American Cancer Society on recommendation of the Committee on Growth of the National Research Council. Thanks are due Dr. Irby Bunding of Armour and Company, and Dr. Augustus Gibson of Merck and Company for generously supplying the hormones. The authors wish to thank Dr. C. M. Breder, Jr., of the American Museum of Natural History for suggestions during the course of the work and for criticism of the manuscript. were divided into six groups of 16 fish each and were treated as shown in the schedule of Table 1 . Four groups received daily doses respectively of ACTH, two different doses of cortisone ace- tate and control injections of physiological saline. Each hormone dose was given in 0.06cc of 0.6% saline; controls received 0.06cc of 0.6% saline. Daily doses of hormone for these groups were 0.2mg cortisone acetate, 0.05mg cortisone ace- tate and 0.1 mg ACTH. Two groups received identical doses of ACTH and physiological sa- line divided into two injections per day. The ACTH was given as 0.05mg in 0.03cc saline, corresponding to the volume given to the con- trols. The double daily injections were consid- ered necessary because other investigators have demonstrated that ACTH is rapidly inactivated in the body. Van Dyke, Simpson, Li & Evans (1950) have shown that the life of the ACTH molecule in the circulation is extremely short. The cortisone acetate was received as a sa- line suspension. Inasmuch as this was mammal- ian physiological saline, the suspension was di- luted with distilled water from 0.9% to 0.6% for use in fish. To make up the 0.05mg doses, the suspension was further diluted with 0.6% saline so that each dose would be contained in 0.06cc, equivalent in fluid volume to those dos- ages given to the other fishes injected once per day. Injections were made for ten consecutive days and two fish of each group were preserved on the third, fifth, eighth and tenth days after the beginning of the injection period and at similar times after the termination of injections. The animals were killed four hours after their final injection, allowing sufficient time to elapse for any shock effect to be obliterated. Two hours 77 78 Zoologica: New York Zoological Society [37: 7: Table 1. Modifications of Thyroid and Pituitary Induced by ACTH and Cortisone Thyroid Pituitary Mean Total No. Weight Total Thyroid Standard Cells in % Baso- % Acido- % Chromo- Fish No. in Dosage Height in Error Transi- phils phils phobes and Sex grams in m ms. microns tional Lobe Controls (One Injection per Day) 1-M 1.3 3.13 0.0802 5,067 59.4 38.4 2.2 2-F 2.3 — 3.23 0.0819 15,071 46.0 52.5 1.5 3-M 1.1 — 2.78 0.0596 6,671 52.0 45.9 2.1 4-F 2.7 — 3.38 0.0712 25,389 67.6 31.3 1.1 5-M 2.0 — 3.36 0.0713 15,236 63.6 35.5 1.9 6-F 1.4 — 3.58 0.0854 7,482 49.0 48.2 2.8 7-M 2.0 — 2.94 0.0713 13,510 60.4 38.2 1.4 8-F 1.6 — 2.83 0.0701 10,963 56.9 40.8 2.3 9-M 1.3 — 3.20 0.0711 7,970 51.5 47.0 1.5 10-F 1.5 — 3.13 0.0702 7,914 61.6 36.9 1.5 1 1-M 0.8 — 3.13 0.0798 6,800 62.8 34.3 2.9 12-F 3.2 — 2.96 0.0644 14,674 62.0 36.9 1.1 13-M 0.9 — 3.11 0.0675 5,258 52.2 45.4 2.4 14-F 2.5 — 3.14 0.0734 18,016 52.2 46.9 0.9 15-M 2.3 — 2.95 0.0563 18,494 51.9 47.0 1.1 16-F 1.4 — 3.10 0.0686 12,506 61.5 37.3 1.2 Cortisone-injected 17-M 1.9 0.6 2.18 0.0581 7,624 44.3 54.0 1.3 18-F 2.9 0.6 2.78 0.0810 12,729 47.0 51.3 1.7 19-M 2.5 1.0 2.44 0.0619 15,350 54.9 44.3 0.8 20-F 1.7 1.0 2.79 0.0888 11,700 46.1 51.8 2.1 21-M 2.1 1.6 2.58 0.0639 9,471 49.8 49.2 1.0 22-F 1.8 1.6 2.51 0.0541 11,756 42.2 55.9 1.9 23-M 2.1 2.0 2.52 0.0628 16,098 38.8 60.8 0.4 24-F 1.5 2.0 2.56 0.0611 8,698 44.2 54.2 1.6 25-M 2.1 — 2.17 0.0549 12,565 37.9 61.0 1.1 26-M 1.3 — 2.23 0.0526 7,626 42.2 57.0 0.8 27-M 1.3 — 2.55 0.0545 7,865 46.2 53.2 0.6 28-M 1.4 — 2.55 0.0656 5,405 41.6 57.3 1.1 29-M 2.2 — 2.65 0.0670 11,151 50.5 48.5 1.0 30-M 1.4 — 2.52 0.0680 8,015 25.1 73.3 1.6 3 1-M 1.8 — 2.63 0.0698 13,818 40.4 58.5 1.1 32-M 1.5 — 2.41 0.0618 16,122 40.2 57.8 2.0 33-M 2.7 0.15 2.08 0.0487 13,188 65.5 33.5 1.0 34-F 3.3 0.15 2.60 0.0626 30,997 44.2 55.2 0.6 35-M 1.3 0.25 2.36 0.0584 9,774 44.5 54.6 0.9 36-M 2.1 0.25 2.73 0.0634 11,374 49.2 49.8 1.0 37-M 1.8 0.40 2.48 0.0560 6,580 49.8 49.5 0.7 3 8-F 2.0 0.40 2.59 0.0661 11,334 55.9 43.0 1.1 39-M 1.5 0.50 2.18 0.0526 10,136 55.7 43.1 1.2 40-M 1.3 0.50 2.28 0.0580 7,245 49.2 49.7 1.1 41-M 1.4 — 2.39 0.0618 10,877 44.6 54.3 1.1 42-M 1.3 — 2.54 0.0658 9,320 39.9 58.8 1.3 43-M 0.8 — 2.62 0.0640 4,915 50.6 48.2 1.2 44-M 3.1 — 2.75 0.0743 14,947 51.4 48.0 0.6 45-M 1.9 — 2.38 0.0596 14,579 47.3 52.0 0.7 46-M 1.1 — 2.77 0.0664 5,870 46.3 53.0 0.7 47-M 1.6 — 2.77 0.0779 8,994 45.6 53.4 1.0 48-M 1.3 — 2.85 0.0697 9,191 51.6 47.3 1.1 1952] Rasquin & Atz: ACTH and Cortisone in a Teleost 79 Table 1. Modifications of Thyroid and Pituitary Induced by ACTH and Cortisone (Continued) Thyroid Pituitary Mean Total No. Weight Total Thyroid Standard Cells in % Baso- % Acido- % Chromo- Fish No. in Dosage Height in Error Transi- phils phils phobes and Sex grams in mgms. microns tional Lobe ACTH-injected 49-M 1.5 0.3 2.45 0.0680 6,562 45.6 53.0 1.4 50-F 2.5 0.3 2.81 0.0751 13,666 64.2 35.0 0.8 51-M 2.2 0.5 2.30 0.0557 16,766 50.7 47.4 1.9 52-F 2.6 0.5 2.67 0.0767 10,575 52.2 46.0 1.8 53-M 1.5 0.8 2.47 0.0595 5,356 48.8 48.3 2.9 54-F 1.7 0.8 2.35 0.0589 8,100 52.0 45.5 2.5 55-M 1.6 1.0 2.37 0.0560 11,579 40.6 57.7 1.7 56-F 1.5 1.0 2.17 0.0549 14,254 57.1 41.8 1.1 57-M 1.9 — 2.24 0.0626 14,383 43.2 55.2 1.6 58-F 2.5 — 2.37 0.0626 12,928 58.0 40.9 1.1 59-M 1.6 — 2.58 0.0642 9,373 46.0 51.7 2.3 60-F 2.6 — 2.87 0.0757 20,663 56.6 41.8 1.6 61-M 1.7 — 2.80 0.0819 6,776 48.9 49.8 1.6 62-M 1.1 — 2.98 0.0749 5,612 62.4 36.2 1.4 63-M 1.7 — 3.06 0.0734 10,025 52.3 46.6 1.1 64-M 1.3 — 3.10 0.0777 8,599 62.8 36.5 0.7 65-M 1.9 0.3 2.25 0.0559 10,969 66.1 33.7 0.2 66-F 1.7 0.3 2.42 0.0541 13,220 61.0 38.0 1.0 67-M 1.2 0.5 2.22 0.0558 7,396 54.7 44.5 0.8 68-F 2.8 0.5 2.10 0.0488 15,512 57.3 41.7 1.0 69-M 1.4 0.8 2.33 0.0602 9,520 40.1 59.0 0.9 70-F 1.9 0.8 2.18 0.0481 11,524 50.7 48.4 0.9 71-M 1.2 1.0 2.33 0.0580 8,853 39.3 59.2 1.5 72-F 3.0 1.0 2.27 0.0545 23,096 50.3 47.9 1.8 73-M 1.8 — 2.34 0.0599 21,710 43.1 55.5 1.4 74-F 1.8 — 2.30 0.0587 10,498 51.3 47.7 1.0 75-M 1.8 — 2.14 0.0514 15,738 52.9 46.1 1.0 76-F 1.8 — 2.92 0.0783 13,902 55.1 43.4 1.5 Controls (Two Injections per Day) 77-M 1.8 2.57 0.0623 14,603 52.2 45.7 2.1 78-F 3.2 — 2.49 0.0620 19,825 74.5 24.3 1.2 79-M 1.8 — 2.65 0.0634 13,465 53.2 45.6 1.2 80-F 2.7 — 2.88 0.0678 19,506 57.6 41.5 2.9 81-M 1.6 — 2.72 0.0613 12,028 56.3 42.7 1.0 82-F 1.3 — 2.72 0.0915 17,831 50.1 47.8 2.1 83-M 1.3 — 3.03 0.0700 8,579 46.1 52.8 1.1 84-F 1.8 — 2.46 0.0629 13,329 55.8 43.1 1.1 85-M 2.4 — 2.96 0.0682 14,183 51.3 47.7 1.0 86-F 2.9 — 2.85 0.0754 17,575 64.3 34.8 0.9 87-M 1.8 — 3.13 0.0712 13,100 58.6 40.1 1.3 88-F 1.5 — 3.07 0.0703 12,369 61.0 37.9 1.1 after injection with ACTH, shock effects were observed, but no such effects were noted four hours after injection (Rasquin, 1951). An at- tempt was made to use equal numbers of males and females and to sacrifice one of each sex at each experimental period. Actually more males than females were selected. The fish were killed by immersion in Bouin’s fluid. The gonads and the part of the kidney con- taining the corpuscles of Stannius were removed 80 Zoologica: New York Zoological Society [37: 7: and sectioned separately. The remainder of each fish after decalcification was serially sec- tioned transversely at seven microns from the region of the eye posteriorly through the head- kidney. Sections through the regions of the pitui- tary were cut at five microns and stained by Masson’s trichrome method. The other sections were stained with either Masson’s trichrome stain or Harris’s hematoxylin and eosin. The term “acidophil” as used here refers to those cells that stain with acid fuchsin or ponceau, and “basophil” denotes those cells that stain with fast green. Differential cell counts of the tran- sitional lobes of the pituitaries were made by means of a Leitz grid ocular micrometer. All the cells were counted in every fourth section from the beginning to the end of transitional lobe tissue and thus every part of the lobe was represented in the final count. The total number of cells for each fish seen in Table 1 is therefore an index and is not intended to represent the actual numbers of cells. Thyroid epithelial heights were measured after the method of Rawson & Starr (1938), except that 100 instead of 200 measurements were taken on each fish. Five times a week the animals were fed a pre- pared dried food containing dried shrimp, both muscle and shell, liver, chopped lettuce and spinach, Pablum and salt. The formula for mak- ing this food has been published by Aronson (1949). Experimental Results In the transitional lobes of the pituitaries of Astyanax mexicanus the basophils number ap- proximately 60% of the total cells and the acid- ophils number about 40% (Rasquin, 1949). Chromophobes are relatively scarce, making up only 1% or 2%, rarely 3%, of the total. The normal percentage of basophils was lowered by all the injection schedules, including those of the saline used for controls. These results differ from those of Golden & Bondy (1952), who found that in rats cortisone caused an increase in basophil count, while ACTH evoked no sig- nificant change in the count. All the data used for making the graphs and for statistical analysis may be found in Table 1 . The table also gives the total dosages of hor- mone received by each injected fish, and com- parisons may be made between the states of the thyroid and pituitary glands of any individuals. Text-figure 1 illustrates graphically the effects of ACTH and cortisone injection on the per- centage of pituitary basophils. Four fish receiv- ing two ACTH injections per day died before the end of the experiment and their correspond- ing controls were not sectioned. The data indi- cate that there is a significant difference between the male and female reaction to ACTH. Both series of ACTH injections lowered the propor- tion of basophils in the males while the propor- tions of cells in the female pituitaries showed no significant difference from the controls. Graphs A and B show the results of double and single daily injections, respectively, of ACTH on the male compared with the female. In Graph C the mean of the basophil percentages of the two fish given two injections daily of ACTH is compared with the control, and Graph D is similarly plotted for the single daily injection. Both these graphs show a decrease in the pro- portion of basophils in the ACTH-injected fish when compared with the saline-injected con- trols. There is no significant difference in effect between the double and single daily injections. In both groups of fish a tendency to return to the normal ratio is seen after the tenth day when injections ceased. Graphs E and F of Text-figure 1 show the re- sults of large and small doses of cortisone respec- tively on the males compared with the females. Cortisone lowers the proportion of basophils in both males and females and there is no signifi- cant difference in the reactions of the two sexes. Only males were left in these groups of fish after injections were completed. In graphs G and H the means of basophil percentages of the two fish given large and small doses of cortisone respectively have been compared with the con- trols. In both instances there is a decrease in the proportion of basophils, although the larger dose is more effective than the smaller. The sta- tistical analysis indicates that no trend to recov- ery within the post-injection period occurred in the pituitaries of the cortisone-treated fish. Histologically, however, some recovery was evidenced by the appearance of mitotic figures in basophils. The inversion of the normal basophil-acid- ophil ratio is accompanied by profound histolog- ical changes. Plate I, Figure 1, is a photomicro- graph of a part of the transitional region of one of the single daily injected controls. This gland is not different histologically from the normal in which most of the basophils show some vacu- olation. Vacuolation is definitely increased in the basophils of the controls receiving two saline injections per day and some degranulation also occurs so that the staining reaction is weaker (Figure 2). The acidophils, which are the smaller, dark, polygonal cells, appear to be un- affected by the saline injection routines. Baso- phils of the ACTH-injected fish show no loss of vacuolation but no degranulation was evident. The cytoplasm maintains its intense staining affinity, although pycnotic nuclei are sometimes seen, particularly near the periphery of the 1952] Rasquin & Atz: ACTH and Cortisone in a Teleost 81 gland. Basophils of the cortisone-treated fish frequently give evidence of complete exhaustion, with a loss of normal staining reaction and large vacuoles filled with debris. Mitotic figures are occasionally seen in the basophils of this group and these are most frequent in the fish killed in the post-injection period. Plate I, Figure 3, shows a basophil in anaphase in the transitional lobe of a fish killed eight days after cortisone injections began. In both ACTH- and cortisone- injected fish there is a hypertrophy and an in- crease in numbers of acidophils. The same phenomenon is observed in both groups but is more pronounced in the cortisone-treated fish where the condition is maintained to the end of the post-injection period. The acidophils in- crease greatly in size of nucleus and quantity of cytoplasm and are so filled with secretion and so intensely stained that the cell boundaries are often impossible to locate and the tissue has become friable like colloid. Plate I, Figure 4, shows the hypertrophy and hyperplasia of acidophils in a cortisone-treated fish. The indi- cation is that the acidophils are witholding their secretion, and that the injected hormones have in some way interfered with its normal release. The cause of the inversion in the ratio of acidophils to basophils in the treated fish is not clear. In all the pituitaries examined only two mitotic figures were seen in acidophils. It seems probable that most of the new acidophils come from the granulation of chromophobes, which also are occasionally seen in mitosis and often appear with a few acidophilic granules in the cytoplasm. These latter sparsely granulated cells were counted as acidophils. The decrease in ratio of basophils is not merely a numerical function of the increase of acidophils alone, for this de- crease is seen to some extent in the controls where histologically the acidophils show no change from the normal. There is no indication that basophils are changed into any other type of cell. There is no evidence of necrosis in any of these glands nor is any phagocytosis seen. Occasionally lymphocytes are observed in the glands but these do not appear to be engaged in phagocytic activity. It is possible that the technique of counting is responsible for what may be only an apparent loss of basophils. In order that no duplication might occur, every fourth section was counted, because the sections were five microns thick and some of the active basophils measured nearly 20 microns across. However, clusters of nuclei of basophils practically devoid of cytoplasm were seen in many of the pituitaries and many of these would remain uncounted as they would lie between the selected sections. The technique then gives a reliable index only of the function- ing basophils. Increasing numbers of basophils in the recovering gland may have their source in regranulation of the depleted cells as well as in mitotic division of more active ones. Thyroid — Text-figure 2A is a graphic repre- sentation of the effects of ACTH on thyroid epithelial heights. No sex difference is shown in the reaction of the thyroids to either ACTH or cortisone, and therefore the two fish from each experimental period were grouped together and their means compared with those of the two controls. From the graph it can be seen that double daily injections of saline depress the thy- roid height below that of single daily injections. Both ACTH injection series depress the thyroid height below that of the controls, the double daily dose being quicker acting and more effective during the experimental period than the single daily injection. After the tenth day when injec- tions ceased, the single daily injected ACTH group shows recovery of the thyroid and indi- cation of a return to normal is observed in the double daily injected ACTH group. Text-figure 2B shows graphically the effects of large and small doses of cortisone on thyroid epithelial heights. While both dosages of cortisone depress the height of the thyroid epithelium below that of the controls, there is no statistically significant difference between the effects of the two doses of cortisone. With neither dosage of cortisone does the height of the thyroid epithelium return to that of the control fish during the post-in- jection period. Histologically, the thyroids of the control fish show a low cuboidal epithelium with some vac- uoles in both cells and colloid. In both ACTH- and cortisone-treated fish, follicles are larger and the epithelium is so flattened as to be squa- mous in character in some follicles. It is probable that this effect on the thyroid is not mediated by the pituitary through any loss of thyrotropic function, but is rather a more direct effect of cortical hormones. Table 1 shows that some of the lowest thyroid epithelial heights were asso- ciated with pituitaries that were still unimpaired by hormone injections. Gonads.— The ovaries of all the control fish contain numbers of immature and mature eggs as well as an occasional resorbing egg. The fol- licular epithelium surrounding the immature egg is flat and becomes higher as the egg matures. In the normal reproductive cycle, the ripe eggs are resorbed if spawning does not take place. The resorption process is carried on by phago- cytic activity of infiltrating connective tissue and of the follicular cells which become hyperplastic and hypertrophied at this time. Plate II, Figure 5, is a photomicrograph of the ovary of a saline- injected control showing normal conditions. PERCENTS OF PITUITARY BASOPHILS 82 Zoologica: New York Zoological Society [37: 7: AGTH CORTISONE 40 - 30 * 20 1 I I i 1 1 1 1 « I 1 1 1 1 i > ' 3 5 6 to 13 15 10 20 3 5 6 10 13 15 18 20 TIME IN DAYS Text-fig. 1. The effect of ACTH and cortisone on percentages of pituitary basophils. In A and B, solid lines indicate the males; broken lines indicate the females, both injected with ACTH. In C and D, solid lines indicate the ACTH-injected fish; broken lines indicate the saline-injected controls. Heavy lines indicate two injections per day; light lines indicate one injection per day. In E and F, solid lines indicate the males; broken lines indicate the females, both injected with cortisone. In G and H, the solid lines indicate the cortisone-injected fish; broken lines indicate the saline-injected controls. Heavy lines indicate the larger dose of cortisone; light lines indicate the smaller dose. Each point of A, B, E and F represents one fish killed at that time. Each point of C, D, G and H represents the mean of two fish killed at that time. Figure 6 shows the breakdown of the yolk in the mature eggs three days after the start of injec- tions of the larger doses of cortisone. Figure 7 shows the ovary eight days after the start of cortisone injections. Here the tissue seen be- tween the ova is mainly follicular epithelium and remnants of resorbed eggs and some con- nective tissue. Only mature eggs are resorbed and not all of these are affected. The effects of ACTH given twice daily are comparable with those of cortisone on the ovaries. The effects of one daily injection are of the same nature although of lesser degree. The testes of all the control animals show cysts of developing spermatogonia and lobules containing mature sperm suspended in fluid secreted by the sperm duct epithelium (Rasquin & Hafter, 1951b). Figure 8 is a photomicro- graph of the testis of a control. The effects of the experimental procedures can be seen in Figure 9. Cortisone, and to a lesser extent ACTH, stimulated the secretory activity of the sperm duct epithelium to such an extent that the duct and the lobules were distended with fluid. In such glands there was a decrease in the amount of spermatogenesis. In the normal re- productive cycle the amount of spermatogenesis observed in the testis appears to be correlated with the content of the lobules. In gonads in which the lobules are packed with mature sperm, spermatogenic activity is less than that observed when the lobules are more nearly empty. It is possible therefore that the decreased amount of spermatogenesis observed in the hormone- treated fish is related to the excess fluid within the lobules and not to a direct effect of hormone administration. Adrenal Cortex and Lymphoid Organs.— The cells of the anterior inter-renal tissue, which is homologous to the mammalian adrenal cortex (Rasquin, 1951), show no degranulation follow- ing any of the experimental techniques. When Masson’s trichrome staining method is used, the glandular tissue shows fuchsinophile granules, irregular in size and shape in all fishes whether experimental or control. However, both ACTH 1952] Rasquin & Atz: ACTH and Cortisone in a Teleost ACTH CORTISONE Text-fig. 2. The effect of ACTH and cortisone on thyroid epithelial heights. In A, solid lines indicate ACTH-injected fish; broken lines indicate saline-injected controls. Heavy lines indicate two injections per day; light lines indicate one injection per day. In B, solid lines indicate cortisone-injected fish; broken lines indicate saline-injected controls. The heavy line indicates the larger dose of cortisone; the light line indicates the smaller dose. Each point on the graph represents the mean of two fish killed at that time. injection schedules cause hypertrophy and hy- perplasia of the inter-renal tissue. Many mitotic figures were seen in the glandular cells, which spread out from their normal position next to the walls of the cardinal veins into the parenchy- ma of the head-kidney. One of these dividing cells is seen in Figure 10. After cortisone injec- tions hyperchromatic nuclei are seen in the inter-renal cells and some shrinkage is noted in the glands of the fishes killed early in the experi- mental period. The head-kidney is greatly depleted of lym- phoid elements after cortisone treatment. Many edematous spaces are evident as well as areas where the only cellular elements remaining are the reticular fibers of the stroma. There is some evidence of repair ten days after the last injec- tion when the lighter dose of cortisone was used, but no recovery has been initiated in the organ at the end of the experimental period when the larger dose of cortisone was used. The administration of ACTH causes a much less drastic reduction in the head-kidney. There is some loss of lymphocytes and appearance of some edematous spaces but this condition is completely repaired ten days after injec- tions were discontinued. The effect is more pro- nounced and the time before recovery is slightly longer in the fish injected twice a day than in those injected once a day. The lymphoid ele- ments of the head-kidneys of the fish that re- ceived the control saline solutions are also some- what affected, showing in still another way that the control injection schedule also produced a stressful condition. These organs in the control fishes, however, were promptly repaired at the end of the injection period. All the fish show a partly involuted thymus which is normal for this species at this age (Haf- ter, 1952). No further involution is noted as a result of any of the experimental procedures. Evidently the thymus, in this species of teleost at least, is extremely refractory to mammalian cortical hormones. A study of the corpuscles of Stannius, for- merly thought to contain part of the teleost cortical tissue, shows no change that can be attributed to any of the experimental proced- ures. Different degrees of cellular activity were noted and some glands showed a more vascular condition than others but no consistent corre- lations can be made. Discussion An unexpected result of this investigation is the difference in the reaction of the male and female pituitary to ACTH. Explanation of this effect may be provided if it is assumed, first, that the cortical hormone produced by the fish possesses androgenic activity similar to that ob- served in higher vertebrates; and, second, that in the female, endogenous estrogens antagonize this action so that the total inhibitory effect on the female pituitary is less than that observed 84 Zoologica: New York Zoological Society [37: 7: in the male where no such antagonism occurs. No sex difference was observed in the pituitary with either dosage of cortisone, and it seems likely that the mammalian cortical steroid was administered in a large enough dose or was sufficiently androgenic to over-ride any antago- nistic action of endogenous estrogens. The effects of strong masculinizing agents on the teleost gonads have been described by Ever- sole (1941) who treated male and female gup- pies, Lebistes reticulatus, with testosterone pro- pionate and pregneninolone. Early in the injec- tion period hypertrophy of the testicular ducts and precocious maturation of spermatogonia were produced. New development of spermato- gonia, however, was not initiated. Continued in- jection of these hormones left the testes of Lebistes “exhausted and degenerate.” As Ever- sole has also indicated, in the female, ovogene- sis and yolk deposition were effected only after prolonged treatment. In the treated mature fe- males the ovaries showed a return to a “juve- nile” condition with resorption of yolk and em- bryos. At the same time, administration of adrenal cortical extracts, of unreported origin, had no effects on the gonads of the same spe- cies. In the male Astyanax, the administration of ACTH and cortisone brought about the same changes reported by Eversole for his early results in the male guppy. Similarly, in the fe- male Astyanax , only already matured ova un- derwent resorption after treatment; immature eggs were not affected. Although early workers reported that ACTH stimulated the gonads and accessory sex or- gans, these results have been contradicted by investigators using purified hormones. Reviews of this material are given by Ingle (1950) and Baker, Schairer, Ingle & Li (1950). From the work of Ingle (1950) and Winter, Silber & Stoerk (1950), it is seen that cortisone admin- istered in small doses causes no significant change in the reproductive system of mature male rats. Massive doses of cortisone cause some regression of the testes and seminal vesi- cles of mature rats (Ingle, 1950) and mice (Antopol, 1950). Sprague (1951) suggested that the results on the gonads observed with large doses of cortisone may have been a gen- eral reflection of the catabolic effects of corti- sone rather than a specific effect on the repro- ductive tract. Similarly, Baker et al. (1950) sug- gested that a negative nitrogen balance result- ing from the metabolic effects of the C-ll oxygenated steroids released from the adrenals after stimulation with ACTH caused the regres- sive changes observed in the reproductive or- gans of mature rats after ACTH administration. The ACTH used for this investigation was contaminated only with small amounts of oxy- tocin. If contamination by gonadotropins was responsible for the effects on the gonads re- ported here, the same effects could hardly have been produced by cortisone. It seems most prob- able that, in this species of teleost, at least, the cortical hormones are androgenic. However, two other possibilities suggest them- selves. Rasquin & Hafter (1951a) have already intimated that the teleost may not react specifi- cally to mammalian ACTH, and perhaps this hormone acted gonadotropically as well as ad- renocorticotropically. The second possibility concerns a shift in production of tropic hor- mones by the pituitary such as has been pos- tulated by Selye (1946) under conditions of stress. The effects of ACTH and cortisone on the Astyanax reproductive system may have been caused by an increased production of pituitary gonadotropin associated with an inhibi- tion of adrenocorticotropin production. It is in- teresting to note here that Witschi (1939) has reported the luteinizing factor to be predomi- nant in teleost gonadotropin. The changes in the ovaries of Astyanax were certainly correlated with proliferation of follicular epithelium and the break-down of already matured ova. Although several investigators (Soffer, Ga- brilove & Dorrance, 1951; Money, Kraintz, Fager, Kirschner & Rawson, 1951; Antopol, 1950) have reported that ACTH and cortisone depress the activity of the thyroid in mammals, the mechanism by which this is achieved is not yet known. Money et al. believe that this effect is mediated through the pituitary, while Perry (1951) suggests that the action is a direct one on the thyroid. The results of this report indicate that the effect on the thyroid of Asty- anax is a direct one. The lowest thyroid epithe- lial heights, indicative of the least active glands, were not necessarily correlated with pituitaries having the most drastically reduced numbers of basophils. The injection of ACTH twice daily was more effective in reducing thyroid height but was no more effective than one daily injec- tion in changing the pituitary, and conversely, no differences were noted in thyroid response between the two dosages of cortisone, but the pituitaries of the two cortisone groups were statistically different from each other. Were the effect on the thyroid mediated through the pit- uitary, the responses of the two glands should have been consistent. Furthermore, there is no histological evidence of a thyrotropic hormone deficiency. In no ex- perimental group was there any evidence of complete exhaustion of all basophils, and it does not seem likely that the acidophils have a thy- rotropic function. The pituitary glands of some 1952] Rasquin & Atz: ACTH and Cortisone in a Teleost 85 of the fishes that showed extremely low thyroid epithelium had not yet arrived at a point where the acidophils were withholding secretion. In addition, although the thyroid epithelium of the controls injected twice per day was more depressed than those of controls injected once per day, the pituitaries of neither control group showed any cytological change in acidophils. Thus there is no consistent correlation between thyroid depression and masses of acidophils with retained secretion. Because the reciprocal function of the pit- uitary and adrenal is so well known (Sayers, 1950; Tuchmann-Duplessis, 1950, 1951; Ingle, 1950) and because an increase in numbers and secretion of acidophils accompanies ACTH and cortisone administration, it is tempting to as- sign adrenocorticotropic function to the acido- phils in this teleost. Thus the presence of exo- genous ACTH or increased levels of cortical steroids would inhibit the release of ACTH from the pituitary. However, that the acidophils do not elaborate ACTH is shown by the following: the basophils alone showed intense vacuolation and depletion as a result of the stress caused by the control saline injections; among the corti- sone-treated fishes, mitotic figures and other evidences of recovery were shown by the baso- phils after injections were discontinued; other types of stress, such as living in darkness (Rasquin, 1949) bring about activation and exhaustion of the basophils. Li & Evans (1947), Winter, Silber & Stoerk (1950), have shown that in rats there is a fun- damental antagonism between the adrenocor- ticotropic and growth hormones. Winter, Silber & Stoerk have said that the antagonism could be due to suppression of the release of the growth hormone or to suppression of its peri- pheral action. The acidophils of most vertebrates are generally credited with the elaboration of the growth hormone. If this should be true of the acidophils of the transitional lobe of the teleosts, their massed unreleased secretion after ACTH and cortisone administration would indicate that the seat of this antagonism is in the pitui- tary itself. Rasquin & Hafter (1951a) injected a lym- phosarcomatous fish and a control with the identical doses of ACTH used for this report. Only the tumorous fish showed inversion of the ratio of secretory cells in the hypophysis. These fish were much older than those used for the present report and their mean body weight was 7.8 grams. From the results obtained here it seems probable that the dose given the larger fishes was insufficient to cause the effect on the pituitary in spite of having an effect on the lym- phoid tissues. The inversion of the pituitary ratio in the tumorous fish was probably, there- fore, associated in some way with the tumorous condition. Summary 1. Ninety-six fish ( Astyanax mexicanus ( Filippi) ) were divided into six groups. Four groups received daily doses respectively of ACTH, two different amounts of cortisone ace- tate and control injections of physiological sa- line. Two groups received identical doses of ACTH and physiological saline divided into two injections per day. Injections were continued for ten days and two fish from each group were killed at similar intervals during the injection period and for ten days thereafter. 2. In the transitional lobes of the pituitaries the normal ratio of approximately 40% acido- phils to 60% basophils was inverted by ACTH in the males and in both sexes by cortisone. Both hormones lowered the percentages of basophils below that found in the controls. 3. A tendency to return to the normal ratio of secretory cells was noted statistically during the post-injection period in both ACTH groups, but with neither cortisone group, although some histological changes indicating repair were seen in the cortisone group. 4. ACTH and cortisone depressed the height of the thyroid epithelium. Double daily injec- tions of saline depressed the height of the thy- roid epithelium below that found in the single daily injected controls but were not as effective as either hormone. 5. Recovery of the thyroid was noted in the single daily injected ACTH group as well as in both control groups. With neither dosage of cortisone did the height of the thyroid epithelium return to that of the control fish during the post-injection period. 6. ACTH and cortisone stimulated secretory activity of the sperm duct epithelium in the testis; lobules were distended with fluid and spermatogenesis appeared to be depressed. In the ovary, both hormones induced resorption of mature eggs and hypertrophy and hyperplasia of follicular epithelium. 7. Both dosages of cortisone and two doses of ACTH per day were more effective in pro- ducing changes in the thyroid and gonads than was the single daily ACTH injection. References Antopol, W. 1950. Anatomic changes produced in mice treated with excessive doses of cortisone. Proc. Soc. Exp. Biol, and Med., 73: 262- 265. 86 Zoologica: New York Zoological Society [37: 7: Aronson, L. R. 1949. An analysis of reproductive behavior in the mouthbreeding cichlid fish, Tilapia macrocephala (Bleeker). Zoologica, 34: 133-158. Baker, B. L„ M. A. Schairer, D. J. Ingle & C. H. Li 1950. The induction of involution in the male reproductive system by treatment with adrenocorticotropin. Anat. Rec., 106 (3): 345-359. Eversole, W. J. 1939. The effects of androgens upon the fish ( Lebistes reticulatus) . Endocrinology, 25: 328-330. Golden, A. & P. K. Bondy 1952. Cytologic changes in rat adenohypophysis following administration of adrenocortico- trophin or cortisone. Proc. Soc. Exp. Biol, and Med., 79 (2): 252-255. Hafter, E. 1952. Histological age changes in the thymus of the teleost, Astyanax. Jour. Morp., 90 (3): 555-581. Ingle, D. J. 1950. The biologic properties of cortisone: a review. Jour. Clin. Endocrinol., 10: 1312- 1354. Li, C. H. & H. M. Evans 1947. The properties of the growth and adreno- corticotropic hormones. In Vitamins and hormones. Academic Press Inc., New York, 5: 198-227. Money, W. L., L. Kraintz, J. Fager, L. Kirschner & R. W. Rawson 1951. The effects of various steroids on the col- lection of radioactive iodine by the thyroid gland of the rat. Endocrinology, 48: 682- 689. Perry, W. F. 1951. The action of cortisone and ACTH on thyroid function. Endocrinology, 49 (2) : 284-288. Rasquin, P. 1949. The influence of light and darkness on thyroid and pituitary activity of the chara- cin Astyanax mexicanus and its cave de- rivatives. Bull. Amer. Mus. Nat. Hist., 93 (7): 501-531. 1951. Effects of carp pituitary and mammalian ACTH on the endocrine and lymphoid systems of the teleost, Astyanax mexi- canus. Jour. Exp. Zool., 117 (2): 317-358. Rasquin, P. & E. Hafter 1951a. Response of a spontaneous fish lympho- sarcoma to mammalian ACTH. Zoologica, 36 (3): 163-169. 1951b. Age changes in the testis of the teleost, Astyanax mexicanus. Jour. Morph., 89 (3): 397-408. Rawson, R. W. & P. Starr 1938. Direct measurement of height of thyroid epithelium. Arch. Int. Med., 38: 726-738. Sayers, G. 1950. The adrenal cortex and homeostasis. Physiol. Rev., 30 (3): 241-320. Selye, H. 1946. The general adaptation syndrome and the diseases of adaptation. Jour. Clin. Endo- crinol., 6: 117-230. Soffer, L. J., J. L. Gabrjlove & W. R. Dorrance 1951. Effect of adrenocorticotropin on thyroidal collection of I131 in the adrenalectomized and intact rat. Proc. Soc. Exp. Biol, and Med., 76: 763-765. Sprague, R. G. 1951. Effects of cortisone and ACTH. In Vita- mins and hormones. Academic Press Inc., New York, 9: 265-304. Tuchmann-Duplessis, H. 1950. L’action de la surrenale sur la structure et le fonctionnement de l’hypophyse du Rat. Bull. d’Hist. app. et de Tech. Micros., 27: 89-99. 1951. Modifications hypophyso - surrenaliennes du rat traite par la cortisone. C. R. Acad. Sci., 232 (17): 1610-1612. Van Dyke, D. C., M. E. Simpson, C. H. Li & H. M. Evans 1950. Survival in the circulation of the growth and adrenocorticotrophic hormones as evidenced by parabiosis. Amer. Jour. Physiol., 163 (2): 297-309. Winter, C. A., R. H. Silber & H. C. Stoerk 1950. Production of reversible hyperadrenocor- tinism in rats by prolonged administration of cortisone. Endocrinology, 47 (1): 60- 72. WlTSCHI, E. 1939. Quantitative determination of the follicle stimulating and luteinizing hormones of the hypophysis. Anat. Rec., 75 (suppl.): 58. 1952] Rasquin & Atz: ACTH and Cortisone in a Teleost 87 Fig. Fig. Fig. Fig. Fig. 5 Plate I. Section through the transitional lobe of the pituitary of a control fish, saline-injected once per day. Acidophils are small, dark, polygonal cells; basophils are lighter- staining vacuolated cells. Part of the pars nervosa appears at the bottom of the photograph. All pituitary sections were stained in Masson’s trichrome stain. X 850. Section through the transitional lobe of the pituitary of a control fish, saline-injected twice daily, showing increased vacuola- tion caused by the additional stress of the double injection schedule. X 850. Section through the transitional lobe of a cortisone-treated fish eight days after injections began. A basophil in anaphase is seen at left center. X 850. Section through the transitional lobe of the pituitary of a cortisone-treated fish, showing increased numbers of hyper- trophied acidophils. Compare with con- trol in Fig. 1. X 850. Plate II Section of ovary of control fish, showing immature and ripe ova. Hematoxylin and eosin. X 85. Section of ovary of a cortisone-treated fish three days after the start of injections, showing regressive changes in the yolk of mature ova. Hematoxylin and eosin. X 85. Section of the ovary of a cortisone-treated fish eight days after the start of injections, showing resorbed ova, hyperplastic fol- licular epithelium and connective tissue among normal ova. Hematoxylin and eosin. X 85. Fig. 8. Section through the testis of a control fish, showing ducts lined with epithelium and filled with ripe sperm. Cysts of developing sperm in various stages are seen mainly in the upper left quadrant of the photo- graph. Hematoxylin and eosin. X 85. Fig. 9. Section through the testis of a cortisone- treated fish, showing increase in seminal fluid and possible depression of sperma- togenesis. Hematoxylin and eosin. X 85. Fig. 10. Section through the inter-renal tissue of an ACTH-treated fish, showing hyper- trophy and hyperplasia of the cortical cells. A mitotic figure is seen in the lower right quadrant of the photograph. Hema- toxylin and eosin. X 85. EXPLANATION OF THE PLATES Fig. 6. Fig. 7. RASQUIN & ATZ PLATE I EFFECTS OF ACTH AND CORTISONE ON THE PITUITARY. THYROID AND GONADS OF THE TELEOST ASTYANAX MEXICANUS RASQUIN a ATZ PLATE II ■ f I. ' >.t ■ '■ , -v. , > * " ■V • 9 r-_t -cv ‘ IS#' - . # , ’®’l V $ tf s 1 ! - « # * # 7.,_ • . • » , \ * 4 i 10 > ' .-a* (f 1BH * ■ ; EFFECTS OF ACTH AND CORTISONE ON THE PITUITARY, THYROID AND GONADS OF THE TELEOST ASTYANAX MEXICANUS 8 The Effects of Holothurin on Fish, and Mice with Sarcoma 180 Ross F. Nigrelli New York Aquarium, New York Zoological Society EMERSON & Taft ( 1945) have reviewed the literature on the pharmacological activity of substances derived from mar- ine organisms. There is very little information, however, on the possible pharmacodynamic ac- tion of poisons from invertebrates (Taft, 1945). It has been found that the sea-cucumber, Actinopyga agassizi Selenka, from the Bahama Banks off Bimini1, produces a toxic agent which, in very high aqueous dilutions, will kill fish and other invertebrates. This preliminary note deals with the effects of this material (for which the name “Holothurin” is proposed) on fish, and on mice with Crocker mouse sarcoma 180. In earlier studies it was found that sea water suspensions of the various organs of the sea- cucumber were toxic to fish in general. This toxic material was more powerful in the fluid which was emitted during autotomy. Further analysis of the organs showed that the “pink gland” (tentatively identified as the Organ of Cuvier) contained the active toxic agent. This substance was used in the tests reported here. Twenty grams (wet weight) of the “pink gland” were suspended in 50 ml. of sea water in Erhlenmeyer flasks and used as a stock solution. Some of these suspensions were steril- ized by autoclaving. The test animals were as follows: Cyprinodon baconi Breder, a hardy killifish from the mangrove regions of South Bimini; Carapus bermudensis (?), the pearl fish which normally lives in the cloaca and within the respiratory tree of the sea-cucumber; and female Swiss-Webster mice trocared in the right axillary region with sarcoma 1802. 1 This investigation was carried out in the Lerner Marine Laboratory of the American Museum of Natural History, Bimini, Bahama Islands. The writer wishes to thank Dr. C. M. Breder, Jr., Director of the Laboratory, for the use of the facilities. Thanks are also extended to Mr. and Mrs. Marshall Bishop for their assistance. 2 The mice were supplied to the Lerner Marine Labo- ratory by the Sloan-Kettering Institute, New York City. The test fish were placed in large stacking bowls containing two liters of sea water, to which different amounts of stock solution were added. All experiments were carried out at room temperature (30° C.). A dilution of 1:100,000 of the stock solu- tion killed Cyprinodon baconi in 23 minutes. The time necessary to kill this hardy species varied with the dilution. A dilution of 1:1,000 killed pearl fish in eight minutes and 1:1,000,- 000 was lethal to this species after several hours. The mice were given subcutaneous injections of the sterilized material at the site of the tumor growth three days after implantation. One-half ml. of the stock solution killed a mouse in IVz hrs.; 0.25 ml. killed other mice in 3 and 4 hrs., respectively. The mice were able to sur- vive a dose of 0.05 ml. of the stock solution and of 0.25 ml. of the stock solution diluted 1:25 and 1:50 with sterile sea water. The sur- viving mice were sacrificed on the sixth day of tumor growth. Histological sections of the gills of the treated fish showed that death was due to a breakdown of the capillaries. Autopsies on the mice showed massive hemorrhages in the region of the injec- tion. In the mice which received the non-lethal doses, the sarcoma was considerably reduced in size and was necrotic. Histological sections are being prepared for a more detailed report to be published later, together with other pertinent data. Holothurin cannot be extracted with acetone, ether, chloroform or with higher ethanol; neither is it inactivated by these solvents, by 10% form- alin, 5N Hcl or 5N NaOH. The material appar- ently is water soluble, non-volatile and heat stable. Further, it decolorizes Lugol’s solution but not methylene blue. Additional studies on the biochemical and antitumorous properties of Holothurin, as well as of other products from invertebrates, are 89 90 Zoologica: New York Zoological Society [37: 8: 1952] under study by the present author in collabora- tion with Dr. Paul Zahl of the Haskins Labor- atory. References Emerson, George A. & Charles H. Taft 1 945. Pharmacologically Active Agents from the Sea. Texas Reports on Biology and Medi- cine, 3 (3) : 302-338. Taft, Charles H. 1945. Poisonous Marine Animals. Ibid. 3 (3): 339-352. 9 Sex Determination in Xiphophorus ( Platypoecilus ) maculatus. III. Differentiation of Gonads in Platyfish from Broods Having a Sex Ratio of Three Females to One Male Myron Gordon Aquarium, New York Zoological Society 1 (Plates I & II; Text-figure I) THE genetic mechanism for sex determi- nation in the common platyfish, Xipho- phorous ( Platypoecilus ) maculatus, is not uniform for all its members. For example, the members of natural platyfish populations from the rivers of Veracruz, Mexico, have a mecha- nism in which the female is homogametic with reference to the sex chromosomes (XX), and the male is heterogametic (XY). On the other hand, the platyfish from the Belize River, British Honduras, have a sex-chromosomal system in which the male is homogametic (YY) and the female is heterogametic (WY). Since platyfish from Mexico and those from British Honduras are of the same species, it has been possible to mate successfully members of the two geo- graphically isolated and genetically different natural populations in order to evaluate the re- sults obtained from combining the various sex chromosomes. This has been facilitated by use of marker genes for color patterns which are linked to specific sex chromosomes. Among the gametic recombinations that might be expected from the mating of an XX-XY type of platyfish with one having WY- YY, the union of chromosomes W and X is of special theoretical interest. This is particularly so because when a WX female is obtained, it may be tested by mating it to an XY male. Theoretically, the mating of a WX female and an XY male should produce three types of daugh- 1 From the Genetics Laboratory of the New York Zoological Society at the American Museum of Natural History, New York 24, N. Y. This work on the in- heritance of the macromelanophores is supported by a grant from the National Cancer Institute, National In- stitutes of Health of the U. S. Public Health Service, for the project: “Genetic and Correlated Studies of Normal and Atypical Pigment Cell Growth.” ters (WX, WY and XX) to every one male (XY). It is the purpose of this paper to present the experimental verification of the theoretical results expected, first, by documenting the ge- netic ratios obtained, and second, by showing that the histological evidence confirms the iden- tity of the sexes as first revealed by the genetic methods of analysis. The color patterns that serve as indicators of the genes on the various sex chromosomes are composed of large black pigment cells or mac- romelanphores. These pigment cells in geneti- cally imbalanced fishes are potentially capable of neoplastic growth (Gordon, 1951c). The data to be presented provide further evidence of the strong and definite genetic control of the pattern of development of these important pigment cells and the precise method of their inheritance. Material and Methods A word of explanation of the symbols used here for the sex chromosomes may be desirable. The formula XX representing femaleness and XY for maleness is familiar, for it has been used to denote the appropriate sexes in Drosophila or in man. The formula WY denoting female- ness and YY denoting maleness is less familiar. They represent a modification of the terms used in referring to the sex chromosomes of birds or moths in which WZ represents femaleness, and ZZ maleness. The reason for substituting the Y for the Z in the platyfish is that there is ample experimental proof that the Z chromo- some is synonymous with the Y. This has been demonstrated repeatedly (Gordon, 1946b, 1947a, 1951a) in the platyfish ( X . maculatus) by mating a homozygous female (XX) of the 91 92 [37: 9: Zoologica: New York Zoological Society XX-XY stock with a homozygous male (YY) of the YY-WY stock. When the parents are XX and YY, all the Fi individuals are XY and male. Historically our earliest knowledge of the sex- determining mechanism in the platyfish was de- rived from analyses made with members of domesticated strains which were originally taken from “Central America” and shipped to Ger- many, in 1907. A few years later, about 1911- 1912, the aquarium-bred platyfish were export- ed by German aquarists to the United States. The genetic mechanism for sex determination in the early domesticated platyfish was found by Bellamy (1922), and independently by Gor- don (1926, 1927), to be YY (or ZZ) for male- ness and WY (or WZ) for femaleness. This corresponds exactly to the system found in mem- bers of wild populations of the platyfish recent- ly obtained (1949) from the Belize River in British Honduras. And this suggests that our domesticated stocks originated most probably in the Belize River. It was the custom in the early days of the tropical fish hobby for sailors to bring in specimens for aquarists. Belize, in British Honduras, was a port of some importance in the early 1900’s for shipping logwood and mahog- any. Sailors doubling as fish collectors probably shipped platyfish from this locality to Germany. Females of the domesticated stocks of the platyfish (WY) and males of the wild stocks (XY) of the same species from the Rio Jamapa, Veracruz, Mexico, were used in the present analysis. The reaction of each of the various sex chromosomes was followed by “tagging” the Y, X and W by one of a series of dominant genes for macromelanophore patterns. The Sp ( spot- sided) gene was manifested by the presence of large black pigment cells scattered over the sides of the platyfish, the Sb ( spotted-belly ) gene by similar cells along the ventral surface of the body and the Sr (striped) gene by several rows of macromelanophores along the sides. The Sr gene in the presence of Sp or Sb was difficult to detect but this did not interfere with the critical analysis. Sp, Sb, Sr and their multiple recessive gene, +, are alleles. In this series of sex-linked alleles there are two additional genes, N and Sd, which were not used in the present study (Gor- don, 1948). Experimental Results from Genetic Analyses The domesticated spot-sided (Sp) platyfish, obtained in 1942 from a tropical fish supply house, were quite similar to those studied by Fraser & Gordon (1929). These authors first showed that the W chromosome may carry dominant genes and this may result in a matro- clinus type of inheritance. Additional proof for this was obtained in a series of experiments, cfne of which is as follows: Domesticated Platyfish (8B Stock) Female Parent Pi Male Parent Spot-sided N on-spotted (W)5p/(Y)+ (Y)+/(Y) + Fi: 20 Spotted Daughters, (W)Sp/(Y) + 25 Non-spotted Sons, (Y)+/(Y) + Inbred Fi produced F2: 10 Spotted Daughters, (W)Sp/(Y)4- 15 Non-spotted Sons, (Y)+/(Y) + Inbred F2 produced F3: 18 Spotted Daughters, (W)Sp/(Y) + 14 Non-spotted Sons, (Y) +/(Y) + The next step was to synthesize a strain in which the females had two dominant alleles, one on the W and a different one on the Y. This was accomplished by use of two domesticated stocks as follows: Female Parent (8B Stock) Spot-sided (W)Sp/(Y)Sp Pi Male Parent (8Sb Stock) Spotted-belly (Y)Sb/(Y)Sb 24 Fi (Pedigree No. 104) Daughters Sons (W)Sp/(Y)Sb 19 (Y)Sp/(Y)Sb The (W)Sp/(Y)Sb heterogametic female of the domesticated stock (No. 104) was then mated to an (X)5r/(Y)5r heterogamic male of the wild Rio Jamapa stock. Sr is more weakly expressed than the other two genes and this makes the detection of Sp and Sb relatively easy: Female Parent Pi Male Parent Domesticated (Stock Wild (Stock 303) No. 104) Rio Jamapa Spot-sided, Spotted- Stripe-sided belly (W)Sp/(Y)Sb X (X)Sr/(Y)Sr Fi (Pedigree No. 107) 75 Spot-sided 65 Spotted-belly Daughters: Sons: 1. (W)Sp/(X)Sr, or 3. (X) Sr/ (Y) Sb, or 2. (W)Sp/(Y)Sr 4. (Y)Sr/(Y)Sb Since all the spot-sided (Sp) Fi individuals were females it may be assumed that some were WX (marked “1”) with regard to their chromo- somal combinations, and approximately the same number were WY (“2”). Since all the spotted-belly (Sb) Fi fish were male it may be assumed that both XY (“3”) and YY (“4”) sex chromosome combinations represent male- ness. Phenotypically it is impossible to separate either the two genotypes representing the fe- 1952] Gordon: Sex Determination in Xiphophorus maculatus 93 males ( 1 and 2) or the two genotypes represent- ing the males (3 and 4). One of the spot-sided ( Sp ) females from Pedigree No. 107 was taken at random and mated to a striped (Sr) male of a known stock from the Rio Jamapa. The ex- periment may be outlined by indicating the pos- sible mating types as follows: Female (Stock 107) Pi Male (Stock 305) 1. (W ) Sp/ (X) Sr, or Rio Jamapa 2. (W)Sp/(Y)Sr X (X)Sr/(Y)Sr Fi (Pedigree No. 194) 13 Daughters: 9 Spot-sided (W )Sp/(X)Sr, or (W)Sp/(Y)Sr A Stripe-sided (X)Sr/(X)Sr 5 Sons: 5 Stripe-sided (X) Sr/ (Y) Sr The 3:1 sex ratio (13 females to 5 males) may be explained on the assumption that the female parent of the constitution (W )Sp/(X)Sr (listed as “1”) was the fish mated to the (X)Sr/ (Y )Sr male. This would produce the following three combinations of sex chromosomes: WX, WY and XX, all of which would determine femaleness. The remaining combination, XY, would determine maleness. (Text-fig. 1). The type of experiment just reported required further substantiation; it was therefore repeated several times. Two spot-sided (Sp) females of Pedigree No. 194 were selected and bred to their striped (Sr) male siblings. In presenting the results the parental genotypes, listed below as “A” and “B,” will be indicated for conven- ience in advance of the results obtained from evaluation of the Fi but, of course, the geno- types of each Pi female (194-1 and 194-2) were determined by the Fi data: “A” Spot-sided Female Pi Stripe-sided Male (194-1) (194-11) (W)Sp/(Y)Sr X (X)Sr/(Y)Sr Fi (Pedigree No. 195): 14 Spot-sided 16 Stripe-sided Daughters : Sons : (W)Sp/(X)Sr, or (X)Sr/(Y)Sr, or (W)Sp/(Y)Sr (Y)Sr/(Y)Sr The sex ratio was 1:1. “B” Spot-sided Female Pi Stripe-sided Male (194-2) (194-12) (W )Sp/(X)Sr X (X)Sr/(Y)Sr Fi (Pedigree No. 195a) : 1 8 Daughters: 12 Spot-sided (W) Sp/ (X) Sr, or (W)Sp/(Y)Sr 6 Stripe-sided (X)Sr/(X)Sr 8 Sons: 8 Stripe-sided (X)Sr/(Y)Sr The sex ratio was apparently 3:1. One final experiment was conducted in order to obtain sufficient numbers of fish in broods showing a sex ratio of three females to one male for a follow-up study of the gonadal develop- ment in the very young fishes of various pheno- types. The purpose of making this study was to see if the sex of an individual, as recognized by histological methods, corresponds with its phenotype. In other words, do the various chro- mosomal combinations determine sexuality of the fishes very early in their development? A spot-sided female (No. 195a-l), presum- ably (W )Sp/(X)Sr, of the last brood (Pedi- gree No. 195a), was mated to its stripe-sided male sibling (No. 1 95a- 11), which presumably was (X) Sr/ (Y) Sr. The results obtained were essentially the same as before, as follows: Fi (Pedigree No. 1952a) 93 Daughters: 59 Spot-sided, (W ) Sp/ (X) Sr, or (W)Sp/(Y)Sr 34 Stripe-sided, (X)Sr/(X)Sr 33 Sons: 33 Stripe-sided, (X)Sr/(Y)Sr From the above results the genotypes of the Fi female and male selected for mating must have been (W )Sp/(X)Sr and (X)Sr/(Y)Sr, respectively. Among their 1 26 Fi young were 93 females and 33 males, a good three to one ratio. The above data on sex ratios were obtained, as were all others given previously, by rearing the Fi fish to four to seven months of age, at which time their sex may be determined by inspection. Externally the sex of mature fish is easily recognized by the differences in the con- figuration of the anal fin of the male and female. In the male, the single median anal fin is trans- formed into a thin, grooved, rod-shaped organ or gonopodium which serves, at time of copu- lation, to transfer spermatophores to the genital aperture of the female. In the female, the anal fin is fan-like, hardly different from that of the young. Determination of Sex in Immature Fish by Histological Methods In addition to the 126 fish (Pedigree No. 1952a) reared to sexual maturity, 84 additional young were obtained in three later broods from the same parents (No. 195a-l and No. 195a-ll). Among the young, 40 were spot-sided and 44 were unspotted. They ranged in size from 9 to 20 mm; their sex could not be recognized with certainty because their anal fins were in an un- differentiated state. On the basis of their color patterns one could predict that the 40 spot-sided ones would turn out to be females, as would 94 Zoologica: New York Zoological Society [37: 9: XIPHOPHORUS (PLATYPOECiLUS) MACULATUS DOMESTICATED FEMALE RIO JAMAPA MALE SEX RATIO, 3 FEMALES: I MALE f0:' -rggPl, Wsp Ysr Ysr Ysr SEX RATIO, 1 TO 1 Text-fig. 1. Variable Sex Ratios in the Platyfish Xiphophorus ( Platypoecilus ) maculatus. When a spotted female of the domesticated stock WY is mated to a non-spotted (stripe-sided) platyfish from the Rio Jamapa wild stock XY, the sex ratio in the members of the Fi is 1:1 (lines 1 and 2). Some of the Fi females contain the sex chromosomes WX while others have WY (line 2). When a WX Fi female is mated to an XY male (line 3, left), their backcross offspring have a sex ratio of 3 females to 1 male (lines 4 and 5, left). When a WY Fi female is mated to an XY male (line 3, right), their backcross offspring have a sex ratio of 1: 1 (lines 4 and 5, right). about half of the 44 unspotted ones. The remain- ing half of the unspotted ones should be male. In order to determine the sex of the 84 im- mature fish by histological methods, they were fixed in Bouin’s fluid. They were decalcified, sectioned at 10 /x, and stained with Delafield’s hematoxylin and eosin. The Females Microscopic examination of the sectioned gonads of the 84 immature fish revealed that 64 of them, 40 spot-sided (Sp) and 24 unspotted (presumbably Sr), had ovaries and the remain- ing unspotted 20 fish (presumably Sr) had testes (Table 1). Thus the histological results con- firmed the theoretical expectancy of the three females to one male sex ratio, and in their ap- propriate phenotypic groupings: all the spotted ones were female; half the unspotted ones were female and half were male. A special analysis was made to determine the various stages of development of the ovaries in the spot-sided and unspotted females in order to evaluate the relative effectiveness of WX or WY as opposed to that of XX in establishing and regulating the sexual development of the females. A convenient method of rating the developing ovaries was worked out which was based upon (1) the size of the largest ovocyte in the ovary, and (2) the nature of the cyto- plasmic substances in the ovocytes with refer- ence to oil and yolk content. This method of analysis was derived from study of a large series of platyfish ovaries, summarized by Berg & Gor- don (1952) as follows: The ovary in the platyfish arises as a paired organ but fuses shortly after its birth or about the time it reaches a length of 7 to 12 mm. In young fish the epithelial cells of the ovarian duct are high columnar. Between these cells occa- 1952] Gordon: Sex Determination in Xiphophortis maculatus 95 Table 1. The Effect of Various Sex Chromosome Combinations on the Gonadal Development in Siblings of the Platyfish, Xiphophorus maculatus of Pedigree Number 1952a Number of Animals Standard Length-range Phenotype Type of Gonad Probable Genotype 40 12 to 19 mm Spot-sided1 Ovary (W )Sp/(X)Sr or (W)Sp/(Y)Sr 24 12 to 20 mm Not spot-sided3 Ovary (X)Sr/(X)Sr2 20 9 to 15 mm Not spot-sided3 Testis (X)Sr/(Y)Sr2 1 Sp gene for macromelanophores, producing spots on the sides. 2 Sr gene for macromelanophores, producing stripes along the sides. 3 In young fish the striped color pattern is hardly developed. This group was recognized by the absence of the spot-sided pattern. sional germ cells are found. In mature fish the ovary fills the major part of the body cavity. The single ovarian duct originates at the anterior- Jorsal part of the ovary. It remains dorsal and straight as it passes caudad; it then curves sharply ventrad in the posterior part of the ovary. Beyond the ovary, the ovarian duct con- tinues posteriorly straight to the urogenital sinus. In mature fish the part of the ovarian duct that lies within the ovary is lined with a single layer of low, cuboidal epithelium. Ovocytes in various stages of development and degeneration lie in the part of the ovary between the ovarian duct and peritoneal cover- ing. Each ovocyte lies within a follicle which is surrounded by a single-layered theca. A thin layer of well-vascularized stroma lies between the follicles. Oil droplets are seen in ovocytes that are approximately 250 p. in diameter, or greater. Yolk first appears in ovocytes that are 600 p. in diameter. A mature, unfertilized ovum measures 1,500 p. in diameter. Fishes having ovocytes measuring no more than 200 p. and containing neither oil nor yolk were classified as being in their earliest stage of ovarian development, or group I (Table 2). Those having ovocytes measuring no more than 600 p., and containing oil droplets only were classified as being in Stage II. Those having ovo- cytes measuring up to 1,200 p, and containing oil and yolk droplets were regarded as almost mature and placed in Stage III. In group I there were four females; in II there were 18; and in III there were 14. The fishes in each group were separated according to their phenotype, spotted or unspotted. Sta- tistical analyses of the relationship between the phenotype of the fish and the condition of their ovaries revealed no significant difference be- tween the spot-sided females and the unspotted ones. It can be inferred, therefore, that in the platyfish the WX or WY chromosome combina- tion has the same effect on the rate of develop- ment of the ovary as the XX chromosome com- plex. The Males Twenty unspotted immature platyfish of Pedi- gree No. 1952a, measuring 9 to 15 mm, studied histologically, had testes; they were classified according to their stage of spermatogenesis (Table 3). The state of development of the testes varied from the earliest to the latest. In none was there any evidence of atypical devel- opment of the gonad. The variation in the state of development may be attributed to the circum- stance that the group represented three differ- Table 2. Stage of Ovocyte Development in 36 Females Siblings of Xiphophorus maculatus with Varying Chromosomal Sex Determining Mechanisms State of Ovocytes Re: Cytoplasm Size of Ovocytes Number of Females Spot-sided Not Spot-sided1 (W) Sp/(X) Sr, or (X)5r/(X)5r (W)5p/(Y)Sr I No oil, no yolk 20 to 200 p 3 1 II Oil droplets 200 to 600 p 7 11 III Yolk droplets 600 to 1200 p 9 5 1 The Sr fish, as indicated in Table 1, may not show their appropriate stripe-sided color pattern, hence they are referred to as “Not spot-sided.” 96 Zoologica: New York Zoological Society [37: 9: Table 3. Stages of Spermatogenesis of 20 Male Siblings of Xiphophorus maculatus of Pedigree Number 1952a Stages1 Characteristics Number of Males 1 : Primordial Germ Cells Gonad primordia largely composed of stroma Primordial germ cells discrete or in small groups at periphery of testes 5 2: Spermatogonial Acini Few In each primordium some stroma cells form one large duct Germ cells (spermatogonia) at the periphery of gonad primordia; not more than 8 cells in small acini 2 3: Spermatogonial Acini Numerous Many branching ducts. Many spermatogonia in each of many acini 7 4: Primary Spermatocytes Acini contain primary spermatocytes. Comparative- ly few acini contain spermatogonia 2 5 : Secondary Spermatophores Cords of primary and secondary spermatocytes 0 6: Spermatophores Spermatophores present in sperm ducts and tubules 4 1 Stages as defined by Chavin & Gordon (1951). ently aged broods of fish. This, of course, was also true of the females from the same broods. Since the various stages in spermatogenesis of platyfish were presented by Chavin & Gordon (1951) in the first of this series of papers on sex determination, in this journal, they will not be reviewed here. Discussion It is a remarkable fact that two opposing ge- netic mechanisms for sex determination are found in a single species of platyfish, Platypoe- cilus maculatus (now known as Xiphophorus maculatus) according to Gordon & Rosen, 1951, and Gordon 1951b. The acceptance of this fact hinges upon this point: that some members of a species follow the WY female, YY male mech- anism, while other members of the same spe- cies follow the XX female, XY male system. Bellamy & Queal (1951) have raised the ques- tion as to whether these two groups really belong to the same species. They do. In papers by the author (1944a, b; 1946a, b; 1947a, b; 1950) the identity of both the wild and domesticated platyfish in question has been established and clearly stated as Platypoecilus maculatus. In this connection, it may be of interest to record the type of genetic mechanism for sex determination in the various wild and domesticated popula- tions that have been analyzed from time to time: Female Heterogamety (WY), Male Homo- gamety (YY) in Xiphophorus maculatus 1. Domesticated Stocks (Bellamy, 1922 to 1951; Kosswig, 1927 to 1939; Breider, 1936 to 1949; Gordon, 1926 to 1951). 2. Natural Populations from the Belize River (Gordon, 1950). Female Homogamety (XX), Male Heterogam- ety (XY) in Xiphophorus maculatus 1. Natural Populations from Rio Jamapa (Gordon, 1944a, b; 1946a, b; 1947a). 2. Natural Populations from Rio Papaloapan (Gordon, 1944a, b; 1946a, b; 1947a). 3. Natural Populations from Rio Coatzacoal- cos (Gordon, 1951a). Many reviewers, confronted with the unique fact that both male heterogamety and female heterogamety occur in closely related species of poeciliid fishes (or as in X. maculatus, in the same species), have, in a speculative mood, de- clared that the latter probably evolved from the former. This idea attained some degree of ac- ceptance owing to Winge’s (1934) discovery that in Lebistes reticulatus, which has the XX female, XY male system, it is possible by arti- ficial selection to get XX males and XY females. Gordon (1951a) pointed out the fallacy of the conclusion based upon this sort of evidence, because when exceptional XX male guppies are mated with equally rare XY females, XX fe- males and XY males are again produced. These results might be expected because in the off- spring of sex-reversed parents, the balance between the genes for sex determination in sex chromosomes and the sex genes in autosomes is restored. A somewhat similar example of the genic imbalance in a sexually abnormal male platyfish was described in a prior paper in this series (Gordon & Aronowitz, 1951). A partially 1952] Gordon: Sex Determination in Xiphophorus maculatus 97 functional male from Rio Jamapa stock that had the two X chromosomes usually character- istic of a female was found by Gordon ( 1947a) . When the exceptional XX male was mated to a normal XX female of a similar stock, all their offspring were normal females, XX, as one would expect on theory. These sex-reversed individuals are important in establishing the type of sex-determination in the members of a particular population but they have little bear- ing upon the evolution of the system itself. Bellamy & Queal (1951) have joined others in concluding that in the maculatus platyfish, female heterogamety “evolved from male heter- ogametic forms . . . if . . . male heterogametic P. maculatus now exist in certain Mexican river systems.” The facts, of course, are that both sex-deter- mining systems are represented in isolated natural populations of the maculatus platyfish. It is more likely that each specialized system (WY-YY and XX-XY) arose independently from an undifferentiated condition, perhaps like the one which exists in the related species, X. hellerii. The chief problem is to determine how the force of natural selection has established the XX female, XY male system in the natural populations of the maculatus platyfish in three rivers of Mexico, while at the same time it has molded the WY female, YY male system in the Belize River population of the same platyfish species. Perhaps an analysis of the distribution of the frequences of five sex-linked and seven autosomal genes in the various natural popula- tions of X. maculatus may throw some light upon this problem (Gordon & Gordon, unpub- lished). From the experimental results indicated in this paper the ratio of three females to one male was obtained by the mating of a “domes- ticated”—“wild” hybrid female, WX, to a Rio Jamapa, XY, male. A similar three to one sex ratio was obtained by Breider (1942) and re- cently by Bellamy & Queal (1951) by mating exceptionally rare genetically sex-reversed “do- mesticated” male platyfish (WY) to normal “domesticated” females (WY). These authors obtained from such matings: three female off- spring (1 WW, 2 WY) to one male (YY). Breider did not test the WW females, but Bellamy & Queal did. When WW females were mated to normal YY males, they produced all female platyfish, WY. When WW females were mated to genetically sex-reversed males WY, they, too, produced all female broods, WY and WW. To understand how an individual with the genetic constitution of a female (XX in the case of the Rio Jamapa stock) is able to function as a male, the gonad of a genetically sex-reversed XX male was studied by Gordon & Aronowitz (1951). The gonad was abnormal at the time it was examined, but it had been functional for a short period. Bellamy & Queal (1951) found that their exceptional WW females were mostly sterile, and that some had significantly fewer offspring. These details indicate an ab- normally functioning gonad. They did not re- port any follow-up histological studies of the gonads in any of their fishes. It is interesting that both in the Rio Jamapa stock (XX female, XY male) and in the “do- mesticated” stock (WY female, YY male) the genetic sex reversals discovered were males only. These exceptional males had the chromosomal constitution of the appropriate type of female, that is, XX male in the former stock and WY male in the latter. Stated another way, no XY females were discovered in the Rio Jamapa stock or YY females in the “domesticated” stocks. (Bellamy & Queal’s exceptional females, which he labels “YY” are equivalent to “WW” accord- ing to the terms used throughout this paper; WW females are, of course, not sex reversals, but the offspring of a genetically sex-reversed male WY and a normal WY female). Anyone assigned the task of reviewing the work on genetic mechanisms for sex deter- mination in the maculatus platyfish faces a for- midable and contradictory array of terms which refer to the various sex chromosomes. For this reason a list of synonyms used by Bellamy, Kosswig, Breider, Castle and Gordon is given in Table 4. It will be noticed from Table 4, that accord- ing to Gordon’s terminology, the “domes- ticated” and the Belize River stock (items 5 and 6) have identical sex-determining mechan- isms. This suggests that the “domesticated” stocks may be descendants of fishes originally taken from the Belize River in British Honduras. The history of the “domesticated” platyfish as traced by Gordon (1927) contains nothing which could be used against such a supposition. Gordon indicated that the platyfish most gene- rally studied in the past by geneticists in Europe and in this country were unquestionably those derived from an importation made in 1907 to Germany for aquarium purposes from some unspecified locality in Central America. In 1912, or soon thereafter, the German aquarium- bred platyfish were imported into the United States. The American derivatives of the German stocks of aquarium-bred platyfish were the only ones available to geneticists up to 1932. Bellamy, Kosswig, Breider and Gordon, who analyzed the sex-determining mechanism in the aquarists’ 98 Zoologica: New York Zoological Society [37: 9: Table 4. Sex-chromosome Terminology in the Platyfish, Xiphophorus maculatus Female Male Author Beginning Stock 1. XY XX Bellamy, Breider 1922- Domesticated 2. WZ zz Gordon, Kosswig, Breider, Bellamy 1926- Domesticated 3. XY YY Castle 1936 Domesticated 4. XY YY Gordon 1946 Domesticated 5. WY YY Gordon 1947- Domesticated 6. WY YY Gordon 1950- Belize River 7. XX XY Gordon 1947- Rio Jamapa, Rio Papaloapan, Rio Coatzacoalcos platyfish, all found that the male was homo- gametic and the female heterogametic. In two expeditions in 1930 and 1932, Gordon (1940) was successful in bringing living platy- fish of all known species ( couchianus , xiphi- dium, variatus and maculatus ) directly from Mexico to the United States. To distinguish the two stocks of maculatus, the early laboratory- bred platyfish were called “domesticated,” while those brought directly from Mexico to the United States were designated as “wild” stock. Now, however, since there are seven “wild” platyfish populations, each from a different river system, each stock is called by the river of its origin. A study of gene frequencies dis- tributions (Gordon & Gordon, unpublished) of the sex-linked and autosomal patterns has re- vealed the fact that the Belize River population contains all the color patterns represented by the 1907-1909 importation of platyfish. The platyfish population from the Rio Jamapa at Veracruz and the one from Rio Coatzacoalcos at Puerto Mexico contain genes that were not found in the early German imported platyfish, making it even more unlikely that the fish came from the latter two areas. If the assumption that the “domesticated” platyfish originally came from the Belize River population is valid, and all the available evi- dence points to this probability, it is no longer necessary to pay any further attention to Gor- don’s (1947a) former speculation that in the platyfish the heterogamic female system of sex determination was derived from the heteroga- metic male system by a process that involved hybridization with the swordtail. In this con- nection, Bellamy & Queal (1951) also point out that obtaining heterogametic males (WY) and homogametic females (WW) in domes- ticated stocks does not provide the elements necessary for the reconstruction of the opposite sex-determining system. This is so because when a WY sex-reversed male is mated to a WW fe- male, all the offspring, both WY and WW, are female. Again, the genetic balance between the sex-linked and autosomal genes for femaleness is restored. The genic balance theory of sex determination originally proposed by Bridges and later elabo- rated for fishes by Winge has merit in explain- ing the sex reversals reported by Bellamy & Queal, as well as those previously found by Breider and by Gordon. Under normally balanced conditions of sex genes in autosomes and in sex chromosomes, fe- maleness is determined ( 1 ) by XX in wild platy- fishes from three Mexican rivers, (2) by WY in those from Belize River, and (3) by WX in some intervarietal hybrids. Maleness is deter- mined (1) by XY in wild Mexican platyfishes and (2) by YY in those from British Honduras. The three female to one male sex ratio is obtained in intervarietal platyfish hybrids not as a result of sex reversals but rather of the orderly recombinations of appropriate sex chromo- somes. The differentiation of the sexes as re- vealed by histological sections indicates normal processes in action. Platyfishes with WY, WX and XX chromosomes are females, and those with XY are males as early in development as may be determined by histological techniques. Summary Members of platyfish ( Xiphophorus macu- latus) populations from the Mexican rivers Jamapa, Papaloapan and Coatzacoalcos, have a genetic mechanism for sex determination in which XX represents femaleness, XY maleness. Members of the same species from the Belize River in British Honduras and those from “do- mesticated” stocks have a mechanism in which WY represents femaleness and YY maleness. When a domesticated or British Honduras female platyfish, WY, is mated to a Mexican male, XY, two types of hybrid females, WY and WX, are produced as well as two types of hybrid males, XY and YY. The sex ratio is one to one. When an intervarietal hybrid female of the constitution WY is backcrossed to a Mexican 1952] Gordon: Sex Determination in Xiphophorus maculatus 99 male XY, the sex ratio remains one to one, being the same as from the original mating. When, however, an intervarietal hybrid fe- male of the constitution WX is backcrossed to a Mexican male XY, the sex ratio is three fe- males (WY, WX, XX) to one male (XY). Histological studies of the differentiating gonads in WY, WX and XX fishes revealed normal ovarian elements. In XY fish, normal testicular features were present. No evidence Was found for sex reversals in any intervarietal or backcross hybrids in the present series of experiments. Acknowledgments I am indebted to Dr. Olga A. Berg for studying the histological sections of the gonads of 84 immature fishes and classifying them for sex. I also thank her and Mr. James W. Atz for read- ing the manuscript. The photographs were taken by Mr. Sam C. Dunton. Bibliography Bellamy, A. W. 1922. Sex-linked inheritance in the teleost, Platy- poecilus maculatus. Anat. Rec., 24: 419- 420. 1936. Interspecific hybrids in Platypoecilus, one species ZZ-ZW, the other XY-XX. Proc. Nat. Acad. Sci., 22: 531-535. Bellamy, A. W., & Marion L. Queal 1951. Heterosomal inheritance and sex determi- nation in Platypoecilus maculatus. Genet- ics, 36(1): 93-107. Berg, Olga, & Myron Gordon 1952. The relationship of atypical pigment cell growth to gonadal development in hybrid fishes. Pigment Cell Growth (In press). Breider, Hans 1936. Eine Allelenserie von Genen verschied- ener Arten. Zeit. induk. Abstamm. Ver- erbgsl. 72: 80-87. 1942. ZW-Mannchen und WW Weibchen bei Platypoecilus maculatus. Biol. Zentralblat. 62: 187-195. 1949. Der Geschlechtsbestimmungsmechanis- mus in der Artdifferenzierung. Biol. Zen- tralblat, 68(1/2): 32-49. Castle, W. E. 1936. A simplified explanation of Bellamy’s ex- periments concerning sex determination in tropical fishes. Proc. Nat. Acad. Sci., 22: 679-682. Chavin, Walter, & Myron Gordon 1951. Sex determination in Platypoecilus mac- ulatus. I. Differentiation of the gonads in members of all-male broods. Zoologica, 36(2): 135-145. Fraser, A. C., & Myron Gordon 1929. The genetics of Platypoecilus. II. The link- age of the two sex-linked characters. Genetics 14: 160-179. Gordon, Hugh & Myron Gordon Mss. Speciation in fishes. II. Distribution of seven autosomal and five sex-linked genes in the platyfish from seven Central Ameri- can rivers. Gordon, Myron 1926. Melanophores of Platypoecilus, the top- minnow of geneticists. (Abstract) Anat. Rec., 34: 138. 1927. The genetics of a viviparous top-minnow Platypoecilus; the inheritance of two kinds of melanophores. Genetics, 12: 253-283. 1937. Genetics of Platypoecilus. III. Inheritance of sex and crossing over of the sex chro- mosomes in the platyfish. Genetics, 22: 376-392. 1940. The fish that was jungle bom and city bred. Natural History, 45: 96-106. 1944a. The XY sex-chromosomes determine maleness in wild populations of Platypoe- cilus maculatus while WZ determine fe- maleness in domesticated breeds of the same species. (Abstract). Rec. Genetics Soc. Amer., 13: 18-19. 1944b. Similar sex-determining mechanisms in wild populations of Platypoecilus xiphi- dium, variatus and maculatus. (Abstract). Rec. Genetics Soc. Amer., 13:19. 1946a. Genetics of Platypoecilus maculatus. Sex determining mechanism in two wild pop- ulations of the Mexican platyfish. (Ab- stract). Genetics, 31: 641. 1946b. Interchanging genetic mechanisms for sex determination in fishes under domestica- tion. Jour. Heredity, 37: 307-320. 1947a. Genetics of Platypoecilus maculatus. IV. The sex determining mechanism in two wild populations of the Mexican platyfish. Genetics, 32: 8-17. 1947b. Speciation in fishes. Distribution in time and space of seven dominant multiple alleles in Platypoecilus maculatus. Ad- vances in Genetics, 1: 95-132. 1948. Effects of five primary genes on the site of melanomas in fishes and the influence of two color genes on their pigmentation. In “The Biology of Melanomas.” Spec. Publ. N. Y. Acad. Sci., 4: 216-268. 1950. Genetics and speciation in fishes. Amer. Philo. Soc. Year Book 1949: 158-159. 1951a. Genetics of Platypoecilus maculatus. V. Heterogametic sex determining mechan- ism in females of a domesticated stock originally from British Honduras. Zoolog- ica, 36(2): 127-134. 100 Zoologica: New York Zoological Society [37: 9: 1952] 1951b. Platypoecilus now becomes Xiphophorus. The Aquarium, 20 (11): 277-279. 1951c. Genetic and correlated studies of normal and atypical pigment cell growth. Growth, Symposium 10: 153-219. Gordon, Myron, & Olga Aronowitz 1951. Sex determination in Platypoecilus mac- ulatus. II. History of a male platyfish that sired all-female broods. Zoologica, 36(2) : 147-153. Gordon, Myron & Donn E. Rosen 1951. Genetics of species differences in the mor- phology of the male genitalia in xipho- phorin fishes. Bull. Amer. Mus. Nat. Hist., 95(7): 409-464. Kosswig, Curt 1927. Uber bastarde der Teleostier Platypoe- cilus und Xiphophorus. Zeit. induk. Ab- stamm. Vererbgsl., 44: 253. 1939. Die Geschlechsbestimmungsanalyse bei Zahnkarpfen. Rev. Fac. Sci. Univ. Istan- bul, 4: 239-270. Winge, O. 1934. The experimental alteration of sex chro- mosomes into autosomes and vice versa, as illustrated by Lebistes. Compt. Rend. Travaux Lab. Carlsberg Ser. Physiol., 21(1): 1-49. EXPLANATION OF THE PLATES Plate I Fig. 1. The spotted platyfish, Xiphophorus (Platy- poecilus) maculatus of the domesticated stock 8B. Above, male (Y)Sp(Y)Sp. Be- low, female (W)Sp/(Y)Sp. Fig. 2. The stripe-sided platyfish of the same spe- cies, but of the wild stock from the Rio Jamapa, Veracruz, Mexico. Above, male (X)Sr/ (Y)Sr. Below, female ( X)Sr / (X)Sr. Plate II Fig. 1. Above, spot-sided female platyfish of the genetic constitution (W )Sp/(X)Sr, Pedi- gree No. 195a-l. Below, its mate, a stripe- sided male, (X)Sr/(Y)Sr, Pedigree No. 195a-ll. They produced the young shown in Figure 2. Fig. 2. Members of brood 1952a produced by the pair of platyfish shown in Figure 1. The following frequencies of color patterns and sexes were observed: left, 1 stripe-sided female (X)Sr/(X)Sr; center, 2 spot-sided females (W )Sp/(X)Sr, or (W)Sp/(Y)Sr; right, 1 stripe-sided male (X)Sr/(Y)Sr. Sex ratio: three females to one male. GORDON PLATE I FIG 1 FIG. 2 SEX DETERMINATION IN XIPHOPHORUS (PLATYPOECILUS) MACULATUS. Ill GORDON PLATE II FIG. 1 FIG. 2 SEX DETERMINATION IN XIPHOPHORUS (PLATYPOECILUS) MACULATUS. Ill 10 Four New Species of Geometridae (Moths) from Rancho Grande, North-central Venezuela1 D. S. Fletcher Department of Entomology, British Museum ( Natural History) (Plate I; Text-figures 1-7) [This is one of a series of papers resulting from the 45th, 46th and 47th Expeditions of the De- partment of Tropical Research of the New York Zoological Society, made during 1945, 1946 and 1948, under the direction of Dr. William Beebe, with headquarters at Rancho Grande in the Na- tional Park of Aragua, Venezuela. The expeditions were made possible through the generous coopera- tion of the National Government of Venezuela and of the Creole Petroleum Corporation. [The characteristics of the research area are in brief as follows: Rancho Grande is located in north- central Venezuela (10° 21' N. Lat., 67° 41' W. Long.), 80 kilometers west of Caracas, at an eleva- tion of 1,100 meters in the undisturbed montane rain forest which covers this part of the Caribbean range of the Andes. The migration flyway of Portachuelo Pass, which is also the water-shed between the Ca- ribbean and Lake Valencia, is 200 meters from Rancho Grande. Adjacent ecological zones include seasonal forest, savanna, thorn woodland, cactus scrub, the fresh-water lake of Valencia and various marine littoral zones. The Rancho Grande area is generally subtropical, being uniformly cool and damp throughout the year because of the prevalence of the mountain cloud cap. The dry season extends from January into April. The average humidity dur- ing the expeditions, including parts of both wet and dry seasons, was 92.4%; the average temperature during the same period was 18° C; the average annual rainfall over a five-year period was 174 cm. The flora is marked by an abundance of mosses, ferns and epiphytes of many kinds, as well as a few gigantic trees. For further details see Beebe & Crane, Zoologica, Vol. 32, No. 5, 1947.] THE Geometridae described as new in this paper are four of five species collected by Dr. William Beebe of the New York Zoological Society. The fifth species was a single female of Eupithecia purpureoviridis Warren, known previously only from the unique type which came from Paramba in Ecuador. All five species were taken at Rancho Grande near Maracay, Venezuela, and were sent for study to the Department of Entomology of the British Museum (Natural History). The types, unless otherwise indicated, are in the British Museum (Natural History). The color names are taken from Ridgway’s “Color Standards and Color Nomenclature” and hence are capitalized in the text. 1 Contribution No. 920, Department of Tropical Re- search, New York Zoological Society. Dr. Beebe informs me that these moths formed part of the great insect migration through Portachuelo Pass at Rancho Grande2. GEOMETRINAE Racheospila beebei, new species Text-figures 3, 4; Plate I, figure 2 Description of the Holotype.— Expanse of male holotype 17 mm. Palpus Pale Pinkish Buff, equal in length to half the diameter of the eye. Antenna bipectinate, the pectinations four times as long as the diameter of the shaft, the upperside of which is White. Face Pale Pink- ish Buff; between the antennae is a bar of White. Head, thorax and abdomen Pale Olivine, the abdominal segments edged posteriorly with White. The wings are Pale Olivine, the costa is 2 101 Zoologica, 36 (4): 243-254 (1951). 102 Zoologica: New York Zoological Society [37: 10: Text-figs. 1-7. 1. Melanoptilon collinsi, new species. Valves. 2. Melanoptilon collinsi, new species. Aedeagus. 3. Racheospila beebei, new species. Valve. 4. Racheospila beebei, new species. Aedeagus. 5. Melanoptilon collinsi, new species. Female genitalia. 6. Oospila zamaradensis, new species. Valves. 7. Oospila zamaradensis, new species. Aedeagus. (All drawings standard magnification, about X 45). 1952] Fletcher: New Geometridae (Moths) from Venezuela 103 lightly irrorate with Drab, the sinuous postme- dial fascia is White and the fringes are Chalce- dony Yellow; the cell spots are Fuscous and well marked on both wings. The undersides of both wings are White and glossy; the anterior half of the forewing is irrorate with Drab; the cell spots are minute and Fuscous. Male genitalia weakly sclerotised. Uncus broadly rounded; strong hair tufts arise from the dorsal side of its base. Valve with a flap on the basal half of the distal margin. Aedeagus with a lightly sclerotised patch near the apex. Vesica with two scobinate cornuti in the apical half. Material.— Holotype, male No. 481604, Vene- zuela, Rancho Grande near Maracay, (W. Beebe). Oospila zam arad aria, new species Text-figures 6, 7; Plate I, figure 1 Description of the Holotype.— Expanse of the male holotype 22 mm. Palpus equal in length to the diameter of the eye. Basal two-thirds of an- tenna bipectinate, the length of the pectinations shortening basad and apicad; at their greatest length the pectinations are ten times as long as the diameter of the shaft, which is Fuscous. Face and head Drab; thorax and first abdominal seg- ment Peacock Green, remaining abdominal segments Drab. On each of the four abdominal segments from two to five there is a glossy, Fus- cous-Black crest and there is a similar but mi- nute crest on segment six. The wings are Peacock Green with a broad sinuous Fuscous band at the distal margin. On the forewing the cell spots and the costa are Fuscous; on the hindwing the cell spots are wanting. The ground color of the underside is Pale Glaucous-Green; the pattern differs from the upperside only on the hindwing, where the Fus- cous band at the distal margin is weak and narrowed. The wing pattern of this species is reminiscent of the African genus Zamarada (Ennominae). Male genitalia with uncus short and truncate. Socii well developed and extending beyond the uncus. Juxta bilobed. Valve truncate, the outer margin folded inwards; the costa is produced beyond the truncate apex of the valve and broad- ly rounded, the inner surface toothed apically. The aedeagus is tapered apically and tipped with a single tooth. The vesica has one cornutus, a single spine equal in length to the width of the aedeagus. Material.— Holotype, male No. 481605, Vene- zuela, Rancho Grande near Maracay, (W. Beebe) ; Paratype, male No. 481616, Venezuela, Rancho Grande near Maracay, (W. Beebe). ENNOMINAE Melanoptilon collinsi, new species Text-figures 1, 2, 5; Plate I, figures 4, 5 Description of the Species.— Expanse of male and female types 24-30 mm. Palpus twice as long as the horizontal diameter of the eye, which is ovate. Male antenna bipectinate almost to the apex. At their longest the pectinations, which decrease in length apicad, are four times as long as the diameter of the shaft. Female antenna simple. Palpus, head, thorax and dorsal half of the abdomen Fuscous-Black. Face, pectus and ventral half of the abdomen White. Wings Fuscous-Black with two Orange patches on the forewing and one on the hindwing. The underside of the forewing is similar to the upperside, but with the costal edge White for three-quarters of its length and there is a White spot at the distal margin between veins M3 and Cui. The basal third of the underside of the hindwing is White, the Orange patch of the upperside is marked in White and there are two White spots near the apex, one be- tween the costal vein and Sc, the other between veins Sc and Mi; there is a further White patch extending to the distal margin between veins M3 and Cut. The remainder of the wing is Fuscous- Black and in the White areas the Fuscous-Black veins contrast sharply. Male genitalia with uncus simple and tapered. Apex of gnathos spatulate and spinose. Furca one half as long as the valve, the apical half spinose on the inner surface. Valve simple and partially sclerotised, rounded at the apex. Apical half of the aedeagus twice as broad as the basal half, except at the tip, which is tapered. Vesica slightly scobinate. Female genitalia with genital plate almost horse-shoe shaped. Posterior half of the bursa copulatrix cylindrical, the anterior half globular and the whole is membranous. The single sig- num, in the anterior half of the bursa copula- trix, is discoid with three stout spines on the inner surface. Most closely related to M. simulans Walker (1854, List. Lep. Ins. B. M., 2:373) and M. timidaria Herrich-Schaffer (1856, Samml. aus- sereurop. Schmett., 49, pi. 94: 538, 539). Material.— Holotype, male No. 481607; Al- lotype, female No. 481608; paratypes, 31 males, 7 females, Nos. 481609 to 481647 inclusive. Taken from May 21 to luly 6 at Rancho Grande near Maracay, Venezuela. Of this series, 16 male and 4 female paratypes (Nos. 481628- 481647) are in the collection of The American Museum of Natural History. 104 Zoologica: New York Zoological Society [37: 10: 1952] Nelo glaucata, new species Plate I, figure 3 Description of the Holotype.— Expanse of fe- male holotype 36 mm. Head, thorax and abdo- men Fuscous-Black; basal half of the patagia Orange. Wings Fuscous-Black; in the center of the posterior half of the forewing is an irides- cent blue-green spot, extending in diameter from the inner margin to the lower median. The underside of the forewing is Fuscous-Black, except for the costa and for a broad band at the distal margin, which are Drab. At the base of the hindwing is an Orange spot; the remain- der of the wing is Drab, with the Fuscous-Black veins contrasting sharply. Most closely related to N. coelisigna Walker (List Lep. Ins. B. M., 2:384), from which it differs in the color of the underside and the differently colored and much reduced iridescent spot on the upperside of the forewing. Material— Holotype, female No. 481648, Venezuela, Rancho Grande near Maracay, July 21, 1946. (W. Beebe). EXPLANATION OF THE PLATE Plate I Fig. 1. Oospila zamaradensis, new species. Fig. 2. Racheospila beebei, new species. Fig. 3. Nelo glaucata, new species. Fig. 4. Melanoptilon collinsi, new species. Upperside. Fig. 5. Melanoptilon collinsi, new species. Underside. (Plate figures twice natural size). FLETCHER PLATE I FOUR NEW SPECIES OF GEOMETRI DAE (MOTHS) FROM RANCHO GRANDE, NORTH-CENTRAL VENEZUELA NEW YORK ZOOLOGICAL SOCIETY GENERAL OFFICE 30 East Fortieth Street, New York 16, N. Y. PUBLICATION OFFICE The Zoological Park, New York 60, N, Y. VICE-PRESIDENTS Alfred Ely Laurance S. Rockefeller Donald T, Carlisle SECRETARY TREASURER Harold J. O’Connell Cornelius R. Agnew OFFICERS PRESIDENT Fairfield Osborn SCIENTIFIC STAFF: Zoological Park and Aquarium John Tee-Van Director Leonard J. Goss Assistant Director ZOOLOGICAL PARK Robert M. McClung . . Acting Curator, Mammals and Birds Grace Davall. Assistant Curator Mammals and Birds James A. Oliver. .... Curator of Reptiles Leonard J. Goss Veterinarian Charles P. Gandal. . . Assistant Veterinarian John V. Quaranta. . .Research Associate, Animal Behavior Lee S. Crandall. .... General Curator Emeritus William Beebe Honorary Curator, Birds AQUARIUM Christopher W. Coates. Curator & Aquarist James W. Atz Assistant Curator Ross F. Nigrelli ...... Pathologist Myron Gordon Geneticist C. M. Breder, Jr Research Associate in Ichthyology Homer W. Smith. . . . .Research Associate in Physiology GENERAL William Bridges Editor & Curator, Publications Sam Dunton ....... Photographer DEPARTMENT OF TROPICAL RESEARCH Jocelyn Crane Assistant Director Henry Fleming ....... Entomologist John Tee-Van . .Associate William K. Gregory .Associate William Beebe ........ Director Emeritus AFFILIATES C. R. Carpenter. .... Co-ordinator, Animal Behavior Research Programs James R. Simon Director, Jackson Hole Wildlife Park SCIENTIFIC ADVISORY COUNCIL A. Raymond Dochez Caryl P. Haskins Alfred E. Emerson K. S. Lashley W. A. Hagan John S. Nicholas EDITORIAL COMMITTEE Fairfield Osborn, Chairman James W. Atz Lee S. Crandall William Beebe Leonard J. Goss William Bridges James A. Oliver Christopher W. Coates John Tee-Van *:Aa — — - ■ . %pj / I ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME 37 • PART 3 • OCTOBER 31, 1952 • NUMBERS 11 AND 12 PUBLISHED BY THE SOCIETY The ZOOLOGICAL PARK, New York Contents Page 11. The Mutillidae (Wasps) of British Guiana. By Clarence E. Mickel. . 105 12. A Revision of the Fishes of the Subfamily Alfarinae in the Family Poeciliidae. By Donn Eric Rosen. Text-figures 1-10 151 11 The Mutillidae (Wasps) of British Guiana12 Clarence E. Mickel University of Minnesota Introduction THE Department of Tropical Research of the New York Zoological Society, under the direction of Dr. William Beebe, made extensive and intensive collections of in- sects in a small area near its station at Kartabo, British Guiana, over a number of years. Some ten years ago their collections of Mutillidae were sent to the writer for determination with the request that a manuscript be prepared setting forth the results. Later when it was pointed out that a number of other collections of Mutil- lidae from British Guiana were available for study, permission was given to include this material and to make the report as comprehen- sive as possible. The additional material includes specimens from the Zoologisk Museum, Copen- hagen, Denmark, labeled “Smidt,” which were probably collected before 1800 and which Fab- ricius may have seen; collections made by Dr. O. W. Richards, London, England; Dr. F. X. Williams, entomologist for many years with the Hawaiian Sugar Planters Association; Mr. D. Vesey-Fitzgerald, formerly of the College of Tropical Agriculture, Trinidad, B. W. I., and specimens from numerous museums in Europe and North America. The types of all the species involved were studied, redescribed and keyed during 1930-31 when the writer was a John Simon Guggenheim Memorial Fellow. Six new genera, twenty-nine new species and one new subspecies are de- scribed. Keys are presented to the genera occur- ring in British Guiana, and to the species recorded from that country as well as related species in some instances in order to facilitate identification. New synonymy has been included wherever relevant. Numerous other species have been described 1 Contribution No. 926, Department of Tropical Re- search, New York Zoological Society. 2 Paper No. 2828 of the Scientific Journal Series of the Minnesota Agricultural Experiment Station. and remain to be described from Venezuela, Dutch Guiana (Surinam), French Guiana, Trin- idad, B. W. I., and Brazil north of the Amazon River. Some of these will undoubtedly be ul- timately recorded from British Guiana, so the present report cannot be regarded as complete. Key to the Genera Females 1. Eyes strongly ovate with distinct facets; thorax rectangular, subrectangular or ovate, not conspicuously broader an- teriorly than posteriorly 2 Eyes circular to subovate, polished, the facets usually indistinct, thorax vari- ously shaped but always conspicuously broader anteriorly than posteriorly. ... 3 2. First abdominal segment completely ses- sile with the second; thorax rectangu- lar Timulla subgen. Timulla Ashmead First abdominal segment cylindrical or subcylindrical, distinctly petiolate; tho- rax ovate Ephuta Say 3. Antennae with 13 segments, the third seg- ment about four times the length of the fourth; head armed with four dentiform or tuberculate processes, one pair on the genae and a second pair near the pro- boscidal fossa Hoplocrates Mickel Antennae with only 12 segments, the third segment not more than three times the length of the fourth 4 4. Mandibles tomentose throughout Pappognatha Mickel Mandibles glabrous, sparsely pubescent proximally 5 5. Pygidial area well defined, with distinct lateral boundaries, usually sculptured . . 6 Pygidial area not defined, the posterior third to half of the last tergum glabrous 8 6. First abdominal segment completely ses- sile with the second . Pseudomethoca Ashmead First abdominal segment disciform (i.e., 105 106 Zoologica: New York Zoological Society [37: 11: with a distinct narrow dorsal face) or distinctly petiolate 7. Second abdominal tergum with longitudi- nal rows of interrupted carinae, and with one pair of yellow or ferruginous integumental spots (rarely the integu- mental spots replaced with pubescent maculations) Hoplomutilla Ashmead Second abdominal tergum without rows of longitudinal carinae, usually with two pairs of yellow or ferruginous integu- mental spots (rarely with one pair) Traumatomutilla Andre 8. Head large, subquadrate, the genal carina extending posteriorly on to the vertex; proboscidal fossa small, its width about equal to half the distance between the insertion of the mandibles; proboscidal fossa not connected with the genal car- ina by a transverse carina; lamellate process between and above hind coxae rounded at the tip; first abdominal seg- ment completely sessile or subsessile with the second segment 9 Head not distinctly subquadrate, the genal carina not extending posteriorly on to the vertex; proboscidal fossa large, its width much greater than half the dis- tance between the insertion of the man- dibles; proboscidal fossa connected with the genal carina by a transverse carina; lamellate process between hind coxae acute at the tip; first abdominal segment either subnodose or disciform. 10 9. Mandibles slender, with a distinct, obtuse tooth within near the base; anterior margin of clypeus with a distinct tooth laterad of the insertion of the antennae Calomutilla, n. gen. Proximal two-thirds of mandibles broad, then narrowed, forming a conspicuous tooth on the inner margin, the distal third attentuated and edentate; anterior margin of clypeus not dentate laterad Pertyella, n. gen. 10. Mandibles tridentate distally, the distal and inner tooth large, the intermediate tooth small; inner margin of mandible with a distinct tooth intermediate be- tween insertion and inner distal tooth; first abdominal segment nodose Lophomutilla, n. gen. Mandibles edentate distally, slender, slightly broadened and angulate at the middle within, but the inner margin not dentate; first abdominal segment disci- form Lophostigma, n. gen. Males 1. Eyes deeply emarginate on their inner margins 2 Eyes entire, not emarginate within 3 2. First abdominal segment entirely sessile with the second; abdominal terga with- out, or only the last one with, a longi- tudinal median keel Timulla subgen. Timulla Ashmead First abdominal segment cylindrical, not at all sessile with the second; third to seventh abdominal terga with a longi- tudinal median keel Ephuta Say 3. Head armed beneath with four dentiform processes, one pair on the genae and one pair near the proboscidal fossa Hoplocrates Mickel Head unarmed beneath 4. First abdominal segment completely ses- sile with the second First abdominal segment not sessile with the second, either disciform, or petiolate 5. Last segment of tarsi with a lamellate process extending over and beyond the base of the claws .... Hoplomutilla Ashmead Last segment of tarsi not produced beyond the base of the claws Pseudomethoca Ashmead 6. Mandibles tomentose throughout Pappognatha Mickel Mandibles glabrous, clothed with sparse hairs 7. Scape bicarinate beneath Traumatomutilla Andre Scape with a single carina beneath 8 8. Proboscidal fossa not extending to base of mandibles; mandibles tridentate at tip; antennal tubercles widely separated Ancipitotilla, n. gen. Proboscidal fossa large, extending laterad to insertion of mandibles; mandibles edentate at the tip; antennal tubercles almost contiguous. . .Mammomutilla, n. gen. Genus TIMULLA Ashmead Subgenus TIMULLA Ashmead Key to the Females 1 . Pygidium longitudinally striate rectangula (Spinola) Pygidium not longitudinally striate, sculp- tured otherwise 2 2. Head entirely ferruginous, .discontinua Mickel Head entirely black 3 3. Thorax entirely ferruginous. . . .Valeria Mickel Thorax maculated on the sides or above with black 4 4. Dorsum of thorax entirely ferruginous; either the posterior face of propodeum or the meso- and metapleura maculated with black 5 Dorsum of thorax maculated with black. 6 5. Thorax broader posteriorly than anterior- ly; pygidium unsculptured, glabrous on posterior half eriphyla Mickel 1952] Mickel: Mutillidae (Wasps) of British Guiana 107 Thorax not broader posteriorly than an- teriorly; pygidium irregularly rugose with the posterior margin finely granu- late mediata (Fabricius) 6. Posterior marginal, pale pubescent band of second tergum complete mediata var. pexsa Mickel Posterior margin of second tergum black, without a pale pubescent band bitaeniata (Spinola) Key to the Males 1 . Posterior margin of last tergum emargi- nate medially 2 Posterior margin of last tergum not emar- ginate medially 4 2. Proximal segments of flagellum yellowish beneath - 3 Flagellum entirely black, .bitaeniata (Spinola) 3. Glabrous area of clypeus with the poste- rior margin evenly arcuate, the poste- rior half to two-thirds of the area almost flat, or evenly, shallowly concave; cell R.5 slightly to distinctly less than three times as long as broad, .mediata (Fabricius) Glabrous area of clypeus with the poste- rior margin elevated and obtusely an- gulate medially, the posterior half to two-thirds of the area distinctly, trans- versely concave; cell R5 slightly to dis- tinctly less than three times as long as broad obtusata Mickel 4. Median, impunctate area of last tergum broadly emarginate posteriorly when viewed from above; posterior, inner an- gle of middle coxae with a small dis- tinct tooth rectangula (Spinola) Median, impunctate area of last tergum not emarginate posteriorly when viewed from above; posterior inner angle of middle coxae toothed or not 5 5. Abdomen entirely ferruginous; posterior, elevated margin of clypeus evenly arc- uate or slightly subangulate medially, and slightly reflexed medially rufogastra (Lepeletier) Last abdominal segment black, the first segment more or less black; clypeus ele- vated posteriorly, the posterior, elevated • margin evenly arcuate Valeria Mickel Timulla (Timulla) mediata (Fabricius) 1805. Dorylus mediatus Fabricius, Syst. Piez., p. 428, male. 1805. Mutilla lineola Fabricius, Syst. Piez., p. 437, female. 1912. Mutilla moorei Cameron, Timehri; Journ. Roy. Agri. Com. Soc. British Guiana, (3), vol. 2, p. 413, male. 1912. Mutilla longilineata Cameron, Timehri; Journ. Roy. Agri. Com. Soc. British Guiana, (3), vol. 2, p. 414, female. 1938. Timulla (Timulla) mediata Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 560, male and female. Specimens Examined.— Male, Kartabo, Bar- tica district, British Guiana, June 15, 1924. Also previously recorded from the following locali- ties in British Guiana: Blairmont, Georgetown and Kamakusa. Distribution— British Guiana, Colombia, Ec- uador, Trinidad, B. W. I., Surinam, French Guiana and Brazil. Timulla (Timulla) mediata var. pexsa Mickel 1938. Timulla ( Timulla ) mediata var. pexsa Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 563, male and female. Previously recorded from: Blairmont, August, 1923 (H. E. Box). Distribution— Brazil, French Guiana and British Guiana. Timulla (Timulla) obtusata Mickel 1937. Timulla ( Timulla ) obtusata Mickel, Rev. Ent., Rio, vol. 7, pp. 169-171, male. 1938. Timulla ( Timulla ) obtusata Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 563, male. Not known from British Guiana, but may occur there as it is recorded from Surinam. Distribution.— Surinam, French Guiana and Brazil. Timulla (Timulla) bitaeniata (Spinola) 1841. Mutilla bitaeniata Spinola, Ann. Soc. Ent. France, vol. 10. p. 91, female. 1938. Timulla (Timulla) byblis Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 566, male. (New synorxymy). 1938. Timulla (Timulla) bitaeniata Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 595, female. Not known from British Guiana but probably occurs there. Distribution— French Guiana, Surinam and Trinidad, B. W. I. A male and female taken in coitu in Trinidad, B. W. I., show that byblis Mickel is the male of bitaeniata Spinola. The specimens are in the collection of the University of Minnesota. Timulla (Timulla) rectangula (Spinola) 1841. Mutilla rectangulum Spinola, Ann. Soc. Ent. Fr., vol. 10, p. 91, female. 108 Zoologica: New York Zoological Society [37: 11: 1938. Timulla ( Timulla ) rectangulum Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 584, female and male. Specimens Examined— Five females and five males, Kartabo, Bartica district, March 6, 1924; March 10, 1924; June 30, 1919; July 5, 1922; July 8, 1922; September 6, 1922; October 6, 1922; and without date. Also previously recorded from the following localities in British Guiana: Blairmont and Es- sequibo river. Distribution— Colombia, Venezuela, British Guiana, Surinam, French Guiana and Trinidad, B. W. I. Timulla (Timulla) rufogastra (Lepeletier) . 1845. Mutilla rufogastra Lepeletier, Hist. Nat. Ins., Hymen., vol. 3, p. 629, male. 1938. Timulla ( Timulla ) rufogastra Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 598, male. Not recorded from British Guiana but prob- ably occurs there. Distribution— French Guiana, Venezuela and Trinidad, B. W. I. Timulla (Timulla) Valeria Mickel 1938. Timulla ( Timulla ) Valeria Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 600, male. 1938. Timulla ( Timulla ) herse Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 625, female, (New synonymy). Specimens Examined.— Male and female (in coitu), Kartabo, Bartica district, May 12, 1924 [in collection of University of Minnesota]; two females, Kartabo, Bartica district, April 5, 1922, and September 26, 1922; female, Bartica, Bar- tica district; female, Tropica! Research Station, No. 21115. Also previously recorded from the following localities: Kamakusa, Tumatumari, Kerikabaru, and Essequibo river. Distribution. — British Guiana, Surinam, French Guiana, Brazil, Venezuela, Colombia and Ecuador. The above pair, taken in coitu, establishes herse Mickel as the opposite sex (female) of Valeria Mickel (male). Timulla (Timulla) discontinua Mickel 1938. Timulla ( Timulla ) discontinua Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 594, female. Not recorded from British Guiana, but prob- ably occurs there. Distribution — French Guiana and Venezue- la. Timulla (Timulla) eriphyla Mickel 1938. Timulla ( Timulla ) eriphyla Mickel, Trans. R. Ent. Soc. London, vol. 87, p. 591, female. Not recorded from British Guiana, but prob- ably occurs there. Distribution.— French Guiana, Trinidad, B. W. L, Venezuela and Colombia. Genus EPHUTA Say KEY TO THE FEMALES 1. Second abdominal tergum with a deep fo- vea near each lateral margin 2 Second abdominal tergum uniformly punc- tate near the lateral margins, without a deep fovea 3 2. Lateral fovea small, inconspicuous, ap- proximately one puncture in width and two punctures in length, not filled with dense pubescence; second tergum with- out a pair of anterior, pale pubescent spots erichtho, n. sp. Lateral fovea conspicuous, approximately six times longer than wide, filled with dense pubescence; second tergum with a pair of anterior, subtriangular, pale pubescent spots weberi, n. sp. 3. Thorax with a very broad, transverse, ele- vated, glabrous scutellar scale aurulenta (Fabricius) Thorax without a scutellar scale 4 4. Head entirely black 5 Front ferruginous, vertex and genae black; thorax ferruginous, except the prono- tum maculated with black. . . .roxane, n. sp. 5. Thorax uniformly bright ferruginous egeria, n. sp. Thorax ferruginous, but the dorsum more or less maculated with black 6 6. Lateral margins of dorsum of thorax, or the pronotum and propodeum, more or less infuscated, the infuscated spots not sharply defined 7 Lateral margins of dorsum of thorax with distinct spots or lines of black, the mac- ulations confluent medially on both the pronotum and the propodeum 8 7. Propodeum with a pair of lateral, elon- gate, infuscated spots, sometimes very indistinct, almost absent copia, n. sp. Anterior two-thirds of dorsum and poste- rior face of propodeum infuscated elvina, n. sp. 8. Abdominal terga three to five clothed throughout with pale golden pubes- cence limbata, n. sp. Abdominal terga three to five, each with a median, transverse, pale pubescent 1952] Mickel: Mutillidae (Wasps) of British Guiana 109 spot, and with small lateral spots of fus- cous to black pubescence, .hylonome, n. sp. Key to the Males 1 . Second abdominal segment entirely black 2 Second abdominal segment almost entire- ly ferruginous rubrocincta, n. sp. 2. Postero-lateral angles of dorsum of propo- deum with a prominent tooth, or spine, or veiy prominently angulate 3 Postero-lateral angles of dorsum of pro- podeum serrate or crenulate, but not toothed, nor spined, nor prominently angulate 3. Tegulae transversely rugoso-punctate. . . 4 Tegulae not transversely rugoso-punctate, usually with fine, separated punctures 6 4. Postero-lateral angles of scutellum pro- duced into prominent teeth; dorsal half of mesopleura elevated into a promi- nent tooth singulars (Spinola) Postero-lateral angles of scutellum not at all produced; mesopleura evenly con- vex, the dorsal half not elevated into a tooth 5 5. Anterior half of disc of second tergum strongly flattened and distinctly con- cave, closely punctate perfica, n. sp. Anterior half of disc of second tergum slightly flattened, but not at all concave, sparsely punctate trifid a (Gerstaecker) 6. Posterior margin of scutellum distinctly emarginate medially; anterior half of disc of second tergum not conspicuous- ly flattened fugax (Smith) Posterior margin of scutellum not at all emarginate; anterior half of disc of sec- ond tergum conspicuously flattened, slightly concave richardsi, n. sp. 7. Tegulae transversely rugoso-punctate abadia (Cresson) Tegulae not transversely rugoso-punctate, with fine, separated punctures 8 8. Scutellum elevated medially into a con- spicuous subconical tubercle 9 Scutellum fiat or convex, not elevated medially into a subconical tubercle. ... 10 9. Second tergum evenly convex throughout; last sternum largely yellow, .subconica, n. sp. Anterior half of disc of second tergum dis- tinctly flattened; last sternum largely dark ferruginous depressa, n. sp. 10. Mandibles entirely yellow, except the tips castaneous 12 Mandibles sometimes yellow medially, usually dark ferruginous to black .... 11 11. Scutellum with a dorsal face and a poste- rior face almost perpendicular to the former, and with a median, shallow emargination at the junction of the two faces emarginata, n. sp. Scutellum evenly convex from anterior to posterior margins, without a posterior face and without a median emargina- tion infract a, n. sp. 12. Hypopygium conspicuously yellow; sides of propodeum reticulate from anterior to posterior margin indiscrete, n. sp. Hypopygium black or concolorous with other abdominal sterna; sides of propo- deum glabrous, unsculptured on ante- rior half, reticulate on posterior half flavidens, n. sp. Ephuta rubrocincta, new species Male— Holotype. Black, except the second abdominal segment red, and the anterior two- thirds of the last sternum pale yellow; scutellum convex; postero-lateral angles of dorsum of pro- podeum not dentate nor tuberculate, but the dorsum separated from the posterior face of propodeum by a strong, transverse carina elevated at the midline into a distinct tubercle; tegulae with sparse, fine punctures; disc of sec- ond tergum evenly convex. Length, 9 mm. Head black, the distal half of the mandibles ferruginous with the tips dark; lateral areas of clypeus with long, subappressed, pale, silky pu- bescence; anterior half of front with thick, ap- pressed, pale pubescence, the posterior half and area through ocelli with sparse, pale pubescence; vertex with the pale, appressed pubescence thin and with sparse, erect, pale hairs; genae with thick, appressed, pale pubescence and sparse, erect, pale hairs; mandibles edentate at the tip and with a small tooth within; clypeal carinae dilated posteriorly to form an incomplete circle, then for a short distance parallel, then each flaring laterally and continuous to the anterior margin; scape strongly bicarinafe beneath; first and second flagellar segments approximately equal in length; antennal scrobes strongly car- inate above; front, vertex and genae with mod- erate, contiguous to confluent punctures, those on the vertex more distinct than on the front, those on the genae almost concealed by the thick, pale pubescence. Thorax entirely black; dorsum of pronotum clothed with thin, appressed, pale pubescence and sparse, erect, pale hairs; mesonotum with sparse, inconspicuous, fuscous hairs, except the postero-lateral angles with a tiny spot of dense, pale pubescence; scutellum with sparse, erect, pale hairs; dorsum and posterior face of propo- deum with thin, subappressed, pale pubescence; lateral aspect of pronotum with sparse, pale pubescence; mesopleura with thick, appressed, pale pubescence, and sparse, erect, pale hairs; 1 10 Zoologica: New York Zoological Society [37: 11: metapleura with sparse, appressed, pale pubes- cence; sides of propodeum with sparse, ap- pressed, pale pubescence and sparse, erect, pale hairs; dorsum of pronotum with moderate, dense, more or less confluent punctures; mesono- tum with six more or less distinct, longitudinal rows of large, shallow, confluent punctures; scutellum evenly convex, with moderate, dense, somewhat confluent punctures; dorsum of pro- podeum separated from the almost perpendicu- lar, posterior face of propodeum by a strong, transverse carina elevated medially into a dis- tinct tubercle; postero-lateral angles of dorsum of propodeum not dentate nor tuberculate; dor- sum and posterior face of propodeum reticulate, the former with a median, enclosed area almost its entire length; lateral aspect of pronotum gla- brous, very indistinctly, shallowly punctate; mesopleura evenly convex, with moderate, con- tiguous punctures, somewhat concealed by the thick, pale pubescence; metapleura glabrous, somewhat punctate ventrally; sides of propo- deum reticulate; tegulae convex, glabrous, finely punctate, the punctures sparse at the margins, clothed with sparse, fuscous pubescence, except the anterior and inner margins with the pubes- cence pale. Abdomen black, except the second segment almost entirely red, only the posterior margin blackish, and the anterior two-thirds of the last sternum pale yellow; first segment distinctly broader than long; first tergum with moderate, contiguous punctures almost concealed by the thick, appressed, pale pubescence and sparse, erect, pale hairs; second tergum evenly convex, with moderate, contiguous, distinct punctures, the disc clothed with short, inconspicuous, ferru- ginous hairs, the lateral and anterior areas with sparse, erect, pale hairs, and the posterior mar- gin with a band of dense, appressed, pale pubes- cence, slightly narrowed medially; terga three to six with small punctures and sparse, erect, pale hairs; last tergum with small, contiguous to confluent punctures and sparse, erect, pale hairs; first sternum with a median, longitudinal carina, not elevated anteriorly, and clothed with sparse, pale pubescence; second sternum with moderate, contiguous, distinct punctures, and with sparse, pale hairs, the lateral thirds of the posterior mar- gin with a band of thick, appressed, pale pubes- cence; sterna three to six with small punctures, and sparse, erect, pale hairs; anterior two-thirds of last sternum with small, contiguous punctures and sparse, erect, pale hairs, the posterior third with sparse, fuscous hairs. Wings subhyaline; cell 2nd R1+R2 three times as long as wide, obliquely truncate at the tip; cell R3 receiving vein M3+4 two-thirds the dis- tance from base to apex; cell Ri much less dis- tinct than R5 and receiving vein M2 distinctly beyond the middle. Legs black, clothed with sparse, pale pubes- cence; calcaria pale. Holotype.— Male, Blairmont, British Guiana, September, 1923 (F. X. Williams), in collection of University of Minnesota. Paratypes.— Male, Blairmont, British Guiana, October, 1923 (F. X. Williams); male, Blair- mont, British Guiana, August 6, 1923 (H. E. Box) ; male, Blairmont, British Guiana, October 15, 1923 (H. E. Box); male, Essequibo, British Guiana (Smidt); male, Jabaty, Para, Brazil, May, 1924 (F. X. Williams); and male, Ara- pary, Para, Brazil, May, 1924 (F. X. Williams). Easily distinguished from all other males of this genus in British Guiana by the red second abdominal segment. Ephuta singularis (Spinola) 1841. Mutilla singularis Spinola, Ann. Soc. Ent. France, vol. 10, p. 95, male. 1902. Mutilla catinga Cresson, Trans. Amer. Ent. Soc., vol. 28, p. 64, male. (New synonymy). 1937. Ephuta singularis Mickel, Rev. Ent., Rio, vol. 7, p. 175, female. Specimens Examined.— Male, Bartica, British Guiana, February 18, 1913; male, Blairmont, British Guiana, September, 1923 (F. X. Wil- liams) . Distribution. — British Guiana, Surinam, French Guiana, Para, Brazil and Santarem, Brazil. Ephuta perfica, new species Male.— Holotype. Entirely black, except the last sternum yellow; scutellum evenly convex and densely punctate; dorsum of propodeum with a low, blunt, longitudinally compressed tooth at the postero-lateral angles and a median, small, longitudinally compressed acute tooth at the tip of the enclosed area; tegulae transversely rugoso-punctate and with a distinct, median, longitudinal ridge; disc of second tergum de- pressed, distinctly concave. Length, 10 mm. Head black; front anterior to emargination of eyes, vertex posterior to lateral ocelli and genae, clothed with thick, appressed, pale, glit- tering pubescence, and sparse, erect, pale hairs; remainder of front and vertex with very sparse, appressed, pale, glittering pubescence and sparse, erect, black hairs; distal half of mandi- bles yellow, the tips reddish; mandibles edentate at the tip and with a distinct tooth within, clothed with pale hairs throughout, thick proximally, sparse at the tip; clypeus with a pair of parallel, median carinae on the posterior half, the ante- 1952] Mickel: Mutillidae (Wasps) of British Guiana 111 rior half depressed, and the broad, anterior mar- gin thin and translucent, clothed throughout with thick, long, subappressed, pale, silky hairs; antennal scrobes distinctly carinate above; scape strongly bicarinate beneath; first and second seg- ments of flagellum approximately equal in length; front, vertex and genae with moderate, close, distinct punctures visible through the thick pubescence as well as elsewhere; genae elevated into a very high carina at the articulation of the mandible, the carina extending caudad a dis- tance equal to half the length of the mandible; ventral aspect of head between the carinae and surrounding the proboscidal fossa glabrous, im- punctate. Thorax entirely black; posterior margin of pronotum, mesonotum entirely, and scutellem with sparse, erect, black hairs, the pronotum also with sparse, appressed, pale pubescence and scattered, erect, pale hairs, the scutellum also with sparse, short, inconspicuous, pale, erect, hairs, and long, erect, pale hairs at the lateral and posterior margins, remainder of thorax clothed with pale pubescence, thick and ap- pressed on the elevated area of mesopleura, thick and irregularly semi-erect on the propo- deum, thin and appressed elsewhere; pronotum with large, close, contiguous punctures, the latter almost obsolete at the midline; mesonotum with longitudinal rows of large, contiguous to con- fluent punctures; scutellum evenly convex, with moderate, contiguous, deep punctures; dorsum of propodeum with a narrow, median, en- closed area four times as long as broad and terminating at a distinct, longitudinally, compressed, acute tooth; postero-lateral angles of propodeum with a low, blunt, longitu- dinally compressed tooth, the elevation longer than high; surface of dorsum, posterior face, and sides of propodeum otherwise reticu- late; lateral aspect of pronotum coarsely punc- tate at the upper margin, otherwise glabrous, impunctate; mesopleura with moderate, deep, contiguous punctures throughout, evenly convex throughout, not elevated anywhere into a tu- bercle or tooth; metapleura with a small area of moderate, contiguous punctures at the ventral margin, otherwise impunctate; tegulae large, with a distinct, median, longitudinal ridge, finely, transversely, rugoso-punctate throughout and clothed with short, sparse, black hairs. Abdomen black, except the last sternum al- most entirely pale yellow, only the posterior margin of this sternum fuscous to black; first tergum entirely, small antero-lateral and pos- tero-lateral areas of second tergum, anterior half of second sternum, and posterior margin of second sternum broadly interrupted medially, all with thick, appressed, pale, glittering pubes- cence; posterior margin of second tergum with a thin fringe of pale, glittering hairs, very nar- rowly and inconspicuously interrupted medially with a very few black hairs; remainder of abdo- men clothed with very sparse, semi-erect and erect, pale hairs, except the posterior half of the disc of the second tergum and the median area of terga three to five with sparse, inconspicuous, black hairs; first abdominal segment viewed from above approximately as broad as long; first tergum with moderate, distinct punctures visible through the thick pubescence; anterior three-fourths of disc of second tergum strongly depressed, concave; second tergum with mod- erate, deep, distinct, contiguous punctures throughout; felt lines of second tergum absent; terga three to six with small, shallow, separated punctures; last tergum with small, shallow, close punctures; first sternum with a distinct but not prominent, median, longitudinal carina slightly elevated anteriorly, and with small, deep, dis- tinct punctures; second sternum with moderate, distinct, contiguous to slightly separated punc- tures; sterna three to six with small, shallow punctures; last sternum with small, close, dis- tinct punctures. Wings subhyaline; cell 2nd Ri+Re three and one-half times as long as wide, subacute at the tip; cell R.5 receiving vein M3+4 at the middle; cell R4 receiving vein M2 at the middle. Legs black, sparsely clothed with pale hairs; posterior coxae with a weak, longitudinal ca- rina on the inner surface; calcaria white. Holotype.— Male, Kamakusa, British Guiana (H. Lang), in collection of University of Min- nesota. Paratypes.— Male, Kartabo, Bartica district, British Guiana, March 29, 1924 (Wm. Beebe); male, Kartabo, British Guiana, July-August, 1920 (W. M. Wheeler); male, Bartica, British Guiana, February 4, 1913; three males, Mora- balli Creek, Essequibo River, British Guiana, September 19, 1929, in Wallaba bush = Iperna sp. (O. W. Richards) [Oxford Univ. Expedi- tion]; male, Moraballi Creek, Essequibo River, British Guiana, September 25, 1929, on trail (O. W. Richards); male, Paramaribo, Surinam (J. Michaelis); and two males, Mana River, French Guiana, May, 1917. Related to trifida Gerstaecker, but differs from that species in the strongly concave second tergum, the prominently elevated carina on the genae at the articulation of the mandibles, the less prominent and blunt elevations at the postero-lateral angles of dorsum of propodeum and the entirely black last abdominal tergum. The paratypes vary in length from 9 mm to 12 mm. 112 Zoologica: New York Zoological Society [37: 11: Ephuta trifid a (Gerstaecker), (new combination) 1874. Mutilla trifda Gerstaecker, Arch. f. Naturg., vol. 40, p. 327, male. Specimens Examined.— ¥ our males, Kartabo, British Guiana, July-August, 1920 (W. M. Wheeler); male, Bartica, British Guiana, May 17, 1901; male, Demerara River, British Gui- ana, March 12, 1913; male, Kamakusa, British Guiana, January, 1923 (H. Lang); two males, Kamakusa, British Guiana (H. Lang); two males, Tukeit, British Guiana, July 17 and 18, 1911; two males, Oko River, British Guiana, June 26, 1936 (N. A. Weber); and six males, Moraballi Cr., Essequibo River, British Guiana, September, 1929 (O. W. Richards). Distribution— Venezuela, British Guiana and Para state, Brazil. Ephuta fugax (Smith), (new combination) 1879. Mutilla fugax Smith, Descr. New Spe- cies Hymen., p. 220, male. 1902. Mutilla morada Cresson, Trans. Amer. Ent. Soc., vol. 28, p. 63, male. (New synonomy) . Specimens examined— Male, Bartica, Bartica district, British Guiana; two males, Moraballi Creek, Essequibo River, British Guiana; Sep- tember 25, 1929, dark forest (O. W. Richards). Distribution — British Guiana, French Gui- ana, Brazil. The type of morada has been compared with the above specimens which also answer to the original description, and to a description drawn by the writer from Smith’s type of fugax; mo- rada is undoubtedly a synonym. Ephuta richardsi, new species Male— Holotype. Entirely black, except the anterior two-thirds of the last sternum yellow; scutellum convex; postero-lateral angles of dorsum of propodeum prominently spinose; posterior margin of dorsum of propodeum not dentate nor tuberculate medially; tegulae with fine sparse punctures throughout; disc of second tergum conspicuously flattened. Length. 10 mm. Head black, clypeus clothed throughout with thick, long, subappressed, pale hairs; front an- terior to emargination of eyes with thick, pale pubescence; remainder of front with sparse, pale pubescence and a few, scattered, black hairs posteriorly; transverse area through ocelli with sparse, erect, fuscous to black hairs; vertex with thin, appressed, pale pubescence and sparse, erect, pale hairs; genae with thick, ap- pressed, pale pubescence and sparse, erect, pale hairs; distal half of mandibles castaneous, ex- cept the tips blackish; mandibles edentate at the tip and with a small tooth within, clothed with pale pubescence; clypeal carinae subpar- allel, strongly elevated anteriorly; scape strong- ly bicar inate beneath; first and second segments of flagellum approximately equal in length; front, vertex and genae with moderate, dense, contiguous punctures; genal carina strongly elevated into a conspicuous tooth slightly pos- terior to the insertion of the mandibles. Thorax entirely black; dorsum of pronotum with sparse, appressed, pale pubescence and sparse, erect, black hairs posteriorly; mesono- tum with sparse, erect, black hairs throughout, except the postero-lateral angles with a small spot of dense, pale pubescence; scutellum with sparse, erect, black hairs on the disc, the nar- row, lateral and posterior faces with thin, pale pubescence; dorsum and posterior face of pro- podeum with thick, subappressed, pale pubes- cence and sparse, erect, pale hairs; lateral aspect of pronotum with thin, appressed, pale pubescence; mesopleura with thin, appressed, pale pubescence and sparse, erect, pale hairs; metapleura with thin, appressed, pale pubes- cence; sides of propodeum with thick, sub- appressed, pale pubescence and sparse, erect, pale hairs like that on the dorsum of pro- podeum; tegulae with sparse, fuscous to black pubescence throughout; dorsum of pronotum with moderately large, dense, contiguous to confluent punctures; mesonotum with moder- ately large, more or less confluent punctures, not arranged in rows, and with a weak, median, longitudinal, glabrous line; scutellum convex, with moderate, more or less confluent punc- tures, and a weak, median, longitudinal, gla- brous line on the anterior two-thirds; dorsum and posterior face of propodeum not separated, rounded into one another, the dorsum without a median tooth or tubercle at its posterior mar- gin, but with a conspicuous, spinose process at each postero-lateral angle; dorsum and pos- terior face of propodeum reticulate; lateral aspect of pronotum glabrous, micropunctate, except dors ally punctate like the dorsum of pronotum; mesopleura evenly convex through- out and with moderately small, distinct, con- tiguous punctures; metapleura glabrous, micro- punctate, except moderately punctate ventrally; anterior third of sides of propodeum glabrous, micropunctate, the remainder reticulate like the dorsum; tegulae evenly convex, with a median ridge on the anterior fourth, glabrous, sparsely, finely punctate throughout. Abdomen black, except the anterior two- thirds of the last sternum yellow; first segment cylindrical, distinctly longer than broad; first 1952] Michel: Mutillidae (Wasps) of British Guiana 113 tergum with moderately small punctures almost concealed by the thick, appressed, pale pubes- cence and sparse, erect, pale hairs; anterior disc of second tergum distinctly flattened and slightly concave; second tergum with moderate, distinct, contiguous punctures, except at the an- terior middle the punctures sparse, shallow and indistinct and at the posterior margin medially obsolete, the surface glabrous; second tergum clothed with sparse, black hairs, except a few, erect, pale hairs at the anterior middle, the lat- eral areas with thin, pale, appressed pubescence, the lateral margin with a narrow, thin fringe of pale pubescence, and the posterior margin with a band of pale pubescence, the latter broad at the sides, narrowed toward the middle and interrupted at the midline; third tergum without a median carina, glabrous, almost impunctate medially, with small, separated punctures to- wards the sides anteriorly, and clothed with sparse, black hairs, except laterally with sparse, pale hairs; fourth and fifth terga with sparse, small punctures, with sparse, black hairs medi- ally, and sparse, pale hairs laterally; sixth ter- gum with sparse, small punctures and sparse, pale hairs; last tergum with small, distinct, slightly separated punctures and sparse, pale hairs; first sternum with a median, longitudinal carina distinctly elevated anteriorly, clothed throughout with thin, pale pubescence; second sternum with moderate, distinct, separated punctures and clothed throughout with thin, appressed, pale pubescence; sterna three to six with sparse, small punctures and sparse, pale hairs; last sternum with separated, small, in- distinct punctures and sparse, erect, pale hairs. Wings subhyaline; cell 2nd R1+R2 approxi- mately three times as long as wide, truncate at the tip; cell Rr, receiving vein M3+4 distinctly beyond the middle; cell R4 much less distinct than Rs and receiving vein M2 distinctly beyond the middle. Legs black, sparsely clothed with pale pubes- cence; calcaria pale. Holotype — Male, Moraballi Creek, Essequi- bo River, British Guiana, September 21, 1929 (O. W. Richards), in collection of University of Minnesota. Related to diacanthci Mickel from Brazil but lacks entirely the dense, pale pubescence on the anterior third of the second tergum and sternum, the pronotum is less thickly clothed with pale pubescence and the dorsum of the propodeum is not so densely pale pubescent. Ephuta abadia (Cresson), (new combination) 1902. Mutilla abadia Cresson, Trans. Amer. Ent. Soc., vol. 28, p. 64, male. Specimens Examined— Three males, Bartica, British Guiana, January 29, 1913; three males, Blairmont, British Guiana, September, 1923 (F. X. Williams); two males, Kamakusa, Brit- ish Guiana. Distribution— Santarem, Brazil and British Guiana. Ephuta subconica, new species Male— Holotype. Entirely black, except the anterior three-fourths of the last sternum yel- low; anterior half of scutellum with a median, longitudinal, elevated line, terminating in a conspicuous, subconical elevation at the mid- dle; postero-lateral angles of dorsum of pro- podeum not spinose or tuberculate; dorsum of propodeum not dentate or tuberculate me- dially at the posterior margin; tegulae evenly convex, with fine, sparse punctures; disc of second abdominal tergum evenly convex, not at ail depressed. Length, 8.5 mm. Head black; lateral areas of clypeus clothed with thick, long, subappressed, pale, silky hairs, the median area very sparsely so; front clothed with thick, appressed, pale pubescence and sparse, erect, pale hairs; ocellar area with only sparse, erect, pale hairs; vertex with sparse, erect, pale hairs and thin, appressed, pale pu- bescence; genae with thick, appressed, pale pu- bescence and sparse, erect, pale hairs; man- dibles dark castaneous on the proximal half and the tips, yellowish just distad of the middle, edentate at the tip and with a small tooth within; clothed with sparse, pale hairs; median carinae of clypeus subparallel; scape strongly bicarinate beneath; first and second segments of flagellum approximately equal in length; front, vertex and genae with moderate, con- tiguous punctures; carina separating anterior half of genae from proboscidal fossa not con- spicuous, only moderately developed. Thorax entirely black; dorsum of pronotum with thin, appressed, pale pubescence and sparse, erect, pale hairs; anterior and lateral margins of mesonotum with thin, appressed, pale pubescence, the postero-lateral angles with a small spot of dense, appressed, pale, pu- bescence; mesonotum otherwise with sparse, erect, pale fuscous to black hairs; lateral and posterior marginal areas of scutellum with sparse, erect, pale hairs, the median area with sparse, erect, fuscous to black hairs; dorsum and posterior face of propodeum with thick to dense, subappressed, pale pubescence; lat- eral aspect of pronotum with thin, sub- appressed, short, pale pubescence; mesopleura with thick, appressed, pale pubescence; meta- pleura and sides of propodeum with thin, 114 Zoologica: New York Zoological Society [37: 11: short, subappressed, pale pubescence; tegulae with sparse, suberect, pale fuscous to fuscous hairs, except the anterior and inner margins with pale pubescence; dorsum of pronotum with moderate, contiguous to confluent punc- tures; mesonotum with six longitudinal rows of moderately large, shallow, confluent punc- tures; scutellum with a median, longitudinal, glabrous, elevated line on the anterior half terminating at the middle of scutellum in a large, conspicuous, subconical tubercle, the re- mainder of scutellum with moderate, dense, confluent punctures; dorsum of propodeum rounded into the posterior face and not dentate nor tuberculate medially at the posterior mar- gin; postero-lateral angles of dorsum of pro- podeum not spinose, dentate nor tuberculate; dorsum and posterior face of propodeum re- ticulate; lateral aspect of pronotum glabrous, micropunctate, except the dorsal margin punc- tate like the dorsum of pronotum; mesopleura pvenly convex throughout and with moderate, contiguous punctures; metapleura glabrous and micropunctate, except a few contiguous punc- tures ventrally; sides of propodeum reticulate, tegulae large, convex, except a short, anterior, median carina glabrous, and with sparse, fine punctures. Abdomen black, except the anterior three- fourths of the last sternum yellow; first abdomi- nal segment cylindrical, about as broad as long when viewed from above; first tergum clothed with thick, appressed, pale pubescence and sparse, long, erect, pale hairs; second tergum evenly convex throughout, the disc not de- pressed, clothed with sparse, erect, pale hairs, and posteriorly with very inconspicuous, ap- pressed, sparse, black hairs, the lateral margins narrowly with thick, short, pale pubescence and the posterior margin with a moderately broad band, somewhat narrowed medially, of thick, appressed pale pubescence; third to sixth terga with sparse, erect, pale hairs; last tergum with sparse, pale hairs; first tergum with moderate, dis- tinct punctures more or less concealed by the pale pubescence; second tergum with moderate, distinct, contiguous, very deep punctures; third to sixth terga with small, shallow, separated punctures; last tergum with small, dense, con- tiguous to confluent punctures; first sternum with a distinct, median, longitudinal carina, not no- ticeably elevated anteriorly, and with sparse, pale pubescence; second sternum with moderate, dis- tinct, contiguous punctures and sparse, erect and suberect pubescence throughout, the pos- terior margin with a fringe of pale pubescence, thin medially, but broadened and thickened to- wards the lateral margins; third to sixth sterna with small, shallow punctures; last sternum with moderately small, contiguous punctures. Wings subhyaline; cell 2nd R.1+R2 about two and one-half times as long as wide, truncate at the tip; cell R5 receiving vein Mh+r about three- fifths the distance from base to apex; cell Rr much less distinct than R5 and receiving vein M2 at the middle. Legs black, sparsely clothed with pale hairs; posterior coxae without a longitudinal carina on the inner surface; calcaria pale. Holo type.— Male, Flores, Manaos, Amazon- as, Brazil, July 29, 1924 (J. Bequaert), in col- lection of University of Minnesota. Paratype.— Male, Kartabo, British Guiana, July-August, 1920 (W. M. Wheeler). Resembles infracta and emarginata but differs from both in the conspicuous, subconical tu- bercle on the scutellum and the absence of a median tubercle at the posterior margin of dor- sum of propodeum. Ephuta depressa, new species Male— Holotype. Entirely black, except the anterior two-thirds of last sternum dark ferru- ginous; anterior third of scutellum with a me- dian, longitudinal, elevated line terminating in a subconical tubercle just anterior to the middle of the scutellum; postero-lateral angles of dor- sum of propodeum not spinose nor dentate; pos- terior margin of dorsum of propodeum not den- tate nor tuberculate medially; tegulae with small, close, somewhat indistinct punctures on the disc, becoming sparse towards the margins; disc of second tergum distinctly flattened. Length, 8.5 mm. Head black; lateral areas of clypeus clothed with thick, long, subappressed, pale, silky hairs, the median area very sparsely so; front anterior to emargination of eyes with thick, appressed, pale pubescence, the posterior area between emargination of eyes and anterior ocellus with sparse, erect, pale hairs; ocellar area with sparse, erect, pale hairs; genae with thick, appressed, pale pubescence and sparse, erect, pale hairs; mandibles castaneous, darker at the tip, edentate at the tip and with a small tooth within, and clothed with pale pubescence almost to the tip; clypeal carinae slightly diverging anteriorly; scape strongly bicarinate beneath; first segment of flagellum approximately equal in length to the second; front, vertex and genae with mod- erate, contiguous punctures; carinae separating anterior half of genae from proboscidal fossa distinct, but not high and conspicuous. Thorax entirely black; dorsum of pronotum with thin, appressed, pale pubescence and sparse, erect, pale hairs; mesonotum with sparse, 1 952] Mickel: Mutillidae (Wasps) of British Guiana 115 erect, fuscous hairs, except the postero-lateral angles with a tiny spot of dense, appressed, pale pubescence; scutellum with sparse, long, erect, pale hairs, except the subconical elevation with sparse, black hairs; dorsum and posterior face of propodeum with thick to dense, short, sub- appressed, pale pubescence and sparse, erect, pale hairs; lateral aspect of pronotum with thin, short, appressed, pale pubescence; mesopleura with thick, short, appressed, pale pubescence and sparse, erect, pale hairs; metapleura and sides of propodeum with thin, appressed, pale pubescence; dorsum of pronotum with mod- erate, contiguous to confluent punctures; mes- onotum with six longitudinal rows of moderately large, shallow, confluent punctures; anterior third of scutellum with a median, longitudinal glabrous line, rising abruptly and terminating in a subconical tubercle just anterior to the middle; scutellum othewise with moderate, dense, contiguous punctures; postero-lateral angle of dorsum of propodeum not spinose or dentate, the lateral margins somewhat crenu- late, the posterior margin without a median tubercle; dorsum and posterior face of propo- deum reticulate; lateral aspect of pronotum gla- brous and micropunctate, except the dorsal margin punctate like the dorsum of pronotum; mesopleura evenly convex throughout and with moderate, contiguous punctures; metapleura glabrous, micropunctate, except for a few mod- erately large punctures ventrally; sides of pro- podeum reticulate; tegulae convex, the disc with small, close, somewhat confluent punctures, be- coming sparse towards the margins. Abdomen black, except the anterior two- thirds of last sternum dark ferruginous; first abdominal segment cylindrical, slightly broader than long when viewed from above; first tergum clothed with thick, appressed, pale pubescence and sparse, long, erect, pale hairs; disc of second tergum with sparse, inconspicuous, pale hairs anteriorly, the hairs gradually changing to fus- cous towards the middle and then to black pos- teriorly, the lateral areas of the tergum with sparse, erect, pale hairs, the lateral margins with a line of thin, appressed, pale pubescence, and the posterior margin with a band of thick, ap- pressed, pale pubescence, broad at the sides and narrow medially; third to sixth terga with sparse, pale hairs and each with sparse, inconspicuous, black hairs submarginally posteriorly; last ter- gum with sparse, pale hairs and scattered, in- conspicuous, black hairs; first tergum with mod- erate, separated punctures visible through the pale pubescence; second tergum with the ante- rior three-fourths of the disc distinctly flattened, the entire tergum with moderate, distinct, close, deep punctures; third to sixth terga with small, sparse punctures; last tergum with small, close punctures; first sternum with a strong, median, longitudinal carina very slightly elevated ante- riorly, the lateral areas distinctly punctate and clothed with sparse, pale pubescence; second sternum with a large, anterior area clothed with thin, appressed, pale pubescence extending from the proximal margin to distinctly beyond the middle on the midline, but only one-third the length of the sternum at the lateral margins; remainder of second sternum with sparse, pale, subappressed hairs, except the lateral thirds of the posterior margin with an indistinct band of thin, appressed, pale pubescence; second ster- num with moderate, deep punctures throughout, the punctures separated anteriorly, close pos- teriorly; third to sixth sterna with small, sparse punctures and clothed with sparse, pale hairs; last sternum with small, close punctures and sparse, pale hairs. Wings subhyaline; cell 2nd Ri + Re slightly more than three times as long as wide, truncate at the tip; cell R5 receiving vein M3+4 three-fifths the distance from base to apex; cell Ri less dis- tinct than R5 and receiving vein M2 beyond the middle. Legs blackish, sparsely clothed with pale pu- bescence; calcaria pale. Holotype. — Male, Bartica, British Guiana, May 17, 1901, in collection of American En- tomological Society, Philadelphia, Pa. Paratypes.— Two males, Para, Brazil, March 6 and 8, 1902 (Ducke). Related to subconica but differs in the smaller subconical tubercle of the scutellum, the dis- tinctly depressed disc of the second tergum, the appressed pubescent area on the second ster- num, and the largely ferruginous last sternum. Ephuta emarginata, new species Male.— Holotype. Entirely black, except the anterior two-thirds of the last sternum yellow; scutellum not evenly convex, with a dorsal, pos- terior and lateral faces; dorsal face of scutellum very large, flat, the posterior face short and almost perpendicular, the two separated by a weak, crenulate ridge interrupted medially by a shallow emargination; dorsum of propodeum separated from the sides of propodeum on the posterior half by a weak, crenulate ridge, the postero-lateral angles not dentate nor spinose, but with a distinct tubercle at the tip of the en- closed area; tegulae evenly convex, with small punctures, the latter somewhat close on the disc, but very sparse at the outer lateral and posterior margins; disc of second abdominal tergum evenly convex, not depressed. Length, 9 mm. 116 Zoologica: New York Zoological Society [37: 11: Head black; lateral areas of clypeus clothed with thick, long, subappressed, pale, silky hairs, the median area sparsely so; head clothed with appressed, and sparse, erect, pale hairs, the ap- pressed pubescence thick to dense on the ante- rior half of front, the posterior half of vertex, and the genae, sparse on the posterior half of front and anterior half of vertex; a transverse area through the ocelli and between the com- pound eyes with the sparse, erect hairs fuscous; mandibles black on the proximal half to two- thirds, the distal third to half light brownish- yellow, with the tips castaneous; mandibles clothed with sparse, pale hairs, edentate at the tip and with a small tooth within; median carinae of clypeus subparallel anteriorly but roundly, slightly dilated on the posterior third and then joined in a subangulate loop at the posterior margin; dilated sector of each median carina with a carina extending laterad from it on to lateral areas of clypeus; antennal scrobes carinate above; scape strongly bicarinate be- neath; first and second segments of flagellum approximately equal in length; front, vertex and genae with moderate, distinct, contiguous punc- tures; anterior half of genae separated from the proboscidal fossa by a high carina extending caudad from the articulation of the mandible. Thorax entirely black; dorsum of pronotum clothed with moderately thick, appressed, pale pubescence and sparse, erect, pale hairs; ante- rior and lateral margins of mesonotum with thin, pale pubescence, the small postero-lateral angles with dense, pale pubescence; mesonotum otherwise with sparse, erect, fuscous hairs; scu- tellum with sparse, erect, pale hairs except the lateral and posterior faces with the erect hairs thicker than elsewhere, and the disc of the dorsal face with sparse, erect, fuscous hairs; dorsum of propodeum with thick, subappressed, pale pubescence; posterior face and sides of propo- deum with thin, similar pubescence; pleural areas with thin, short, subappressed, pale pubes- cence and sparse, erect, pale hairs; pronotum with moderate, contiguous punctures visible through the pale pubescence; mesonotum with six longitudinal rows of large, shallow, contig- uous to confluent punctures; scutellum with moderate, contiguous to confluent punctures on the dorsal face, the punctures somewhat deep- er and smaller on the posterior face, and small on the lateral faces; dorsal and posterior faces of scutellum separated by a weak crenulate ridge, interrupted medially by a shallow emar- gination; dorsum of propodeum reticulate, with a median, enclosed space terminating posterior- ly at the base of a conspicuous tubercle; dor- sum and posterior faces of propodeum separ- ated by a distinct, transverse ridge extending laterad each side from the median tubercle; posterior face of propodeum reticulate; lateral aspect of pronotum micropunctate, except dor- sally the punctation of the dorsal face extend- ing somewhat onto the lateral aspect; meso- pleura evenly convex throughout, and with moderate, contiguous to slightly separated punctures visible through the pubescence; metapleura glabrous, micropunctate, except ventrally with large, shallow, contiguous punc- tures; sides of propodeum reticulate through- out; tegulae large, glabrous, with small punc- tures, close on the disc and inner margins, sparse at the outer and posterior margins, clothed with sparse, fuscous pubescence, except the inner and anterior margins slightly pale pubescent. Abdomen black, except the anterior two- thirds of the last sternum yellowish; first ab- dominal tergum broader than long, and with moderate, contiguous punctures on the dorsal face visible beneath the thick, appressed, pale pubescence and sparse, erect, pale hairs; sec- ond tergum evenly convex throughout, the disc not at all depressed; disc of second tergum with moderate, distinct, slightly separated punctures, the latter becoming smaller and closer toward the lateral and posterior margins; second tergum clothed with sparse, erect, fuscous hairs on the disc, and sparse, erect pale hairs elsewhere, ex- cept the lateral margins with a very narrow band of thin, appressed, pale pubescence and the posterior margin with a conspicuous, com- plete narrow band of thick, appressed pale pu- bescence, the band slightly broader laterally than at the middle; third to sixth terga with small distinct punctures and each clothed with sparse, erect, pale hairs, except a few, incon- spicuous black hairs at the posterior margin of each; last tergum with small, close, distinct punctures and sparse, erect, pale hairs; first sternum with a distinct, median, longitudinal carina, only slightly elevated anteriorly, the sternum clothed with sparse, pale hairs; second sternum with moderate, close to contiguous punctures and sparse, erect, pale hairs, except the anterior third also with thin, appressed pale pubescence; sterna three to six with small punc- tures and sparse, erect, pale hairs; last sternum with moderate contiguous punctures and sparse, erect, fuscous hairs. Wings subhyaline; cell 2nd R1+R2 three times as long as wide, truncate at the tip; cell Rr, receiving vein M3+4 distinctly beyond the middle; cell Rt less distinct than Rs and receiv- ing vein M2 distinctly beyond the middle. Legs black, sparsely clothed with pale hairs; posterior coxae with a weak, longitudinal car- ina on the inner surface; calcaria pale. 1952] Mickel: Mutillidae (V/asps) of British Guiana 117 Holotype — Male, Blairmont (near New Am- sterdam), British Guiana, September, 1923 (F. X. Williams), in collection of University of Minnesota. Paratypes.— Two males, Blairmont, British Guiana, August, 1923 and September, 1923 (F. X. Williams); male, Kartabo, Bartica dis- trict, British Guiana, July 3, 1922 (Wm. Beebe); male, Brasso, Trinidad, B. W. I., No- vember 23, 1940 (E. McC. Callan) ; male, Mont- serrat, Trinidad, B. W. I., June 30, 1905 (Aug- ust Busck) . Very similar in appearance to infracta but differs in the slightly emarginate scutellum, the presence of a carina on the lateral areas of clypeus extending out from the dilated sector of the median carinae and the slightly less prom- inent crenulate ridge separating the posterior half of the dorsum of propodeum from the sides of propodeum. Ephuta infracta, new species Male.— Holotype. Entirely black, except the anterior two-thirds of the last sternum yellow; scutellum evenly convex, without a posterior face and without a median, posterior emargina- tion, densely punctate throughout; dorsum of propodeum separated from the sides of the pro- podeum on the posterior half by a crenulate ridge, the postero-lateral angles not dentate or spinose, but with a distinct, median, blunt tu- bercle at the tip of the enclosed area; tegulae evenly convex, with small, shallow punctures close on the disc and sparse posteriorly; disc of second abdominal tergum evenly convex, not depressed. Length, 10 mm. Head black; clypeus clothed with thick, long, subappressed, pale, silky hairs; front clothed with appressed, pale pubescence, thick ante- riorly on the antennal tubercles, becoming gradually thinner towards the anterior ocellus; vertex with sparse, erect, pale hairs and a few, erect, fuscous hairs on the ocellar area; genae clothed with thick appressed, and sparse, erect, pale pubescence; mandibles black, the tips cas- taneous, clothed with thick, pale hairs, except at the tip, edentate at the tip and with a small tooth within near the tip; median carinae of clypeus parallel, extending two-thirds the length of the clypeus from the posterior margin; an- tennal scrobes carinate above; scape strongly bicarinate beneath; first and second segments of flagellum approximately equal in length; front, vertex and genae with moderate, distinct, contiguous punctures; anterior half of genae separated from the proboscidal fossa by a high carina extending caudad from the articulation of the mandible. Thorax entirely black; dorsum of pronotum clothed with thin, appressed, pale pubescence, sparse, long, erect, pale hairs, and the posterior margin with a few, long, erect, fuscous hairs; anterior and lateral margins of mesonotum with thin, appressed, pale pubescence, the remainder with sparse, erect, black hairs; scutellum clothed with thin, erect, pale hairs and with a few, erect, black hairs at the middle anteriorly; re- mainder of thorax clothed with pale pubescence, sparse and short on lateral aspect of pronotum, dense and appressed on mesopleura, sparse, ap- pressed and scattered, erect on the metapleura and sides of propodeum, thin, appressed and erect pubescence on the dorsum and posterior face of propodeum; dorsum of pronotum with moderately large, contiguous to confluent punc- tures; mesonotum with longitudinal rows of large, shallow, confluent punctures; scutellum evenly convex, without a posterior face and without a posterior median emargination, and with moderately large, contiguous punctures throughout; dorsum of propodeum with a narrow, median enclosed space about three times as long as broad and terminating at the base of a blunt, median tubercle; remainder of dorsum of propodeum and posterior face of propodeum reticulate; posterior half of dor- sum of propodeum separated from sides of propodeum by a low, crenulate ridge; postero- lateral angles of dorsum of propodeum not spinose nor tuberculate; lateral aspect of pronotum for the most part glabrous and mi- cropunctate; mesopleura with moderate, dis- tinct, contiguous punctures, mostly concealed by the dense pubescence; metapleura glabrous and micropunctate, except a few, large, shallow, confluent punctures ventrally; sides of propo- deum reticulate; tegulae large, glabrous, with fine punctures, the latter close but distinctly separated anteriorly and becoming sparse pos- teriorly; tegulae clothed with sparse, fuscous pu- bescence, except the lateral margins with the pubescence pale. Abdomen black, except the anterior two- thirds of the last sternum yellow; first tergum with moderately large, contiguous punctures, more or less concealed by dense, appressed, pale pubescence and sparse, erect, pale hairs; second tergum evenly convex, the disc not at all de- pressed, with moderate, distinct, slightly sep- arated punctures throughout, clothed with sparse, erect, pale hairs on the anterior half, and sparse, fuscous to black hairs on the pos- terior half, except the broad, lateral margins with thin, appressed, pale pubescence, and the posterior margin with a band of dense, pale, appressed pubescence, the posterior band broad at the sides and gradually narrowed towards the 118 Zoologica: New York Zoological Society [37: 11: middle to a mere, marginal fringe; terga three to six with small, distinct separated punctures and sparse, appressed and scattered, erect pale hairs, except the lateral margins with thick, ap- pressed, pale pubescence and the posterior mar- gin of each with scattered, inconspicuous, erect, black hairs; last ter gum with small, distinct, sep- arated punctures and clothed with sparse, sub- appressed, pale pubescence; first sternum with a distinct, median, longitudinal carina only slightly elevated anteriorly, and clothed with thin, appressed and erect, pale pubescence; sec- ond sternum with moderate punctures, well separated anteriorly, becoming contiguous on the posterior half, clothed throughout with very thin, inconspicuous, appressed, pale pubescence and sparse, erect, pale hairs, the appressed, pale pubescence thick at the posterior margin, ex- cept at the middle; sterna three to six with small, scattered punctures and sparse, erect, pale hairs; last sternum with moderately small, contiguous punctures and sparse, erect, pale hairs. Wings subhyaline to subfuscous; cell 2nd Ri+Ra three times as long as wide, truncate at the tip; cell Rj receiving vein M3+4 slightly beyond the middle; cell R4 less distinct than Rr, and receiving vein M2 slightly beyond the mid- dle. Legs black, sparsely clothed with pale hairs; posterior coxae with a weak, longitudinal car- ina on the inner surface; calcaria pale. Holotype. — Male, Bartica, British Guiana, May 17, 1901, in collection American Entomo- logical Society, Philadelphia, Pa. Paratypes. — Male, Bartica, British Guiana, May 17, 1901; male, Bartica, British Guiana, February 12, 1913; male, Bartica, British Guiana, December 23, 1912; and male, Deme- rara River, British Guiana, March 12, 1913. Differs from perfica in the absence of a tooth at the postero-lateral angles of the dorsum of propodeum, the latter being separated on the posterior half from the sides of propodeum by a low, crenulate ridge; and in the evenly convex second tergum, the disc of the latter not at all flattened. Ephuta indiscreta, new species Male.— Holotype. Black, except the mandibles and the anterior two-thirds of the last sternum yellow; scutellum convex; postero-lateral angles of dorsum of propodeum not dentate or tuber- culate; posterior margin of dorsum of propo- deum distinctly tuberculate medially; tegulae with sparse, fine punctures throughout; disc of second tergum evenly convex, not at all flat- tened. Length, 7 mm. Head black, except the mandibles almost en- tirely yellow with the tips castaneous; lateral areas of clypeus clothed with long, subappressed, silky, pale pubescence; front and vertex clothed with thin, appressed, pale pubescence and sparse, erect, pale hairs, except the antennal tubercle area with the appressed, pale pubes- cence thick; genae with thick, appressed, pale pubescence and sparse, erect, pale hairs; man- dibles edentate at the tip and with a small tooth within; clypeal carinae subparallel; scape strong- ly bicarinate beneath; flagellum broken off of holotype, but presumably the first and second segments approximately equal in length; front vertex and genae with moderate, contiguous punctures; genae sharply separated from pro- boscidal fossa by a carina. Thorax entirely black; dorsum of pronotum clothed with moderately thick, appressed, pale pubescence, and sparse, erect, pale hairs; meso- notum clothed with sparse, intermixed, pale fuscous and black hairs, except the postero- lateral angles with a very small spot of dense, pale pubescence; scutellum clothed with sparse, erect, pale hairs; dorsum and posterior face of propodeum clothed with moderately thick, sub- appressed, pale pubescence; lateral aspect of pronotum with sparse, appressed, pale pubes- cence; mesopleura with thick, appressed, pale pubescence and sparse, erect, pale hairs; meta- pleura with very sparse, appressed, pale pubes- cence; sides of propodeum with very sparse, erect, pale hairs; dorsum of pronotum with moderate, contiguous punctures; mesonotum with six longitudinal rows of large, shallow, con- fluent punctures; scutellum convex, its posterior margin not emarginate, with moderate, contigu- ous to confluent punctures throughout; dorsum and posterior face of propodeum reticulate, the two separated by a weak, transverse ridge ele- vated medially into a distinct tubercle; lateral aspect of pronotum glabrous, micropunctate, except the dorsal margin punctate like the dor- sum of pronotum; mesopleura evenly convex, with moderate, contiguous to slightly separated punctures more or less concealed by the thick, pale pubescence; metapleura glabrous, micro- punctate, except ventrally with a few moderate, close punctures; sides of propodeum reticulate; tegulae glabrous, with sparse, fine punctures throughout, clothed with sparse, fuscous pubes- cence, except the anterior and inner margins more or less sparsely, pale pubescent. Abdomen black, except the anterior two- thirds of last sternum yellow; first segment cy- lindrical, distinctly broader than long; first ter- gum with moderate, contiguous punctures, largely concealed by the thick, appressed, pale pubescence and sparse, erect, pale hairs; second 1952] Mickel: Mutillidae (Wasps) of British Guiana 119 tergum evenly convex throughout, not at all flattened, with moderate, distinct, contiguous to slightly separated punctures throughout; second tergum clothed with sparse, erect, pale hairs, except the lateral margins with a narrow fringe of appressed, pale pubescence, and the posterior margin with a band of thick, pale pubescence, broad at the sides and gradually narrowed to a mere fringe at the middle; terga three to six with sparse, small punctures and sparse, erect, pale hairs, the fifth and sixth terga also with a few inconspicuous black hairs at the posterior margin; last tergum with small, distinct punc- tures and sparse, erect, pale hairs; first sternum with a median, longitudinal carina, not elevated anteriorly, clothed with thin, pale pubescence; second sternum with moderate, contiguous to slightly separated punctures, the anterior third and the posterior margin with thin to sparse, appressed, pale pubescence, the remainder with sparse, erect, pale hairs; sterna three to six with sparse, small punctures, and sparse, erect, pale hairs; last sternum with small, contiguous punc- tures and sparse, erect, pale hairs on the anterior two-thirds, the posterior third with sparse, fus- cous hairs. Wings su’ohyaline; cell 2nd Ri-fRs approx- imately two and one-half times as long as wide, truncate at the tip; cell R5 receiving vein M3+4 approximately three-fifths the distance from base to apex; cell R4 much less distinct than Rs and receiving vein M2 slightly beyond the mid- dle. Legs black, clothed with sparse, pale pube- scence; calcaria pale. Holotype — Male, Blairmont, British Guiana, September, 1923 (F. X. Williams), in collection of University of Minnesota. Related to infracta, but differs in having the mandibles almost entirely pale lemon yellow, with the tips castaneous. Ephuta flavidens, new species Male.— Holotype. Entirely black, except the mandibles; mandibles lemon yellow except the extreme base and tip castaneous; hypopygium entirely black; postero-lateral angles of dorsum of propodeum not dentate or prominently angu- late; posterior margin of dorsum of propodeum distinctly tuberculate medially; tegulae with sparse, fine punctures throughout; sides of pro- podeum glabrous, unsculptured on anterior half, reticulate on posterior half; disc of second ter- gum evenly convex, not at all flattened. Length, 7.5 mm. Head entirely black, except the mandibles as above; mandibles slender, edentate at the tip and with a small tooth within near the tip; cly- peal carinae subparallel, slightly diverging an- teriorly, terminating before the anterior margin in a pair of small tubercles; scape bicarinate beneath; first and second flagellar segments approximately equal in length, but each shorter than the third; genae separated from ventral aspect of head by a distinct carina, the ventral aspect between the two carinae glabrous, im- punctate; clypeus laterad of the medial carinae closely micropunctate and clothed with thick, recumbent, pale pubescence; front, vertex and genae with moderate, close, deep punctures, thick, appressed, pale pubescence (except sparse in the ocellar area) , and sparse, erect, pale hairs. Thorax entirely black; dorsum and sides of pronotum with moderately thick, appressed, pale pubescence and sparse, erect, pale hairs; mesonotum with only very sparse, erect, pale hairs; scutellum with erect, pale hairs slightly thicker than on mesonotum; dorsum and post- terior face of propodeum with thick, appres- sed, pale pubescence; mesopleura with thick, appressed, pale pubescence; metapleura and sides af propodeum very sparsely, pale pube- scent; dorsum of pronotum with moderately large, contiguous punctures; sides of pronotum closely micropunctate; mesonotum with six lon- gitudinal rows of large, shallow, confluent punc- tures; scutellum with moderately large, contig- uous, deep punctures throughout; scutellum slightly elevated posteriorly and with a short, transverse posterior face; postero-lateral angles of dorsum of propodeum not produced or angulate; dorsum and posterior face of propo- deum broadly reticulate, not separated by a transverse carina, but the posterior margin of the dorsum with a distinct, median tubercle; mesopleura with moderate, close punctures throughout; metapleura and anterior half of sides of propodeum glabrous; posterior half of sides of propodeum reticulate. Abdomen, including the hypopygium, entire- ly black; first segment cylindrical, distinctly ly broader than long; first tergum with moder- ately large, contiguous punctures and thick, ap- pressed, pale pubescence, as well as sparse, erect, pale hairs; second tergum evenly convex throughout, not at all flattened, with moderate, slightly separated punctures on the disc becom- ing close and contiguous laterally and poster- iorly, clothed with sparse, pale hairs, except the lateral margins with a narrow band of dense, appressed, pale pubescence, and the posterior margin with a band of similar pubescence, the band broad at the sides and narrowed to a mere fringe at the middle; terga three to six with sparse, very small punctures and sparse, erect, pale hairs; last tergum with small, close punc- tures (distinctly larger than on terga three to 120 Zoologica: New York Zoological Society [37: 11: six), and sparse, pale pubescence; first sternum with the median carina strongly elevated anter- iorly; second sternum with moderately large, close punctures, the anterior two-thirds with thin, appressed, pale pubescence and sparse, erect, pale hairs, the posterior third with only sparse, erect, pale hairs; sterna three to six with sparse, small punctures and sparse, erect, pale hairs; hypopygium with small, close punctures and sparse, pale hairs. Wings subhyaline; cell 2nd R1+R2 approx- imately two and one-half times as long as wide, subtruncate at the tip; cell Rs receiving vein M3+4 approximately three-fifths the distance from base to apex; cell R4 obsolete. Legs black, clothed with sparse, pale pub- escence; calcaria pale. Holotype.—Ma.\Q, Moraballi creek, Essequibo river, British Guiana (1742), September 7, 1929 (O. W. Richards), in collection of Uni- versity of Minnesota. Parcitypes— Male, Moraballi creek, Essequibo river, British Guiana (3019), September 15, 1929 (O. W. Richards), three males, Trinidad Rio, Panama, March 19, 1912, March 20, 1912 and March 29, 1912 (A. Busck) ; male, San- tarem, Brazil. Resembles indiscreta in the yellow mandibles, but differs in having the hypopygium entirely black, only the posterior half of the sides of propodeum reticulate, the scutellum slightly ele- vated posteriorly, and the median carina of the first sternum strongly elevated anteriorly. Ephuta erichtho, new species Female— Holotype. Head, abdomen and legs black, the thorax ferruginous with the ventral margins of the pleural areas slightly infuscat- ed; first abdominal segment broader than long; second tergum immaculate, except the lateral margins narrowly and the posterior margin broadly, pale pubescent; second tergum with a small, inconspicuous fovea near the lateral mar- gin, the fovea approximately one puncture wide and two punctures long, not filled with pube- scence. Length, 10 mm. Head black, except the mandibles medially and the flagellum beneath tinged with ferrugi- nous; mandibles edentate at the tip and with a small tooth within near the tip; posterior margin of clypeus elevated, arcuate and serrate, the serrations formed by five emarginations, one at the median line and two on each lateral, area; mandibles and clypeus with sparse, long, pale hairs; antennal scrobes not carinate above; first segment of flagellum short, equal in length to the second segment from some aspects, slight- ly longer than the second from other aspects; front, vertex and genae with dense, very deep punctures; anterior half of front, and vertex and genae entirely with sparse, short, appres- sed, pale pubescence, and scattered, erect, pale hairs; posterior half of front with sparse, erect, fuscous hairs; genae defined posteriorly by a strong carina, the latter terminating anteriorly in a prominent tooth ventrad of the articulation of the mandible where the genal carina unites with the carina bounding the proboscidal fossa. Thorax entirely ferruginous, except the vent- ral margins of the pleural areas somewhat infus- cated; dorsal and pleural areas of thorax with dense, deep punctures throughout; dorsum clothed with sparse, erect, fuscous hairs; anter- ior and propodeal spiracular tubercles only slightly elevated, not forming prominent teeth; scutellar scale entirely absent; pleural areas clothed with sparse, appressed, pale, pubes- cence, and scattered, erect, pale hairs. Abdomen black; first segment viewed from above, broader than long; first tergum with dense, deep punctures, excepting the posterior margin, and clothed with thick, appressed, pale pubescence, as well as scattered, long, erect, pale hairs; second tergum with dense, deep punc- tures, clothed with sparse, black, erect and ap- pressed pubescence, except narrow lateral mar- gins, a broad, complete band at the posterior margin, and the postero-lateral areas with thick, appressed, pale pubescence; second tergum with a small, inconspicuous fovea near each lateral margin, the fovea approximately one puncture wide, two punctures long, and not filled with pu- bescence; terga three to five with moderate, shallower, elongate, contiguous punctures, sparsely clothed with pale fuscous hairs, except each with an inconspicuous, median spot of thin, appressed, pale pubescence; last tergurn with shallow, dense, contiguous to confluent punc- tures, and a distinct, but not prominent, median, longitudinal ridge extending its entire length; pygidial area defined laterally by a distinct ca- rina, the pygidial area thus not evenly convex, but with a median, longitudinal ridge, a longitu- dinal depression each side, and then the lateral carinae; last tergum clothed with sparse, erect, pale fuscous hairs; first sternum with a very weak, longitudinal, median carina produced an- teriorly into a small tooth; second sternum with moderately large, close, distinct, deep punctures, clothed with scattered, erect, pale hairs and with sparse, appressed, pale pubescence, the latter absent on the disc; sterna three to five with small, shallow, close punctures and sparse, pale pubes- cence; last sternum tinged with ferruginous, and with a pair of distinct tubercles anteriorly. Legs black, clothed throughout with sparse, pale pubescence; posterior tibiae slightly pro- duced externally at the tip to form an acute 1952] Mickel: Mutillidae (Wasps) of British Guiana 121 angle, and bearing a single, short, weak spine; calcaria pale. Holotype.— Female, Kartabo, Bartica district, British Guiana, March 12, 1924 (Wm. Beebe), in collection of University of Minnesota. Related to the following species, but differs • in the very small inconspicuous, lateral foveae of the second tergum, the foveae not filled with pubescence, and in the absence of anterior pale pubescent spots on the second tergum. Ephuta weberi, new species Female— Holotype. Head and abdomen black; thorax ferruginous, the lateral aspects with obscure blackish markings ventrally; sec- ond tergum with a pair of lateral, narrow, elon- gate pits, one near each lateral margin; second tergum maculated with a pair of anterior, sub- triangular, pale pubescent spots and a broad, posterior marginal, pale pubescent band; second sternum with a distinct, median carina on the anterior fourth; last sternum with a pair of small, anterior tubercles; intermediate coxae each with a small tubercle at the outer, distal margin. Length, 12 mm. Head black, except the mandibles dark fer- ruginous with blackish tips, and the flagellum ferruginous beneath; mandibles edentate at the tip and with a small tooth within near the tip; clypeus elevated posteriorly, the elevated margin distinctly emarginate laterally, not quadriden- tate; first segment of flagellum almost twice the length of the second; antennal scrobes not cari- nate above; genal carina elevated immediately posterior to insertion of the mandibles into a very prominent tooth; front, vertex and genae with moderate, distinct, contiguous punctures, the latter deeper on the vertex and genae; man- dibles clothed with sparse, pale hairs; clypeus anterior to elevation clothed with thick, erect, fulvous hairs; scape clothed with sparse, pale pubescence; front and vertex clothed with thick, appressed, pale golden pubescence, the an- terior half of front also with sparse, erect, pale hairs, the remainder of front and vertex with sparse, erect, fuscous hairs; genae with thick, appressed, pale pubescence and sparse, erect, pale hairs. Thorax ferruginous, the ventral area of the sides of propodeum and mesopleural areas black, a stripe of black extending dorsally from the an- tero-ventral angle of mesopleura to beneath the anterior spiracular tubercles and continuing across the dorsum at the anterior margin of pro- notum as a narrow obscure stripe; dorsum and lateral aspects of thorax with moderate, conti- guous punctures, slightly smaller and much dens- er on the dorsum than on the lateral aspects; dor- sum clothed with sparse, short, suberect, fuscous hairs, and sparse, long, erect, black hairs; lateral aspects clothed with thick, very short, appressed, pale pubescence, and sparse, erect, pale hairs, the punctation not obscured by the pubescence; propodeal spiracular tubercles large, prominent, yellowish. Abdomen black, except the first segment fer- ruginous; first segment distinctly broader than long; first tergum with moderate, dense, deep punctures, clothed with sparse, pale golden pu- bescence and sparse, erect, pale hairs, and with a posterior marginal band of thick, appressed, pale golden pubescence; second tergum with a very narrow, elongate pit (six times as long as wide) each side near the lateral margin; second tergum with moderate, close but slightly separ- ated, deep punctures, clothed with thick, sub- appressed, black pubescence and sparse, erect, black hairs, except the erect hairs on the an- terior third pale, a pair of anterior, subtriangular spots of pale golden pubescence, the lateral margins with a band of thick, appressed, pale golden pubescence, and a very broad, posterior marginal band of pale golden pubescence; terga three to five with moderate, close, deep punc- tures anteriorly and medially, clothed with thick, pale golden pubescence and sparse, erect, pale to fuscous hairs, except each with a pah of lateral spots of black pubescence and sparse, erect, black hairs, thus isolating a broad, median, pale spot and a pair of lateral, marginal, pale spots; last tergum ferruginous along the mid- line on the posterior half, with small close punc- tures laterad and anterior to pygidial area, and clothed with sparse, erect, golden hairs, except at the midline the pubescence thick; pygidial area narrow, elongate, glabrous on the anterior half, shagreened and margined laterad on the posterior half, the anterior half concealed by the pubescence; first sternum with the median longi- tudinal carina almost obsolete on the posterior half, elevated and conspicuous on the anterior third; second sternum with a median, longitu- dinal carina on the anterior fourth, with well separated, moderate punctures, except at the lateral and posterior margins the punctures close, and clothed with sparse, erect, pale hairs, ex- cept the anterior, lateral, and posterior margins with thin, appressed, pale pubescence; sterna three to five with small punctures and sparse, erect, pale hairs; last sternum with a pair of small, anterior tubercles, with small, close punc- tures and sparse, erect, pale hairs. Legs black, clothed with sparse, pale pubes- cence; middle coxae with a small tubercle at the outer distal margin; calcaria pale. Holotype.— Female, Oko River, British Gui- ana, June 25, 1936 (N. A. Weber), in collection of University of Minnesota. 122 Zoologica: New York Zoological Society [37: 11: Related to teapae Cameron and tolteca Blake from Mexico, but differs from both in the black head, abdomen and legs, and the ferruginous thorax, teapae being entirely black and tolteca entirely ferruginous; both the latter also have the dorsum of thorax somewhat marked with lines or spots of pale pubescence. Ephuta aurulenta (Fabricius) , (new combination) 1805. Mutilla aurulenta Fabricius, Syst. Piez., p. 436, female. Specimens Examined— Female, Kartabo, Bar- tica district, British Guiana, October 11, 1920; female, Kartabo, British Guiana; female, Bar- tica district, British Guiana, November 5, 1924; female, Tropical Research Station, N. Y. Zoolo- gical Soc., British Guiana, no. 2266; female, Essequibo, British Guiana; female, Blairmont, British Guiana, November, 1923 (F. X. Wil- liams) . Ephuta roxane, new species Female— Holotype. Ferruginous, except the vertex, genae, a transverse stripe on the prono- tum, the disc and anterior area of second tergum black, and terga three to five very dark ferrugi- nous or blackish; clypeus elevated posteriorly, the elevated margin quadridentate; first and sec- ond flagellar segments approximately equal in length; first abdominal segment broader than long; second abdominal tergum with a posterior marginal band of thick, appressed, pale golden pubescence, sub interrupted medially with black; terga three to five, each with a large, transverse, median spot of thick, appressed, pale golden pubescence. Length, 6 mm. Head ferruginous, except the vertex and genae black; mandibles blackish at the tip, eden- tate at the tiD and with a minute tooth within a near the tip; clypeus elevated posteriorly, the elevated margin quadridentate; antennal scrobes not carinate above; first and second segments of flagellum approximately equal in length; front, vertex and genae with small, distinct, deep, con- tiguous punctures, more or less concealed by the pubescence; genal carina extending posteriorly from insertion of mandibles and elevated into a prominent tooth one-third its length from mandibles; mandibles and clypeus with sparse, erect, pale hairs; scape with thin, appressed, pale golden pubescence; front, vertex and genae with thick, appressed, pale golden pubescence and sparse, erect, pale hairs, the pubescence becoming whitish towards the genal carina. Thorax entirely ferruginous, except a trans- verse, black stripe on the pronotum; dorsum and lateral aspects of prothorax with small, distinct, deep, contiguous punctures, except those on the posterior area of mesonotum, the dorsum of propodeum and posterior face of propodeum very dense and somewhat confluent; scutellar scale absent; propodeal spiracular tu- bercles small, yellow; pronotal area, anterior half of mesonotal area, and lateral aspects of thorax with thin, appressed, pale pubescence, , and sparse, erect, very pale golden hairs; re- mainder of dorsum of thorax with sparse, erect, inconspicuous, very pale golden hairs. Abdomen ferruginous, except all the second tergum, excluding the broad, lateral and pos- terior margins, blackish, and terga three to five very dark ferruginous, almost blackish; first abdominal segment distinctly broader than long; first and second terga and anterior half of terga three to five with small, distinct, deep, contigu- ous punctures; first tergum clothed with thin, appressed, pale pubescence and sparse, erect, pale hairs; anterior half and lateral fifths of second tergum with thin, appressed, pale pubes- cence and sparse, erect, pale hairs; posterior margin of second tergum with a broad band of thick, appressed, pale golden pubescence, the anterior margin of the band deeply emarginate medially, subinterrupted; remainder of disc of second tergum with thin, appressed, black pu- bescence, and sparse, erect, dark fuscous hairs; terga three to five each with a median, trans- verse spot of thick, appressed, pale golden pu- bescence occupying almost the median half of each tergum, the lateral fourths with dark fus- cous pubescence; last tergum with sparse, erect, fuscous hairs, except the margins of the pygidial area with a few pale hairs and a small median spot of pale pubescence anterior to the pygidial area: pygidial area glabrous, the posterior fourth of the lateral margins defined by a carina; first sternum with the median, longitudinal carina almost obsolete on the posterior half, but strongly elevated and very prominent on the anterior half; first sternum clothed with thin, mostly erect, pale pubescence; second sternum with small, distinct, deep, contiguous punctures throughout, clothed with thin, appressed, pale pubescence and sparse, erect, pale hairs; lateral areas of sterna three to five with distinct, deep, contiguous punctures and clothed with thin, appressed, pale pubescence, the median area of each glabrous, with only a few, fine punctures and a few, erect, pale hairs; last sternum with small punctures and sparse, erect, pale golden hairs. Legs ferruginous, sparsely clothed with pale hairs; calcaria pale. Holotype— Female, Moraballi creek, Esse- quibo river, British Guiana, September 25, 1929 (O. W. Richards) , in collection of University of Minnesota. 1952] Miclcel: Mutillidae (Wasps) of British Guiana 123 Related to elvina, copia and limbata, but dif- fers in having the front ferruginous, the black maculation of the thorax limited to a transverse stripe on the pronotum, and in the pubescent maculation of the abdominal terga. The thorax is shorter than in the above species and the posterior face of propodeum is subperpendicular to the dorsum. Ephuta egeria, new species Female.— Holotype. Head, abdomen and legs very dark ferruginous, blackish, the thorax en- tirely bright ferruginous; first and second seg- ments of flagellum equal in length; first ab- dominal segment broader than long; second abdominal tergum without a pair of anterior, pale pubescent spots, but with a complete, pos- terior marginal band of pale pubescence; ab- dominal terga three to five with a small, median spot of pale pubescence. Length, 4 mm. Head very dark ferruginous, blackish, except the mandibles and scape ferruginous, the tips of the mandibles castaneous; mandibles eden- tate at the tips and with a minute tooth within; clypeus elevated posteriorly, the elevated margin quadridentate; first and second segments of flagellum approximately equal in length; an- tennal scrobes not carinate above; front, vertex and genae with small, deep, contiguous punc- tures; genal carina extending posteriorly from the insertion of the mandibles and elevated into a prominent tooth one-third of its length from the mandibles; mandibles and clypeus clothed with long, sparse, pale hairs; scape clothed with sparse, pale pubescence; antennal scrobes and anterior margin of front with sparse, pale pu- bescence; remainder of front and vertex with sparse, short, fuscous pubescence, and scattered, erect, fuscous hairs; genae with sparse, pale pubescence and sparse, erect, pale hairs. Thorax entirely bright ferruginous; lateral and dorsal aspects of thorax with small, deep, con- tiguous punctures; lateral aspects of thorax with sparse, short, pale pubescence and sparse, erect, pale hairs; dorsal aspect of thorax with sparse, short, fulvous inconspicuous pubescence, as well as scattered, long, erect, dark fulvous hairs; pro- podeal spiracular tubercles small, concolorous with the thorax. Abdomen very dark ferruginous, blackish; first segment distinctly broader than long; first tergum ferruginous, with small, deep, contigu- ous punctures, clothed with sparse, short, pale pubescence and sparse, erect, pale hairs; second tergum with small, deep, contiguous punctures throughout, clothed with sparse, short, recum- bent, blackish pubescence, the anterior third also with sparse, erect, pale hairs, the posterior two-thirds, except the posterior margin, with sparse, erect, dark fuscous hairs, with an elon- gate spot of pale pubescence near the lateral margins, and the posterior margin with a nar- row, complete band of thick, pale, appressed pubescence slightly narrowed medially; terga three to five with small, close punctures an- teriorly, each clothed with sparse, short, dark fuscous pubescence and sparse, erect, fuscous hairs, except a small, median spot of thick, pale pubescence; last tergum with small, deep, con- tiguous punctures laterad of the pygidial area and with sparse, erect, pale hairs; pygidial area elongate, glabrous, the posterior third weakly shagreened and very weakly margined laterally; first sternum with a complete median, longitu- dinal carina, not elevated anteriorly; second sternum with small, deep, contiguous punctures and sparse, erect, pale hairs; posterior margin of sterna three to five with small, close punctures and sparse, erect, pale hairs; last sternum gla- brous, closely punctate at the lateral margins, and with a pair of small, distinct, anterior tu- bercles. Legs dark ferruginous, blackish, clothed with sparse, erect, pale hairs; calcaria pale. Holotype— Female, New Amsterdam, British Guiana, July, 1923 (F. X. Williams), in collec- tion of University of Minnesota. Distantly related to hylonome. Differs from that and other small British Guiana females of this genus in the entirely ferruginous thorax, the small, median spots of pale pubescence on abdominal terga three to five, the dark ferrugi- ous, blackish legs, and the anterior tubercles of the last sternum. Ephuta copia, new species Female— Holotype. Head and second abdom- inal tergum black; posterior abdominal terga and second sternum more or less infuscated; thorax ferruginous with lateral fuscous mark- ings; clypeus elevated posteriorly, the elevated margin dentate; first and second segments of flagellum approximately equal in length; first abdominal segment distinctly broader than long; second abdominal tergum without a pair of pale pubescent spots, the posterior marginal, pale pubescent band narrowly interrupted medially with black. Length, 8 mm. Head black, except the mandibles with dark tips, clypeus, antennal tubercles and antennae, all ferruginous; mandibles edentate at the tip and with a small tooth within, clothed with sparse, pale hairs; clypeus elevated posteriorly, the elevated margin quadridentate; first and second segments of flagellum approximately equal in length; antennal scrobes not carinate above; front, vertex and genae with moderate, distinct, deep, contiguous punctures, front and 124 Zoologica: New York Zoological Society [37: 11: vertex with sparse, appressed, pale pubescence, and sparse, erect, blackish hairs; genae with thick, appressed, pale pubescence and sparse, erect, pale hairs; genal carina extending pos- teriorly from the insertion of the mandibles and elevated into a distinct small tooth at one-third its length from mandible. Thorax ferruginous, with three pairs of lateral, obscure, fuscous spots on the pronotal area, mesonotal area and propodeum respectively, the propodeal spots less distinct than the others; dorsum and lateral aspects of thorax with mod- erate, distinct, deep, contiguous punctures throughout; prothorax with a narrow band of thick, appressed, pale pubescence at its anterior margin, the band more conspicuous on the lat- eral aspects than on the dorsum; dorsum of thorax with sparse, erect, dark hairs; lateral aspects of thorax with very sparse, inconspicu- ous, appressed, pale pubescence and sparse, erect, pale hairs; scutellar scale absent; propo- deal spiracular tubercles small, yellow. Abdomen ferruginous, except the second ter- gum black, and the following terga and second sternum more or less infuscated; first abdominal segment distinctly broader than long; first ter- gum with moderate, distinct, deep, contiguous punctures, clothed with thick, appressed, pale pubescence and sparse, erect, pale hairs; lateral and posterior margins of the black second ter- gum ferruginous, but the color obscured by thick, appressed, pale pubescence, the pale pu- bescent, posterior marginal band narrowly in- terrupted medially with black; second tergum with moderate, distinct, deep, contiguous punc- tures throughout, the bottoms of the punctures showing ferruginous in certain lights, the disc clothed with sparse, erect, blackish hairs, the lateral and anterior areas with sparse, erect, pale hairs; terga three to five with moderately small, deep punctures anteriorly, with sparse, erect, pale hairs, and each with a pair of small, obscure, median spots of thick, appressed, pale pubescence, the spots separated at the middle by sparse, blackish pubescence; last tergum with small, distinct punctures, sparse, erect, fuscous hairs laterally, and pale, erect hairs towards the middle; pygidial area narrow, very slightly ele- vated at its anterior terminus; second sternum with moderate, distinct, close punctures, clothed throughout with thin, appressed, pale pubes- cence, and sparse, erect, pale hairs; sterna three to five with small, close punctures and sparse, erect, pale hairs at the posterior margin; last sternum with shallow, indistinct punctures and erect, pale golden hairs, without anterior lateral tubercles. Legs ferruginous, sparsely clothed with pale hairs; calcaria pale. Holotype.— Female, Blairmont, British Gui- ana, October, 1923 (F. X. Williams), in collec- tion of University of Minnesota. Paratypes.— Female, Blairmont, British Gui- ana, October, 1923 (F. X. Williams); female, Blairmont, British Guiana, August, 1923 (H. E. Box). Related to elvina, but differs in the infuscated markings of the thorax, the interrupted, pos- terior marginal band of the second abdominal tergum, the paired pale pubescent spots on the third to fifth terga, the complete pale pubescent band at the anterior margin of the pronotum, its darker ferruginous color, and larger size. Ephuta elvina, new species Female— Holotype. Head and second abdom- inal tergum blackish, the remainder of the body pale ferruginous, except the thorax with a pair of lateral, vague, elongate, fuscous spots ex- tending from the pronotum to the propodeum, and the anterior two-thirds of the dorsum and posterior face of propodeum entirely infuscated; clypeus elevated posteriorly, the elevated margin dentate; first and second segments of flagellum equal in length; first abdominal segment dis- tinctly broader than long; second abdominal tergum without a pair of pubescent spots. Length, 3.5 mm. Head blackish, except mandibles yellow with castaneous tips, the antennal tubercles, scape, pedicel and first two flagellar segments yellow, and the remainder of the flagellum fuscous; front, vertex and genae clothed with thin, ap- pressed and subappressed, very pale golden pu- bescence, thinner on the posterior half of front and anterior half of vertex than elsewhere, the front and vertex also with sparse, erect, fuscous hairs; mandibles edentate at the tip and with a slight tooth within near the tip; clypeus elevated posteriorly, the elevated margin quadridentate; antennal scrobes not carinate above; scape with distinct punctures distally and clothed with sparse, pale pubescence; first and second seg- ments of flagellum approximately equal in length; front, vertex and genae with small, dis- tinct, deep, contiguous punctures, the bottoms of the punctures showing ferruginous in certain lights; genal carina extending from the insertion of the mandibles posteriorly and elevated into a distinct tooth at one-third its length from man- dible. Thorax pale ferruginous, except the dorsum with a pair of lateral, elongate, vague, infus- cated spots extending from the anterior margin to the propodeal spiracular tubercle, and the anterior two-thirds of the dorsum and posterior face of propodeum entirely infuscated; thorax clothed throughout with very sparse, appressed. 1952] Mickel: Mutillidae (Wasps) of British Guiana 125 inconspicuous, pale pubescence, and sparse, erect, pale hairs, except the dorsum of the thorax with sparse, erect, fuscous hairs, and the an- terior margin of the lateral aspect of pronotum with a narrow stripe of thick, appressed, pale pubescence; dorsum and lateral aspects of thorax with small, distinct, deep, contiguous punctures throughout; propodeal spiracular tu- bercles small, yellow; scutellar scale entirely absent. Abdomen pale ferruginous, except the second tergum infuscated, but with narrow, ferrugi- nous, posterior and lateral margins; first segment distinctly broader than long; first tergum clothed with sparse, appressed, pale pubescence and sparse, erect, pale hairs, and with small, distinct, deep, contiguous punctures throughout; second tergum with small, distinct, deep, contiguous punctures, except the latter shallow at the ferru- ginous posterior margin, the bottom of the punc- tures showing ferruginous in certain lights; second tergum clothed with very sparse, ap- pressed, pale pubescence, the large infuscated area also with sparse, erect, fuscous hairs, the ferruginous lateral margins with the pale pubes- cence thin but distinct, and the posterior ferru- ginous margin with thick, appressed, very pale golden pubescence; terga three to five with small, shallow punctures, and with thin, ap- pressed, very pale golden pubescence and sparse, erect, pale golden hairs; last tergum with small, shallow punctures and sparse, erect, pale golden hairs, except the large pygidial area glabrous, impunctate, bare; pygidial area not defined la- terally by a carina, except the posterior fifth weakly shagreened and with carinate lateral margins; first sternum with a median, longitu- dinal carina, slightly elevated at its anterior terminus; second sternum with small, distinct, deep, contiguous punctures throughout, and clothed with thin, appressed, pale pubescence and sparse, erect, pale hairs, the pale pubescence somewhat thicker at the posterior margin; sterna three to five with small punctures and sparse, erect, pale hairs; last sternum with small punc- tures and sparse, erect, pale hairs, apparently not tuberculate anteriorly. Legs pale ferruginous, sparsely clothed with pale hairs; calcaria pale. Holotype — Female, Moraballi creek, Esse- quibo river, British Guiana, September 24, 1929 (.O. W. Richards) , in collection of University of Minnesota. Distinguished from copia by the fuscous markings of the thorax, the complete pale gold- en, posterior marginal pubescent band of second tergum, and the ferruginous abdomen, except- ing the fuscous second tergum. Ephuta limbata, new species Female.— Holotype. Head and all abdominal terga, except the first, black; abdominal sterna ferruginous; thorax and first abdominal segment pale ferruginous, the dorsum of thorax with distinct, lateral, black lines united anteriorly on the pronotum and posteriorly on the propo- deum; antennal tubercles, antennae, clypeus, mandibles and legs yellow; first abdominal seg- ment distinctly broader than long; first and sec- ond segments of flagellum approximately equal in length; posterior marginal band of second tergum and all the following terga clothed with dense, appressed, pale golden pubescence. Length, 5 mm. Head black, except the mandibles, clypeus, antennal tubercles and antennae yellow; man- dibles edentate at the tip and with a small tooth within; clypeus elevated posteriorly, the elevated margin quadridentate; antennal scrobes not car- inate above; first and second segments of flagel- lum approximately equal in length; front, vertex and genae with small, distinct deep, contiguous punctures more or less obscured by thick, ap- pressed, pale golden pubescence and sparse, erect, pale golden hairs; genal carina extending posteriorly from insertion of mandibles and ele- vated into a prominent tooth one-third its length from mandibles. Thorax pale ferruginous, the dorsum with a pair of distinct, longitudinal, lateral stripes sub- interrupted at the anterior spiracles and propo- deal spiracles, and united medially on the pro- notum and propodeum, thus enclosing a median, longitudinal, ferruginous, integumental stripe on dorsum of thorax; median ferruginous stripe with sparse, erect and suberect, pale golden hairs; remainder of thorax with thin, appressed, pale pubescence and sparse, erect, pale hairs; dorsum and lateral aspects of thorax with small, distinct, deep, contiguous punctures; propodeal spiracular tubercles small and yellow; scutellar scale absent. Abdomen black, except the first segment pale ferruginous, and the sterna dark ferruginous; first and second terga with small, distinct, deep, contiguous punctures; first tergum with sparse, appressed, pale pubescence, and sparse, erect, pale hairs; second tergum with very sparse, ap- pressed, pale pubescence together with sparse, erect, pale hairs on anterior third, the median half with sparse, appressed, fuscous pubescence and sparse, erect, fuscous hairs, the lateral areas with thin pale, appressed pubescence and sparse, erect, pale hairs, and the posterior margin with a band of thick, appressed, pale golden pubes- cence, the anterior margin of the band slightly sinuate; remaining terga clothed throughout with thick, appressed, pale golden pubescence 126 Zoologica: New York Zoological Society [37: 11: and sparse, erect, pale golden hairs; pygidial area narrow, elongate, glabrous; second sternum with small, close, distinct, deep punctures throughout, clothed with thin, appressed, pale pubescence and sparse, erect, pale hairs; sterna three to five with small, indistinct punctures, thin, appressed, pale pubescence and sparse, erect, pale hairs at the posterior margin; last sternum with small, sparse, shallow punctures, and sparse, erect, pale hairs, but without an- terior, lateral tubercles. Legs yellow, clothed with sparse, pale yellow hairs; calcaria pale. Holotype.— Female, Kartabo, British Guiana, July-August, 1920 (W.M. Wheeler), in Mu- seum of Comparative Zoology, Cambridge, Mass. Related to elvina and copia, but with the black maculations of the thorax very distinct and united anteriorly and posteriorly. Ephuta hylonome, new species Female.— Holotype. Ferruginous, except the head and abdomen above blackish, and the dor- sum of thorax with broad, lateral stripes of black, the stripes united medially on the prono- tum and propodeum and distinctly interrupted at the propodeal spiracular tubercles; clypeus elevated posteriorly, the elevated margin quadri- dentate; first and second flagellar segments ap- proximately equal in length; first abdominal segment broader than long; second abdominal tergum with a posterior marginal band of thick, appressed, pale golden pubescence; abdominal terga three to five each with a transverse, median spot of pale golden pubescence. Length 5.5 mm. Head black, except the mandibles, clypeus, antennal tubercles and antennae pale ferrugi- nous; mandibles with dark tips, edentate at the tip and with a minute tooth within; clypeus ele- vated posteriorly, the elevated margin quadri- dentate; first and second segments of flagellum approximately equal in length; antennal scrobes not carinate above; front, vertex and genae with small, distinct, deep, contiguous punctures; genal carina extending posteriorly from inser- tion of mandibles and elevated into a prominent tooth one-third of its length from mandibles; mandibles and clypeus clothed with sparse, erect, pale golden hairs; scape clothed with thin, appressed, pale golden pubescence; anterior third of front with thick, appressed, pale pu- bescence and sparse, erect, pale hairs; median transverse third of front with sparse, erect, pale fuscous hairs; posterior third of front, vertex entirely and genae entirely with thick, appressed, pale golden pubescence, and sparse, erect, pale golden hairs, that on the genae paler than on the vertex, almost white. Thorax ferruginous, except a pair- of very broad lateral stripes on the mesonotal area ex- tending down on the pleura and forward on to the pronotum where they are confluent medi- ally, the stripes interrupted at the propodeal spiracular tubercles, and then continued on the propodeum and confluent medially posteriorly thus forming a U-shaped black mark on the propodeum; black markings of thorax almost but not quite enclosing a very broad, median, longitudinal, ferruginous stripe on the dorsum of thorax; dorsum and lateral aspect of thorax with small, distinct, deep, contiguous punctures, except the latter becoming dense and somewhat confluent on the posterior part of mesonotal area and on the propodeum; scutellar scale absent; propodeal spiracular tubercles small and yellow; thorax clothed with thin, appressed, pale pubescence and sparse, erect, pale hairs, except the pale, appressed pubescence thick on lateral aspect of pronotum, and the median, longitudi- nal, ferruginous stripe on dorsum with only sparse, erect, pale golden hairs. Abdomen above, including the first segment, blackish or very dark ferruginous, beneath fer- ruginous; first segment distinctly broader than long; first and second terga with small, distinct, deep, contiguous punctures, the bottoms of the punctures showing ferruginous in certain lights; anterior half of third to fifth terga with small, distinct, contiguous punctures; lateral areas of last tergum with very small, close, shallow punc- tures; first tergum with erect, pale hairs; anterior half and lateral fourths of second tergum with thin, appressed, pale pubescence and sparse, erect, pale hairs; post-medial area of second tergum with thin, appressed, black pubescence and sparse, erect, fuscous hairs, the posterior margin with a broad band of thick, appressed, pale golden pubescence, the anterior margin of the band emarginate medially, subinterrupted; terga three to five each with a transverse spot of thick, appressed, pale golden pubescence cover- ing approximately the median half of the ter- gum, the lateral fourths fuscous pubescent; la- teral areas of last tergum with sparse, erect, pale golden hairs; pygidial area glabrous; first sternum with the median, longitudinal carina almost obsolete on the posterior half, but strong- ly elevated and conspicuous on the anterior half; second sternum with small, distinct, deep, con- tiguous punctures throughout and with thin, appressed, pale pubescence and sparse, erect, pale hairs; lateral thirds of sterna three to five with small, distinct, contiguous punctures and thin, appressed, pale pubescence, the posterior margin of each with a very thin fringe of sparse, erect, pale hairs; lateral areas of last sternite 1952] Mickel: Mutillidae (Wasps) of British Guiana 127 with small punctures and sparse, erect, pale ferruginous hairs. Legs entirely ferruginous, clothed with sparse, pale hairs; calcaria pale. Holotype — Female, Moraballi creek, Esse- quibo river, British Guiana, August 30, 1929 (O. W. Richards), in collection of University of Minnesota. Somewhat similar to limbata, but the thorax is shorter and the posterior face of propodeum almost perpendicular; the black markings of the pronotum and mesonotum are broader than in limbata, the head black but the punctures of the front showing ferruginous at the bottom, and abdominal terga three to five maculated with black laterally. Genus TRAUMATOMUTILLA Andre Key to the Females 1. Postero-lateral angles of head rounded, not prominent, not carinate or tuber- culate 2 Postero-lateral angles of head prominent, distinctly or conspicuously carinate or tuberculate 7 2. Second abdominal tergum with four yel- lowish integumental spots; lateral mar- gins of mesonotum not toothed 3 Second abdominal tergum with only two ferruginous integumental spots; lateral margins of mesonotum with a promi- nent tooth 6 3. Dorsum of thorax and propodeum with- out a median, longitudinal carina; scu- tellar scale broad, well developed indica (Linnaeus) Dorsum of thorax and propodeum with a distinct, median, longitudinal carina; scutellar scale absent 4 4. Vertex with an arcuate band of dense, appressed, pale, glittering pubescence extending from eye margin to eye mar- gin (Bolivia and Peru). . .indicoides Mickel Vertex entirely black pubescent 5 5. Lateral, pale pubescent stripes on dorsum of thorax extending anteriorly only two- thirds the length of the mesonotum (Brazil and Bolivia) parallela (Klug) Lateral, pale pubescent stripes on dorsum of thorax extending anteriorly to the posterior margin of the pronotum (Ven- ezuela, British Guiana) . . . .gausapata, n. sp. 6. Posterior two-thirds of pygidial area transversely rugose; pale pubescent markings of head and thorax faint, in- conspicuous (British Guiana, French Guiana and Brazil) incerta (Spinola) Pygidium longitudinally rugose, the rugae not reaching the posterior margin; pale pubescent markings of head and thorax strong and conspicuous (Peru) weyrauchi Mickel 7. Pygidial area about as broad as long, oc- cupying nearly all the surface of the last tergum; terga three to five, each with a median spot of pale, glittering pubes- cence 8 Pygidial area decidedly longer than broad, occupying only the median area of the last tergum; terga three to five without a definite, median spot of pale, glittering pubescence latona, n. sp 8. Mesonotum with a pair of lateral spots of dense, pale, glittering pubescence 9 Mesonotum without such spots, entirely black pubescent. . . .dubia dubia (Fabricius) 9. Anterior pair of integumental spots on second abdominal tergum small and weak, or entirely absent (Brazil; Ama- zon valley) dubia albata (Cresson) Anterior pair of integumental spots on second abdominal tergum well devel- oped (British Guiana and Dutch Gui- ana) dubia spilota (Gerstaecker) Key to the Males 1. Mesopleura protuberant above 2 Mesopleura convex, but not protuberant above A 2. Proximal two-thirds of wings hyaline, the distal third fuscous; second abdominal sternum with a pubescent fovea sphegea (Fabricius) Wings fuscous throughout; second abdom- inal sternum without a pubescent fovea 3 3. Clypeal fringe pale, glittering; lateral scu- tellar processes rounded at tip; fourth sternum black pubescent and with a black posterior marginal fringe; sec- ond tergum with scattered, small punc- tures intermixed with fine, separated punctures (Brazil) dentata (Smith) Clypeal fringe fuscous; lateral scutellar processes obliquely truncate; fourth sternum with very sparse, erect, pale hairs and with a pale posterior marginal fringe; second tergum with fine punc- tures throughout dignitosa, n. sp 4. Proximal half of wings hyaline, the distal half fuscous; dorsum of abdomen clothed throughout with dense, glitter- ing, golden pubescence; second abdom- inal sternum without a pubescent fovea dubia dubia (Fabricius) Wings fuscous throughout; dorsum of ab- domen largely black pubescent; second abdominal sternum with a pubescent fovea bartica, n. sp 128 Zoologica: New York Zoological Society [37: 11: Traumatomutilla indica (Linnaeus) 1758. Mutilla indica Linnaeus, Syst. Nat., ed. x, vol. 1, p. 583, female. 1787. Mutilla diadema Fabricius, Mant. Ins., vol. 1, p. 311, female. 1821. Mutilla diadema Klug, Nov. Acta Acad. Nat. Curios., vol. 10, P. 2, p. 314, T. 22, F. 12, female, (in part; specimens from Para, Brazil). 1874. Mutilla indica Gerstaecker, Arch. f. Na- turg., vol. 40, p. 72, female (in part; nec specimens from Bogota, and Bolivia). 1912. Traumatomutilla indica Cameron, Ti- mehri, Journ. Roy. Agri. Com. Soc. Brit. Guiana, (3), vol. 2, p. 415, female. 1937. Traumatomutilla indica Miclcel, Rev. Ent., Rio, vol. 7, p. 196, female. Type.— Female, India, in Natural History Mu- seum, Stockholm, Sweden. A specimen from Parimaribo, Surinam, was sent to Dr. A. Roman in 1931. He compared the specimen with Lin- naeus’s type and found them to be identical. Linnaeus erroneously cited the locality as India. Three specimens labeled diadema by Fabri- cius were examined in 1931, one from the Kiel collection, one from the Copenhagen collection, and one from the Zoological Museum Academy, Leningrad, U. S. S. R. The latter specimen is labeled “diadema Fbr. Hb., Surin.” in Germar’s handwriting, and has been labeled “lectotype” for the following reasons: Fabricius described diadema from the Hiibner collection. One part of Fliibner’s collection was sold to Germar. Germar’s collections were acquired through in- heritance by Schaurn (a nephew of Germar). The Zoological Museum der Universitat, Berlin, Germany, bought part of the Schaum collection from Stein and other parts were dispersed before Schaum’s death. No specimen in the collection of the Berlin Museum fulfills the conditions necessary for a lectotype of diadema. Schaum sold or gave part of his collection to Kraatz before his death and the specimen designated above as lectotype was a part of the material acquired by Kraatz. This specimen was acquired by the Zoological Museum Academy, Lenin- grad, from Kraatz. It fulfills all the necessary conditions for a type specimen: it agrees with the description; it comes from the type locality, Surinam; and it was originally a part of the Hiibner collection. All three of the above speci- mens are the same as indica Linnaeus. Distribution— Surinam, French Guiana, Brit- ish Guiana, Brazil (states of Bahia, Parahyba, Rio Grande do Norte, Ceara, Para and northern Amazonas), Venezuela and Colombia. British Guiana Specimens Examined —Fe- male, Penal settlement, Bartica district, 1917; three females, New Amsterdam, August, 1923 (F. X. Williams); female, Blairmont, August 17, 1923 (H. E. Box); two females, Blairmont, September 6, 1923 (H. E. Box); nine females, Blairmont, September, 1923 (F. X. Williams); two females, Blairmont, November, 1923 (F. X. Williams); three females, Blairmont, 1923- 24 (H. E. Box); female, Georgetown, July, 1921 (A. Busck); female, Georgetown (De- merara), spring, 1878; female, Demerara (Harper); female (1586), Berbia savannah, January 25, 1933 (D. Vesey-Fitzgerald) ; fe- male, Tumatumari, July, 1923 (F. X. Wil- liams) ; female, mouth of Courontyne river, July 5, 1936 (N. A. Weber); female, Rupu- nuni river, September, 1913 (Thurston collec- tion). Traumatomutilla sphegea (Fabricius) 1804. Mutilla sphegea Fabricius, Syst. Piez., p. 435, male. 1811. Mutilla fasciata Olivier, Encycl. Meth- od. Insect, vol. 8, p. 55, male. 1841. Mutilla argyra Spinola, Ann. Soc. Ent. France, vol. 10, p. 85, male. 1845. Mutilla argentea (Spinola) Lepeletier, Hist. Nat. Insect. Hymen., vol. 3, p. 636, male. 1912. Ephuta sphegea Cameron, Timehri, Journ. Roy. Agri. Com. Soc. British Guiana, (3), vol. 2, p. 414, male. 1937. Traumatomutilla argyra Mickel, Rev. Ent., Rio, vol. 7, p. 197, male. (New synonymy) . Ty pes— sphegea Fabricius.— Essequibo (Schmidt), Mus. Sehestadt, in Zoological Museum, University of Copenhagen. argyra Spinola.— Lectotype. Ca- yenne, Museo Zoologia et Ana- tomia Comparata della R. Uni- versity, Torino, Italy. argentea (Spinola) Lepeletier. Lo- cation unknown. fasciata Olivier. Location unknown. The types of sphegea Fabricius and argyra Spinola have been examined and found to be identical. Lepeletier states that argentea was described from a specimen in the Spinola col- lection bearing that name. When the writer studied the Spinola collection in 1931 no speci- men labeled argentea could be found. British Guiana Specimens Examined— Two males, Blairmont, September, 1923 (F. X. Wil- liams); two males, Blairmont, November, 1923 1952] Mickel: Mutillidae (Wasps) of British Guiana 129 (F. X. Williams); male, Blairmont, September 6, 1923 (H. E. Box); male, Blairmont, 1922-24 (H. E. Box); male, Rockstone, June 2, 1929 (A. Mackie); male, Rockstone, July 9, 1911; male, Mahaica, Dem., June 20, 1927; male, Camaria, July 30, 1924 (William Myers); male. Distribution.— British Guiana, Surinam, Ca- yenne and Brazil. This may be the male of indie a (Linnaeus) . Traumatomutilla gausapata, new species 1874. Mutilla parallela var. Gerstaecker, Arch, f. Naturg., vol. 40, p. 75, female (Ori- noco specimen). (New synonymy). Female— Holotype. Black, the second abdom- inal tergum with two pair of yellow integumen- tal spots, the anterior pair linear, the posterior pair subrectangular, transverse, slightly ob- lique, and separated medially by a space slightly less than their greatest width; dorsum of propodeum with a median, longitudinal carina; dorsum of thorax with a pair of lateral, linear stripes of pale, glittering pubescence ex- tending from the anterior margin of the meso- trotum to the posterior margin of the propo- deum; tarsi above clothed with pale, glittering pubescence; tibiae externally with black hairs and with a brush of black hairs at the proximal end externally. Length, 18 mm. Head entirely black, and clothed with sparse, black pubescence, except the genae an- teriorly near the insertion of the mandibles, the caudal aspect of the head and the scape, all with pale, glittering pubescence, the scape with the pubescence thick, elsewhere very sparse, appressed and erect; mandibles edentate and with a small tooth within near the tip; an- tennal tubercles contiguous, rugose; first seg- ment of flagellum much longer than the second, almost as long as two and three combined; an- tennal scrobes strongly carinate above; front, vertex and genae with moderately large, dense, contiguous to confluent punctures; posterior margin of genae strongly carinate, the carina extending almost to a groove at the base of a high, sharp carina margining the proboscidal fossa; postero-latera! angles of head not carinate or tuberculate. Thorax entirely black, clothed with black pu- bescence, except a pair of lateral, linear stripes of dense, appressed, pale, glittering pubescence extending from the anterior margin of the meso- notum to the posterior margin of the propodeum and converging slightly towards the rear, and the ventral third of the lateral aspect of the pro- notum, ventral half of mesopleura and meta- pleura, as well as a small spot at the ventral margin of the sides of the propodeum, all with dense, appressed, pale, glittering pubescence; humeral angles not dentate; anterior spiracles prominent; lateral margins of mesonotum weak- ly angulate medially and strongly constricted before the prominent propodeal spiracles; meso- notum and dorsum of propodeum with a median longitudinal carina; scutellar scale absent; pro- notum, mesonotum and dorsum of propodeum with large, dense, contiguous and sometimes confluent punctures; posterior face of propo- deum finely, closely punctate; lateral aspect oi pronotum separated from dorsum of pronotum by a glabrous ridge, interrupted by deep punc- tures; lateral aspect of pronotum with moderate, close punctures, much less coarsely punctate than the dorsum; mesopleura micropunctate on anterior half, the remainder coarsely punctate; micropunctate area of mesopleura with thick, appressed, pale pubescence, the ventral two- thirds of the coarsely punctate area with thick, appressed, pale pubescence and sparse, long, erect, pale hairs, the dorsal third with sparse, black, appressed hairs, and sparse, long, erect, black hairs; ventral half of metapleura micro- punctate and with thick, appressed, pale pubes- cence, the dorsal half glabrous, impunctate; sides of propodeum with large, close, coarse punc- tures, clothed with scattered, erect, black hairs, except a few, erect hairs near the anterior mar- gin pale. Abdomen black, except the second tergum with two pairs of yellow, integumental spots, the anterior pair linear, contiguous with the an- terior margin of the tergum and approximately one-fourth the length of the tergum, the pos- terior pair transverse, slightly oblique, and separated medially by about two-thirds their greatest width; all the yellow spots glabrous, the anterior pair each with one or two setiferous punctures, the posterior pair impunctate; first abdominal segment disciform; first tergum with a pair of longitudinal stripes of dense, appressed, pale pubescence, the stripes contiguous poster- iorly with the anterior, yellow spots of the sec- ond tergum and distinctly broader at the pos- terior margin than the yellow spots; interstripe area of first tergum clothed with thick, ap- pressed, black pubescence and scattered, erect, black hairs; areas of first tergum laterad of the stripes with only scattered, erect, pale hairs; second tergum with lateral margins, felt line, and a median spot at the posterior margin all densely, pale pubescent, otherwise, except the yellow spots, with thick, appressed, black pu- bescence and scattered, erect, black hairs; sec- ond tergum coarsely punctate beneath the black pubescence; third and fourth terga with thick, recumbent, black pubescence and scattered, erect, black hairs, except each with the lateral 130 Zoological New York Zoological Society [37: 11: margins and a median, elongate spot densely, pale pubescent; third, fourth and fifth terga all closely punctured, the fifth with the lateral margins black pubescent, but with a median, elongate spot of dense, appressed, pale pubes- cence; last ter gum with a large pygidial area well defined and transversely rugose, the an- terior half of the tergum with a median, pale pubescent spot, the lateral areas, black pubes- cent; median, pale pubescent spots on terga two, three, four, five and last, forming a longitudinal pale line, narrowest anteriorly and slightly but distinctly broader at its posterior terminus; all the sterna with pale hairs or pubescence, except the posterior marginal fringe of the fifth and all the pubescence of the last, black; broad, pos- terior marginal, pale, appressed pubescent bands on second, third and fourth sterna, otherwise the pale pubescence scattered or sparse and erect; first sternum with a median, elevated, longitudinal carina; second sternum with large, separated punctures; posterior margin of second sternum and third, fourth, fifth and last sterna with moderate, close punctures. Legs black, clothed with pale hairs, except the middle and hind tibiae externally with black spines and scattered black hairs, and the tarsal segments beneath with coarse, black hairs; cal- caria pale. Holotype.— Female, Rupununi savannah, Brit- ish Guiana (794), October 13, 1932 (D. Vesey- Fitzgerald), in collection of University of Min- nesota. P aratypes— Female, Orinoco (Moritz) (this is the specimen designated as “parallela var.” by Gerstaecker) ; female, Akuriman, Venezuela, October, 1940 (Paul Anduze); female, Ara- bapo, Mt. Roraima, Venezuela (1284), No- vember 24, 1932 (D. Vesey-Fitzgerald) ; female, Caroni-Amazon watershed, Brazil-Venezuela boundary (1353), November 29, 1932 (D. Vesey-Fitzgerald); female, Waranama, British Guiana, November 20, 1936 (Mrs. L. Ogilvie); female, Rupununi savannah, British Guiana (795) , October 13, 1932 (D. Vesey-Fitzgerald) ; female, Ichilibar, Pakarima foothills, British Guiana (863), October 20, 1932 (D. Vesey- Fitzgerald) ; female, Kerikabaru, Pakarima foothills, British Guiana (914), October 22, 1932 (D. Vesey-Fitzgerald); female, Minas Geraes, Brazil (Reinhardt), Mus. Drewsen. Related to parallela Klug, borba Cresson and indicoides Mickel; all have a median, longitudi- nal carina on the dorsum of the propodeum, and the dorsum sloping towards the lateral margins from the median carina. The three differ from each other in pale pubescent markings and in geographical distribution. Traumatomutilla incerta (Spinola) 1841. Mutilla incerta Spinola, Ann. Soc. Ent. France, vol. 10, p. 93, female. 1937. Traumatomutilla incerta Mickel, Rev. Ent., Rio, vol. 7, p. 196, female. British Guiana Specimens Examined.— Six fe- males, Kartabo, Bartica district: April 16, 1922; April 23, 1919; May 16, 1924; July 22, 1924; and July 23, 1922; two females, Kartabo, July- August, 1920 (W. M. Wheeler) ; female (2075), Moraballi creek, Essequibo river, September 14, 1929 (dark forest) (O. W. Richards); female (2553), Moraballi creek, Essequibo river, Sep- tember 22, 1929; female, source Essequibo river (J. Ogilvie); female, Tumatumari, July, 1923 (F. X. Williams); female (514), Oko river, June 25, 1936 (N. A. Weber); female, August 14, 1911. Distribution.— British Guiana, French Guiana and Amazonas, Brazil. Closely related to weyrauchi Mickel from Peru. Traumatomutilla latona, new species Female— Holotype. Black, the second ab- dominal tergum maculated with a single pair of large, subquadrate, postmedian, red, integu- mental spots, and posterior margin of the second tergum with a median, pale pubescent spot. Length, 11 mm. Head entirely black, very sparsely pubescent, that on the ventral aspect, the genae and the anterior half of front, pale glittering and semi- erect, elsewhere black; clypeal fringe fuscous; mandibles slender, edentate at the tip, apparent- ly not toothed within; antennal tubercles con- tiguous; scape clothed with sparse, pale glitter- ing hairs; first segment of flagellum distinctly longer than the second; flagellum black, suf- fused with reddish beneath; antennal scrobes distinctly carinate above, the carina not reaching the eye margins; front, vertex and genae with moderate, deep, contiguous to confluent punc- tures; postero-lateral angles of head, each with a small, triangular, glabrous tubercle; posterior margin of genae defined by a prominent, sharp carina, the latter extending anteriorly to the carina bounding the proboscidal fossa; eyes very large, prominent, hemispherical, the least dis- tance between the margin of the eyes and the genal carina equal to about one-sixth the greatest diameter of the eyes; ventral aspect of head glabrous, with small, sparse punctures. Thorax black, the dorsum immaculate, clothed throughout with sparse, erect and semi- erect, black hairs; all the pleural areas, except sides of propodeum, with thick, appressed, fine, pale, glittering pubescence, and a few, sparse, 1952] Mickel: Mutillidae (Wasps) of British Guiana 131 erect, pale, glittering hairs; sides and posterior face of propodeum with sparse, long, erect, pale glittering hairs; dorsum of thorax, with moderately large, deep, contiguous to confluent punctures; scutellar scale present and distinct; lateral margins of mesonotal area not prom- inently dentate; lateral aspect of pronotum mi- cropunctate and with a few, distinct punctures at the anterior margin; mesopleura micropunc- tate throughout and with moderate, shallow, close punctures at the posterior margin and postero-ventral area; metapleura micropunctate throughout and with scattered, shallow punc- tures ventrally; sides of propodeum rounded into the posterior face, with close to separated, moderate punctures; posterior face of propo- deum with moderately large, deep, contiguous to confluent punctures. Abdomen black, the second tergum macu- lated with a single pair of large, postmedian, subquadrate, red, integumental spots, the dis- tance between the two equal to slightly more than one-third the transverse diameter of one of them; first segment distinctly nodose; anterior face of first tergum with moderate, close punctures posteriorly, clothed throughout with very sparse, long, erect, pale, glittering hairs, and with thick, appressed, pale, glittering pubescence at the posterior median margin; dor- sal face of first tergum almost impunctate, but with a few, moderate, close punctures at the anterior margin, the median half clothed with dense, appressed, pale, glittering pubescence; second tergum with dense, confluent, elongate punctures, the latter becoming smaller and sep- arated on the lateral areas; red, integumental spots of the second tergum with similar elongate punctures, but the latter distinct and separated slightly from one another; second tergum clothed with sparse, erect, black hairs, except the integumental spots with very sparse, erect, fulvous hairs; lateral areas of second tergum with very sparse, erect, pale, glittering hairs, the felt lines pale, the lateral margins with a fringe of appressed, pale, glittering hairs, the lateral thirds of the posterior margin with a pale fringe, and a broad spot of pale, glittering hairs medi- ally on the posterior margin separated from the lateral, pale fringe by a cupreous to black fringe of hairs; third, fourth and fifth terga clothed with sparse, semi-erect, pale, glittering hairs in- terspersed with very sparse, erect, black hairs, each tergum with moderate, contiguous to con- fluent punctures; pygidial area long, narrow, triangular, occupying only the median area of last tergum, the area longitudinally rugose on the anterior two-thirds, glabrous, unsculptured on the posterior third; last tergum otherwise with contiguous to confluent punctures and sparse, erect, pale, glittering hairs; first sternum glabrous, almost impunctate, with very sparse, erect, pale, glittering hairs, and with the longi- tudinal, median carina very weakly developed only on the anterior third; second sternum with small punctures near the anterior margin, but elsewhere with moderate, distinct, close punc- tures, clothed with very sparse, erect, fine, pale, glittering hairs, and with a thin, pale, posterior marginal fringe; sterna three to five with mod- erate, distinct, contiguous punctures, very sparse, erect, fine, pale, glittering hairs, and each with a thin, pale, posterior marginal fringe. Legs black, sparsely clothed with pale, glitter- ing hairs; calcaria pale. Holotype — Female, Bartica, Bartica district, British Guiana (Wm. Beebe), in collection of University of Minnesota. Paratypes — Female, Kartabo, Bartica district, British Guiana, May 20, 1919; female, Kartabo, Bartica district, British Guiana, April 14, 1922; female, British Guiana, without other data; fe- male, Maracas, Trinidad, B. W. I., February 22, 1941 (D. J. Billes); female, Trinidad, B. W. I., May, no. 413 (D. Vesey-Fitzgerald) ; female, Port of Spain, Trinidad, B. W. I., January 30, 1912 (G. A. K. Marshall); female, Manaos, Brazil, August, 1929 (G. Arnold). Related to trochanterata Gerstaecker from which it differs by the larger integumental spots of the second tergum, the absence of pale pu- bescence on the vertex, and the much less sculp- tured .pygidial area (longitudinally rugose throughout in trochanterata) . The integumental spots of the second tergum in trochanterata are separated by a distance equal to approximately four-fifths the transverse diameter of one of them. The paratypes range in length from 8 to 11 mm and the color of the legs varies from dark mahogany red to black. Traumatomutilla dignitosa, new species Male— Holotype. Black, the pronotum, dor- sum of propodeum, dorsal face of first tergum, band on anterior fourth of second tergum, pos- terior marginal fringe of second tergum inter- rupted medially with black, and third tergum entirely, all with dense, appressed, pale, glitter- ing pubescence. Length, 14 mm. Head black, the ventral aspect, genae, anten- nal tubercles, anterior margin of front and ver- tex, all with pale, glittering pubescence, thick, appressed and sparse, erect on the ventral as- pect, genae, and antennal tubercles, sparse, ap- pressed and erect on anterior part of front and on vertex; mandibles black, slightly reddish sub terminally, edentate at the tip, and apparent- ly with two small teeth within near the tip; clypeus with small, dense punctures throughout, 132 Zoologica: New York Zoological Society [37: 11: the anterior margin bidentate medially, clothed with sparse, fuscous pubescence; antennal tu- bercles contiguous; scape bicarinate beneath, clothed with black pubescence; first segment of flagellum only slightly shorter than the second; antennal scrobes strongly carinate above; front with moderate, close to separated punctures; vertex with small, close to separated punctures; genae with small punctures more or less con- cealed by the pubescence; postocellar area slightly elevated, the lateral posterior margins of the vertex therefore slightly concave. Thorax black, the dorsal aspect of pronotum, dorsum of propodeum and mesopleura clothed with dense, appressed, pale, glittering pubes- cence, the lateral aspects of pronotum and the metapleura with sparse, appressed, pale, glitter- ing pubescence, the posterior face of propodeum with sparse, appressed and erect, pale, glittering pubescence, the sides of propodeum with very sparse, erect, pale, glittering hairs, and the meso- notum and scutellum with sparse, black pubes- cence, except the lateral and posterior areas of scutellar elevation with sparse, erect, pale, glit- tering hairs; anterior face of pronotum not emarginate medially; dorsal aspect of pronotum densely punctate, but concealed by the pale pubescence; mesonotum with moderate, con- tiguous to confluent punctures; parapsidal fur- _rows present on posterior two-thim^f mesono- tum; lateral processes of scutellum densely "punctate, obliquely truncate at the tip, two- fifths the length of the mesonotum on the me- dian line; scutellum strongly, conically elevated, deeply reticulo-punctate, dorsum of propodeum broadly reticulo-punctate, but the sculpture con- cealed by the pubescence; posterior face of pro- podeum reticulate; lateral aspects of pronotum micropunctate and with a very few, small, scattered punctures; mesopleura coarsely reti- culo-punctate, except the anterior marginal area less coarsely sculptured with shallow punctures; dorsal area of mesopleura elevated into a con- spicuous, blunt tooth; metapleura glabrous, closely punctate ventrally; sides of propodeum reticulate, except the anterior marginal area glabrous, impunctate; tegulae black, clothed with sparse, black pubescence. Abdomen black; first segment strongly no- dose; anterior face of first ter gum glabrous with scattered, small punctures and very sparse, erect, pale, glittering hairs; second tergum with fine, separated punctures throughout, the punctures slightly larger laterally and posteriorly than an- teriorly; second tergum clothed with sparse, black pubescence, except a transverse band at the anterior margin of dense, appressed, pale, glittering pubescence, the width of the band at the midline about two-thirds the width of that on the dorsal face of first tergum, the width at the sides slightly more than twice the width at the midline, thus the posterior margin of the band concave; felt lines pale; lateral margins of second tergum with a fringe of pale pubescence; posterior marginal fringe of second tergum pale, except interrupted medially with black; third tergum clothed throughout with thick, ap- pressed, pale, glittering pubescence, and scat- tered, erect, black hairs, the posterior marginal fringe also pale and not interrupted medially; fourth, fifth, sixth and pygidial terga finely punc- tate and clothed with black pubescence, except the lateral margins of the fourth with a few, erect, pale hairs; pygidial area impunctate, ex- cept for small, close punctures at the posterior margin, the latter with a fringe of long, black hairs; first sternum moderately punctate, with very sparse, erect, pale hairs, and with a very weak, median, longitudinal carina; second ster- num without a pubescent fovea, with small, sparse punctures, very sparse, erect and semi- erect, pale hairs, and with the posterior marginal fringe pale; third and fourth sterna with small, well separated punctures, very sparse, erect, pale hairs, and with the posterior marginal fringe pale; fifth sterna with small, sparse punc- tures, very sparse, erect, pale hairs and with the thin, posterior marginal fringe black; sixth sterna with small, sparse punctures, very sparse, erect, black hairs and a black, posterior marginal fringe; last sternum rectangular, longer than broad, with moderately small punctures later- ally, the median, longitudinal area almost im- punctate, clothed with very sparse, erect, black hairs. Wings dark fuscous throughout; cell 2nd Ri + R.2 about three times as long as broad, square- ly truncate at the tip; cell R5 receiving vein M3+4 about one-fourth the distance from the base to the tip; cell R4 very indistinct, receiving the indistinct vein M2 very close to the base. Legs black, clothed with pale, glittering pu- bescence, except the tibiae externally with black pubescence, and the tarsi with fuscous to bronze pubescence; calcaria pale. Holotype — Male, Bartica, Bartica district, British Guiana (Wm. Beebe), in collection of University of Minnesota. Related to dentata Smith from which it differs as expressed in the key. Traumatomutilla bartica, new species Male— Holotype. Black, the posterior half of dorsum of propodeum, anterior half of posterior face of propodeum, dorsal face of first tergum, and an anterior marginal band on the second tergum (the latter broad at the sides, equaling 1952] Mickel: Mutillidae (Wasps) of British Guiana 133 half the length of the tergum and gradually nar- rowed toward the midline to one-fifth the length of the tergum), all with dense, appressed, pale, glittering pubescence; front, anterior margin of pronotum, pleural areas, and third and fourth terga entirely, all with sparse, appressed, pale, glittering pubescence; second sternum with a median, longitudinal line of appressed, pale pu- bescence passing through the pubescent fovea. Length, 15 mm. Head black, clothed throughout with sparse, pale, glittering pubescence, except that on the malar space dense, and a transverse band an- teriorly on the vertex with subappressed and erect, sparse, black hairs; mandibles edentate at the tip and apparently with a tooth within near the tip, the proximal half with thick, pale pu- bescence; clypeus finely, densely punctate, its anterior margin bidentate medially; antennal tubercles contiguous; scape bicarinate beneath; first segment of flagellum distinctly shorter than the second; antennal scrobes strongly carinate above; front and vertex with moderate, conti- guous to confluent punctures; genae with small, contiguous punctures, more or less concealed by the pubescence; postocellar area slightly but distinctly elevated, the lateral posterior margin of the vertex therefore slightly concave. Thorax black, the posterior half of dorsum of propodeum and anterior half of posterior face of propodeum with a large area of dense, appressed, pale, glittering pubescence; anterior margin of pronotum with appressed, pale pu- bescence; remainder of thorax with sparse, pale, glittering pubescence, except the most of pro- notum, mesonotum entirely, and scutellum en- tirely, with sparse, black pubescence; anterior face of pronotum glabrous, distinctly, shallowly emarginate medially; dorsal face of pronotum, mesonotum entirely and scutellum with moder- ate, dense, confluent punctures; parapsidal fur- rows present on posterior half of mesonotum; lateral processes oFscuteTIu m den seTy p u n c tat e , their outer distal margin rounded; scutellum moderately convex, not conically elevated; dor- sum and posterior face of propodeum reticulate; lateral aspects of pronotum micropunctate and with a few, moderate punctures anteriorly and dorsally; anterior fourth of mesopleura micro- punctate, the remainder with moderate, dense, confluent punctures; mesopleura evenly convex, not elevated dorsally; metapleura very weakly micropunctate and micropubescent, and with a few, moderate, close punctures ventrally; sides of propodeum reticulate posteriorly, the reticu- lations becoming shallower anteriorly and al- most obsolete at the anterior margin; tegulae black, glabrous, impunctate, except the anterior and inner margins punctate and black pubes- cent. Abdomen black with pale pubescent mark- ings; first segment-strongly nodose, the tergum with an anterior and dorsal face; a broad, pos- terior band of dense, appressed, pale, glittering pubescence covering the dorsal face entirely, except for a very narrow, median, nonpubescent line, and extending anteriorly on the lateral areas of the anterior face about one-third its length; first tergum with moderate punctures visible through the pale pubescence; second tergumwith an anterior marginal band of dense, appressed, pale pubescence, broad at the sides (equal to half the length of the tergum) and gradually narrowed to the midline (equal to one-fifth the length of the tergum); second ter- gum otherwise black pubescent, except the felt lines, lateral margins, and lateral extremes of posterior margin pale; sjcond tergum with fine and small punctures intermixed; third tergum clothed throughout with sparse, appressed, pale pubescence, except the posterior marginal fringe black; fourth tergum clothed with sparse, ap- pressed, pale pubescence somewhat interrupted medially with black, and the posterior marginal fringe black; fifth, sixth and last terga black pu- bescent, except the fifth and sixth with some pale pubescence at the lateral margins; pygidial area finely granulate; first sternum with a low, median, longitudinal carina and very sparsely clothed with erect, pale hairs; second sternum with moderate, well separated punctures, a mod- erately large, postmedian, pale pubescent fovea, a median, longitudinal line of appressed, pale pubescence passing through the fovea, and with sparse, pale pubescence; postero-lateral areas of second sternum with fine punctures interspersed among the moderate ones; posterior marginal fringe of sixth sternum and sparse pubescence of last sternum, black. Wings dark fuscous; cell 2nd R i + R2 acute at the tip; cell R5 receiving vein M3+4 one-fourth the distance from the base to the tip; cell R4 receiving vein M2 at the extreme tip. Legs black, clothed throughout with sparse, pale pubescence, except the tips of the middle and hind femora, and the extreme base of the middle and hind tibiae, black pubescent. Holotype — Male, Kartabo, Bartica district, British Guiana, March 10, 1922 (Wm. Beebe), in collection of University of Minnesota, Related to picada Cresson, but differs in the mostly black pubescent pronotum, the posterior marginal fringe of the second tergum black, the third and fourth terga only sparsely, pale pu- bescent, and the median, longitudinal, pale pubescent line of the second sternum. 134 Zoologica: New York Zoological Society [37: 11: Traumatomutilla dubia dubia (Fabricius) 1804. Mutilla dubia Fabricius, Syst. Piez., p. 435, gynandro morph. 1879. Mutilla midas Smith, Descr. New Spe- cies Hymen., p. 221, male. 1912. Traumatomutilla nitidipustulata Camer- on, Timehri: Journ. Roy. Agri. Com. Soc. Brit. Guiana, (3), vol. 2, p. 415, female. 1931. Traumatomutilla dubia Bischoff, Mitt. D. Ent. Ges., Jahrg. 2, pp. 54-56, gynan- dro morph type. 1931. Traumatomutilla nitidipustulata Bisch- off, Mitt. D. Ent. Ges., Jahrg. 2, p. 55, female. Type— dubia Fabricius, Zoologischen Museum der Universitat, Kopenhagen, Denmark. midas Smith, British Museum (Nat. Hist.). nitidipustulata Cameron, British Muse- um (Nat. Plist.). Distribution— British Guiana, Brazil. British Guiana Specimens Examined— Fe- male, Rockstone, Essequibo river, March 23, 1913 (G. E. Bodkin); female, Rockstone, June 2, 1929 (A. Mackie); two females, Rockstone, July 9, 1911; seven females, Rockstone, Esse- quibo river, June 26, 1927; female, Rockstone, November 30, 1913 (G. E. Bodkin); female, Essequibo (Smidt). Dr. H. Bischoff, Zoological Museum, Univer- sity of Berlin, and the writer examined Fabri- cius’s type of dubia in 1931 with the above re- sults. It was determined from the gynandro- morph type that the male must be the same as midas Smith and that the female was one of the group of subspecies including spilota Gerstaeck- er (Surinam), albata Smith (Ega), simulans Smith (Venezuela), and nitidipustulata (Cam- eron) (British Guiana). The Fabrician type bears the locality designation “Essequibo river.” The above thirteen specimens are all from the Essequibo river and are the same as nitidipus- tulata Cameron, with the type of which two of them have been compared. Three specimens, noted hereafter, from Essequibo river are the same as spilota Gerstaecker. However, the typi- cal subspecies of female must be chosen from the same type locality as the male and since the type locality of spilota is Surinam, it is excluded from consideration. Only one specimen of male is known to me. It is from Vista Alegre, Rio Branco, Amazonas, Brazil, September 6, 1924. Midas Smith has for type locality, Para, Brazil. Without more ma- terial it is impossible to distinguish subspecies in the males. If material of the male sex from Para, Vista Alegre, Essequibo river and other locali- ties should prove to represent recognizable sub- species, then the Para specimens and those con- subspecific with them will have to be designated as midas Smith. Traumatomutilla dubia spilota (Gerstaecker) 1874. Mutilla spilota Gerstaecker, Arch. f. Na- turg., vol. 40, p. 73, female. Lectotype — Female, no. 6689, Surinam (Cor- dua) , in Zoologisches Museum der Universitat, Berlin. Distribution— Surinam, British Guiana, Ama- zonas. British Guiana Specimens Examined— Fe- male, Rockstone, Essequibo river, June 26, 1927; female, Rio Essequibo (Source) (J. Ogil- vie); female, Ikuraka Lake, Essequibo, March, 1918 (G. E. Bodkin); female 1523, Rukununi savanna, Jan. 12, 1933 (D. Vesey-Fitzgerald) . The above Rockstone specimen was collected at the same time and place as seven specimens of dubia dubia (Fabricius). Gerstaecker had four specimens, all labeled the same, from which he described this species. The writer labeled one of these a lectotype, and the remaining three as paratypes in 1931. Genus HOPLOMUTILLA Ashmead Key to the Females 1 . Carinae of second tergum entirely absent on the integumental spots, the latter sometimes sparsely punctured; meso- notum with only antero-lateral spots of dense, pale pubescence, never with a transverse, posterior, pale pubescent spot between the antero-lateral spots. . 2 Carinae of second tergum present on the integumental spots, although sometimes reduced to tubercular elevations; meso- notum usually with a more or less well developed transverse spot of pale pu- bescence between and posterior to the antero-lateral spots 3 2. Vertex with a pair of pale pubescent spots lanata (Lepeletier) Vertex entirely black pubescent atricapilla Mickel 3. Genae clothed for the most part with very sparse, black pubescence, sometimes a few pale hairs at the posterior margin. 4 Genae clothed for the most part with very sparse, pale pubescence; integumental spots of second tergum ferruginous; dis- tal spine of hind tibiae at the tip of a cylindrical process, the spine and the process equal in length .. derasa (Fabricius) 4. Sides of propodeum entirely glabrous, al- most impunctate, with only a few, scat- 1952] Mickel: Mutillidae (Wasps) of British Guiana 135 tered, inconspicuous punctures; only the dorsal third of lateral aspect of prono- tum punctate, the remainder glabrous, impunctate; broad, lateral margins of first tergum bare, glabrous, impunctate glabella Mickel Sides of propodeum distinctly, closely punctate posteriorly; lateral aspects of pronotum punctate throughout; lateral margins of first tergum pubescent, punc- tate bequaerti Mickel 4 Hoplomutilla lanata (Lepeletier) 1845. Mutilla lanata Lepeletier, Hist. Nat. Ins., Hymen., vol. 3, p. 644, female. 1939. Hoplomutilla lanata Mickel, Rev. Ent., Rio, vol. 10, p. 647, female. Three females have been recorded from Brit- ish Guiana: Bon Successo, Rio Takatu, Rupu- nuni Savanna, January 1-2, 1933 (D. Vesey- Fitzgerald) . Distribution.— Northern Brazil, British Gui- ana and Venezuela. Hoplomutilla atricapilla Mickel 1939. Hoplomutilla atricapilla Mickel, Rev. Ent., Rio, vol. 10, p. 648, female. Known only from the unique type. Type lo- cality label “British Guiana.” Hoplomutilla glabella Mickel 1939. Hoplomutilla glabella Mickel, Rev. Ent., Rio, vol. 10, p. 655, female. Recorded from British Guiana: Rio Esse- quibo (source) (J. Ogilvie). Distribution.— Brazil, Amazon river region and British Guiana as above. Hoplomutilla bequaerti Mickel 1939. Hoplomutilla bequaerti Mickel, Rev. Ent., Rio, vol. 10, p. 665, female. Recorded from British Guiana.— Blairmont, 1923 (H. E. Box). Distribution— Colombia, British Guiana (as above), Surinam, French Guiana and Ama- zonas, Brazil. Hoplomutilla derasa (Fabricius) 1805. Mutilla derasa Fabricius, Syst. Piez., p. 429, female. 1939. Hoplomutilla derasa Mickel, Rev. Ent., Rio, vol. 10, p. 685, female. British Guiana Specimens Examined.— Thir- teen females, Kartabo, Bartica district: March 23, 1922; April 13, 1924; April 28, 1924; May 15, 1922; May 23, 1922; May 26, 1922; June 22, 1924; July 24, 1920; August 27, 1922; No- vember 5, 1922; and 1919; three females, Penal Settlement, Bartica district, May 6, 1924, and without date; female, Kalacoon, Bartica district, 1916. Distribution.— British Guiana, French Gui- ana, Surinam and northern Brazil. Hoplomutilla caerulea Mickel 1939. Hoplomutilla caerulea Mickel, Rev. Ent., Rio, vol. 10, p. 707, male. Recorded from British Guiana: Thirteen males (including holotype), Kamakusa, Janu- ary, 1923. Distribution— Known only from type locality. Genus PAPPOGNATHA Mickel Pappognatha radiata (Andre) 1906. Ephuta ( Tilluma ) radiata Andre, Zeit. Hymen. Dipt., vol. 6, p. 35, female. 1939. Pappognatha radiata Mickel, Ann. Ent. Soc. Amer., vol. 32, p. 339, female. Recorded from British Guiana: Moraballi creek and Essequibo river. British Guiana Specimens Examined.— Three females, Kartabo, Bartica district: May 8, 1924; May 30, 1919; and June 16, 1919; two females, Bartica district, April 23 and April 24, 1924. Distribution.— French Guiana and British Guiana. Genus HOPLOCRATES Mickel Key to the Females 1. Median, pale pubescent spot of second abdominal tergum fulvous, more or less elongate, subovate . centromaculata (Cresson) Median, pale pubescent spot of second ab- dominal tergum pale, glittering white, linear pompalis Mickel Hoplocrates centromaculata (Cresson) 1902. Mutilla centromaculata Cresson, Trans. Amer. Ent. Soc., vol. 28, p. 27, female. 1941. Hoplocrates centromaculata Mickel, Rev. Ent., Rio, vol. 12, p. 369, female. Recorded from British Guiana: Georgetown, July, 1921 (A. Busck). British Guiana Specimens Examined— Three females, Kartabo, Bartica district: February 26, 1922; March 12, 1922; and June 4, 1919. Distribution— Brazil, French Guiana, British Guiana, Venezuela and Peru. Hoplocrates pompalis Mickel 1941. Hoplocrates pompalis Mickel, Rev. Ent., Rio, vol. 12, p. 378, female. Not recorded from British Guiana, but prob- ably occurs there. Distribution.— Colombia, Venezuela, Trini- dad, B. W. I., and Surinam. 136 Zoologica: New York Zoological Society [37: 11: Genus PSEUDOMETHOCA Ashmead Key to the Females 1 . Abdomen above with a narrow, median line of pale, glittering pubescence extending from first to last segments; second ab- dominal tergum with a postmedian, transverse, narrow line of pale, glitter- ing pubescence; thus the abdomen with the pale pubescent markings forming a cross 2 Abdomen without such pale, glittering pubescent markings, the latter not in the form of a cross 4 2. Head entirely black; legs ferruginous; in- tegument ferruginous beneath trans- verse, pale pubescent line on second tergum (Colombia) cruciatella Mickel Front and anterior part of vertex bright ferruginous; legs black; integument of second tergum entirely black through- out 3 3. Dorsum and pleural regions of thorax en- tirely ferruginous (British Guiana) virgata, n. sp. At least mesopleura and metapleura large- ly black (Venezuela, Trinidad, French Guiana) plagiata (Gerstaecker) 4. Integumental, ferruginous, transverse markings of second tergum confluent medially forming a postmedian trans- verse band; pygidium granulate 5 Integumental, ferruginous, transverse markings of second tergum well sep- arated spots; pygidium either granulate or glabrous 6 5. Clypeus, antennal tubercules black; legs ferruginous with the tips of femora black; length 6.5 to 10 mm. (British Guiana) pulchella, n. sp. Clypeus and antennal tubercules ferrugin- ous; legs black, the tibiae and tarsi ob- scure ferruginous; length, 4 mm. (Ara- gua, Venezuela) . . . .pulicaria (Gerstaecker) 6. Pygidium granulate; lateral margins of dorsum of propodeum with a conspicu- ous tubercle immediately behind the propodeal spiracle; integumental spots beneath a complete, transverse band of pale pubescence (Para, Brazil) crepera (Cresson) Pygidium glabrous; dorsum of propodeum not conspicuously tuberculate; second tergum without a complete, transverse, postmedian band of pale pubescence (British Guiana) diligibilis, n. sp. Pseudomethoca plagiata (Gerstaecker), (new combination) 1874. Mutilla plagiata Gerstaecker, Arch. f. Naturg., vol. 40, p. 65, female. 1879. Mutilla crenulata Smith, New Descr. Hymen., p. 211, female. (New synon- ymy). Types. — plagiata Gerstaecker, female, no. 6626, Aragua (Moritz) in Zoological Museum of University of Berlin, Germany. Gerstaecker gives the locality in his description as “Aragua Columbiae.” The specimen was collected by Moritz, and the records of the Berlin Museum show that Moritz collected in South America before 1829 when the country now known as Venezuela was a part of Colombia. The type locality, then, is Aragua, Venezuela. Type of crenulata : Female, Venezuela, in British Museum of Natural History. Both types have been examined and found to be the same. Distribution.— Venezuela., Trinidad, B. W. I., and French Guiana. Not recorded from British Guiana but will probably be found there. Pseudomethoca virgata, new species Female— Holotype. Front, anterior part of vertex, genae in part, and thorax entirely, fer- ruginous; abdomen black, with a median, nar- row line of pale, glittering pubescence extending from the first segment to the last, and a trans- verse, postmedian line of pale, glittering pubes- cence on the second tergum, thus the narrow, pale pubescent markings forming a cross; pygi- dial area well defined, granulate; legs black. Length, 9 mm. Head black, except the front, anterior part of vertex, and genae in part, bright ferruginous, the genae not so brightly ferruginous as the front and vertex; genae, clypeus and scape, clothed with sparse, pale hairs; ferruginous area of front and vertex with sparse, recumbent, bright ferruginous hairs; black area of vertex With sparse, black hairs; scape and flagellum black; front and vertex with moderate, dense punctures; genae with moderate, close punc- tures; anterior half of clypeus glabrous, trans- versely concave, the posterior half finely, dense- ly punctate, the margin separating the two halves produced laterally into a small tubercle; posterior margin of genae carinate, the carina not extending on to the vertex, not elevated anteriorly into a tooth and not reaching the carina margining the proboscidal fovea; first segment of flagellum almost as long as the second and third united; antennal scrobes dis- tinctly carinate above; head slightly narrower than the thorax. Thorax entirely ferruginous; dorsum of tho- rax with moderate, dense punctures anteriorly, the punctures increasing in size posteriorly and 1952] Mickel: Mutillidae (Wasps) of British Guiana 137 merging into reticulate on the dorsum and pos- terior face of propodeum; scutellar scale absent; lateral areas of pronotum with moderate, dense punctures; mesopleura micropunctate anterior to the sharp pleural ridge, and glabrous, im- punctate posterior to the latter; metapleura and sides of propodeum glabrous, impunctate; an- terior half of posterior face of propodeum reticulate, the posterior half glabrous, with only a few scattered punctures; dorsum of pronotum with sparse, black hairs; remainder of dorsum of thorax with sparse, bright ferruginous hairs; lateral areas of pronotum, mesopleura anterior to pleural ridge, ventral margin of metapleura, and postero-ventral area of sides of propodeum, all with pale, micropubescence; posterior face of propodeum with an elongate area of long, erect, thick, pale hairs at the lateral margins medially; humeral angles angulate but not den- tate; lateral margins of dorsum of thorax with a distinct tooth midway between the anterior and propodeal spiracles; anterior margin of propodeal spiracles produced; lateral margins of dorsum and posterior face of propodeum ser- rate. Abdomen entirely black, the terga clothed with sparse, recumbent, black pubescence and a few, erect, black hairs, except as follows: a median spot at the posterior margin of first ter- gum, a median narrow line the entire length of the second tergum, and a median spot on terga three to five, all forming a longitudinal, median line, a transverse, postmedian line on the second tergum, lateral fifths of first and second terga forming an obscure, longitudinal stripe, and lateral spots on terga three to five, all of pale, glittering pubescence; the lateral fifths of the second terga not uniformly pale pubescent throughout, but sparse, except at the inner and outer margins where the pubescence is dense, thus forming the above obscure, lateral stripes; sterna clothed throughout with sparse, pale hairs, the latter slightly thicker at the posterior margins of the sterna; all the terga finely, densely punctate; pygidial area occupying most of the last tergum, distinctly margined laterally, strongly granulate throughout; first sternum with a low, poorly developed, median, longitu- dinal carina; second sternum with moderate, contiguous to separated punctures; remaining sterna with small, close punctures. Legs black, except all the coxae distinctly ferruginous; legs clothed throughout with sparse, pale hairs; calcaria pale. Holotype.— Female, Rupununi savanna, Brit- ish Guiana, October 2, 1932 (D. Vesey-Fitz- gerald), in collection of University of Minne- sota. Related to plagiata Gerstaecker, but differs from that species in the entirely ferruginous tho- rax, the partly ferruginous genae, the partly ferruginous vertex, and the ferruginous coxae. Pseudomethoca DiLiGiBius, new species Female— Holotype. Head and abdomen most- ly black; thorax and legs bright ferruginous; first abdominal segment sessile with the second; second tergum with a pair of postmedian, trans- verse, integumental, ferruginous spots distinctly separated medially; posterior marginal fringe of second tergum black; terga three to five clothed with sparse, pale pubescence; pygidium glabrous, impunctate, distinctly margined lat- erally. Length, 6 mm. Head black, except mandibles, excluding tips, clypeus, antennal tubercles, scape, pedicel and first three or four segments of flagellum, bright ferruginous, under side of head very dark fer- ruginous; genal carina present and distinct, but not terminating anteriorly in a tooth and not extending on to the postero-lateral angles; man- dibles edentate at the tip, their inner margins not dentate; elevated, median area of clypeus with a small tubercle at each antero-lateral angle, a narrow rectangular area between the tubercles glabrous, the subtriangular area pos- terior to the latter, finely, densely punctate; an- tennal scrobes carinate above; front, vertex and genae with moderate, dense punctures and sparse, erect and recumbent, pale hairs; eyes large, the distance between the posterior margin of eyes and the postero-lateral angles about one- third the greatest diameter of the eyes. Thorax entirely bright ferruginous, subpyri- form, somewhat constricted at the propodeal spiracles; dorsum of thorax, including sides of pronotum with moderate, dense punctures an- teriorly, the punctures becoming progressively larger posteriorly and merging into reticulate on the dorsum and posterior face of propodeum; mesopleura glabrous and micropunctate; meta- pleura and sides of propodeum glabrous, mostly impunctate; lateral margins of dorsum of thorax with two small teeth each side between the an- terior and propodeal spiracles; lateral margins of dorsum and posterior face of propodeum ser- rate; dorsum of thorax clothed with recumbent, ferruginous hairs and erect, pale hairs; pleural areas of thorax with inconspicuous, pale micro- pubescence, more evident on the mesopleura than elsewhere; metasternal process short, ex- tending between the hind coxae anteriorly but not over them, acute at the tip. Abdomen black, except the first segment, the posterior margins of the third, fourth and fifth terga, the last segment, the second sternum, the posterior margins of the fourth and fifth sterna, and a pair of postmedian, transverse, integu- 138 Zoologica: New York Zoological Society [37: 11: mental spots well separated at the midline on the second tergum, all ferruginous; integumental spots of second tergum extending to the lateral margins of tergum and then produced poste- riorly to the posterior margin; first tergum with a median, posterior darkened area, finely, close- ly punctate, the posterior third interspersed with moderate, setiferous punctures, each bear- ing a long, pale hair; second tergum with mod- erate, dense punctures throughout, with sparse, recumbent, dark pubescence, except at the sides and on the integumental ferruginous spots, the sparse pubescence pale; posterior marginal fringe of second tergum black throughout; third to fifth terga with small, dense punctures, clothed with sparse, erect and recumbent pale pubescence; pygidial area well defined, glabrous, the remainder of the pygidial tergum with small, close punctures, and sparse, erect, pale hairs; median, longitudinal carina of first sternum well developed; second sternum with moderate, dis- tinct punctures and sparse, pale hairs; remain- ing sterna with small, close punctures and sparse, pale hairs. Legs entirely bright ferruginous, calcaria pale. Holotype — Female, Blairmont, British Gui- ana, September, 1923 (F. X. Williams), in col- lection of University of Minnesota. Paratypes.— Female, Blairmont, British Gui- ana, September, 1923 (F. X. Williams); female Blairmont, British Guiana, October, 1923 (F. X. Williams); female, Blairmont, British Guiana, 1923-24 (H. E. Box) (135); female, Blairmont, British Guiana, August, 1923 (135); female, Blairmont, British Guiana, October 1, 1923; female, New Amsterdam, British Guiana, Au- gust, 1923 (F. X. Williams). Similiar in apearance to pulchella, but is smal- ler, the paired integumental spots of the second tergum well separated, not confluent, and the pygidium glabrous, not granulate. The paratypes vary in length from 4 to 6 mm. Pseudomethoca puchella, new species Female.— Holotype. Head black, clothed with moderately thin, pale golden pubescence, the proximal two-thirds of mandibles, and the scape and flagellum entirely, ferruginous; thorax en- tirely ferruginous; abdomen black, except the first segment largely ferruginous, and the second with a transverse, slightly postmedian, integu- mental ferruginous band slightly constricted at the middle; posterior marginal fringe of first ter- gum, ferruginous integumental band of second tergum and terga three to five entirely, with very pale golden pubescence; legs ferruginous, except the tips of the femora blackish, and the tibiae and tarsi darker ferruginous than the femora. Length, 10 mm. Head black, the proximal two-thirds of man- dibles, and the scape and flagellum entirely, fer- ruginous; head clothed throughout with moder- ately thin, semi-appressed, pale golden pubes- cence, not obscuring the black integumental color; mandibles apparently edentate at the tip and with a small tooth within near the tip; med- ian area of clypeus a broad, low triangle with the anterior margin as its base, the two sides defined by a distinct carina, and each carina terminating at the anterior margin in a small tooth; antennal tubercles approximate at their base, glabrous; first segment of flagellum equal to or slightly shorter than segments two and three united; antennal scrobes distinctly carinate above; front, vertex and genae with small, dense, contiguous to confluent punctures; posterior margin of ge- nae defined by a distinct carina terminating ante- riorly at the carina bounding the proboscidal fossa and posteriorly on the postero-lateral an- gles of head; ventral surface of head with small distinct, separated punctures. Thorax ferruginous, the pronotal and propod- eal areas with sparse, inconspicuous, pale hairs, the mesonotal area with sparse, black hairs; tho- rax distinctly longer than broad, violin shaped, narrowest at the propodeal spiracular tubercles; pronotal and mesonotal areas with small, dense, contiguous to confluent punctures, the latter be- coming larger on the posterior half of meso- notal area and merging into the distinct reticulations of the dorsum and posterior face of propodeum; lateral margins of dorsum of thorax denticulate, i. e., a small tooth just posterior to the anterior spiracles, a second small tooth midway between the first and the propodeal tubercles, the latter subdentate, and the lateral margins of the posterior face of propodeum with a series of short, small teeth; lateral margins of dorsum of propodeum with- out a prominent tubercle immediately behind the propodeal spiracle; scutellar scale absent; posterior face of propodeum joining the dorsal face at an obtuse angle, not perpendicular; lateral aspect of pronotum with small, close, distinct punctures; remainder of pleural areas glabrous, impunctate, but clothed with sparse, appressed, fine, inconspicuous, pale pubescence. Abdomen black, except the first segment (excluding the posterior margin of the tergum) and a slightly postmedian integumental band slightly constricted medially on the second ter- gum, ferruginous; first tergum with the narrow, posterior margin black, clothed with sparse, erect, pale hairs and a posterior marginal fringe of pale golden hairs; greatest width of trans- verse, ferruginous, integumental band of second tergum approximately one-third the length of the second tergum on the midline, its width 1952] Mickel: Mutillidae (Wasps) of British Guiana 139 gradually narrowing towards the lateral margins of the tergum, and the ferruginous color extend- ing anteriorly along the narrow lateral margin of the tergum almost to, but not reaching the an- terior margin; second tergum with small, dense, contiguous to confluent punctures throughout, clothed with sparse, black pubescence and the apical fringe black, except the ferruginous, trans- verse band and lateral fifths with sparse, very pale golden pubescence; terga three to five with small, dense, contiguous to confluent punctures, clothed throughout with very pale golden pub- escence; pygidial area well defined, granulate; first sternum with small, distinct punctures, and a uniform, median, longitudinal carina; second sternum suffused with ferruginous on median anterior area, with small, close, sometimes con- tiguous and confluent punctures throughout, and clothed with sparse, pale hairs; sterna three to five with small, dense, contiguous to confluent punctures and clothed with sparse, pale hairs; last sternum with small, distinct punctures, and with erect, very pale golden hairs. Legs ferruginous, except the tips of the fe- mora blackish, and the tibiae and tarsi a dis- tinctly darker ferruginous than the femora, clothed throughout with sparse, very pale golden pubescence; calcaria pale. Holotype.— Female, Kartabo, Bartica district, British Guiana, August 13, 1920 (Wm. Beebe), in University of Minnesota collection. Paratypes.— Two females, Kartabo, Bartica, district, British Guiana, August 13, 1920 and June 9, 1924 (Wm. Beebe); female, Tropical Research Station, N. Y. Zool. Soc., British Gui- ana, no. 21252; female, Kurupukari, R. Esse- quibo, British Guiana, January 15, 1933 (D. Vesey-Fitzgerald) . Apparently related to pulicaria Gerstaecker, which is smaller (length, 4 mm), has the legs black, and the clypeus and antennal tubercles ferruginous. The specimen from Kurupukari differs slightly in that the transverse band of the second tergum is reduced to two large trans- verse spots, narrowly separated medially by a black line, and which are widely separated later- ally from the ferruginous color of the lateral tergal margins, and also in the reduced amount of ferruginous on the first abdominal segment. The paratypes vary in length from 6.5 mm to 10 mm. Genus CALOMUTILLA, new genus Genotype— Mutilla temporalis Gerstaecker. Generic Characters— Female; head large, subquadrate, the distance between the posterior margin of the eyes and the postero-lateral angles of head distinctly greater than the greatest di- ameter of the eyes; proboscidal fossa small, sub- triangular, not extending laterad to the insertion of the mandibles;_genal carina extending almost to insertion of mandibles, not connected with the proboscidal fossa by a transverse carina, extending posteriorly on to the vertex; mandibles slender, slightly sinuate, i. e., curved slightly ven- trad beyond the middle, edentate jit the tip, and with a small, distinct, obtuse tooth within near the base; anterior margin of clypeus with a tooth anterior to and slightly laterad of the insertion of the antennae; first segment of flagellum dis- tinctly longer than the length of the second and third segments combined; antennal scrobes strongly carinate above; eyes hemispherical, glabrous. Thorax subpyriform, constricted at the pro- podeal spiracles; lateral margins of dorsum of thorax denticulate~or dentate, two small teeth or tubercles between the anterior and~'prGpodeaT sjfffacTes and the lateral margins of dorsum and posterior face of propodeum with six or seven small teeth; mesosternal lamellate process be- tween and above hind coxae rounded at the tip; scute liar scale absent. First abdominal segment entirely sessile with the second; antero-lateral areas of second ter- gum not elevated and without parallel, short, interrupted carinae; no definite pygidial area present, the posterior third or half of the tergum glabrous. External, distal spine of hind tibiae inserted on a conical or cylindrical process; calcaria pale. Male unknown. The genus includes in addition to the geno- type: crucigera Burmeister, cruciata Smith and williamsi, n. sp. Key to the Females 1. Elevated median area of clypeus distinctly trituberculate; genal carina terminating anteriorly in a distinct elevation 2 Elevated median area of clypeus dis- tinctly bituberculate; genal carina ter- minating anteriorly without an eleva- tion (Novo Friburgo, Brazil) crucigera (Burmeister) 2. Second abdominal tergum with a com- plete, postmedian, transverse band of pale pubescence 3 Second abdominal tergum without a transverse band of pale pubescence, but with three postmedian spots (one median and two lateral) of pale pubes- cence (S. Joao del Rey, Brazil). temporalis (Gerstaecker) 3. Head mostly ferruginous; integument be- neath postmedian, pubescent band of second tergite ferruginous (Para, Brazil) cruciata (Smith) 140 Zoologica: New York Zoological Society [37: 11: Head black; integument beneath post- median pubescent band of second ter- gum black (British Guiana) .williamsi, n. sp. Calomutilla williamsi, new species Female— Holotype. Head and abdomen black, thorax ferruginous; antennae, mandibles and legs mahogany red; genal carina terminating an- teriorly in a small, but distinct, rounded eleva- tion; light pubescent markings of abdominal seg- ments cruciate, the vertical bar interrupted just anterior to the cross bar; external, distal spine of hind tibiae inserted on a cylindrical process. Length, 7 mm. Head quadrate, black (very dark mahogany red when viewed through binocular with micro- scope light), sparsely clothed with recumbent, pale pubescence and erect, dark hairs; median elevated area of clypeus trituberculate, the small tubercles in a transverse line; front, vertex and genae finely, densely punctate; ventral aspect of head with fine, sparse, shallow punctures; genal carina high, sharp, terminating anteriorly in a rounded elevation. Thorax bright ferruginous; dorsum finely, densely punctate; posterior face of propodeum reticulate; lateral aspect of pronotum with fine, shallow, close punctures; mesopleura glabrous, with a line of vertical punctures along the pos- terior margin; metapleura and sides of propo- deum glabrous; dorsum and posterior face of propodeum with sparse, dark, erect hairs; pleural areas with sparse, recumbent, pale pubescence: lateral margins of thoracic notum and of pos- terior face of propodeum, denticulate. Abdomen black, clothed with sparse, erect black hairs and recumbent black pubescence, ex- cept first tergum and broad lateral margins of terga two to five with sparse, recumbent and erect, pale pubescence, and a cruciate mark of recumbent, pale pubescence on terga one to five, the vertical bar of the cross made up of a med- ian spot on the margin of tergum one, a narrow, elongate, median mark on the anterior third of tergum two, a median posterior extension to the posterior margin of the transverse bar on tergum two, and median spots on terga three to five, the transverse bar on the second tergum post- median in position, thus the cruciate pale mark strongly interrupted just anterior to the trans- verse bar; first tergum with sparse, small to fine punctures; second tergum with small, close punc- tures, very minutely, shallowly punctate be- tween the punctures, the area behind the trans- verse pale band less strongly punctured than that anterior to it; terga three to five finely, shallowly, closely punctate; last tergum without a definite pygidial area, the anterior % finely punctate, posterior % glabrous, impunctate. Holotype— Female, Blairmont, British Gui- ana, October, 1923 (F. X. Williams), in collec- tion of University of Minnesota. Genus PERTYELLA, new genus Genotype— Mutilla inyestigatrix Smith. Generic Characters.— Female; head large, sub- quadrate, the distance between posterior mar- gin of eyes and postero-lateral angles of head about equal to the greatest diameter of the eyes; proboscidal fossa small, subtriangular, not extending to insertion of mandibles; genal carina terminating -anteriorly before the in- sertion of the mandibles, not conriecfe'd“Wi!Ti ' the proboscidal Fossa by a transverse carina, extending posteriorly on to the vertex; proxi- mal two-thirds of mandibles broad, then narrowed, forming a conspicuous tooth on the inner margin (visible with the mandibles closed), the distal third slender and attenuated to the edentate tip; anterior margin of clypeus sometimes weakly tuberculate, but not dentate anterior to and laterad of the insertion of the antennae; first segment of flagellum distinctly longer than segments two and three combined; antennal scrobes carinate above; eyes hemi- spherical, glabrous. Thorax subpyriform, strongly constricted at the propodeal spiracles; lateral margins of the dorsum of thorax strongly dentate, two teeth between the anterior and propodeal spiracles, four to five teeth on lateral margins of dorsum and posterior face of propodeum; anterior mar- gin of dorsum of pronotum carinate laterally, the carina extending up on to the humeral angles, but almost absent between the humeral angles; lamellate process of mesosternum ex- tending above and between the posterior coxae, the process broadly rounded at the tip; scutellar scale absent. First abdominal segment entirely sessile or subsessile with the second; antero-lateral areas of second tergum slightly elevated and with short, parallel, interrupted carinae, the latter inconspicuous and almost obsolete in some spe- cies; no definite pygidial area present, the pos- terior third or half of the last tergum glabrous. External, distal spine of hind tibiae inserted on a conical or cylindrical process. Calcaria pale. Male unknown. The genus includes in addition to the geno- type: inimica Cameron, salutatrix salutatrix Smith, salutatrix euryclea Cameron, holmbergi E. Lynch-Arribalzaga, and decora, n. sp. Key to the Females 1. Fourth abdominal tergum clothed en- tirely with pale pubescence 2 1952] Mickel: Mutillidae (Wasps) of British Guiana 141 Fourth abdominal tergum with only a me- dian or linear quadrate spot of pale pubescence 3 2. Thorax bright ferruginous (Mexico) salutatrix salutatrix (Smith) Thorax very dark mahogany red, almost black (Mexico) salutatrix euryclea (Cameron) 3. Median, longitudinal, pale pubescent line of second tergum complete 4 Median, longitudinal, pale pubescent line of second tergum incomplete 5 4. Thorax and legs bright ferruginous, con- colorous (Panama) inimica (Cameron) Thorax bright ferruginous, legs dark fer- ruginous, blackish (Venezuela; Trini- dad) decora, n. sp. 5. Head and abdomen black; second tergum with a complete, transverse, postme- dian, pale pubescent band (Ega, Brazil) investigatrix (Smith) Entirely ferruginous, except the terminal abdominal segments infuscated; second tergum without a transverse, postme- dian, pale pubescent band (Argentina) holmbergi (E. Lynch-Arribalzaga) Pertyella decora, new species Female.— Holotype. Plead, abdomen and legs black, the dorsum and upper parts of the sides of the thorax bright ferruginous, the venter and lower parts of the sides of the thorax blackish; vertex and front with a pair of broad, lateral stripes adjacent to the eyes, the stripes bent inwardly both anteriorly and posteriorly and confluent at the midline, thus forming a circle; anterior portions of stripes not as conspicuous as elsewhere; abdomen marked with three narrow, longitudinal stripes of dense, appressed, pale pubescence, one stripe at each lateral margin and one at the midline, and a narrow, postmedian, transverse stripe of similar pubescence on the second tergum inter- secting the median, longitudinal stripe and join- ing the two lateral longitudinal stripes. Length, 9 mm. Head black, except the mandibles and the flagellar segments beneath, dark ferruginous; genal carina high, thin and lamellate, obtusely angulate at its anterior termination, rectangulate and prominent at the postero-lateral angles; head clothed throughout with sparse, pale pub- escence, except for the lateral stripes on front and vertex as described above, and the circular area enclosed by the stripes with dark, sparse pubescence; ventral aspect of head between the genal carinae weakly, transversely rugose and with scattered, fine punctures; median area of clypeus obscurely punctate, the anterior and posterior margins slightly raised; front vertex and genae with moderate, dense, confluent punctures. Thorax bright ferruginous, except the venter, ventral third of mesopleura and ventral half of metapleura blackish; pleural areas glabrous, micropunctate, clothed with sparse, short, pale micropubescence, except a row of moderate punctures adjacent to the posterior margin of the mesopleura, each with a long, pale hair, and scattered, moderate punctures on the lateral aspects of the pronotum; dorsum of thorax with moderate, dense, confluent punctures anteriorly, the punctures becoming progressively large to- ward the posterior, merging with reticulate on the dorsum of the propodeum and broadly retic- ulate on the posterior face of propodeum; dor- sum clothed with scattered, pale, recumbent, and scattered, dark, long, erect hairs; lateral margins of dorsum of mesothoracic area and metathoracic area, each with a conspicuous tooth, thus two such teeth on each lateral margin between the anterior and propodeal spiracles; lateral margins of dorsum and posterior face of propodeum conspicuously toothed. Abdomen black, maculated with pale pubes- cence as described above, the median, longitud- inal stripe composed of a posterior marginal spot on the first tergum, a complete narrow stripe on the second, and longitudinal spots on terga three to five; first tergum with small, scattered punc- tures bearing long, erect, pale hairs; second ter- gum densely, finely punctate with interspersed, moderate punctures, and the antero-lateral areas very slightly elevated and with inconspicuous, short, sparse, parallel carinae; due to the slight elevation of the antero-lateral areas the anterior half of the median, longitudinal and the post- median, transverse pubescent stripes appear to be depressed; second tergum clothed with sparse, recumbent, black pubescence and scattered, erect, dark hairs; terga three to five finely, densely punctate, clothed with recumbent and erect, blackish hairs, except for the pale, median spots and lateral margins; anterior half to two- thirds of last tergum with small, close punctures and sparse, fuscous to pale hairs, median, long- itudinal carina of first sternum moderate, spar- sely, pale pubescent; second tergum with moder- ate to large, separated punctures interspersed at the sides with fine punctures, clothed with pale hairs; sterna three to five with small, close punctures medially, becoming dense at the sides, all clothed with sparse pale hairs; last sternum with small, close punctures and sparse, pale hairs. Legs very dark ferruginous to black, clothed with pale hairs; calcaria pale. Holotype.— Female, Brasso, Trinidad, B.W.I., 142 Zoologica: New York Zoological Society [37: 11: January 17, 1941 (E. McC. Callan), in collec- tion of University of Minnesota. Paratypes.— Female (1839), Mt. St. Bene- dict, North Ranges, Trinidad, B. W. I., April 24, 1933 (D. Vesey-Fitzgerald) ; female, St. Aug- ustine, Trinidad, B. W. I., December 21, 1943 (E. McC. Callan); female (414), Trinidad, May (D. Vesey-Fitzgerald); female (1216), Arabopo, Mt. Roraimo, 4,000 ft., Venezuela, November 18, 1932 (D. Vesey-Fitzgerald); fe- male, Los Canales, Naiguata, Venezuela, July 23, 1939, 720 meters (G. Vivas-Berthier) , female, Caracas, Venezuela, July, 1891 (Meinert 1,); female, Caracas, October, 1891 (Meinert 1,); female, Carapito, Venezuela, March 26, 1942; female (9148), Venezuela; female (55.89), Venezuela; female, (AYN) Venezuela; female, Macuto, Venezuela, September, 1925 (G. N. Wolcott) . The specimen above numbered “1216” from Mt. Roraimo bears a label “N. Brazil 1216,” but the list of numbers and correspond- ing localities sent me by Mr. Vesey-Fitzgerald cites the locality data as given above for 1216. All of the Venezuela paratypes have the thorax, mandibles and legs more ferruginous than the Trinidad specimens and may warrant at a later time separating them as a subspecies of the Trinidad form. While no specimens of this species are known from British Guiana, the fact that it occurs both in Trinidad, B. W. I., and in Venezuela indicates the possibility of it occurring in British Guiana and therefore it is included here. Similar to investigatrix in many respects, but the latter has the median, longitudinal, pale pubescent stripe of the abdominal terga dis- tinctly interrupted on the anterior half of the second tergum. Genus LOPHOMUT1LLA, new genus Genotype.— Mutilla denticulata Smith. Generic Characters.— Female; proboscidal fos- sa moderate in size, but not extending to base of mandibles; genai carina not extending on to vertex, extending anteriorly to near proboscidal fossa, then parallel to margin of latter to near insertion of mandible, then laterad paralleling margin of mandibular fossa; genai carina con- nected to posterior margin of proboscidal fossa by a transverse carina; mandibles tridentate distally, the distal and inner tooth large, the in- termediate tooth small; inner margin of mand- ible with a distinct tooth intermediate between insertion of mandible and the inner tooth of the distal three; anterior margin of clypeus with a distinct tooth in front of each antennal tubercle; antennal tubercles separated by a distance ap- proximately equal to the diameter of one of them; first segment of flagellum as long or slight- ly longer than the length of segments two and three combined; antennal scrobes strongly car- inate above; head only slightly broader than the thorax; eyes hemispherical, glabrous. Thorax subpyriform, slightly constricted at the prominent, propodeal spiracles; humeral an- gles with a transverse carina approximately equal in length to the first segment of flagellum; lateral margins of dorsum and posterior face of propodeum serrate or dentate; lamellar process between posterior coxae short and acute at the tip. First abdominal segment slightly but distictly nodose, not completely sessile with the second segment; antero-lateral areas of second tergum slightly to strongly elevated and with moderate to exceedingly strong, short, interrupted, lon- gitudinal carinae; last tergum without a definite pygidial area, the posterior third to half of the tergum glabrous, impunctate. Calcaria pale. Male unknown. Distribution.— South America. The genus includes in addition to the geno- type: tuberculata (Fabricius), nigripes (du Buysson), prionophora (Burmeister) , staphy- loma (Gerstaecker), and triguttata, n. sp. Some of the species ( denticulata , tuberculata, priono- phora and nigripes) have the antero-lateral areas of the second abdominal tergum strongly ele- vated and conspicuously crested; others (sta- phyloma and chry Somalia) have these areas only slighty, inconspicuously elevated, but with short, parallel, interrupted carinae. Key to the Species 1. Antero-lateral areas of second abdom- inal tergum strongly elevated and con- spicuously crested with interrupted, parallel carinae 2 Antero-lateral areas of second abdominal tergum only slightly and inconspicu- ously elevated, and with short, parallel, interrupted carinae 7 2. Legs yellow-ferruginous; head yellow- ferruginous, maculated or not with darker areas; thorax yellow-ferrugin- ous, maculated dorsally with darker areas; abdomen yellow-ferruginous, with at least the antero-lateral crests of the second tergum dark ferruginous to black 3 Legs entirely very dark ferruginous or black 4 3. Head unicolorous, pale rufo-testaceous; crested areas of second abdominal ter- gum narrowly marked with ferrugin- ous to black (Para, Brazil) denticulata denticulata (Smith) 1952] Mickel: Mutillidae (Wasps) of British Guiana 143 Head pale rufo-testaceous, marked with ferruginous to black on the front and genae; each crested area of second ab- dominal tergum in a subtriangular, deep black spot (British and Dutch Guiana) . . .denticulata guianensis, n. subsp. 4. Second abdominal tergum black, macu- lated with only a pair of postmedian, ferruginous spots, the latter sometimes confluent medially 5 Second abdominal tergum black, macu- lated with three ferruginous spots, an anterior, median one between the an- tero-lateral crests, and two posterior ones, laterad and posterior to the an- tero-lateral crests 6 5. Postmedian, ferruginous spots of second abdominal tergum confluent medially, thus forming a narrow, transverse, post- median band; head and thorax mostly dark ferruginous (British Guiana) tuberculata (Fabricius) Postmedian, ferruginous spots of' second abdominal tergum separated by about half their diameter; head and thorax black (Venezuela) . .nigripes (du Buysson) 6. Dorsum and posterior face of propodeum clothed throughout with thick, pale pubescence; ferruginous spots of second tergum subequal in size, very large (Novo Friburgo, Brazil) prionophora (Burmeister) Dorsum and posterior face of propodeum mostly black pubescent, but with a long, narrow, cuneiform spot or line of dense, pale pubescence; anterior, median, ferruginous spot of second ter- gum distinctly smaller than the poste- rior ones (British Guiana) . .triguttata, n. sp. 7. Thorax entirely ferruginous, except the humeral angles tinged with black; pale, postmedian, integumental spots of sec- ond tergum confluent medially form- ing a transverse band, or the posterior two-fifths of the tergum entirely fer- ruginous (Brazil meridionale) staphyloma (Gerstaecker) Pronotum almost entirely deep black; re- mainder of thorax ferruginous; integu- mental, postmedian spots of second tergum separated medially (Brazil) chrysomala (Gerstaecker) Lophomutilla denticulata denticulata (Smith), (new combination) 1855. Mutilla denticulata Smith, Cat. Hymen. Brit. Mus., vol. 3, p. 48, female. Redescription of Smith’s type: Female.— Pale rufo-testaceous, the thorax and second abdominal tergum marked with black; mandibles broad at the tip, with two teeth within; antero -lateral areas of second ter- gum greatly elevated and crested with short, high, parallel, thin, dentiform carinae; pygidial area not margined at the sides, entirely glabrous. Length, 8 mm. Head entirely rufo-testaceous, clothed with sparse, erect and recumbent, pale golden pubes- cence; mandibles rufo-testaceous, edentate at the tip, with a small tooth within near the tip, and with a second, larger, obtuse tooth posterior to the first; anterior margin of clypeus with a sharp, distinct tooth each side, the distance be- tween the teeth equal to the length of the pedicel and the first and second segments of flagellum united; clypeus slightly elevated posteriorly at the middle, between the antennal tubercles, the clypeal fringe thin and pale; antennal tubercles distinctly separated; antennae rufo-testaceous, somewhat fuscous above; scape with small, close punctures and sparse, pale pubescence; first seg- ment of flagellum equal in length to the second and third united; antennal scrobes distinctly car- inate above; front, vertex and genae with small, dense, somewhat confluent punctures; genae with a high, sharp carina at the posterior margin, the carina not elevated anteriorly into a tooth or extended posteriorly on to the vertex; postero- lateral angles rounded; head fairly well devel- oped behind the eyes, the distance between the posterior margin of the eyes and the postero- lateral angles slightly but distinctly less than the greatest diameter of the eyes; relative widths of head and thorax, 4.0 - 3.3. Thorax rufo-testaceous, the pronotum, a spot on the posterior part of the mesonotum extend- ing anteriorly along the sides and posteriorly along the midline of the propodeum, black, the areas adjacent to the black spots more or less ferruginous; dorsum of thorax and propodeum with moderate, dense, contiguous punctures, clothed with sparse, erect, blackish pubescence, except the anterior and lateral margins of the dorsum, and the propodeum throughout, with sparse, long, erect, pale hairs; humeral angles distinctly angulate; lateral margins of mesono- tum with two teeth between the anterior and propodeal spiracles, the anterior small and blunt, the posterior larger and acute; anterior margin of propodeal spiracles elevated into a prominent rounded tubercle; thorax narrowed behind the middle, slightly constricted at the propodeal spir- acles; scutellar scale entirely absent; propodeum with a median, longitudinal carina from base to apex; lateral margins of propodeum prominently spinose; lateral aspect of pronotum not defined anteriorly by a carina, with moderate, dense, contiguous punctures and sparse, pale pubes- cence; mesopleura, metapleura and sides of pro- podeum glabrous, micropunctate, clothed with sparse, inconspicuous, pale pubescence, the 144 Zoologica: New York Zoological Society [37: 11: mesopleura defined posteriorly by a ventro- dorsal carina. Abdomen rufo-testaceous, except the elevated antero-lateral areas of second tergum, apical margin of second tergum at the sides, and a small, median spot near the lateral margins, black; first tergum with small punctures, sparse anteriorly, dense towards the posterior margin, clothed with sparse, long, erect, pale hairs, and sparse, appressed, very pale golden pubescence; antero-lateral areas of second tergum greatly elevated and crested with short, high, parallel, thin, dentiform carinae; second tergum with small, dense, confluent punctures throughout, clothed with sparse, erect and recumbent, pale golden pubescence, except the black areas with blackish pubescence; terga three to five with fine, shallow, dense punctures, clothed with sparse, erect and appressed, pale golden pubescence; anterior and lateral margins of last tergum with sparse, erect, pale golden pubescence; pygidial area not margined laterally, entirely glabrous; first sternum with a median, longitudinal, sharp carina on the anterior two-thirds, glabrous, with sparse, fine, pale pubescence; second sternum with moderate, distinct, close punctures, with sparse, erect, pale pubescence, and a thin, pos- terior marginal fringe of pale hairs; sterna three to five finely, densely punctate at the posterior margin, and each with a thin, posterior marginal fringe of pale hairs; last sternum with fine, close punctures and sparse, erect, pale hairs. Legs rufo-testaceous, sparsely clothed with pale hairs; calcaria pale; external, distal tibial spine at the apex of a slender process. Type— Para, Brazil, in British Museum (Nat- ural History), London. Lophomutilla denticulata guianensis, n. subsp. Female— Holotype. Differs from the type sub- species as follows: head not unicolorous, the front with a pair of moderately large, ferrugin- ous spots, and the genae marked with ferrugin- ous to black anteriorly adjacent to the mandibles, the dark color continuing along the genal carina and curving inward at the posterior end of the latter to a ferruginous spot behind the eye; crested areas of second abdominal segment broadly black anteriorly, attenuated posteriorly, thus the black area subtriangular. Length, 7 mm. Holotype.— Female, Courantyne river, Mora Forest, July 9, 1936 (N. A. Weber) (550), in collection of University of Minnesota. Paratypes.— Female, Rio Essequibo (Source), British Guiana ( J. Ogilvie) ; female, Pakaraima Mts., British Guiana (Meyers) (3448). Lophomutilla tuberculata (Fabricius). (new combination) 1805. Mutilla tuberculata Fabricius, Syst. Piez., p. 438, female. Type.— Female, Essequibo (Smidt), Dom. Sehestedt, in Zoological Museum, University of Copenhagen. Only the unique type seen. Lophomutilla triguttata, new species Female— Holotype. Head, thorax, abdomen and legs black, the second abdominal tergum marked with three ferruginous spots, one small, anterior, median and subrectangular extending to the anterior margin between the antero-lat- eral crests, and two very large, postmedian ones posterior to the crests and almost confluent me- dially; dorsum of metathorax and propodeum with a long, narrow, median, cuneiform line of dense, pale pubescence. Length, 10 mm. Head, antennae and mandibles black; mandi- bles, clypeus, scape, genae, under parts of head, and front with sparse, pale pubescence; head between and posterior to compound eyes with sparse, erect, intermixed dark and pale hairs; front, vertex and genae with dense, deep punc- tures, the bottoms of the punctures showing ob- scure ferruginous in certain lights. Thorax entirely black, the dorsum, clothed with sparse, erect, dark hairs, except a long, me- dian, narrow, cuneiform line of dense, pale pub- escence on dorsum of metathorax and propo- deum; thoracic pleura and venter clothed with sparse, recumbent, pale pubescence; dorsum of thorax and propodeum and lateral aspect of pro- notum with dense, deep punctures like those of the head; posterior half of mesopleura and sides of propodeum with strong, close punctures, but not as dense and deep as those on dorsum of thorax; anterior half of mesopleura and all of metapleura micropunctate, shining; humeral angles dentate; sides of dorsum of thorax with a small, obtuse tooth half the distance between the mesothoracic and propodeal spiracles; lat- eral margins of dorsum of propodeum with five conspicuous teeth each side. Abdomen entirely black, except for ferrug- inous spots of second tergum; first tergum clothed with sparse, pale, erect hairs and a me- dian, posterior, marginal, subtriangular spot of thick, recumbent, pale pubescence; second ter- gum with sparse, erect, pale hairs, except the posterior margin black pubescent, and the black areas of tergum also with sparse, intermixed, erect, dark hairs; terga three to five with recum- bent, pale pubescence obscurely interrupted medially with dark pubescence, and with scat- tered, erect, dark hairs intermixed throughout; 1952] Mickel: Mutillidae (Wasps) of British Guiana 145 venter with sparse, pale hairs; crested, carinate, antero-lateral areas of second tergum conspic- uous; posterior half of first tergum and all of second tergum with dense, deep punctures, be- coming less dense and deep towards the lateral margins of the second tergum, and with very small, dense punctures at the posterior margin of the second tergum; terga three to five with very small, very close punctures throughout; anterior third to half of pygidial tergum with moderate, separated punctures; first sternum with a strong, high, median carina; second sternum with large, close, mostly distinct punctures; remaining ster- na with small, close punctures. Legs black, except the tarsi obscurely dark ferruginous, clothed with sparse, pale hairs; calcaria pale. Holotype — Female, Blairmont, British Gui- ana, September, 1923 (F. X. Williams), in col- lection of University of Minnesota. Paratype.— Female, New Amsterdam, British Guiana, August, 1923 (F. X. Williams). Related to prionophora (Burmeister) , but the latter has the propodeum clothed through- out with dense, pale pubescence, the median, anterior, ferruginous spot of the second tergum subequal in size to the posterior pair of spots, and the latter distinctly separated. Also, prio- nophora has abdominal terga three to five clothed throughout with thick, pale pubescence. Genus ANCIPITOT1LLA, new genus Genotype.— A ncipitotilla anceps , new species. Generic Characters.— Male; proboscidal fossa moderate in size, but not extending to base of mandibles; genal carina absent; mandibles tri- dentate d i s taTTyT" fife disl aTafi'crin n e r tooth large, the intermediate tooth small; inner margin of mandible with a small, distinct tooth interme- diate between insertion of mandible and inner tooth of the distal three; anterior margin of cly- peus with a small, but distinct tubercle in front of each antennal tubercle; antennal tubercles widely separated, the distance between them ap- proximately equal to the distance between an antennal tubercle and the inner margin of the compound eye; scape with a single sharp carina beneath; first segment of flagellum slightly short- er to subequal to the second; front with a me- dian, longitudinal impressed line extending from the anterior ocellus to space between antennal tubercles; antennal scrobes not carinate above, but with a small, median tubercle above; humer- al angles of pronotum rounded; tegulae glab- [•ousTmiiunrcrac; sc u tell u m slightly evenly con- vex; dorsum and posterior face of propodeum rounded into one another, without an enclosed space, reticulate throughout; mesosternal lam- ellate process erect in front of posterior coxae, but not extended caudad between them, angu- late at the tip; first abdominal segment slightly nodose, not completely sessile with the second; second tergum with felt lines; second sternum without felt lines; calcaria pale; mar^n^cell of Tron't wings not truncate distally; third sub- marginal cell present but less distinct than the second. The species included here may very probably be the male sex of the females included in the preceding genus Lophoinutilla. Both genera have the proboscidal fossa reduced, not reaching the insertion of the mandibles, the latter are exactly alike in both, the anterior margin of the clypeus is tuberculate or dentate in front of each antennal tubercle in both, and the an- tennal tubercles are considerably separated in both. All the evidence indicating that these males are congeneric with the females of Lophomu- tilla is morphological however, and they are otherwise so different in appearance that it appears best to treat the males as a separate taxonomic unit until more positive biological evidence of their identity is available. Two ad- ditional species to the genotype will be described at a later date. Ancipitotilla anceps, new species Male.— Holotype. Body and legs entirely piccous (black with a distinct reddish tinge), except mandibles, antennal tubercles and integu- ment beneath felt lines of second tergum, yellow- ish testaceous; body and legs clothed throughout with sparse, pale, glittering, mostly erect hairs and the posterior margin of the second, third, fourth and fifth abdominal terga with a sparse fringe of pale, glittering, coarser hairs; front, vertex and genae glabrous, with very small punc- tures well separated on the front and vertex, close on the genae; pronotum and mesonotum with small, separated punctures; scutellum with small, close, more or less contiguous punctures; dorsum and posterior face of propodeum reticu- late; mesopleura with small, separated, weak punctures interspersed with micropunctures; metapleura glabrous, impunctate; sides of propo- deum with small, weak punctures; first tergum with small, sparse punctures; anterior three- fourths of second tergum with small sparse, well separated punctures, the posterior fourth with small, close punctures; terga three to six with small, close punctures; last tergum with small separated punctures; median, longitudinal carina of first sternum low and weakly developed; sec- ond sternum with moderately small, separated punctures, except at the lateral margins with 146 [37: 11: Zoologica: New York Zoological Society small close punctures; last sternum with small, separated punctures. Length, 5 mm. Holotopye— Male, Blairmont, British Guiana, September, 1923 (F. X. Williams), in collec- tion of University of Minnesota. This is the only described species in this genus and probably is the male sex of some species of Lophomutilla from British Guiana. The two species of that genus recorded from British Guiana are denticulata (subsp. guian- ensis) and triguttata. Two other undescribed species of this genus are known to me. These differ from anceps in the puncturation of the integument and color of pubescence. Genus LOPHOSTIGMA, new genus Genotype— Mutilla iracunda Cresson. Generic Characters— Female; proboscidal fos- sa moderate in size, but not extending to base of mandibles; genal carina not extending on to vertex, extending anteriorly to near proboscidal fossa, then parallel to latter to near insertion of mandible, then laterad paralleling margin of mandibular fossa; genal carina connected to posterior margin of proboscidal fossa by a transverse carina; mandibles edentate at the tip_ somewhat broadened at the middle and weakly angulate on the inner margin near the middle, but inner margin not dentate; anterior margin of clypeus very strongly dentate in front of insertion of antennae; first segment of flagel- lum as long or slightly longer than the length of segments two and three combined; antennal scrobes strongly carinate above; head only slightly broader than the thorax; eyes hemi- spherical, glabrous. Thorax subpyriform, slightly constricted at the prominent, propodeal spiracles; humeral angles with a transverse carina approximately equal in length to the first segment of flagellum; lateral margins of dorsum and posterior face of propodeum serrate or dentate; lamellar process between posterior coxae short and acute at the tip. First abdominal segment nodose, disciform, with a narrow, dorsal face and a large, sub- circular anterior face; margin between anterior and dorsal faces dentate or serrate; antero- lateral areas of second tergum slightly to strong- ly elevated; if strongly elevated, the elevation crested with close, parallel, high, longitudinal carinae; if only slightly elevated, at least with short, parallel, interrupted, longitudinal carinae; last tergum without a definite pygidial area, the posterior third to half of tergum glabrous, impunctate. Calcaria pale. Male unknown. Distribution.— Central and South America. The genus includes in addition to the geno- type: cayennensis (Andre), acanthophora ( Dalle Torre) , simoni (du Buysson) , subgracilis (Cameron), /eZ>asT(Mickel) , cincta-' ( du Buys- son), caenodonta1 (Cameron) and alopha, n. sp. Some of the species f iracunda , cayennensis, acanthophora, caenodonta and simoni) have the antero-lateral areas of the second tergum strongly elevated and conspicuously crested; others ( subgracilis , alopha, lebasi and cincta ) have these areas only slightly elevated, but with short, parallel, interrupted carinae. Key to the Females 1. Antero-lateral areas of second abdominal tergum strongly elevated and con- spicuously crested with interrupted, parallel carinae 2 Antero-lateral areas of second abdominal tergum only slightly and inconspicu- ously elevated, and with short, parallel, interrupted carinae 6 2. Second abdominal tergum with a median, anterior spot and either a pair of post- median spots or a transverse post- median band of ferruginous or yellow- ish integumental color 3 Second abdominal tergum without a me- dian, anterior integumental spot, with only a postmedian pair of ferruginous or yellowish integumental spots 4 3. Thorax black; anterior, median, integu- mental spot of second abdominal ter- gum large, almost as large as either one of the postmedian spots (Santarem, Brazil) iracunda (Cresson) Thorax ferruginous; anterior, median, in- tegumental spot of second abdominal tergum small, elongate; second tergum with a transverse, postmedian, integu- mental yellow band (Cayenne) cayennensis (Andre) 4. Thorax without pale pubescent markings; sides of propodeum glabrous, impunc- tate, at the most micropunctate 5 Dorsum and posterior face of propodeum with a pair of broad, lateral stripes of appressed, pale pubescence; sides of propodeum with distinct, deep punc- tures (Venezuela) simoni (du Buysson) 5. Head and thorax black; length 10 mm. (Para, Brazil) . .acanthophora (Dalle Torre) Head and thorax ferruginous; length 6 mm. (British Guiana) caenodonta (Cameron) 6. Sides of propodeum with strong, dis- tinct, deep punctures 7 Sides of propodeum glabrous, impunc- tate, except for a few, scattered punc- 1952] Mickel: Mutillidae (Wasps) of British Guiana 147 tures near the posterior margins (Panama) subgracillis (Cameron) 7. Thorax entirely bright ferruginous, with- out darker markings; lateral margins of dorsum of thorax and dorsum and pos- terior face of propodeum without pale pubescent stripes (British Guiana) alopha, n. sp. Thorax maculated with dark ferruginous to black; lateral margins of dorsum of thorax and dorsum and posterior face of propodeum with pale pubescent stripes 8 8. Vertex with a large, yellow integumental spot (Colombia and Panama) lebasi (Mickel) Vertex entirely ferruginous, not macu- lated with yellow (Venezuela) cincta (du Buysson) Lophostigma caenodonta (Cameron), (new combination) 1912. Ephuta? caenodonta Cameron, Timehri: Journ. Roy. Agri. Com. Soc. Brit. Gui- ana, (3), vol. 2, p. 415, female. Redescription of type: Female— Head and thorax ferruginous, the abdomen black, except the second tergum with a pair of narrowly transverse, preapical, pale yellow, integumental spots; mandibles slender, not toothed within near the tip; parallel carinae of antero-lateral areas of second tergum high and conspicuous; sides of propodeum glabrous; pygidium not margined laterally, entirely gla- brous. Length, 6 mm. Head ferruginous, the genae towards the insertion of the mandibles, and the front an- teriorly at the sides, somewhat blackish, the front and vertex medially with sparse, erect, blackish pubescence, the vertex laterally and the genae with sparse, pale, glittering pubes- cence; mandibles very dark ferruginous, black- ish, slender, edentate at the tip and not toothed within near the tip; anterior margin of clypeus bituberculate, the tubercles separated by about the distance between the antennal tubercles; clypeal fringe thin, pale fuscous; antennal tu- bercles glabrous, distinctly separated; antennae very dark ferruginous, blackish, somewhat paler beneath, scape with small, indistinct punctures and sparse, fine, pale pubescence, first segment of flagellum equal in length to the second and third segments united; antennal scrobes dis- tinctly carinate above; front, vertex and genae with moderate, dense, contiguous and slightly confluent punctures throughout; posterior mar- gin of genae sharply, distinctly carinate, not elevated into a tooth anteriorly, nor extending on to the vertex posteriorly; postero-lateral an- gles rounded; head weakly developed behind the eyes, the distance between the posterior mar- gin of the eyes and the postero-lateral angles equal to half the greatest diameter of the eyes; relative widths of head and thorax, 3.4 -2.6. Thorax entirely ferruginous; dorsum of thorax, and propodeum with moderate, dense, contiguous and somewhat confluent punctures, clothed with sparse, inconspicuous dusky pubes- cence, and scattered, long, erect, pale hairs; thorax widest anteriorly, gradually narrowed towards the rear, slightly constricted at the pro- podeal spiracles; humeral angles distinctly angu- late; lateral margins of mesonotum with a pair of small, blunt teeth between the anterior and propodeal spiracles; scutellar scale entirely ab- sent; anterior margins of propodeal spiracles elevated to form a transverse tubercle; lateral margins of propodeum prominently spinose; sides of pronotum defined anteriorly by a carina, with moderate, close punctures, and sparse, inconspicuous, pale pubescence; anterior half of mesopleura glabrous, micropunctate, the posterior half with moderate, close punc- tures, clothed with sparse, inconspicuous, pale pubescence; metapleura and sides of propodeum glabrous, micropunctate and with sparse, in- conspicuous, pale pubescence. Abdomen black, except the second tergum with a pair of narrowly transverse, preapical, pale yellow, integumental spots; first tergum with small, dense punctures, sparse, erect and appressed, pale, glittering pubescence, the pos- terior margin pale pubescent, the narrow dorsal face only slightly, angulately separated from the anterior face; second tergum with moderate, dense, contiguous and somewhat confluent punctures throughout, the antero-lateral areas strongly elevated and crested with short, parallel, interrupted, dentiform carinae, clothed with sparse, very pale golden pubescence at the sides and posterior third, the elevated areas with dusky pubescence, elsewhere with very pale golden, inconspicuous pubescence, the pos- terior marginal fringe black; terga three to five with fine, close punctures and sparse, blackish pubescence, except laterally pale pubescent; anterior and lateral margins of last tergum with sparse, blackish pubescence; pygidial area not margined at the sides, entirely glabrous; first sternum pale ferruginous, with a very high, thin, sharp carina its entire length, glabrous, clothed with sparse, short, inconspicuous, pale pubescence; second sternum with moderate, distinct, close punctures throughout, with sparse, erect, pale fuscous pubescence, and a very thin posterior marginal fringe of pale fuscous hairs; sterna three to five towards the posterior 148 Zoologica: New York Zoological Society [37: 11: margin, and last sternum throughout, with fine close punctures, three to five each with a very thin, posterior marginal fringe of pale fuscous hairs, the last with sparse, erect, pale fuscous hairs. Legs very dark ferruginous, sparsely clothed with pale pubescence; calcaria pale; external, apical, tibial spine at the apex of a slender prominence. Type.— Female, British Guiana, in British Museum (Natural History), London. Known only from the unique type. Lophostigma alopha, new species Female. — Holotype. Head, abdomen and legs dark ferruginous to black, thorax and first abdominal segment bright ferruginous; second abdominal tergum with a narrow, longi- tudinal, pale pubescent line on the anterior third, and a pair of large, transversely subovate, yellow, integumental, postmedian spots, sep- arated medially. Length, 7 mm. Head dark ferruginous, except antennal tu- bercles, antennae, clypeus and mandibles, bright ferruginous, almost unicolorus with the thorax; head clothed with sparse, recumbent and erect, pale hairs, except the erect hairs on the front dark; front, vertex and genae with moderate, dense, deep punctures, the bottoms of the punc- tures showing ferruginous in certain lights. Thorax entirely bright ferruginous; dorsum of thorax clothed with sparse, recumbent and erect, dark hairs, dorsum of propodeum, and lateral margins of posterior face of propodeum, with very inconspicuous, sparse, appressed, pale pubescence; dorsum and posterior face of pro- podeum also with sparse, long, erect, dark hairs; lateral aspect of pronotum, mesopleura, meta- pleura, except ventral third, and sides of pro- podeum with moderate, distinct punctures, clothed with sparse, inconspicuous, appressed, pale pubescence; lateral margins of thorax with two teeth between the anterior and propodeal spiracles; lateral margins of dorsum and pos- terior face of propodeum each with four longer and stronger teeth. Abdomen dark ferruginous, except the first abdominal segment bright ferruginous like the thorax; anterior face of first tergum shining, micropunctate, except for a very few, scattered, moderate punctures, clothed with sparse, pale pubescence; dorsal face of first tergum micro- punctate and with dense, appressed, pale pu- bescence; margin between anterior and dorsal faces of first tergum denticulate and with a transverse row of very long, erect, dark hairs; antero-lateral areas of second tergum only slight- ly, inconspicuously elevated, but with short, parallel, interrupted carinae; second tergum with moderate, dense, deep punctures, the bot- tom of the punctures on the dark areas showing ferruginous in certain lights; second tergum maculated as described above, the anterior two- thirds with sparse, recumbent and erect, dark hairs, the posterior third with sparse, erect, dark hairs and sparse, recumbent, pale hairs; pos- terior margin of second tergum with a median, more or less indefinite, pale pubescent spot, the remainder of the margin dark, but not black, pubescent; lateral margins of second tergum pale pubescent; terga three to 'five and anterior third of last tergum finely punctate, the punc- tures dense on the third tergum and becoming progressively less dense posteriorly; these same terga with sparse, appressed, pale pubescence, and scattered, erect, dark hairs; median carina of first sternum elevated anteriorly; second sternum with moderate, distinct, close punc- tures and sparse, pale pubescence; sterna three to five finely punctate and sparsely pale pubescent. Legs dark ferruginous, clothed with sparse, pale hairs; calcaria pale. Holotype— Female, Blairmont, British Gui- ana, September, 1923 (F. X. Williams), in col- lection of University of Minnesota. Paratype— Female, Blairmont, British Gui- ana, August, 1923 (F. X. Williams). Related to subgracilis, cincta and lebasi from which it can be readily distinguished by the characters given in the key. Genus MAMMOMUTILLA, new genus Genotype— Mammomutilla pectorosa, n. sp. Generic Characters.— Male; proboscidal fossa large, extending laterad to the insertion of the mandibles; genal carina absent; mandibles moderately slender, suddenly narrowed near the tip, the distal sixth slender and acute at the tip, the angle formed on the inner margin by the sudden narrowing, rounded; antennal tubercles not separated, almost contiguous; scape with a single sharp carina beneath; first segment of flagellum approximately subequal in length to the second; front with a median, impressed, longitudinal line extending from the anterior margin between the antennal tubercles pos- teriorly about three-fourths the distance to the anterior ocellus; antennal scrobes not carinate above, but with a small, median tubercle above; lateral areas of pronotum with an anterior.,., strong, transverse carina terminating at the hu- meral angles in a small, but distinct tooth; scutel- lum slightly, evenly, convex; dorsum of propo- deum rounded into the posterior face, both reti- culate throughout; mesosternum conspicuously 1952J Mickel: Mutillidae (Wasps) of British Guiana 149 convex each side of midline and with a distinct arcuate carina in front of each middle coxae; mesosternum declivous between the carina and the coxal fossa; mesosternal lamellate process short and erect in front of posterior coxae, but not extending cauaad between them, angulate at the tip; first abdominal segment slightly no- dose; second tergum with felt lines; second ster- num without felt lines; calcaria paleTmafghral ceTTSubacirte^af the tip, not truncate; third sub- marginal cell absent. Female unknown. Mammomutilla pectorosa, new species Male— Holotype. Piceous to black, the man- dibles, antennal tubercles and tegulae testace- ous, the legs and antennae slightly lighter in color than the body; body and legs clothed throughout with sparse, pale, glittering hairs, the fringes of the abdominal terga not coarser than other body hairs, except the last two terga with largely fuscous hairs; front and vertex with small, distinct, separated punctures, those on the vertex slightly larger than those on the front; genae with small, close, more or less contiguous punctures; pronotum and mesonotum with mod- erate, distinct punctures; scutellum with mod- erate, close punctures; dorsum and posterior face of propodeum broadly reticulate; meso- pleura with moderately large, close punctures dorsally and ventrally, with moderate, shallow punctures interspersed, and micropunctate else- where; mesosternum with a deep, broad median sulcus, rugoso-punctate anterior to the arcuate carina and becoming much less coarsely sculp- tured anteriorly; metapleura mostly glabrous, impunctate; sides of propodeum with large, close, shallow punctures; tegulae glabrous, im- punctate; first tergum glabrous, with scattered, very small punctures; second tergum with mod- erate, distinct, close to contiguous punctures on the anterior three-fourths, the posterior fourth with the punctures becoming small and very close at the posterior margin; terga three to seven with very small, distinct punctures; first sternum with a distinct, median, longitudinal carina, strongly elevated anteriorly into a conspicuous tooth; second sternum with moderate, distinct, well separated punctures; sterna three to seven with very small, distinct punctures. Length, 6 mm. Holotype— Male, Blairmont, British Guiana, August, 1923 (F. X. Williams), in collection of University of Minnesota. Easily recognized by the form of the mandi- bles and the form of the mesosternum in front of the middle coxae. No satisfactory evidence is available as to the female sex of this species. The obvious possibilities seem to be species of the preceding genus Lophostigma, or Pseudome- thoca diligibilis described herein. However, this is purely conjecture and one can only await some positive information as to what the female may be. Literature Cited Andre, Ernest 1905-06. Nouvelles especes de Mutillides d’Amerique. Zeitschr. fur Hymen, und Dipt., vol. 5, pp. 361-376; vol. 6, pp. 33-48, 65-80, 161-169. Bischoff, H. 1931. Der typus der Mutilla dubia F.— ein Gynander. Mitt. D. ent. Ges., Jahrg. 2, pp. 54-56. Cameron, Peter 1912. The Hymenoptera of the Georgetown Museum, pt. 4. Timehri: Journ. Roy. Agri. and Commercial Soc. British Guiana, ser. 3, vol. 2, pp. 413-416. Cresson, E. T. 1902. Descriptions of some Mutilla from Brazil. Trans. Amer. Ent. Soc., vol. 28, pp. 1-82. Fabricius, J. C. 1787. Mantissa Insectorum, vol. 1, pp. 311-313. 1805. Systema Piezatorum, pp. 428-439. Gerstaecker, C. E. A. 1874. Multillarum Americae meridionalis in- digenarum synopsis systematica et syn- onymica. Arch, fur Naturgeschichte, vol. 40, pp. 41-77, 299-328. Klug, J. C. F. 1821. Entomologiae brasilianae specimen. Nova acta academie Caesarae Leopoldino- Carolinae Germanicae naturae curiosum, vol. 10. pt. 2. pp. 305-324. Lepeletier de Saint-Fargeau 1845. Histoire Naturelle des Insects. Hymenop- teres. vol. 1, p. 14; vol. 3, pp. 517, 589- 646. Linnaeus, Carl 1758. Systema Naturae per regna tria naturae secundum classes, ordines, genera, spe- cies, cum characteribus, differentiis, syn- onymis, locis. Edit. 10 Holmiae. Vol. 1, pp. 582-583. Mickel, Clarence E. 1937. New World Mutillidae in the Spinola col- lection at Torino, Italy (Hymenoptera). Rev. de Entomologia, Rio de Janeiro, vol. 7, pp. 165-207. 150 Zoologica: New York Zoological Society [37: 11: 1938. The neotropical Mutillid wasps of the genus Timulla Ashmead (Hymenoptera, Mutillidae). Trans. Roy. Ent. Soc. Lon- don, vol. 87, pp. 529-680. 1939. A monograph of the Neotropical Mutillid genus Hoplomutilla Ashmead (Hymenop- tera: Mutillidae). Rev. de Entomologia, Rio de Janeiro, vol. 10, pp. 337-403, 641-717. 1939. Monograph of a new neotropical Mutillid genus, Pappognatha (Hymenoptera; Mu- tillidae). Ann. Ent. Soc. Amer., vol. 32, pp. 329-343. 1941. Monograph of the South American Mu- tillid genus Hoplocrates Mickel (Hymen- optera: Mutillidae). Rev. de Entomol- ogia, Rio de Janeiro, vol. 12, pp. 341-414. Olivier, A. G. 1811. Encyclopedic methodique. Histoire natu- relle. Insectes, vol. 8, pp. 51-66, 129-130. Smith, F. 1855. Catalogue of Hymenopterous Insects in the collection of the British Museum, pt. 3, Mutillidae and Pompilidae, pp. 1-63. London. 1879. Descriptions of New Species of Hymen- optera in the Collection of the British Museum, pp. 189-227. London. Spinola, M. 1841. Hymenop teres recueillis a Cayenne en 1839 par M. Leprieur. Ann. Soc. Ent. France, vol. 10, pp. 85-98. 12 A Revision of the Fishes of the Subfamily Alfarinae in the Family Poeciliidae Donn Eric Rosen New York Aquarium, New York Zoological Society1 (Text-figures 1-10) WHILE examining specimens of the Costa Rican poeciliid, Alfaro cul- tratus (Regan), their strong resem- blance to the related species, Furcipenis huberi (Fowler), was noted. A comparison of two geo- graphical populations of Furcipenis huberi , col- lected by Dr. Myron Gordon in Honduras in 1951, with some Alfaro cultratus in the Museum of Zoology, University of Michigan (UMMZ 159157), suggests that the two genera are more closely related than is indicated by their present systematic positions. In his extensive revision of the poeciliid fishes, Regan (1913) indicated that the affinities of Alfaro lay with Tomeurus. Among other details he pointed out that both possess a keel of scales on the ventral margin of the caudal peduncle. Hubbs (1924) agreed with Regan in relating the genera Alfaro and Tomeurus, for he erected the tribes Alfarini and Tomeurini to receive them, and then included these two tribes in the subfamily Tomeurinae Eigenmann (1909). In 1926, however, Hubbs reversed his former esti- mate of the taxonomic positions of the alfarins and tomeurins when he discovered that the ven- tral keels in these two genera are not homolo- gous. In accordance with this view, Hubbs (1926) placed the tribe Alfarini in the subfamily Poeciliinae to which he thought it bore a closer relationship, leaving the single species Tomeur- us gracilis in the subfamily Tomeurinae. The separation of these two groups is well taken, as may be appreciated by an examination of their gonopodia (Text-figs. 1, 2). 1 From the Genetics Laboratory of the New York Zoological Society at the American Museum of Natu- ral History, New York 24. N. Y. Aided in part by a grant from the American Philosophical Society to the New York Zoological Society for Dr. Myron Gordon. Furcipenis huberi was originally described by Fowler (1923) as Priapichthys huberi because of its superficial resemblance to P. annectens (Regan). Hubbs (1931) claimed, however, that Priapichthys huberi Fowler is more closely re- lated to the Heterandriini, and that it possesses certain poeciliin features such as a gonopodial hood on the anterior margin of ray 3. This latter diagnosis was confirmed by Howell Rivero & Hubbs ( 1936) , but they thought that the genera Alfaro and Furcipenis compose a compact group which should be placed apart from the poeciliins as a separate subfamily, the Alfarinae. According to the same authors, the Al- farinae may be characterized as follows : Ventral margin of the caudal peduncle with a well-de- veloped or incipient keel of scales. Pelvic fins relatively unspecialized in adult males. Gono- podium short; the paired halves of ray 5p either joined at the median plane of the fin or separated near the edges of an open trough formed by the flat, fused segments of ray 5a; a well-developed gonopodial hood present (see Clark & Aronson, 1951, for a description of the gonopodial hood in Lebistes ) , surrounding the anterior margin of the gonopodium below ray 3; segments of ray 3 angulated antrorsely into the base or region of attachment of the gonopodial hood; gonopodial rays without serrae. Gonopodial suspensorium simple; gonapophyses without uncinatoid proc- esses. To the above list may be added some addi- tional features which distinguish the known alfarins. 1. The body is strongly compressed, with the dorsal fin origin closer to the caudal base than to the supra-occipital bone of the skull (Text-figs. 4, 5, 6) . These details give the alfarin a streamlined, gambusia-like appearance. 2. The three interorbital pores of the supra-orbital canal 151 152 Zoological New York Zoological Society [37: 12: Text-fig. 1. Distal tip of the gonopodium of Tomeurus gracilis. X 25. are not separated; together they form a deep furrow over the frontal (supra-orbital) bone. The postorbital pores also are confluent, forming major depressions above and slightly behind the orbit (Text-fig. 7). 3. In the gonopodium, a se- ries of lateral wings on the outer faces of the bases of the spines extends from the fifth or ninth spine proximally to the last member of the series. Each lateral wing is perpendicular to the vertical axis of a spine and projects backward away from the tip (Text-fig. 8). Together the lateral wings form a ledge which may function as a partial support for the enlarged gonopodial hood. The gonopodial hood articulates with the two eccentric grooves on the anterior margins of ray 3. These anterior or ventral grooves are the structures that Howell Rivero & Hubbs (1936) refer to as antrorse segments. They are present in most poeciliid gonopodia but they are especially large in the alfarins (Text-figs. 2, 3, 9). Analysis of Differences in the Genera Alfaro and Furcipenis These two nominal genera share many fea- tures, as indicated above. This is especially true with reference to their general body plan and the general configurations of their gonopodial elements. But certain details (of taxonomic importance in this group) have been shown to deviate significantly. Urosomal Keel— The ventral margin of the caudal peduncle of Alfaro cultratus is strongly keeled. Meek (1912) observed that the keel in Alfaro is composed of a double row of scales with their inner surfaces in close apposition. Howell Rivero & Hubbs (1936) showed that the alfarin keel represents a downward extension of the ventro-lateral scales and a loss of the median scale row that normally rounds over the lower edge of the urosome. They regarded the modified squamation in Furcipenis merely as an incip- ient keel because the median scale row still per- sists, although pinched in between the lateral scales. In general, this is true of specimens of Furcipenis liuberi from the inland Department of Cortez, Honduras, in which the keel is present as a slender ridge. But in the Furcipenis collected in the coastal Department of Atlantida, Hon- duras, about 10 kilometers west of Ceiba (Text- Text-fig. 2. Distal tip of the gonopodium of Alfaro cultratus. X 25. Text-fig. 3. Distal tip of the gonopodium of Alfaro huberi ( Furcipenis huberi as formerly constituted). X 25. 1952] Rosen: Revision of Fishes of the Subfamily Alfarinae 153 Text-fig. 4. Male of Alfaro cultratus showing posi- tion and extent of development of the urosomal keel of scales. X 1. Text-fig. 5. Male of Alfaro huberi from inland areas of the Department of Cortez, Honduras. Note the presence of an incipient keel. X 1. Text-fig. 6. Male of Alfaro huberi from the coastal areas of the Department of Atlantida, Honduras. Note enlarged keel with the rudimentary scales of the median row present. X 1. fig. 10), the keel is fully expressed, particularly in adult males, and the median scale row is represented by a few loose scale remnants, or is entirely lacking. The urosomal keel apparently is a poor criterion for the taxonomic separation of these two genera. Coloration— There are two layers of melano- phores in Furcipenis huberi : sub-epidermal or scale melanophores and dermal melanophores. The scale melanophores are present as a fine dusting over the entire body surface and appear to be continuous in distribution with the melano- phores of the fins. The dermal melanophores line the scale pockets and are deeper; they form a distinct, sharply defined reticular pattern, con- forming to the outlines of the scale bases. This pattern fades abruptly and is faint below the mid-lateral line. Occasionally the dermal mel- anophores are grouped in a linear series forming, in addition to reticulations, one or more lateral streaks. There is usually a series of small black blotches on the belly, at the ventral margin of the pectoral fin. The caudal fin has a smooth wash of an even gray which extends to its subdistal limits. The other fins are finely stippled. The coloration of Alfaro cultratus is unlike that of Furcipenis huberi. There are no reticula- tions. Only a fine dusting of sub-epidermal mel- anophores is present in addition to a thin lateral streak that extends from the pectoral to the caudal base. Gono podium.— The gonopodium of Alfaro cultratus differs from that of Furcipenis huberi in only three very minor respects. 1. The small serrations on the anterior margin of ray 4a in the gonopodium of Furcipenis are not present in the gonopodium of Alfaro. 2. The spines of Alfaro are not angular proximally, while the proximal spines of Furcipenis are angular. The shafts of the spines of Furcipenis are attenuate and they overlap. 3. The gonopodial hood of Furcipenis is larger and extends farther out at the tip than the comparable structure in Alfaro (Text-figs. 2, 3). Axial Skeleton— According to Howell Rivero & Hubbs (1936) , Alfaro has six caudal vertebrae which bear ribs, whereas Furcipenis has only four caudal vertebrae which bear ribs. They at- tribute this to the more advanced position of the anal fin and the greater penetration of the urosome by the air bladder in the genus Alfaro. Gordon & Benzer (1945) indicate that compa- rable variations are found among xiphophorin species, but they do not attribute special syste- matic importance to them. Taxonomic Conculsions The nominal genera Alfaro and Furcipenis, of the subfamily Alfarinae, agree in most of their morphological details. This is particularly true of their genitalic elements where differences, al- though constant, are of an extremely minor character. Some other differences not previously mentioned are summarized in Table 1. Alfaro has two or three fewer dorsal fin rays and an average of two more lateral scales. But because of their extreme variability, these details have never constituted a basis for generic separation in poeciliid fishes. The most trenchant remaining argument advanced for recognizing the two gen- era is the circumstance that the marginal keel is incipient or small in Furcipenis, and prominent and well developed in Alfaro. The keel, however, is as strongly expressed in some geographical populations of Furcipenis as it is in the related Alfaro. Therefore the urosomal keel can no longer be regarded as critical for a comparison of the alfarin genera. Since the two species, A. cultratus and F. huberi, differ from each other only superficially, or at most statistically, there is no justification 154 Zoologica : New York Zoological Society [37: 12: Table 1. A Comparison of the Nominal Genera Alfaro and Furcipenis Furcipenis Alfaro Body Compressed Compressed Keel on caudal peduncle Usually present Always present Dorsal origin Closer to caudal base than to posterior margin of skull Closer to caudal base than to posterior margin of skull Head pores Well developed and confluent Well developed and confluent Lateral scales 30 or 31 31 to 33 Dorsal fin rays Gonopodium 9 or 10 7 a. Serrae on ray 4a Present Wanting b. Spines on ray 3 Present; proximal members angular Present; proximal members not angular c. Gonopodial hood Present; large, extending to tip of fin Present; smaller, not quite reach- ing to tip of fin Coloration Heavily reticulated; black belly blotch No reticulations or other dark markings Habitat Clear, shaded, rocky streams, and lakes (Lake Yojoa, Cortez, Honduras) Clear, shaded, rocky streams only for maintaining two monotypic genera to con- tain them. The need of joining closely related genera follows the principles used to synonymize Xiphophorus and Platypoecilus. Gordon & Rosen (1951) and Rosen & Gordon (MS.) show that although there is a wide range and variety of differences among the seven species of the genus Xiphophorus, each species possesses at least several features in common with other Text-fig. 7. Dorsal view of the head of Alfaro huberi showing the position of the head pores. X 4. members of the genus. For example, four of the seven species possess a caudal appendage or sword, while six out of seven species, excluding one of the swordtails, have large, numerous, dis- tal serrae on ray 4p of their gonopodia. In ad- dition, Rosen & Gordon (1951) point out that there are at least five well-defined species groups within the genus Gambusia. Each of these groups is composed of forms with similar gono- podial details and each has a definite geograph- ical range. Among the morphologically different species groups in the genus, there is little inter- group geographical overlap. In conformity with present taxonomic prac- tice, it is desirable to suppress the genus Furci- Text-fig. 8. Diagram of the spines of ray 3 in the gonopodium of Alfaro showing position of lateral wings. X100. 1952] Rosen: Revision of Fishes of the Subfamily Alfarinae 155 Text-fig. 9. Stereogram of the gonopodium of Alfaro huberi showing the position of one of the two eccentric grooves, located on the left ventro- lateral surface of ray 3. The gonopodial hood has been cut at its base; its extent is indicated by the stippled line. X 25. penis. This will serve to lessen the danger of ex- aggerating what are in reality small differences. Assigning A. cultratus and F. huberi to the same genus will also serve to emphasize their seem- ingly close phylogenetic relationship. Of the two generic terms, Alfaro is the older and it thereby enjoys priority in the present synonymy: Alfaro Meek Petalosoma Regan, Ann. Mag. Nat Hist., (8), II, 1908, p. 462 (P. cultratum Regan, name pre-occupied) . Alfaro Meek, Field Mus. Nat. Hist., Zool., X, September, 1912, p. 72 (A. acutiv entrails Meek). Petalurichthys Regan, Ann. Mag. Nat. Hist., X, November, 1912, p. 494 ( Petalosoma cultra- tum Regan; substitute for Petalosoma, pre- occupied). Alfaro cultratus (Regan) Petalosoma cultratum Regan, Ann. Mag. Nat. Hist., XI, 1908, p. 458. Alfaro acutiv entrails Meek, Field Mus. Nat. Hist., Zook, X, 1912, p. 72. Alfaro huberi (Fowler) Priapichthys huberi Fowler, Proc. Acad. Nat. Sci. Phila., 75, 1923, pp. 27-28. Text-fig. 10. Distribution of the genus Alfaro in Central America. Alfaro huberi is repre- sented by solid dots, and Alfaro cultratus by open circles. 156 Zoologicci : New York Zoological Society [37: 12: 1952] Furcipenis huberi Hubbs, Univ. Mich., Mus. Zool., Occ. Pap., 230, 1931, pp.1-3. The following key may be used as a method for the separation of these two species: Key to the Alfarin Species A. Gonopodial hood originating on the ante- rior margin of ray 3. Proximal serrae of ray 4a usually wanting (rudimentary when present). Spines of ray 3 flanked by large, horizontal lateral wings. Eccen- tric grooves on ray 3 large and flaring Alfaro B. Dark reticulations present on body above mid-lateral line. Lower edge of caudal peduncle with a well-developed or an incipient keel of scales. Spines of ray 3 of gonopodium angulated antrorsely. Gonopodial hood extending out to tip of fin or slightly beyond. . . .A. huberi BB. No dark reticulations present on body; only a thin lateral streak present. Lower edge of caudal peduncle al- ways sharply keeled. Spines on ray 3 of gonopodium always vertical. Gonopodial hood not quite reaching to tip of fin A. cultratus AA. Gonopodial hood originating on the an- terior margin of ray 3. Proximal serrae on ray 4a always present. Spines on ray 3 without lateral wings. Eccentric grooves on ray 3 small and not flaring Mollienesia, Poecilia, Micropoecilia, Parapoecilia, AUopoecilia, Limia, Lebistes i Acknowledgments For aid in the preparation of the manuscript and for their many helpful suggestions, 1 wish to thank Dr. Myron Gordon, Mr. James W. Atz, Drs. Reeve M. Bailey, Charles M. Breder, Jr., and Robert Rush Miller and Miss Carmela Ber- ritto. I also thank the American Museum of Natural History for use of their laboratory and library facilities. References Clark, Eugenie, & Lester R. Aronson 1951. Sexual Behavior in the guppy, Lebistes reticulatus (Peters). Zoologica, vol. 36, pp. 49-66. Eigenmann, Carl H. 1909. Reports on the expedition to British Guiana of the Indiana University and the Carnegie Museum. Report no. 1. Some new genera and species of fishes from British Guiana. Ann. Carnegie Mus., vol. VI, pp. 4-54. Fowler, Henry W. 1923. Fishes from Nicaragua. Proc. Acad. Nat. Sci. Phila., vol. 75, pp. 23-32. Gordon, Myron, & Paul Benzer 1 945. Sexual dimorphism in the skeletal elements of the gonopodial suspensoria in xipho- phorin fishes. Zoologica, vol. 30, pp. 57- 72. Gordon, Myron, & Donn Eric Rosen 1951. Genetics of species differences in the mor- phology of the male genitalia of xipho- phorin fishes. Bull. Amer. Mus. Nat. Hist., vol. 95, art. 7, pp. 413-464. Howell Rivero, Luis, & Carl L. Hubbs 1936. Studies of the fishes of the order Cyprino- dontes. XV. The characters and relation- ships of Furcipenis huberi and Alfaro cul- tratus. Occ. Pap. Mus. Zool. Univ. Mich., no. 339, pp. 1-11. Hubbs, Carl L. 1924. Studies of the fishes of the order Cyprino- dontes. II. An analysis of the genera of the Poeciliidae. Misc. Publ. Mus. Zool. Univ. Mich., no. 13, pp. 1-31. 1926. Studies of the fishes of the order Cyprino- dontes. VI. Material for a revision of the American genera and species. Misc. Publ. Mus. Zool. Univ. Mich., no. 16, pp. 1-87. 1931. Studies of the fishes of the order Cyprino- dontes. IX. A new primitive genus of Poeciliidae from Central America. Occ. Pap. Mus. Zool. Univ. Mich., no. 230, pp. 1-3. Meek, Seth Eugene 1912. New species of fishes from Costa Rica. Field Mus. Nat. Hist., Publ. 163, Zool. Ser., vol. X, no. 7, pp. 69-75. Regan, C. Tate 1908. A collection of freshwater fishes made by Mr. C. F. Underwood in Costa Rica. Ann. Mag. Nat. Hist., ser. 8, vol. 11, pp. 455- 464. 1913. A revision of the cyprinodont fishes of the subfamily Poeciliinae. Proc. Zool. Soc. London, vol. 11, pp. 977-1018. Rosen, Donn Eric, & Myron Gordon 1951. A new fish of the genus Gambusia from southern Veracruz, Mexico, with a discus- sion of the tribe Gambusiini Hubbs. Zoologica, vol. 36, pp. 267-272. [MSI Functional anatomy and evolution of the male genitalia in poeciliid fishes. NEW YORK ZOOLOGICAL SOCIETY GENERAL OFFICE 30 East Fortieth Street, New York 16, N. Y. PUBLICATION OFFICE The Zoological Park, New York 60, N. Y. OFFICERS PRESIDENT VICE-PRESIDENTS Fairfield Osborn Alfred Ely Laurance S. Rockefeller Donald T. Carlisle SECRETARY TREASURER Harold J. O’Connell Cornelius R. Agnew SCIENTIFIC STAFF: Zoological Park and Aquarium John Tee- Van Director Leonard J. Goss Assistant Director DEPARTMENT OF TROPICAL RESEARCH Jocelyn Crane Assistant Director Henry Fleming Entomologist John Tee-Van Associate William K. Gregory Associate William Beebe Director Emeritus AFFILIATES C. R. Carpenter Co-ordinator, Animal Behavior Research Programs James R. Simon Director, Jackson Hole Wildlife Park SCIENTIFIC ADVISORY COUNCIL A. Raymond Dochez Caryl P. Haskins Alfred E. Emerson K. S. Lashley W. A. Hagan John S. Nicholas EDITORIAL COMMITTEE Fairfield Osborn, Chairman James W. Atz Lee S. Crandall William Beebe Leonard J. Goss William Bridges James A. Oliver Christopher W. Coates John Tee-Van ZOOLOGICAL PARK Robert M. McClung. . Acting Curator, ‘ Mammals and Birds Grace Davall. Assistant Curator Mammals and Birds James A. Oliver Curator of Reptiles Leonard J. Goss Veterinarian Charles P. Gandal. . .Assistant Veterinarian John V. Quaranta. . .Research Associate, Animal Behavior Lee S. Crandall General Curator Emeritus William Beebe Honorary Curator, Birds AQUARIUM Christopher W. Coates . Curator & Aquarist James W. Atz Assistant Curator Ross F. Nigrelli Pathologist Myron Gordon Geneticist C. M. Breder, Jr Research Associate in Ichthyology Homer W. Smith Research Associate in Physiology GENERAL William Bridges Editor & Curator, Publications Sam Dunton Photographer ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME 37 • PART 4 • DECEMBER 31, 1952 • NUMBERS 13 TO 21 PUBLISHED BY THE SOCIETY The ZOOLOGICAL PARK, New York Contents Page 13. Introduction to the Ecology of the Arima Valley, Trinidad, B.W.I. By William Beebe. Plates I-V; Text-figures 1-5 157 14. Spontaneous Neoplasms in Fishes. VI. Thyroid Tumors in Marine Fishes. By Ross F. Nigrelli. Plates I-IX 185 15. Further Comparisons of Length and Voltage in the Electric Eel, Electro- phones electricus (Linnaeus). By M. Vertner Brown & C. W. Coates. Text-figures 1-7 191 16. A Contribution to the Life History of Colob ura ( Gynaecia auct.) dirce dirce (Linnaeus). (Butterfly). By William Beebe. Plates I & II 199 17. The Saturnioidea (Moths) of Rancho Grande, North-central Venezuela. By Henry Fleming 203 18. The Functional Morphology of the Egg-eating Adaptations in the Snake Genus Dasypeltis. By Carl Gans. Plates I-IV; Text-figures 1-15 209 19. An Annotated List of the Mantids (Orthoptera, Mantoidea) of Trinidad, B.W.I. By William Beebe, Jocelyn Crane & Sally Hughes-Schrader. Plates I- VIII; Text-figures 1&2 245 20. A Comparative Study of Innate Defensive Behavior in Trinidad Mantids (Orthoptera, Mantoidea). By Jocelyn Crane. Plates I-X; Text-figures 1 & 2 259 21. An Experimental Study of the Color Vision of the Giant Tortoise. By John V. Quaranta. Plates I & II; Text-figures 1&2 295 13 Introduction to the Ecology of the Arima Valley, Trinidad, B.W.L’ William Beebe Department of Tropical Research, New York Zoological Society (Plates I-V; Text-figures 1-5) Contents Page I. Introduction 157 II. Location 159 III. Meteorology 159 IV. Biotic Zones 162 V. Botany 165 VI. Zoology 168 VII. References 182 I. Introduction THE object of this paper is to deal briefly with various ecological aspects of the Arima Valley, Trinidad, the location of a research field station of the New York Zoo- logical Society. The valley lies in the middle of the Northern Range, an area which includes large stretches of undisturbed forest reserve. The property which has recently been acquired there for the Zoological Society comprises 200 acres and includes the neighboring estates of Simla and St. Patrick. Buildings on the former have been equipped to serve as a practical laboratory for research in tropical ecology and animal behavior. Throughout all the first three seasons of oc- cupancy, our endeavors have been made pos- sible by the sympathetic and intelligent coopera- tion of all branches of the Trinidad Govern- ment and of innumerable civilian friends. His Excellency the Governor Sir Hubert and Lady Ranee have given freely of their interest and help. In special departments, our appreciation goes particularly to Mr. A. T. Shill and his staff of Her Majesty’s Customs for their courtesy in the entry of our scientific equipment; to Mr. Ludolf Wehekind of the Department of Works and Hy- draulics for the loan of rain-recording instru- 1 Contribution No. 927, Department of Tropical Re- search, New York Zoological Society. ments and for assistance in many other ways; to Mr. A. F. A. Lamb, the Conservator of For- ests, for his close cooperation, and to Mr. Ray Johnson of the Cartographical Department for his aid with government maps. The Imperial College of Tropical Agriculture has placed its library, study collections and other facilities at our disposal. In addition, we wish particularly to thank Dr. T. W. Kirkpat- rick, Dr. F. J. Simmonds, Major R. Senior- White and Dr. Angus Simmons. We also appreciate heartily the friendly assist- ance of members of the Caribbean Commission, of the American Consulate and of successive commanders of the United States Naval Base. We are indebted to the United States Weather Bureau in Washington for the loan of meteoro- logical instruments. To Mr. John Probst and the Trinidad staff of Pan-American World Airways go our thanks for expediting the safe and rapid shipment of live animals. The generous cooperation of the Alcoa Steamship Co., through President W. C. White and Vice-president F. K. Bell in New York and Mr. Edward Collins in Trinidad, has placed us of the New York Zoological Society under a host of obligations. Very special mention must also be made of the constant friendly help of our neighbors in the Arima Valley, Dr. and Mrs. H. Newcome Wright and Squadron Leader and Mrs. John A. Carmichael. I gratefully recognize the assistance of Mr. Charles M. Bogert, Dr. James A. Oliver and Mr. Edwin McConkey in the identification of amphibians and reptiles. Finally, thanks are due members of my staff, including Jocelyn Crane, Henry Fleming and Ellen Ordway, for insect data and for recording and compiling the meteorological and phenolog- ical data presented in the following pages. 157 158 Zoologica: New York Zoological Society [37: 13 Text-fig. 1. Arima Valley, to show general biotic zones. Forest Reserves are shaded, their boundaries approximate. Physical boundaries of valley, north of Arima, include approximately the vertical middle third of this map. See Plate I for contour lines and PI. V for airplane view. 1952] Beebe: Ecology of Arima Valley, Trinidad 159 II. Location Simla, the permanent Field Station of the Department of Tropical Research of the New York Zoological Society, is situated in the north- central mountains of the island of Trinidad, British West Indies. Trinidad itself is a small rectangular chip close to the northeast shoulder of South America. Geologically and biolog- ically, Trinidad and its small sister island of Tobago are typically South American. Trinidad measures roughly 30 by 50 miles with extensive peninsulas projecting from three of the four corners. It contains a total of 1,862 square miles, which makes it slightly larger than Rhode Island and somewhat smaller than Lan- cashire. Trinidad is just north of the delta of the Orinoco, separated from the continent by the Gulf of Paria. At the north and south limits of the gulf the insular peninsulas extend to within eight miles of the Venezuelan coast. The Field Station as a whole consists of the Simla estate of 22 acres, and the adjoining estate of St. Patrick of 176 acres. Simla Laboratory is four miles by road north of the town of Arima, in the southern third of the Arima Valley on the Blanchisseuse Road. Its geographical position is North Latitude 10° 41' 1", and West Longitude 61° 17'. Its elevation above the sea is almost exactly 800 feet. Simla’s geographical relationship to the loca- tions of former stations of the department covers relatively short distances: the two Venezuelan stations, Caripito and Rancho Grande, are, re- spectively 100 and 400 miles to the west (Beebe, 1942; Beebe & Crane, 1947), while Kartabo, British Guiana, lies 350 miles to the southeast (Beebe, 1925). The Northern Range is composed of Jurassic and Cretaceous rocks. These were probably folded in the Middle Eocene. Since then erosion has been proceeding steadily. Omitting the three westernmost drowned valleys which are now marked only by open waterways in the Dragon’s Mouth, the southern slopes of Trinidad’s north- ern mountains are furrowed by 15 more or less parallel valleys each with its respective river or stream flowing along its floor. The Arima is about the center of these tumbled ranges, and ends at the northern end of the valley in a blind gorge. The river’s opposite fellow, on the north- ern slope, is the Marianito, which flows north and empties into the Marianne. This in turn flows into the Atlantic at the little village of Blanchisseuse. III. Meteorology A. General Weather Conditions In spite of its varying altitude, the general climate of the Arima valley is moist and tropical throughout. Between the seasonal forest of its nearly sea-level southern end and the lower mon- tane rain forest of the 1,800-foot northern Pass, there is of course climatic variation. However, rainfall records from three stations, covering three to five years, give averages ranging from only 102 to 121 inches (2,591 to 3,073 mm) per year, although the stations’ range in altitude is from 200 to 1,200 feet. No station has yet been established at the 1,800-foot Pass. Temperature, humidity and sunshine records are known only for Simla during parts of two years, but are probably typical of the valley, its geographical position, at 800 feet, being about midway between the mouth and the Pass. Here the monthly temperature, including parts of both wet and dry seasons, averages in the mid- dle and upper seventies, the percentage of hu- midity in the upper seventies and throughout the eighties, and the sunshine from more than three to less than eight hours daily. At the Pass itself a considerable decrease of temperature and sunshine and an increase of rain and humidity is of course highly probable. There we would expect an approach to the Rancho Grande cli- mate, although the altitude is only one-half that of the Venezuelan location. (See Beebe & Crane 1947, p. 49 ff.) Table 1 gives general weather data for parts of the years 1951 and 1952, when the Simla Station was occupied. The periods included most of the dry and part of the wet seasons of each of the years. The comparison is of special inter- est, since these two seasons were among the wettest and driest, respectively, on record in the colony. They may be regarded therefore as the extremes of any weather likely to be met by future workers in the valley. Each of these years reflected conditions which prevailed throughout the southern Caribbean area. They are a re- minder of the immense variation in tropical weather from year to year, and hence of the caution with which meteorological averages should be accepted in connection with the planning of ecological work. It will be seen that associated with the high rainfall of 1951 are, as would be expected, high humidity records, low temperatures and reduced sunshine. Enveloping cloud or fog is practically un- known at Simla, although, as under similar con- ditions at Rancho Grande, it is prevalent on the mountains surrounding the valley and, more rarely, in pockets between the ridges. B. Rainfall The year at Simla, as in Trinidad as a whole, is divided into only two seasons, dry from Jan- Table 1. Weather at Simla, Arima Valley, Trinidad. February-June, 1951 and 1952 160 Zoologica: New York Zoological Society [37: 13 . 6 pH o .S £ to ON no co (N On r- «o r-H c4 On (N CO to r- o a> CO r-H a a i—i . s P, o •S £ to ON »o •O Tf c4 r-J F- CO OO G O £ Tf- NO r- 01) C3 C/3 a «H-H o P- Ui $ C3 Q Q t-H O 4) < o P CO c/3 c r> CO -D C >S.S rp X H P 53 tw o O E ° I o o a> S-H u •a § .5 tn OQ 03 OQ a *c3 rD > Ui .9 <3 s P S a u O Ph O < < Pi z z < < uary to late May or early June, wet the rest of the year. There is, however, usually a “little dry season,” the petit careme, in October. This is not nearly as pronounced as in British Guiana and elsewhere, but similar to that found in many parts of Venezuela. The rainy season in Trinidad usually begins a full month later than on the adjacent mainland. Text-fig. 2 shows that in the valley most rain falls in August and Sep- tember, least in February and March. Complete rainfall records are available for the Verdant Vale Estate, of which Simla used to be a part, for the years from 1935 to 1940. These are the only ones made so far for an equal pe- riod on the northern range. They are given in Text-figure 2 and Table 2. It will be seen that the six-year average was 108.1 in. (2,746 mm). For comparison, in the year 1951, the first half of which was extremely wet, the Simla total was 117.4 in. (2,982 mm). A three-year average at Torrecilla Estate (1949-1951), altitude 200 feet, was 101.83 in. (2,586 mm); at Spring Hill Estate for the same years, altitude 1,200 feet, 121.24 in. (3,079 mm). Showers do not occur regularly at certain hours of the day or night. Except for the highly abnormal 1951, completely rainy, overcast days are very rare. The maximum rainfall occurring at Simla during 1951 and 1952, between Feb- ruary and June, was 2.76 in. (70.1 mm). It fell on February 10, 1951, a date which nor- mally is well into the dry season. Rainless days during the seasons at Simla ranged from three in February of 1951 to 21 in April of 1952. The total for the year 1951 was 85, in contrast to 133 for the five-year Verdant Vale average. A government rain gauge was established at Simla, thanks to Mr. Ludolf Wehekind of the Department of Works and Hydraulics. During the months when Simla was unoccupied, the records of Mr. J. A. Carmichael are used. The site of his instrument is on a ledge opposite, immediately adjacent to Simla to the south, at an altitude of about 650 feet, approximately 400 yards across a ravine, and in plain sight of the Simla gauge. Variations in the monthly rainfall totals in the two localities are negligible, although moderate differences sometimes occur in the daily totals. C. Humidity As will be seen from Table 1, the average humidity per month varied, in the two years during our occupancy of Simla, from 77.1 to 89%. The saturation point was reached almost every night and sometimes was maintained throughout most of the period of darkness. 1952] Beebe: Ecology of Arima Valley, Trinidad 161 Table II. Rainfall Records at Verdant Vale (1935-40) and Simla (1951 only) Verdant Vale Estate 1935-1940 5-year Average Simla 1951 Only January 4.79 in. (121.7 mm) 8.79 in. (223.3 mm) February 2.65 in. ( 67.3 mm) 12.05 in. (306.1 mm) March 2.99 in. ( 75.9 mm) 5.41 in. (137.4 mm) April 3.77 in. ( 95.8 mm) 3.13 in. ( 79.5 mm) May 7.87 in. (199.9 mm) 12.57 in. (319.3 mm) June 10.86 in. (275.8 mm) 14.70 in. (373.4 mm) July 13.19 in. (335.0 mm) 15.53 in. (394.5 mm) August 16.20 in. (411.5 mm) 8.95 in. (227.3 mm) September 14.50 in. (368.3 mm) 11.89 in. (302.0 mm) October 8.30 in. (210.8 mm) 7.77 in. (197.4 mm) November 12.55 in. (318.8 mm) 10.82 in. (274.8 mm) December 10.43 in. (264.9 mm) 5.79 in. (147.1 mm) TOTALS 108.10 in. (2,745.7 mm) 117.40 in. (2,982.1 mm) Even during the height of the 1952 dry season, it was invariably in the upper nineties. The minimum ever reached was on March 1, 1952, with 43%. The usual monthly minimums ho- vered around 50% and were reached shortly after noon. Through the cooperation of the U. S. Weather Bureau, a Friez hygrothermograph, model 594, was loaned us during 1951 and 1952. Typical weeks in the wet and dry seasons are reproduced in Text-fig. 3. D. Temperature Reference to Table 1 shows that the average temperature per month varied during the 1951- 1952 seasons from 74.4° F (23.6° C) in March of 1951 to 78.5° F (25.8° C) in May of 1952. The usual daily extremes were, very roughly, from around 70° F (21.1° C) (several degrees above or below) before sunrise to the upper middle eighties (approximately 31° C) in the early afternoon. The maximum ever reached was 91° F (32.8° C) on May 23, 1952, between 12 and 1 PM, the minimum, 64° F (17.8° C) on March 30, 1951, between 5 and 7 AM. The highest minimum night temperature recorded was 77° F (25° C) on May 21, 1952, the tem- perature remaining steady between 6 PM and 1 AM; the lowest maximum day temperature was 73° F (22.8° C), on May 12, 1951, a very rainy day during which the temperature re- mained practically unchanged from 10 o’clock in the morning until midnight. Typical weeks in the dry and wet seasons are reproduced in Text-fig. 3. E. Sunshine Sunshine data are given in Table 1. It will be seen that the total hours of sunshine per month during the recorded period varied from 103 to 226, or on the average from about 3 Vi hours daily to more than 7 hours. The fact that Simla is overlooked on east and west by moun- tain ridges reduces the theoretical 12-hour pos- sibility of sunshine to between 9 and 10 hours. Days of full sunshine were practically non-ex- istent. No definite differences were found be- tween the tendency of maximum sunshine to occur in the morning or afternoon. The instruments giving these data were loaned to us by the U. S. Weather Bureau. Two experimental models, one in 1951 and an im- proved version in 1952, were used; both were designed and developed by the Instrument Division. The instrument, known as the U. S. Weather Bureau Photoelectric Sunshine Switch, is designed to record only the periods during which the sun is shining, irrespective of rel- ative intensity. Since, for our biological pur- poses, it was desired to record the hours of maximum diurnal flight activity of insects, the instrument was adjusted to record an impulse whenever the disk of the sun was sufficiently unobscured to cast an easily visible shadow. Basically the sunshine switch consists of an instrument containing two photoelectric cells mounted in an open unshaded area, and a switch box to translate the impulses from the photoelectric cells to a register. The latter pens on a chart the minutes of sunshine or cloudi- ness. One particular merit of this apparatus is 162 Zoologica: New York Zoological Society [37: 13 IN CM Text-fig. 2. Average rainfall at Verdant Vale Estate, close to the site of Simla, 1935-1940. (After data given in Beard, 1946). that it is powered by a six-volt wet cell, making it completely independent of the conventional electric power systems seldom available in the field. The record of a partly cloudy day, totalling 4 Vi hours of sunshine, is reproduced in Text- fig. 4. Because of the newness of the sunshine switch, no special record sheets have yet been printed for it, the Bendix 24-hour Wind Velocity Recorders being used with appropriate altera- tions of hour markings. The sheets were changed after sundown each night, rather than at noon, so that a continuous daytime record of sunshine is obtained on a single sheet. Every small step up or down indicates sunshine during that par- ticular minute. Thus each upward and each downward series of five steps is counted as five minutes of sunshine. F. Wind No wind records were made. As usual in mountainous country, the breezes are variable. The northeast trades are, however, distinctly noticeable as the prevailing wind, although the valley is at most times well protected from their full strength, which is so apparent during the dry season on Caribbean coasts. Only rarely, usually around midday, does the wind become strong enough at Simla to be unpleasent. IV. Biotic Zones (Note: The concepts and terminology of Dr. John Beard, as given in the “Natural Vegetation of Trinidad” (1946) are used throughout this paper. Detailed accounts of all the following zones of vegetation will be found in that refer- ence. See especially pp. 36-42 and 100-115. See Table 3 for systematic lists of botanical names used in this and the following section). Although Rancho Grande in Venezuela is almost four hundred miles away to the west from Simla and the Arima Valley, yet the two places are strikingly alike in physical geographic features. There is a similar general succession of biotic zones from south to north, from low, dry savannas gradually up to a forested pass, and on down the north slope to the sea. The same jungle-bound, winding roads character- ^ ize each place, as also frequent neblinas or fogs and excessive humidity at the higher alti- tudes. 1952] Beebe: Ecology of Arima Valley, Trinidad 163 Text-fig. 3. Hygrothermograph recordings of typical weeks at Simla, Arima Valley. A— Dry Season, March 2-9, 1952. B— Wet Season, June 11-18, 1951. In each case, the upper part of the chart records temperature in degrees Fahrenheit; the lower part, percentage of humidity. The first zone south of the entrance of Arima Valley is savanna country characterized by poor and badly drained soil, covered with a general expanse of bunch grass dotted sparsely with chapparal ( Curatella americana) , serrette (Byr- sonima crassifolia) and a few gnarled shrubs. Closer to the foothills, among secondary sa- vanna resulting from destruction of the forest, are cocorite palms ( Maximiliana elegans), the hardy and prolific survivors of the original cli- max Seasonal Evergreen Forest Formation. The Arima Wood Reserve in the foothills of the Northern Range consists of a transition to Lower Montane Rain Forest. It is composed primarily of Deciduous Seasonal Forest, a for- mation which has arisen following the destruc- 164 Zoologica : New York Zoological Society [37: 13 ITATIOKi Text-fig. 4. Recording of hours of sunshine on March 4, 1952, Simla, Arima Valley. For explanation, see text, p. 161. In reading the chart, the time of day is read from the notation below that of the graph: e.g. above, the first recording was at approximately 7 : 39 AM, the hour at which the sun appeared over the eastern ridge bounding the valley. Sunshine in the early morning and late afternoon is naturally cur- tailed by the surrounding mountains. tion of original vegetation types. Beard describes the general characteristics of this zone as fol- lows: “Tropical temperatures; not exposed to violent winds. Vegetation experiences drought equivalent on free draining soil without human interference to 5 months with less than 4 in. of rain each, of which 2 months have under 1 in. Available moisture 30-50 in. in total amount, derived wholly from precipitation. Human disturbance is active, causing dessica- tion of the habitat. Freedom from water logging and inundation. “A highly discontinuous emergent layer at 40-60 ft. and an almost continuous canopy layer at 10-30 ft. Individuals per 100 acres, 15,000 over 1 ft., 50 over 6 ft., nil over 10 ft. girth. Lianes and epiphytes very scarce, buttressing absent, palms absent. One-half of the species and two-thirds of individuals in upper layer deciduous, one-tenth and one-quarter respec- tively in lower story. Leaves about 50 per cent mesophyllous, 50 per cent microphyllous, pre- dominantly compound in upper story, simple in lower.” (Beard, 1946, p. 39). Yellow poui (Tabebuia serratifolia) is a prom- inent component, and during its extended, spo- radic blooming during the dry season is an ex- cellent diagnostic local guide to the extent of this type of forest. Toward the northern end of the Arima Wood Reserve and on the heights of adjoining St. Patrick Estate, is found a most interesting tran- sitional region. There has been apparently no interference from man, except for rare trail cutting and occasional, supervised lumbering; certainly these slopes have never been cleared. It has elements of both the Deciduous Seasonal Forest to the south and the Lower Montane Rain Forest of the higher altitudes to the east, north and west. In addition, however, it com- prises a varied assortment of extremely local characteristics, including patches of highly de- veloped lianas, and dominants frequently show- ing both buttressing and low branching. These attributes are characteristic of Evergreen or Semi-evergreen Seasonal Forest rather than of either Deciduous Seasonal or Lower Montane. The floral composition does not, however, fit well into any of Beard’s described Semi-ever- green Seasonal formations nor into his Transi- tional Seasonal Forest. It probably bears a close resemblance to the original cover of the foothills, where the climax Evergreen Seasonal Forest of the plains merged into the Lower Montane Rain Forest. The undisturbed heights of most of the re- mainder of the valley are typical Lower Mon- tane Rain Forest, almost all being included in the government’s Forest Reserve. The general characteristics are as follows: “Sub-tropical temperatures. Evaporating ability of the air high due to exposure. No seasonal drought. Available moisture more or less evenly distrib- uted throughout the year and over 70 in. in total amount. Freedom from waterlogging and inun- dation. Exposure causes variations from true rain forest type. Canopy is lowered to between 70 and 100 ft., but is closed at this level. No readily definable lower stratification. Average mature girth 7-8 ft. 16,000 trees over 1 ft. girth per 100 acres, 800 over 6 ft., and 80 over 10 ft. Dominants have long, thin, clear boles. Lianes and epiphytes poorly developed. Palms very rare. Ferns abundant but only small tree-ferns. Buttressing not a character. 100 per cent ever- green, leaves simple, mesophyllous, shiny and slightly leathery. Ground vegetation extremely sparse. Rich tree flora.” (Beard, 1946, pp. 41- 42). However, in certain pockets of specially favorable exposure and humidity, epiphytes and tree-ferns become the equal of those in the Montane Rain Forest of Rancho Grande. 1952] Beebe: Ecology of Arima Valley, Trinidad 165 On the highest slopes and ridges in very lim- ited localities occur patches of true Montane Rain Forest. Its general characteristics are de- scribed as follows: “Temperate climatic condi- tions. Evaporating ability of the air high. No seasonal drought. Available moisture very high in amount, derived both from precipitation and from mist-condensation. Canopy 60 ft., much wind-damaged. Lower stratum 20-40 ft. and under story of tree-ferns and small palms. Average mature girth 4-5 ft., 11,000 trees over 1 ft. girth per 100 acres and 100 over 6 ft. Trees are branching and of bad form. Lianes very abundant, epiphytes tremendously luxuriant. Only small-stemmed palms, but abundant, and several species of large tree-fern. Buttressing not a character. 100 per cent evergreen, leaves simple, mesophyllous, leathery, with epiphyllae. Dense herbaceous ground vegetation. Tree flora extremely poor.” (Beard, 1946, p. 42). On the summit of isolated Morne Bleu, three- fourths of a mile northeast of the Blanchisseuse Road, a small patch of Elfin Woodland is re- ported, with characteristics typical of this for- mation: “Temperate climatic conditions. Evap- orating ability of the air high to very high. No seasonal drought. Moisture very abundant due to almost continuous precipitation and/or con- densation, but possibly not entirely available due to lowered temperature . . . Stunted gnarled thicket, 15-25 ft. in height of stilt-rooted, fleshy- leaved trees with long straggling branches, fes- tooned with moss and lichen. Canopy formed at 6-10 ft. by an under story of tree-ferns and small palms. Flora extremely limited.” (Beard, 1946, p. 42). Beyond the pass, on the north slopes of the Northern Range, the general succession is re- peated. Proceeding north and down, first comes Lower Montane Rain Forest, then Deciduous Seasonal and Evergreen Seasonal. Finally, in- stead of savanna near the coast, is found a kind of thorn woodland, then patches of man- grove marsh and, ultimately, the sand and rocks of the littoral zone. These complete the succes- sion of climax zones. However, an extremely important part of the valley’s zonation is occupied by cultivated land of several kinds. Of primary importance is the cacao land, shaded by tall Erythrina trees. It occupies roughly the middle of the valley, from the Arima Wood Reserve on the south to the beginning of the national Forest Reserve on the heights of the west, north and east. Since much of this land has not been well tended, agriculturally speaking, it forms an excellent habitat for many kinds of wild life. In the valley are also small areas devoted to citrus, tonka and nutmegs, all of which protect the soil and are at the same time good cover for animal life. Hillside field crops, including bananas, yams, tomatoes and corn, are grown on small plots which after two or three seasons are allowed to revert temporarily or semi-permanently to wasteland. The subsequent succession of weeds, bushes and Cecropia and other short-lived trees has not yet been studied floristically in the col- ony (Beard, 1946, p. 139 ff.). As usual, these areas of abandoned cultivation, when supported as they are here by adequate climax forest, are rich locations for animal life. This agricultur- ally ruinous practice of shifting cultivation is not, fortunately, followed extensively in the valley. Furthermore, the government is trying to discourage the system, both by education and by continuing to purchase agriculturally un- suitable lands for the government reserves. On the northern coast are extensive cocoa- nut plantations which, as usual, hold little of general biological interest. V. Botany The floristic composition of the Arima Valley may, from the point of view of practical eco- logy, be thought of in three major divisions. These are, first, the dominant undisturbed Lower Montane Rain Forest; second, the flora of the cultivated tree-crop areas, particularly cacao and citrus; and, third, that of the open areas, confined to forest edges, abandoned clear- ings and trails. According to Beard ( 1946, p. 1 12) , the com- monest trees of the Lower Montane Rain Forest are as follows, in order of their general abun- dance: Canopy Layer— bois gris ( Licania terna- tensis), mahoe ( Sterculia caribaea), serrette (Byrsonima spicata ), wild cacao ( Licania big- landulosa) and wild kaimit ( Micropholis crue- geriana). Lower Story— carimbo ( Guarea glabra Vahl), niaura ( Calliandra guildingii ), bois Tail ( Cassipourea latifolia), wild calabash ( Tube - buia stenocalyx) and red mangue (Mczn'/a gran- diflora). He continues “The diagnostic species are Sterculia and L. biglandulosa in association with either L. ternatensis or Byrsonima; the last two species are almost never both abundant together, the former being dominant on north and east aspects and the moist sites generally, and the latter frequently on the drier south and west aspects.” Two palms, Badris broadwayi and Prestoea pubigera, are occasional. Decidu- ous trees are practically absent. Fortunately for ecological diversity in the valley, the more southern ridges are covered, as has been said, by a transition forest with seasonal 166 Zoologica: New York Zoological Society [37: 13 characteristics. Here in the dry season its bound- aries can be detected from Simla by waves of suddenly, and usually briefly, leafless trees. Here are found many trees typical of lowland and foothill seasonal forest. Although a few of these, such as the hogplum and poui, occur sporadi- cally on the lower slopes of the lower montane region, they are not characteristic there. These seasonal forest trees include silk cotton ( Ceiba occidentalis) , bois d’orme ( Guazuma ulmifo- lia ), incense ( Protium guianense), cedar (Ce- drela mexicana), hogplum ( Spondias mombin), purpleheart ( Peltogyne porphyrocardia) , yellow olivier ( Buchenavia capitata), white olivier ( Terminalia amazonia ), cypre ( Cordia allio- dora), poui ( Tabebuia serratifolia) sandbox ( Hura crepitans), figuier ( Ficus taboganus) and moussara ( Brosimum alicastrum) . The co- corite palm ( Maximiliana elegans) also occurs occasionally in the foothills. Melastomaceae is an important under story family in all the local forests. The tree-crops of the cultivated areas, prin- cipally cacao, citrus and tonka, form the domi- nant characteristics in habitats created by and for them. The citrus is, of course, particularly attractive to many kinds of wild life, from bees to bats. However, of special importance eco- logically is the mountain immortelle ( Erythrina micropteryx), naturalized from Peru, which was originally planted as shade for the cacao. These great emergents have become one of the most characteristic components of the valley, forming the principal habitat of numerous plants and animals. Bromeliads, orchids, night-bloom- ing cereus, termites, beetles, lepidopterous cat- erpillars, ants, frogs, lizards, snakes, humming- birds, caciques, squirrels and silky anteaters are among the diverse groups which find food, shelter or both in this Peruvian species. Since the trees never receive any attention from the cacao cultivators, save to be cut down when they become weakened and hence a danger to the crop, they remain a rich source of wild life, even in the few estates which are carefully cultivated. A great deal of cacao in the valley is not con- sistently well brushed, however, but has been allowed to revert to wilderness. In these areas, and to a lesser extent in all estates, certain quick- growing flowering plants, such as black rod ( Pachystachys coccinea) and cacao lilies ( Hip- peastrum equestre) make the ground layer very attractive to hummingbirds and to many insects. From the point of view of the general en- tomologist, one of the most important divisions of floristic composition is the flora of the forest edges and clearings. Here, in ample light, backed up by abundant forest, are found numerous in- sects and birds around the wide variety of scrub- by trees, shrubs and flowering herbs. The plants of these open areas of the clearings, new and old, resemble those found in similar places throughout the neotropics, with an abundance of Cecropia peltata, black sage ( Cordia cylin- drostachya), Lantana camara, Convolvulus, Bidens pilosa, and, where suitable protection is available, great stands of Heliconia bihai. How- ever much of a pest these “weeds” may be con- sidered by agriculturalists, they are highly ap- preciated by insects, birds, entomologists and ornithologists alike. Typical tropical niches are found along the banks of the Arima River and its tributary streams. Here wild life is found in profusion, among the luxuriant ferns, bamboos, Cyclan- thus, terrestrial arums, and lofty clumps of wild banana (Heliconia) . In the following phenological notes on the seasonal succession of bloom in the Arima val- ley, additional examples of the common shrubs and wildflowers will be included. The basis of choice in the plants selected for observation was purely pragmatic: all are locally common, easily observed, and, in most cases, are important to us because of their being a factor in the lives of certain animals or insects. It will be noted that many of the freely flowering herbs mentioned, such as Bidens and Lantana, are widely dis- tributed in the tropics. As usual with such suc- cessful organisms, they have become a completely integrated part of their Trinidad environment, playing important roles in the ecology of many animals of restricted distribu- tion; often one of their principal characteristics is a practically continuous season of bloom. Casual observers and authorities alike, how- ever, agree that flowering seasons of tropical species often extend throughout the year, al- though, of course, individual plants or branches go through at least brief resting periods. Even with this provision, however, the practice is practically unknown in the north. The swift succession of flowers in a New England meadow, for example, is famous. In the Arima valley, on the other hand, only a few trees and vines, such as Bignonia unguis-cati, rush through their flowering in a few days. Far more common are wild flowers that are in bloom for months, often with several nodes of peak production, or trees, such as cypre, that show blossoms on the same individual for weeks at a time. Except in localities with a severe drought, which cor- responds in the tropics to the winter of the north, the chief regulators of tropical flowering seem to be internal. The details of this tropical physiology are still largely unknown. 1952] Beebe: Ecology of Arima Valley, Trinidad 167 Text-fig. 5A. Flowering seasons of common plants of the Arima Valley, for five months in three suc- cessive years. The shaded areas show relative amounts of rainfall, in respect to the maximum for a monthly period, 16.4", which fell in June, 1950. Notes on flowering were made weekly, by inspection, often by more than one observer. In spite of the obviously large human factor, we consider the general results reliable, showing as they do the general tendency to considerable annual variation, long blooming seasons, and the slight direct dependence on rainfall variation. When the graph line lies flat along the bottom of the species chart, a very slight continuation of bloom is indicated. See text, p. 168, for discussion. 168 Zoologica: New York Zoological Society [37: 13 In the Arima Valley there is only a mild dry season, and blooming activity is intense at this time of the year, between January and June. Most of the forest trees in the valley, as in Trinidad as a whole, tend to have, however, a far shorter and more definite season than do shrubs and herbs. Schimper’s data (1903, p. 252 ff.) showed that the majority of tropical woody plants which had been investigated at that time lost their leaves and flowered in the dry season. Beard’s figures for Trinidad (1946, p. 5) do not agree with this conclusion. He found that one half the deciduous species flower during the rains and fruit during the dry; one- quarter flower during the dry and drop fruit during the dry; one-fourth flower during the dry and drop fruit during the rains. Hence in deciduous Trinidad trees, blossoming is about evenly divided between the two periods. How- ever, virtually all the deciduous species do lose their leaves during the dry season. For this reason it seems possible that some earlier reports of tropical flowering may have risen from the fact that leafless trees in blossom are so much easier to see. Although there seems to be a physiological connection between the dropping of leaves and the appearance of blossoms, there is certainly also an adaptive one, in the tropics as in the north, in connection with pollenization by birds and insects. A comparative study of pollenating agents, whether wind or insects, and the releasing mechanisms of insect attrac- tion of trees blooming in both seasons, would go far toward clearing the matter up. The 3,600-foot cloud forest at Rancho Grande, Venezuela, (Beebe & Crane, 1947), seems to be on a much more definitely seasonal basis than the Arima Valley, showing an ap- proach to the shorter, more circumscribed sea- sons of the north. The lists below divide some common spe- cies of Arima Valley plants into three arbitary categories, in regard to the character of their flowering season. The periods of observation were restricted to the months of February through June, 1950, 1951 and 1952. a. Flowering season definite and annual, rel- atively independent of variations in seasonal drought. (Note: Principal flowering months in parentheses; in general, flowering begins and ends in the preceding and following month, re- spectively) . Trees: Swietinia macrophylla (April); Spon- dias mombin (April) ; Erythrina micropteryx (January-March; Text-fig. 5A); Delonix regia (May-June; Text-fig. 5A); Albizzia caribaea (April); Samanea saman (April); Cordia allio- dora (Feb.-March or April; Text fig. 5A) ; Tab- ebuia pentaphylla (sporadic, March-May) ; Tabebuia serratifolia (sporadic, March-June; Text-fig. 5A). Plants other than trees : Epidendrum fragrans (April-June) ; Oncidium luridum (March; in 1952 a second crest in May); Begonia spp. (Feb.-March); Melastomaceae, numerous gen. et spp. with various moderately well-defined flowering seasons; Bignonia unguis-cati (early May). b. Flowering constant or at frequent intervals, the low ebb definitely correlated with time and degree of seasonal drought. Tree: Cordia cylindrostachya. Plants other than trees: Spathiphyllum cannifolium, Cipura martinicensis (Text-fig. 5B; Mimosa pudica; Desmodium spp.; Ipo- maea spp.; Lantana camara (Text-fig. 5B); Browallia americana (Text-fig. 5C) ; Solanum stramontifolium; Cephaelis tomentosa; Hamelia erecta; Momordica charantia (Text-fig. 5D) ; lsostoma longiflora. c. Flowering constant or at frequent intervals, the resting periods rather irregular and appar- ently little correlated with seasonal drought. Trees: Warscewiczia coccinea; Acnistus arborescens. Plants other than trees: Monstera pertusa; Heliconia bihai (Text-fig. 5B); Heliconia hir- suta; Costus friederichsenii; Costus spiralis (Text-fig. 5B) ; Piper scabnum; Asclepias curas- sivica (Text-fig. 5B); Tussacia pulchella (Text- fig. 5C) ; Pachystachys coccinea (Text-fig. 5D) ; Borreria verticillata; Centropogon surinamensis (Text-fig. 5D) ; Emilia sonchifolia; Bidens pilosa (Text-fig. 5D) ; VI. Zoology In this preliminary paper on the ecology of the Arima Valley, there are presented notes on a few groups of invertebrates and a list of the vertebrates. The only excuse for unannotated lists is for the information of prospective work- ers, for whom a knowledge of mere occurrence is necessary. As such these lists are included. A. Invertebrates To condense into a few sentences even the most casual view of the tropical invertebrate fauna of Arima Valley is a feat of three-ring- circus proportions. As at Rancho Grande, a protozoologist or helminthologist could spend a lifetime invest- igating the fauna of the natural aquaria collected in the bract-pools of bromeliads and heliconias. Land planaria are common, and mollusks, while not in great numbers, are to be found on every walk, some terrestrial, others arboreal or aquatic. Large Crustacea are represented by 1952] Beebe: Ecology of Arima Valley, Trinidad 169 WILD FLOWERS YEAR FEB. MAR. APR. MAY JUNE CIPURA 1950 MARTINICENSIS 1951 « DWARF YELLOW IRIS } 1952 Text-fig. 5B. For explanation, see Text-fig. 5A. 170 Zoologica: New York Zoological Society [37: 13 BROWALLIA 1950 AMERICANA 1951 ( FALSE VIOLET) ■ill lllli 1952 SOLANUM 1950 - ■ STRAM0NIF0L1UM 1951 (PRICKLY NIGHTSHADE) ' k * 1352 tl Text-fig. 5C. For explanation, see Text-fig. 5A. 1952] Beebe: Ecology of Arima Valley, Trinidad 171 MOMORDICA CHARANTIA ( CARILLA) 1950 El 1951 3 1952 m Text-fig. 5D. For explanation, see Text-fig. 5A. 172 Zoologica: New York Zoological Society [37: 13 Table III. Systematic List of Plants Mentioned in the Text* A. TREES Dilleniaceae Curatella americana L. Rough Leaf; Chaparro; Chaparral Guttiferae Marila grandiflora Griseb. Cacao Mangue; Red Mangue; Wild Cacao Bombacaceae Ceiba occidentalis (Spreng.). Burkill. Silk Cotton Sterculiaceae Sterculia caribaea R. Br. Mahoe Guazuma ulmifolia Lam. Bois d’Orme Malpighiaceae Byrsonima crassi folia H.B.K. Serrette Byrsonima spicata (Cav.) Rich. Serrette Burseraceae Protium guianense (Aubl.) March. Incense Meliaceae Guarea glabra Vahl. Carimbo Cedrela mexicana Roem. Cedar; Cigar Box Cedar; Spanish Cedar Swietenia macrophylla King. Honduras Mahog- any Anacardiaceae Spondias inombin L. Hog Plum Papilionatae Erythrina micropteryx Poepp. Anauca Immor- telle; Mountain Immortelle Delonix regia Raf. Flamboyant; Royal Poinciana; Flame Tree Caesalpinieae Peltogyne porphyrocardia Griseb. Purpleheart; Sapatere Mimoseae Calliandra guildingii Benth. Niaure; Niaura Albizzia caribaea (Urb.) Britt. & Rose. Tantakayo Samanea saman (Jacq.) Merrill. Saman; Rain Tree * Sources of identification and nomenclature: Dr. Angus Simmons: Bailey, 1947; Beard, 1946; Marshall, 1939; Williams et al„ 1928-1947; Williams & Williams, 1951. Rosaceae Licania biglandulosa Griseb. Wild Debasse; Wild Cacao; Wild Gasparee Licania ternatensis Hook.f. Bois Gris; Casse Rhizophoraceae Cassipourea latifolia Alston. Bois l’Ail; Garlic Wood Combretaceae Buchenavia capitata (Vahl) Eichl. Yellow Olivier Terminalia amazonia (I. F. Gmel.) Exell. White Olivier Lecythidaceae Eschweilera trinitensis Smith & Beard. Mountain Guatecare Rubiaceae Warscewiczia coccinea (Vahl) Kl. Wakamy; Wild Poinsettia Sapotaceae Micropholis cruegeriana Pierre. Wild Kaimit Hura crepitans L. Sandbox Boraginaceae Cordia alliodora Cham. Cypre Cordia cylindrostachya R. & S. Black Sage Solanaceae Acnistus arborescens Schl. Wild Tobacco Bignoniaceae Tabebuia stenocalyx Sprague & Stapf. Wild Cala- bash Tabebuia pentaphylla Hemsl. Pink Poui Tabebuia serratifolia (Vahl) Nichols. Yellow Poui Moraceae Ficus tobagensis Urb. Figuier Brosimum alicastrum Sw. Moussara Cecropia peltata L. Bois Canon; Bois Canot; Trumpet Wood Palmaceae Badris broadwayi Bailey. Gri-gri Palm Prestoea pubigera Nichols. Prestoe Palm Maximiliana elegans Karst. Cocorite Palm B. PLANTS OTHER THAN TREES Cyclanthaceae Cyclanthus bipartitus Poit. Araceae Monstera pertusa Schott. Hurricane Plant Spathiphyllum cannifolium Schott. Arum Lily Philodendron spp. Amaryllidaceae Hippeastrum equestre (Ait.). Cacao Lily Hymenocallis tubiflora Salisb. Spider Lily Iridaceae Cipura martinicensis Kth. Dwarf Yellow Iris Musaceae Heliconia bihai L., Swartz. Balisier; Wild Plan- tain; Wild Banana Heliconia hirsuta L. Heliconia humilis Jacq. Zingiberaceae Costus friederichsenii Petersen. Snowy Cane Reed Costus spiralis Rose. Scarlet Cane Reed Orchidaceae Epidendrum fragrans Swartz. Purple Streak Orchid Oncidium luridum Lindl. Brown Bee Orchid 1952] Beebe: Ecology of Arinta Valley, Trinidad 173 Table III. Systematic List of Plants Mentioned in the Text ( Continued ) Pipraceae Piper scabrum Swartz. Pepper Bush Mimoseae Mimosa pudica L. Shame Bush; Sensitive Plant Papilionatae Desmodium sp. Blue Pea Begoniaceae Begonia spp. Melastomaceae. Genera et spp. Asclepiadaceae Asclepias curassavica L. Matac; Wild Ipecacuan- ha; Yellow Head; Milkweed Convolvulaceae Ipomaea spp. Verbenaceae Lantana camara L. Wild Sage Solanaceae Browallia americana L. False Violet Solanum stramonifolium Jacq. Prickly Night- shade Bignoniaceae Bignonia unguis-cati L. Cat’s Claw Creeper Gesneraceae Tussacia pulchella Rchb. Harlequin Flower Acanthaceae Pachystachys coccinea Ns. Blackrod; Black Stick Rubiaceae Borreria verticillata (L.) Meyer. White Broom Cephaelis tomentosa Willd. Wild Ipecacuanha Hamelia erecta Jacq. Wild Clove; Coral Stick Cucurbitaceae Momordica charantia L. Carilla; Maiden’s Blush Campanulaceae Isotoma longiflora (L.) Presl. Star Flower Centropogon surinamensis (L.) Presl. Crepe Coq; Deer Meat Compositae Emilia sonchifolia (L.) DC. Dwarf Thistle Bidens pilosa L. Railway Daisy; Spanish Needles good-sized crayfish in the Arima River, and a single land crab, Pseudothelphusia garmani Rathbun. As is usual in the tropics, various orders of Arachnoidea are common, including spiders, scorpions, whip scorpions, chelifers and harvest- men. Ticks are fortunately almost nonexistent in the valley, and mites or bete rouge are only moderately troublesome. At least two species of Peripatus are not rare. We have recorded every order of insects and there is no need to go into any detail as to the vast numbers of Insecta in the tropics. In es- pecial abundance are the various groups of grasshoppers, including Tropidacris with its eight-inch spread of wings, mantids and other stick insects. Termites are less common than would be expected. Odonata are common and include Mecistogaster. Hemiptera and Neuroptera are abundant and among the fulgorids is the remarkable Lanter- naria. Coleoptera and Diptera are in their great tropical numbers, especially as regards quanti- ties of species. Giant harlequin beetles (Acro- cinus) are common, and the interesting, special- ized lymexylonid beetle is occasionally seen. All groups of Hymenoptera are abundant, and ants are legion, but seldom troublesome and easily guarded against in the laboratory. Eciton and Atta are of course present. Lepidoptera are optically dominant. A first walk, or five minutes at the night light will reveal the richness of species of butterflies, or of moths. A quarter of a century ago, more than five hundred Rhopalocera and upwards of one thou- sand Heterocera had been recorded from Trini- dad. During our first two seasons at Simla, nightly collections at a 6' x 6' sheet a few yards from the laboratory, lighted by a 150-watt spot- light, yielded an estimated five hundred species of moths. As a hint for important future study of rel- ative abundance, an 8 PM census of the sheet, from February 5 to 18, gave the following con- sistent percentages of ten families of moths; Pyralidae Microlepidoptera Geometridae Noctuidae Arctiidae Euchromidae Pericopidae Notodontidae Lasiocampidae 53.2 per cent 28 9 8.5 4 3.7 1.7 .36 .27 The richness in species of insect life at Simla is made very apparent in the case of three groups to which, in the first two seasons, we paid especial attention. These groups are Arc- tiidae and Euchromidae among the Heterocera, and Mantoidea of the Orthoptera. Fifty species of Arctiidae moths have, up to the present, been recorded from Trinidad. Within the brief period mentioned at Simla, we took 37 species, 14 of which were identical with already recorded Trinidad species. Twenty-three of our list were new to Trinidad, some of them, doubtless, new to science as well. Eighty-eight species of the family Euchrom- 174 Zoologica: New York Zoological Society [37: 13 idae have, in past years, been collected on the island. During the same two first Simla seasons we captured 42, of which 35 were common to both lists, while 8 were additions to the island fauna. Not only are the unusual numbers taken re- markable, but also the restricted habitat in which they were found, for all the specimens of both the above families of moths were cap- tured within the laboratory or on the lighted screen near by, all within an area whose diam- eter is not more than 15 yards. Before we began our studies in the Arima Valley, ten species of mantids had been re- ported from all of Trinidad. Our collections, made in the valley itself, include all of those previously recorded, and five additional species, bringing the total to fifteen. All except one of these species were taken within 100 yards of Simla laboratory. B. Vertebrates Fish Several seines have been drawn in the waters of the Arima River, and these show that the fish fauna is unexpectedly rich. In such a pre- liminary survey as the present there is nothing to be gained, however, by publishing so inade- quate a representation of the ichthyology of the valley. Future plans include a thorough study of this vertebrate Class. Amphibians Without any thorough or intensive search we have recorded 16 species of frogs and toads from Arima Valley. This is considerably more than half of the amphibian fauna known to occur in Trinidad. Every family except Pipidae is represented, and two interesting frogs, Pipa pipa and Pseudis paradoxa, appear to be absent, although the latter is found in ponds a few miles to the southeast. Eight hylids occur in the valley, two toads, three leptodactylids, one dendrobatid and one ranid. The list is as follows: Hylidae Phyllomedusa burmeisteri burmeisteri Boulenger Hyla maxima (Laurenti) Hyla rubra Daudin Hyla venulosa (Laurenti) Hyla crepitans Wied Hyla punctata (Schneider) Hyla minuta Peters Gastrotheca fitzgeraldi Parker Bufonidae Bufo marinus (Linnaeus) Bufo granulosus Spix Leptodactylidae Eleutherodactylus urichi (Boettger) Leptodactylus peter si (Steindachner) Leptodactylus typhonius (Daudin) Eupemphix pustulosus trinitatis (Boulenger) Dentrobatidae Phyllobates trinitatis Garman Ranidae Rana palmipes Spix Lizards From a total Trinidad lacertilian fauna of about 20 species, we have taken 15, or three- fourths of this number, in Arima Valley. Even the brief periods of observation devoted to lizards reveal considerable complexities of abun- dance and distribution. For example, Anolis and Tupinambis are rare intrusions from the south- ern lowlands, and Cnemidorphorus is confined to lower elevations of St. Pats. The so-called luminous lizard, Proctoporus shrevei Parker, has been taken in some numbers near the Aripo Caves, four miles northeast of Simla. The Arima Valley list is as follows: Gekkonidae Thecadactylus rapicaudus (Houttuyn) House Gecko Common Sphaerodactylus molei Boettger Black-and-white-headed Gecko Common Gonatodes humeralis (Guichenot) Orange-spotted Gecko Common Gonatodes vittatus (Lichtenstein) White-banded Gecko Common Iguanidae Anolis chrysolepis Dumeril & Bibron Stripe-backed Anolis Rare Plica plica (Linnaeus) Spiny Tree Lizard Occasional Iguana iguana iguana (Linnaeus) Iguana Occasional 1952] Beebe : Ecology of Arima Valley, Trinidad 175 Polychrus marmoratus (Linnaeus) Many-colored Tree Lizard Occasional Teiidae Tupinambis nigropunctatus Spix Tegu Rare Cnemidophorus lemniscatus lemniscatus (Linnaeus) Striped Runner; Foot-shaker Rare Ameiva ameiva atrigularis Garman Ameiva; Large Jungle Runner Common Scolecosaurus trinitatis Barbour Worm Lizard Occasional Amphisbaenidae Amphisbaena alba Linnaeus White Legless Lizard Occasional Amphisbaena fuliginosa Linnaeus Black-and-white Legless Lizard Occasional SCINCIDAE Mabuya mabouya aenea (Gray) Trinidad Skink Rare Snakes Trinidad has an ophidian fauna of about 38 species. Seventy per cent of these, or 27 snakes, have been taken in Arima Valley. The present collection is, for the most part, the result of the activities of native men and boys in the valley, and represents only such casual records. Of the poisonous snakes, bushmasters, Lachesis m. muta, are considerably more numerous than ferdelance, Bothrops a. atrox. Coral snakes are more rare. Boa constructors are not uncommon but none reach a large size. The following are all Arima Valley records: Leptotyphlopidae Leptotyphlops tenella Klauber Boidae Epic rates cenchris cenchris (Linnaeus) Corallus enydris cookii Gray Boa constrictor constrictor Linnaeus Colubridae Ninia atrata atrata (Hallowell) Sibon nebulatus (Linnaeus) Pseustes sulphureus sulphureus (Wagler) Pseustes poecilonotus poly le pis (Peters) S pilot es pullatus pullatus (Linnaeus) Drymarchon corals corals (Boie) Chironius carinatus (Linnaeus) Thalerophis richardi coeruleodorsus (Oliver) Leimadophis melanotus (Shaw) Leimadophis reginae reginae (Linnaeus) Liophis cobella cobella (Linnaeus) A tract us trilineatus Wagler Leptodira annulata annulata (Linnaeus) Imantodes cenchoa cenchoa (Linnaeus) Oxyrhopus petola petola (Linnaeus) Pseudoboa neuwiedi (Guenther) Oxybelis fulgidus (Daudin) Oxybelis aeneus aeneus (Wagler) Tantilla melanocephala (Linnaeus) Elapidae Micrurus circinalis (Dumeril & Bibron) Micrurus lemniscatus (Linnaeus) Viperidae Lachesis muta muta (Linnaeus) Bothrops atrox atrox (Linnaeus) Crocodiles and Turtles Caiman crocodilus crocodilus (Linnaeus) Spectacled Caiman Two caimans, about two feet in length, have been reported from the Arima River near the head of the valley. Testudo denticulata Linnaeus Geoemyda punctularia (Daudin) These two species of turtles are not rare in the valley. Birds The period of field work in the Arima Valley, up to the present time, includes the five months of February to June inclusive, throughout three years. During this time 164 species of birds have come under observation, belonging to 14 orders and 41 families. The percentage of this avifauna of the Arima Valley compares favorably with that of the entire island. The carefully edited list of H. Radclyffe Roberts includes 320 species, of which 230 are land birds (Roberts, 1934, List of Trinidad birds with field notes, Tropical Agriculture, 11, No. 4). Our Arima Valley list thus amounts to 70 per cent of the island’s land birds. Correlated with this we must remember that our study area is less than 1 /170th of the square mileage of Trinidad. All details of life histories would be out of place in this preliminary ecological report, and are reserved for future monographic treatment. However, I have added references of monthly breeding records taken from Belcher & Smook- 176 Zoologica: New York Zoological Society [37: 13 er’s “Birds of the Colony of Trinidad and To- bago” (Ibis, Thirteenth Series, 4, 5, 6, 7; 1934- 1937), together with my own observations. Tinamidae A single species of tinamou occurs in Trinidad and this is a common resident in Arima Valley. It is much more often heard than seen. Crypturellus soui andrei (Brabourne & Chubb) Pileated Tinamou (May and October) Fregatidae We have several records of frigatebirds over the valley, straying south from the sea. Fregata magnificens Mathews Frigatebird (March) Cathartidae Two species, the turkey and black vultures, are common in the valley, as elsewhere in Trini- dad. The king vulture seems to have become extinct. Cathartes aura ruficollis Spix Turkey Vulture (November to March) Coragyps atratus (Beckstein) Black Vulture (November to February) Accipitridae Eight out of 17 species of this family, kites, hawks and eagles, have been observed in the valley. Elanoides forficatus yetapa (Vieillot) Swallow-tailed Kite Ictinia plumbea (Gmelin) Plumbeus Kite (March and April) Buteo albicaudatus colonus Berlepsch White-tailed Buzzard (March and April) Buteo nitidus nitidus (Latham) Shining Buzzard Hawk (March and April) Leucopternis albicollis albicollis (Latham) White-collared Hawk Buteogallus ant hr acinus (Lichtenstein) Mexican Black Hawk (March to July) Urubitinga urubitinga urubitinga (Gmelin) Brazilian Eagle (May) Spizaetus o. ornatus (Daudin) Crested Hawk-eagle Pandionidae The osprey occurs as a migrant. Pandion haliaetus carolinensis (Gmelin) American Osprey Falconidae Three out of five falcons occur, including the migrant duckhawk. Falco peregrinus anatum Bonaparte Duckhawk Falco rufigularis rufigularis Daudin Trinidad Bat Falcon (February and July) Falco fusco-caerulescens fusco-caerulescens Vieillot Black-bellied Falcon (April) Scolopacidae The migrant spotted and solitary sandpipers represent this family in Arima Valley. Tringa solitaria solitaria Wilson Solitary Sandpiper Actitis macularia (Linnaeus) Spotted Sandpiper Columbidae Six out of 11 Trinidad pigeons live in the valley. The most common is the gray-fronted dove. Leptotila ruf axilla hellmayri Chapman Gray-fronted Dove (January to July) Oreopeleia montana montana (Linnaeus) Red Dove (May) Oreopeleia linearis trinitatis (Hellmayr & Seilern) Mountain Dove (February) Columba speciosa Gmelin Splendid Pigeon (March and April) Columbigallina minuta minuta (Linnaeus) Gray Ground Dove (March to August) Columbigallina talpacoti rufipennis (Bonaparte) Rufous-winged Ground Dove (May to December) Cuculidae Ten species of cuckoos are found in Trinidad, of which seven occur in our study area. Among these are migrating yellow-billed cuckoos, squir- rel cuckoos and three species of anis. Coccyzus americanus americanus (Linnaeus) Yellow-billed Cuckoo Piaya cayana insulana Hellmayr Squirrel Cuckoo (July) Piaya minuta minuta (Vieillot) Little Cuckoo (July) Taper a naevia naevia (Linnaeus) Brown Cuckoo (June to August) Crotophaga major Gmelin Greater Ani (August to November) Crotophaga ani Linnaeus Common Ani (January to September) Crotophaga sulcirostris sulcirostris Swainson Groove-billed Ani 1952] Beebe: Ecology of Arima Valley, Trinidad 177 Psittacidae Four species of parrots and parrakeets inhabit Arima Valley. The most abundant breeder is the seven-colored parrakeet. Forpus passerinus viridissimus (Lafresnaye) Blue-winged Parrakeet (April and May) Amazona amazonica lobagensis Cory Orange-winged Amazon Parrot (May and June) Pionus menstruus (Linnaeus) Blue-headed Parrakeet Touit batavica (Boddaert) Seven-colored Parrakeet (February and March) Tytonidae The barn owl is found at Simla. Tyto alba hellmayri Griscom & Greenway Barn Owl (February to June) Strigidae Three out of four owls, including screech and pygmy owls, occur. Otus choliba crucigerus (Spix) Screech Owl (April) Ciccaba virgata virgata (Cassin) Variable Wood Owl (April and May) Glaucidium brasilianum phalaenoides (Daudin) Trinidad Pygmy Owl (February to April) Steatornithidae The guacharo or oilbird inhabits a cave near the head of the valley. Steatornis caripensis Humboldt Oilbird (January to April) Nyctiibidae Poor-me-ones are few in number but often heard calling in the valley. Nyctibius griseus griseus (Gmelin) Poor-me-one (April to August) Caprimulgidae Two out of five Trinidad nighthawks are found in the valley. The white-necked is the most common. Nyctidromus albicollis albicollis (Gmelin) White-necked Nighthawk (February to July) Podager nacunda minor Cory Nacunda Nighthawk (April) Micropodidae Six out of seven Trinidad swifts have been recorded from our valley. The most common is the short-tailed swift, the most aberrant the feather-toed palm swift. Chaetura spinicauda spinicauda (Temminck) Spine-tailed Swift. Chaetura cinereiventris lawrencei Ridgway Lawrence’s Swift Chaetura brachyura brachyura (Jardine) Short-tailed Swift (June) Chaeturellus rutilus brunneitorques (Lafresnaye) Rufous-collared Swift (September and November) Streptoprocne zonaris albicincta (Cabanis) Banded Swift Panyptila cayennensis (Gmelin) Feather-toed Palm Swift (April) Trochilidae Fifteen of the 16 of the hummingbirds found on the island inhabit the valley. Eight of these are common or abundant and have been found nesting. Glaucis hirsuta insularum Hellmayr & Seilern Hairy Hermit (February) Phaethornis guy guy (Lesson) Guy’s Hermit Phaethornis longuemareus longuemareus (Lesson) Longuemare’s Hermit (March to May) Florisuga mellivora mellivora (Linnaeus) Great Jacobin Colibri delphinae (Lesson) Brown Violet-ear (February) Anthacothorax viridigula (Boddaert) Green-throated Mango (January) Anthracothorax nigricollis nigricollis (Vieillot) Violet-tailed Mango (April) Chrysolampis mosquitus (Linnaeus) Ruby and Topaz Hummingbird (January to March) Lophornis ornata (Boddaert) Tufted Coquette (February) Chlorestes notatus notatus (C. Reichenbach) Blue-chinned Sapphire (March) Chlorostilbon canivetii caribaeus Lawrence Lawrence’s Emerald (May) Polytmus guainumbi guainumbi (Pallas) Golden-throated Hummingbird (August) Amazilia chionopectus chionopectus (Gould) White-breasted Emerald (March) Amazilia tobaci erythronotus (Lesson) Common Emerald (February) Heliomaster longirostris longirostris (Audebert & Vieillot) Long-billed Star-throat J78 Zoologica: New York Zoological Society [37: 13 Trogonidae All three Trinidad trogons live in Arima Val- ley. By far the most common as to numbers and breeding is the lesser yellow-bellied trogon. Trogon strigilatus strigilatus Linnaeus Greater Yellow-bellied Trogon (April to July) Trogon violaceus violaceus Gmelin Lesser Yellow-bellied Trogon (April to July) Trogonurus collaris collaris Vieillot Collared Trogon; Red-bellied Trogon (April to July) Alcedinidae The little green kingfisher is the only species known to occur. Chloroceryle americana croteta Wetmore Little Green Kingfisher (March to October) Momotidae The single species of Trinidad motmot is often heard and seen. Momotus momota bahamensis (Swainson) King of the Woods (May) Galbulidae Trinidad possesses a single species of jacamar and it is a resident in Arima Valley. Galbula ruf cauda ruficauda Cuvier Jacamar (April to June) Ramphastidae The single species of Trinidad toucan is found in the valley. Ramphastos vitellinus vitellinus Lichtenstein Sulphur-and-white-breasted Toucan (June) Picidae Four out of the five Trinidad woodpeckers live in Arima Valley. None are abundant. Piculus rubiginosus trinitatis (Ridgway) Trinidad Green Woodpecker (May and June) Celeus elegans leotaudi Hellmayr Yellow-headed Woodpecker (April and May) Dryocopus lineatus lineatus (Linnaeus) Big Red-headed Woodpecker (April) Veniliornis kirkii kirkii (Malherbe) Little Red-headed Woodpecker (February) Dendrocolaptidae Two out of the four Trinidad woodcreepers (I prefer this term to the more usual but wholly inappropriate “woodhewers”) have been re- corded. They are not uncommon in the cacao plantations and more open jungle. Dendrocincla fuliginosa meruloides (Lafresnaye) Northern Red-vented Woodhewer (June and July) Xiphorhynchus guttatus susurrans (Jardine) Cocoa Woodhewer (April to June) Furnariidae Four out of five species live in the valley, including both of the spine-tails with their great stick nests. Synallaxis albescens trinitatis Zimmer White-throated Spine-tail (June to December) Synallaxis cinnamonea carri Chapman Trinidad Spine-tail (June to September) Xenops rutilans heterurus Cabanis & Heine Red-tailed Recurved-bill (May) Scleurus albigularis albigularis Sclater & Salvin White-throated Leaf-scraper (October to May) Formicardiidae Trinidad posseses 1 1 species of antbirds. Eight inhabit Arima Valley, and the notes of three, Taraba, Thamnophilus and Grallaria, are char- acteristic of the woods. Taraba major semifasciatus (Cabanis) Para Bush-shrike (May to July) Thamnophilus doliatus fraterculus Berlepsch & Hartert White-barred Bush-shrike (almost every month) Dysithmanus mentalis andrei Hellmayr Andre’s Antbird Myrmotherula axillaris axillaris (Vieillot) White-flanked Antbird (July) Myrmeciza longipes longipes (Swainson) Small Antbird (March to May) Formicarius analis saturatus Ridgway Rufous-necked Ant-thrush (March) Grallaria guatimalensis aripoensis Hellmayr & Seilem Trinidad Long-legged Ant-thrush (May) Cotingidae Four out of five cotingas occur in the valley. The mossy-throated bellbird is by far the most notable, a bird so often heard and not seen, and whose nest and eggs are still a mystery. Attila spadiceus spadiceus (Gmelin) Trinidad Attila (April) Pachyrhamphus polychopterus tristis (Kaup) Black Thick-bill (May to October) Tityra cay ana cay ana (Linnaeus) Cayenne Tityra (March) Procnias averano car nobar ba (Cuvier) Mossy-throated Bellbird 1952] Beebe: Ecology of Arima Valley, Trinidad 179 Pipridae Both of the Trinidad bright-colored manakins are not uncommon. Pipra erythrocephala erythrocephala (Linnaeus) Golden-headed Manakin (February and April) Manacus manacus trinitatis (Hartert) White-breasted Manakin (January to May) Tyrannidae Out of 33 Trinidad flycatchers, 21 have been found in Arima Valley. The commonest are the gray-breasted kingbirds and the kiskadees. The olive-sided flycatcher is a new migrant record for Trinidad. Muscivora tyrannus tyrannus (Linnaeus) Swallow-tailed Flycatcher Tyrannus melancholicus chloronotus Berlepsch Gray-headed Kingbird (January to July) Tyrannus dominicensis subsp. ? White-breasted Kingbird (May to September) Empidonomus varius septentrionalis Todd Guiana Streaked Flycatcher Legatus leucophaius leucophaius (Vieillot) Little White-necked Flycatcher (April) Myiodynastes maculatus tobagensis Zimmer Streaked Flycatcher (May to July) Megarynchus pitangua pitangua (Linnaeus) Broad-billed Kiskadee (April and May) Pitangus sulphuratus trinitatis Hellmayr Kiskadee (December to May) Myiarchus tyrannulus tyr annulus (P.L.S. Muller) Rusty-tailed Flycatcher (April to August) Myiarchus tuberculifer tuberculifer (Lafresnaye & d’Orbigny) Dark-capped Flycatcher (April to June) Contopus cinereus bogotensis (Bonaparte) Caribbean Wood-pewee (May and June) Empidonax euleri lawrencei Allen Lawrence’s Pewee (April to July) Myiophobus fasciatus fasciatus (P.L.S. Muller) Little Brown Flycatcher (April to August) Tolmomyas sulphurescens exortivus (Bangs) Sulphury Flat-bill (April to July) Myiornis ecaudatus miserabilis (Chubb) Short-tailed Pygmy Tyrant Elaenia flavogaster flavogaster (Thunberg) Yellow-vented Crested Flycatcher (February to October) Nuttallornis borealis (Swainson) Olive-sided Flycatcher Camptostoma obsoletum venezuelae Zimmer Little Flycatcher (July) Mionectes olivaceus venezuelensis Ridgway Olivaceous Flycatcher (February to June) Pipromorpha oleaginea pallidiventris (Hellmayr) Oily Flycatcher (February to May) Leptopogon superciliaris pariae Phelps & Phelps Eye-browed Flycatcher (February to July) Hirundinidae Three out of five Trinidad swallows live in the valley. Hirundo rustica erythrogaster Boddaert Barn Swallow Progne chalybea chalybea (Gmelin) Gray-breasted Martin (April and May) Stelgidopteryx ruficolis aequalis Bangs Rough-winged Swallow (April to June) Troglodytidae Both Trinidad species are common in the valley. At Simla Laboratory, both the house and jungle wrens nested within 30 feet of each other. Troglodytes musculus clarus Berlepsch & Hartert Venezuelan House Wren (Almost every month) Thryothorus rutilus rutilus Vieillot Bar-throated Jungle Wren (January to September) Mimidae The mockingbird is a rare stray from the lowland savannas. Mimus gilvus tobagensis Dalmas Mockingbird (February to July) Sylviidae This family is represented only by the Trini- dad ant-wren. Rarnphocaenus melanurus trinitatis Lesson Trinidad Ant- wren Turdidae Four out of the six Trinidad thrushes are found in Arima Valley. Platycichla flavipes melanopleura (Sharpe) Yellow-footed Thrush (May and June) T urdus albicollis phaeopy goides Seebohm White-throated Thrush (January to June) Turdus nudigenis nudigenis Lafresnaye Bare-cheeked Thrush (May to September) Turdus fumigatus aquilonalis (Cherrie) Sabian Thrush (April to July) Vireonidae Three out of four Trinidad species of vireos 180 Zoologica: New York Zoological Society [37: 13 inhabit Arima Valley. The song of one, the yellow-breasted shrike-vireo, is perhaps the most persistent and characteristic bird song ot the valley. Vireo ciltiloquus altiloquus (Vieillot) Antillean Vireo (June) Vireo olivaceus vividior Hellmayr & Seilern Chivi Vireo (May to July) Hylophilus aurantiifrons saturatus (Hellmayr) Golden-fronted Wood-bird (April to August) Cyclarhis gujanensis flavipectus Sclater Yellow-breasted Shrike-vireo (March) Coerebidae Five out of six Trinidad honeycreepers are found in our study area. Coereba flaveola luteola (Cabanis) Common Honeycreeper (almost every month) Cyanerpes caerulea trinitatis (Bonaparte) Purple Honeycreeper Cyanerpes cyaneus cyaneus (Linnaeus) Blue Honeycreeper (July) Chlorophanes spiza spiza (Linnaeus) Green Honeycreeper (June) Dacnis cay ana cay ana (Linnaeus) Turquoise Honeycreeper (June) Parulidae All eight Trinidad warblers occur in the val- ley. Five of these are migrants from North America. Basileuterus culicivorus olivascens Chapman Olive Warbler (April to July) Dendroica petechia aestiva (Gmelin) Yellow Warbler Dendroica striata (Forster) Black-poll Warbler Geothlypis aequinoctialis aequinoctialis (Gmelin) South American Yellow-throat Parula pitiayumi elegans (Todd) Elegant Warbler (June and July) Protonotaria citrea (Boddaert) Prothonotary Warbler Seiurus noveboracensis noveboracensis (Gmelin) Waterthrush Setophaga ruticilla ruticilla (Linnaeus) American Redstart Icteridae Trinidad possesses eight species of this family. The only ones absent from Arima Valley are the yellow-headed Agelaius, and red-breasted Leistes, blackbirds, which are confined to the lowland savanna marshes. They have been seen within two or three miles of our southern valley boundaries. Cacicus cela cela (Linnaeus) Yellow-backed Cacique (January to June) Holoquiscalus lugubris lugubris (Swainson) Boat-tailed Grackle (May to September) Icterus nigrogularis trinitatis Hartert Black-throated Oriole (April to August) Molothrus bonariensis minimus (Dalmas) Glossy Cowbird Psarocolius decumanus insularis (Dalmas) Great-crested Cacique (January onward) Psomocolax oryzivorus oryzivorus (Gmelin) Rice Grackle (January to March) Tersinidae We have a single record of this rare swallow- tanager from Arima Valley, although it has been found breeding on Morne Bleu. Tersina viridis occidentalis (Sclater) Swallow Flycatcher (June) Thraupidae Trinidad possesses 15 tanagers, of which only two have not been found in the valley. Habia rubica rubra (Vieillot) Cardinal Ant-tanager (April) Piranga rubra rubra (Linnaeus) Summer Tanager Ramphocelus carbo magnirostris Lafresnaye Silverbeak Tanager (March to August) Tachyphonus luctuosus flaviventris (Sclater) Lesser White-shouldered Tanager (April and June) Tachyphonus rufus (Boddaert) Greater White-shouldered Tanager (February to August) Tangara gyrola viridissima (Lafresnaye) Red-headed Tanager (March and May) Tangara mexicana vieilloti (Sclater) Sulphur-bellied Tanager (May) Tangara chrysophrys trinitatis (Todd) Larger Spotted Tanager Tanagra trinitatis (Strickland) Yellow-capped Euphonia (April) Tanagra violacea violacea (Linnaeus) Yellow-throated Euphonia (March to May) Thraupis cyanocephala busingi (Hellmayr & Seilern) Blue-headed Tanager (June) Thraupis palmarum melanoptera (Sclater) Palm Tanager (May and June) Thraupis virens nesophila Riley Blue Tanager (February to July) 1952] Beebe: Ecology of Arima Valley, Trinidad 181 Fringillidae Thirteen sparrows and finches inhabit Trini- dad. Of these, three have not been recorded from Arima Valley. Saltator albicollis striatipectus Lafresnaye Stripe-breasted Saltator (May to July) Saltator coerulescens brewsteri Bangs & Penard Gray-breasted Saltator (April to October) Cyanocompsa cyanoides rothschildii (Bartlett) subsp.? Guiana Blue Grosbeak Volatinia jacarina splendens (Vieillot) Glossy Grassquit (Every month) Spermophila bouvronoides Lesson Lesson’s Seedeater (April to September) Spermophila nigricollis nigricollis (Vieillot) Yellow-bellied Seedeater (May to August) Spermophila minuta minuta (Linnaeus) Pygmy Seedeater (April to July) Spermophila intermedia intermedia (Cabanis) Gray Seedeater (May to September) Oryzoborus crassirostris (Gmelin) Thick-billed Seed-finch (April to July) Oryzoborus angolensis torridus (Scopoli) Small-billed Seed-finch (February to August) Mammals The island of Trinidad possesses 70 species of mammals. In our present work in Arima Val- ley we have made no special effort to collect an adequate representation of bats, rats and mice. These groups comprise 49 of Trinidad’s mam- malian fauna and have been omitted from the present preliminary list. Of the remaining 21 mammals, 19 have been collected or observed in the valley; the two omissions include the very rare Carr’s mouse opossum, Marmosa carri Allen & Chapman, of which only a half-dozen specimens are known, and the manatee, Trichechus manatus Linnaeus. This latter creature is not uncommon in the Oropouche River, a few miles east of our valley. The Arima River is far too narrow and shallow to accommodate an animal of such dimensions as the sea-cow. The 19 mammals found in Arima Valley divide well-balanced habitats among them. Nine are terrestrial: armadillo, deer, peccary, aguti, paca, ocelot, raccoon, mongoose and otter. The remaining ten may be classified as arboreal: three opossums, two anteaters, squirrel, porcu- pine, tayra and two monkeys. Another classi- fication reveals that six are vegetarians: deer, peccary, aguti, paca, squirrel and porcupine. Eleven of the remainder are carnivorous, while the two monkeys must be considered om- nivorous. Marsupialia Didelphidae Didelphis marsupialis insularis Allen Large Opossum Occasional to common Philander trinitatis Thomas Trinidad Woolly Opossum Occasional Marmosa mitis chapmani Thomas Trinidad Mouse Opossum Common; taken by hand and in live traps Edentata Myrmecophagidae Tamandua longicaudata Wegner Tamandua Rare; two seen in high trees Cyclopes didactylus (Linnaeus) Silky Anteater Common, but seldom seen in the daytime Dasypodidae Dasypus novemcinctus (Linnaeus) Nine-banded Armadillo Occasionally met with in the woods Artiodactyla Cervidae Mazama rufa (Cuvier) Trinidad Deer Not rare, although being thinned in numbers by hunters Dicotylidae Dicotyles tajacu (Linnaeus) Trinidad Peccary Rare; one seen and others heard near the head of the valley Rodentia Sciuridae Sciurus aestuans hoffmanni (Peters) Trinidad Squirrel Common in cacao plantations Dasyproctidae Dasyprocta aguti (Linnaeus) Aguti Still common in spite of constant shooting Coelogenys paca (Linnaeus) Paca Rare, but still seen in the deeper forests 182 Zoologica: New York Zoological Society [37: 13 Cercolabidae Sy nether es prehensilis (Linnaeus) Tree Porcupine Porcupines are not uncommon but are diffi- cult to see Carnivora Felidae Leopardus pardalis Linnaeus Ocelot Occasional; three have been seen near Simla Procyonidae Procyon cancrivorus (Cuvier) Raccoon Occasional Mustelidae Galictis barbata trinitatis Thomas Tayra Occasional Lutra enudris Cuvier Trinidad Otter Two otters were seen by Mr. Ray Johnson in the Arima River near the head of the val- ley; the only record Herpestidae Her pest es mungo Gmelin Mongoose Common in all kinds of habitats Primates Two species of monkeys are found in Trini- dad, and both occur in Arima Valley. Cebidae Cebus apella Linnaeus Capuchin Monkey Almost extinct in the Northern Range; we have only three records Alouatta seniculus insulans Elliot Trinidad Howling Monkey At least three bands of howlers frequent the valley VII. References Bailey, L. H. 1947. Palmae indigenae Trinitenses et Toba- genses. Gentes Herbarum, 7: Fasc. IV. Beard, J. S. 1944. Climax vegetation in tropical America. Ecology, 25: 127-158. 1946. The natural vegetation of Trinidad. 152 pp. Clarendon Press, Oxford. Beebe, W. 1925. Studies of a tropical jungle; one quarter of a square mile of jungle at Kartabo, British Guiana. Zoologica, 6: 1-193. 1942. Physical factors in the ecology of Cari- pito, Venezuela. Zoologica, 28: 53-59. Beebe, W. & J. Crane 1947. Ecology of Rancho Grande, a subtropical cloud forest in northern Venezuela. Zoologica, 32: 43-60. Marshall, R. C. 1939. Silviculture of the trees of Trinidad and Tobago, British West Indies. 247 pp. Ox- ford University Press, London. Schimper, A. F. W. 1903. Plant-geography upon a physiological basis. 839 pp. Clarendon Press, Oxford. Williams, R. O., E. E. Cheesman, A. W. Hill, & B. L. Burtt 1928- Flora of Trinidad and Tobago. Vol. 1, 1947. Parts 1-7; Vol. 2, Parts 1-3. Department of Agriculture, Trinidad and Tobago. Government Printing Office, Port-of- Spain. Williams, R. O., & R. O. Williams, Jr. 1951. The useful and ornamental plants in Trini- dad and Tobago. Revised fourth edition. 335 pp. Department of Agriculture, Trini- dad and Tobago. Guardian Commercial Printery, Port-of-Spain. 1952] Beebe: Ecology of Arima Valley, Trinidad 183 EXPLANATION OF THE PLATES Plate I Fig. 1. Contour map of the Arima and Marianne Valleys of the Northern Range, Trinidad, B.W.I. Reduced from a section of the Trinidad and Tobago Government Sur- vey map, 1:50,000, Sheet B. Government Reserve boundaries as of 1922-1925. Since then they have been enlarged. See also PI. V, Fig. 9. Plate II Fig. 2. Simla, the field station of the Department of Tropical Research of the New York Zoological Society, from the eastern ridge. Fig. 3. Seining the Arima River, below Simla. Plate III Fig. 4. Simla from the southeast. Fig. 5. Inside the laboratory at Simla. Plate IV Fig. 6. Lower Montane Rain Forest, Government Reserve, Arima Valley. Fig. 7. Roadside, showing tree ferns. Lower Mon- tane Rain Forest. Plate V Fig. 8. Edge of transitional type of forest, St. Patrick Estate, showing figuier ( Ficus tobagensis ) with strangler fig. Stream bed flora in foreground, including Heliconia, Costus, Cyclanthus and bamboo. Fig. 9. Airplane view of Arima Valley, from the town of Arima approximately to Spring Hill Estate. Pale line is the road, which closely follows the Arima River. Arrow points to Simla. , ■ BEEBE PLATE 1 Blanchisseuse 7?«« BlitnclijSsPiisf1 C«* tsg&mg&m !#%r^ FIG. 16 SPONTANEOUS NEOPLASMS IN FISHES. VI. THYROID TUMORS IN MARINE FISHES. NIGRELLI PLATE IX FIG. 17 FIG. 18 SPONTANEOUS NEOPLASMS IN FISHES. VI. THYROID TUMORS IN MARINE FISHES. 15 Further Comparisons of Length and Voltage in the Electric Eel, Electrophorus electricus (Linnaeus) M. Vertner Brown Department of Physics, the College of the City of New York & C. W. Coates New York Aquarium, New York Zoological Society (Text-figures 1-7) A COMPARISON of length and voltage in the electric eel, Electrophorus elec- tricus (Linnaeus), has been published (Coates & Cox, 1945), but at that time only one observation on eels shorter than 40 centi- meters was available. We have recently received shipments of smaller eels and it now seems worth while to supplement the earlier paper. The new data fit smoothly into the previous curves (Text-figures 1 & 2). Although our graphs show no points for eels shorter than 10 centimeters, we did make brief qualitative ob- servations of six fish, four to nine centimeters long. The maximum obtainable voltages ranged from 15 to 30 volts. This is within the range to be expected, as may be seen from Text-figure 2. Reliable measurements were not made, since the fish were received in poor condition and died shortly after arrival. The electric eel breathes atmospheric air and may be kept out of water for extended periods of time. When taking data, we are therefore able to place the eel in a dry trough and so control the currents flowing outside its body. Metallic electrodes are pressed against the eel and leads are carried to the vertical deflection plates of a cathode ray oscillograph. All data in this paper were taken with one electrode at the anterior end of the electric organ, the second being placed at various posi- tions posterior to the first. No matter where the electrodes are placed, however, the anterior electrode is always positive to the posterior. Maximum voltage is obtained with 60 to 70 per cent of the electric organ between the electrodes. Including a larger fraction of the organ pro- vides the same peak voltage but for a longer period of time. A part of the electric organ yields as large a voltage as the whole because all parts of the organ do not discharge simultaneously; excitation begins at the anterior end and pro- gresses rapidly down the eel. The maximum voltage per centimeter is gen- erated by a small segment of the electric organ at the anterior end. This results partly from the lack of simultaneity noted above and partly from the decrease in number of electroplaxes per centimeter which occurs posteriorly (Cox, et al., 1940). For this reason it is necessary to use electrodes separated by no more than two centimeters in measuring the maximum voltage gradient in small eels. In the case of large fish, however, the gradient across the anterior 10 centimeters is as great as that across the first two. The points of Text-figure 3 are plotted so as to distinguish between data taken with various electrode separations. When about 50 centimeters long, eels gener- ate their maximum voltage. We therefore take this length as marking electrical maturity. The decrease in voltage gradient for eels past this maturity has been shown to result from the fact that, as growth continues beyond 50 centimeters, the electroplaxes become thicker, so that there are fewer per centimeter of length (Cox, et al., 1940). A comparison of length and electromotive force per centimeter is shown in Text-figure 4. 191 192 Zoologica: New York Zoological Society [37: 15 Text-fig. 1. Length of electric eel compared with length of the large electric organ. LENGTH OF EEL IN CM 450 1952] 193 Brown & Coates: Length and Voltage in Electric Eeel S o X o o < o 5 * o ^ O 2 o o _J 9 UJ - UJ S110A Text-fig. 2. Length of electric eel compared with maximum voltage of entire organ. 2 CM- 5 CM - 10 CM - DATA PUBLISHED IN 1945* 194 Zoologica: New York Zoological Society [37: 15 <1 o D X □ 1N3I0VU9 39V110A 2 o o - <7» O o in O O o 2 2 Li- O o O o Z ® LlI o 10 © o o Ki o