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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

BREVIORA

MUSEUM OF COMPARATIVE ZOOLOGY

AT

HARVABD COLLEGE, IN CAMBRIDGE

Numbers 67-120 1957-1960

CAMBRIDGE, MASS., U.S.A.

1960

Edited By

Nelda E. Wright

V.^

.\

CONTENTS

BREVIORA

^Museum of Comparative Zoology

Numbers 67-120

1957

No. 67. Notes on Certain Species of Tetragnatha (Araneae, Ar- giopidae) in Central America and Mexico. By Arthur ]\[. Chickering. 4 pp. January 31.

No. 68. The Genus Tetragnatha (Araneae, Argiopidae) in Ja- maica, B.W.I., and other Neighboring Islands. By Arthur M. Chickering. 15 pp. January 31.

No. 69. A New Zodariid Spider from Panama. By Arthur M. Chickering. 7 \)\). January 31.

No. 70. "Anguimorph" Tooth Replacement in Amphishaena alha Linnaeus, 1758, and A. fuliginosa Linnaeus, 1758 (Reptilia : Ampliisbaenidae). By Carl Gans. 12 pp. January 31.

No. 71. Taxonomic Notes on the New World Forms of Troglo- dytes. By Raymond A. Paynter. Jr. 15 pp. March 29.

No. 72. Is the Ant Genus Tetramorium Native in North Amer- ica? By W. L. Brown, Jr. 8 pp. March 29.

No. 73. Additions to the Mammalian Fauna of Peru and Notes on Some Other Peruvian Mammals. By Oliver P. Pearson. 7 pp. March 29.

No. 74. The Discovery of Cerapachyine Ants on New Caledonia, with the Description of New Species of Phyracace.s and Sphinctomyrmex. By E. 0. Wilson. 9 pp. May 1.

No. 75. Oil a New Oetochaetine Earthworm Supposedly from Guatemala. By G. E. Gates. 8 pp. May 1.

No. 7(). Two New Land and Freshwater Mollusks from New Guinea. By William J. Clench. 4 pp. June 18.

No. 77. Dacetinopa, A New Ant Genus from New Guinea. By W. L. Brown, Jr. and E. 0. Wilson. 7 pp. June 21.

No. 78. The Larva of the Ant Genus Dacetinops Brown and Wilson. By George C. Wheeler and Jeanette Wheel- er. 4 pp. Jvme 21.

No. 79. Dasypeltis nicdici lamiiensis, A New Race of Egg-Eating Snake (Ophidia, Reptilia) from Coastal East Africa. Bj'- Carl Gans. 13 pp. August 9.

No. 80. A Collection of Drawings of Fishes Ascribed to J. P. Kirtland (1793-1877), in the Library of Bowdoin College. By James M. Moulton. 4 pp., 2 pis. Sep-^ tember 30.

No. 81. Contributions to a Revision of the Earthworm Family Lumbricidae. I. Allolobophora limicola. By G. E. Gates. 14 pp. September 30.

1958

No. 82. The Trunk ^lusculature of Sanzina and its Beai-ing on Certain Aspects of the ]\Iyological Evolution of Snakes. By Walter Auffenberg. 12 pp. January 31.

Xo. 83. Thamnophis hovaUii Dunn Rediscovered (Reptilia, Ser- })entes). By Benjamin Shreve and Carl Gans. 8 pp. January 29.

No. 84. Rediscovery of the Australian Chelid Genus Pseudemy- dura Siebeurock (Chelidae, Testudines). By Ernest E. Williams. 8 pp., 4 pis. January 30.

No. 85. The Choanal Papillae of the Cheloniidae. By Thomas S. Parsons. 5 pp., 2 pis. January 31.

No. 86. A New Sicistine Rodent from the ^Miocene of Wyoming. By Craig C. Black. 7 pp. May 29.

No. 87. An Enibolonioro J aw from the JMid-Carboniferoiis of Nova Scotia. By Alfred Sherwood Romer. 7 pp., 1 pi. June 20.

No. 88. A New Species of the Genus iirotheca (Serpentes: Colu- bridae) from Venezuela. B}' J. A. Roze. 5 pp. June 30.

No. 89. Remarks on Some Forms of Cinclus (Aves). By James C. Greenway. Jr. and Charles Vaurie. 10 pp. July 15.

No. 90. A Fossil X'ampire Bat from Cuba. By Karl F. Koop- man. 4 pp., 1 pi. July 30.

No. 91. Contribution to a Revision of the Earthworm Family Lumbricidae. II. Indian Species. By G. E. Gates. 16 pp. August 13.

No. 92. A New Genus of Erethizontid Rodents from the Col- huehuapian of Patagonia. By Bryan Patterson. 4 pp. September 17.

No. 93. A New Barylambdid Pantodont from the Late Paleo- cene. Bj^ Bryan Patterson and Elwyn L. Simons. 8 pp. September 18.

No. 94. Affinities of the Patagonian Fossil Mammal Necrolestes. By Bryan Patterson. 14 pp. September 18.

No. 95. A New Bolivian Land Snail of the Genus Dryniaeus. By Juan Jose Parodiz. 3 pp. September 19.

No. 9(j. A New Dichobunid Artiodactyl from the Uinta Eocene. By C. Lewis Gazin. (i pp. September 19.

No. 97. Fusion of Cervical Vertebrae in the Erethizontidae and Dinomyidae. By Clayton E. Ray. 11 pp., 2 pis. October 27.

No. 98. Two New Species of Bathylagus from the Western North Atlantic with Notes on Other Species. By Daniel M. Cohen. 9 pp. December 12.

No. 99. A New Subspecies of Chamadeo jacksoni Boulenger and a Key to the Species of Three-llorned Chamaeleons. By A. Stanley Rand. 8 pp. December 19.

No. TOO. On the Pineal Organ of the Tuna, Thijnnus thynnus L. By Uno Hohngren. 5 pp., 2 pis. December 23.

1959

No. 101. Cervical Ribs in Turtles. By Ernest E. Williams. 12 pp., 1 pi. March 2.

No. 102. A New Jamaican Galliwasp (Sauria, Anguidae). By Garth Underwood. 13 pp. April 9.

No. 108. Two New Species of Eleutheroductyius from Puerto Rico. By Juan A. Rivero. 6 pp., 1 pi. April 10.

No. 104. Studies on Fishes of the Family Ophidiidae. III. A New Species of Lepophidium from Barbados. By C. Richard Robins. 7 pp. April 13.

No. 105. Bufo (jmidlachi, A New Species of Cuban Toad. By Rodolfo Ruibal. 14 pp. April 14.

No. 106. The Occipito-Vertebral Joint in the Burrowing Snakes of the Family Uropeltidae. By Ernest E. Williams. 10 pp. April 28.

No. 107. A Revision of the Dacetine Ant Genus Neostruma. By William L. Brown, Jr. 13 pp. May 6.

No. 108. Some New Species of Dacetine Ants. By William L. Brown, Jr. 11 pp. May 7.

No. 109. On the Pineal Area and Adjacent Structures of the Brain of the Dipnoan Fish, Protopterus annectens (Owen). By Uno Holmgren. 7 pp., 2 pis. May 8.

No. 110. The Spider Genus Coleosoma (Araneae, Theridiidae). By Herbert W. Levi. 8 pp.. 1 pi. June 16.

No. 111. On the Caudal Neurosecretory System of the Teleost Fish, Fundulus heterodiUis L. By Uno Holmgren. 13 pp., 2 pis. June 17.

No. 112. A Mounted Skeleton of the Giant Plesiosaur Kronosau- rus. By Alfred Sherwood Romer and Arnold D. Lewis. 14 i^p.. 1 pi. October 15.

No. 113. A New Phyllomedusa from Bolivia (Salientia, Hyli- dae). By Benjamin Shreve. I^ pp., 1 })1. November 2.

No. 114. Anomalophis bolccnsis (Massalonj^o), A New Genus of Fossil Snake from the Italian Eocene. By Walter Auffenberji'. 16 pp. November 23.

No. 115. The Lemon-Colored Plexaurids from the West Indies and Brazil. By Elisabeth Deiehmann and P. M. Bayer. 12 pp., 5 pis. Noveml)er 25.

1960

No. 116. Insectivores of the Middle Miocene Split Rock Local Fanna, Wyoming-. By Katherine Milmine Reed. 11 pp., 2 pis. January 6.

Xo. 117. Notes on Hispaniolan Herpetology. 1. Anolis christo- phci. New Species, from the Citadel of King Christo- phe, Haiti. By Ernest E. Williams. 7 pp. January 20.

No. 118. A Survey of the Leptodactylid Frogs, Genus Eupso- phus, in Chile. By Jose M. Cei. 13 pp. February 24.

No. 119. Arctic Archibenthal and Abyssal Mollusks from Drift- ing Station Alpha. By Arthur H. Clarke, Jr. 17 pp., 1 pi. March 8.

No. 120. Two Species of Tortoises in Northern South America. By Ernest E. Williams. 13 pp., 3 pis. March 9.

INDEX OF AUTHORS BREVIORA

MUSEUM OP COMPARATIVE ZOOLOGY

Numbers 67-120

1957-60

No.

AUFFENBERG WALTER 82, 114

Bayer, F. M. and Elisabeth Deichmann 115

Black, Craig C 86

Brown, William L., Jk 72, 107, 108

Brown, William L., Jr. and E. 0. Wilson 77

Cei, Jose M 118

Chickering, Arthur M 67, 68, 69

Clarke, Arthur H., Jr 119

Clench, William J 76

Cohen, Daniel M 98

Deichmann, Elisabeth and F. M. Bayer 115

Gans, Carl 70, 79

Gans, Carl and Benjamin Shreve 83

Gates, G. E 75, 81, 91

Gazin, C. Lewis 96

Greenway, James C, Jr. and Charles Vaurie 89

Holmgren, Uno 100, 109, 111

KooPMAN, Karl F 90

Levi, Herbert W 110

Lewis, Arnold D. and Alfred Sherwood Romp:r 112

MouLTON, James M. 80

Parodiz, Juan Jose 95

Parsons, Thomas S 85

Patterson, Bryan 92, 94

Patterson, Bryan and Elwyn L. Simons 93

Paynter, Raymond A.. Jr 71

Pearson, Oliver P 73

Rand, A. Stanley 99

Ray, Clayton E 97

Reed, Katherine Milmine 116

RivERO, Juan A. 103

Robins, C. Richard 104

RoMER, Alfred Sherwood 87

RoMER, Alfred Sherwood and Arnold D. Lewis 112

Roze, J. a 88

RuiBAL, Rodolfo 105

Shreve, Benjamin 113

Shreve, Benjamin and Carl Cans 83

Simons, Elwyn L. and Bryan Patterson 93

Underwood, Garth 102

Vauree, Charles and James C. Green way, Jr 89

Wheeler, George C. and Jeanette Wheeler 78

Wheeler, Jeanette and George C. Wheeler 78

Williams, Ernest E. 84, 101, 106, 117, 120

Wilson, E. 0 74

Wilson, E. 0. and William L. Brown, Jr 77

BREVIORA

Moseiiim of Comparative Zoology

Cambridge, Mass. Januiiry 31, 1957 Number 67

NOTES ON CERTAIN SPECIES OP TETRAGNATHA (ARANEAE, ARGIOPIDAE) IN CENTRAL AMERICA

AND MEXICO

By Arthur M. Chickering

Albion CollpRp, Albion. Michigan

In connection with my study of the ^enns Tefrngnafha Latreille, 1804 in Panama and the West Indies I have also had occasion to examine a number of species from parts of Central America north of Panama, and also from Mexico, types of which are in the Museum of Comparative Zoology at Harvard College. The following notes are offered as a contribution to the further clarification of the genus.

Tetragnatha versicolor Walckenaer, 1841

T. convexa Banks, 1898

T. convexa Petrunkevitch, 1911

T. convexa Eoewer, 1942

A vial labelled T. convexa Banks and marked "cotypes" now contains one male and three females all from San Jose del Cabo, Baja California. The chelicerae, palp, and other characters make it certain that the male belongs to T. versicolor Walckenaer. The females are always more difficult to place with accuracy but I feel certain that these belong in the same species with the male. There is another male in the collection from Sierra Laguna, Baja California, originally identified as a T. convexa Banks l)ut this- is also clearly a T. versicolor Walckenaer.

Tetragnatha guatemalensis 0. P. Cambridge, 1889

T. fraterna Banks, 1898

T. mnndihulata Banks, 1898

BREVIORA

NO. 67

T. fraterna Petrunkevitch, 1911 T. fraterna Eoewer, 1942

The characteristics of the male palp and the male chelicerae definiteh" identify the males collected at San Jose del Cabo, Baja California and described as T. fraterna, and I feel confident that we may be certain of the correct placement of the females as well. Banks identified both sexes collected at Tepic and San Jose del Cabo as specimens of T. mnndihiilata Walckenaer, 1841, known at present only from Ceylon, India, Australia, and Poly- nesia. Re-examination of these shows clearly that they belong to T. guatemalensis 0. P. Cambridge.

Tetragnatha tristani Banks, 1909 (Figures 1-5) There is only a single specimen to represent this species so far as I have been able to determine. This is the holotype very briefly described by its author. I have carefully searched through my collections of Tetragnatha from the regions of Panama closely contiguous to Costa Rica, from which country the original was collected, without discovering any additional specimens. I have also compared the holotype with all other species known to me, with the result that I am compelled to regard it as a valid species.

^ 'y /

e

D kA

i * '

*■«

External Anatomy of Tetragnatha tristani Banks Figures 1, 2. Cheliceral teeth along the fang groove; promarginal nnd

retromarginal teeth, respectively.

Figures 3, 4. Distal ends of cymbiiun, embolus, and eonduetor from two

different views.

Figure .t. Paracymbium.

1957 TETRAGNATHA FROM CENTRAL AMERICA 6

Since the orig:inal description was so brief I have thoup:ht it desirable to furnish a detailed treatment in accord with my usual procedure.

Male holofypc. Total leng-th includinor the chelicerae 4.55 mm. ; exclusive of the chelicerae 4.16 mm. Carapace 1.495 mm. long, .97 mm. wide opposite second coxae where it is widest ; other features as usual in the genus.

Eyes. Eight in two rows as usual in the genus; viewed from above, both rows moderately recurved ; viewed from in front, both rows slightly procurved, both measured by centers. Central ocular quadrangle wider behind than in front in ratio of about 14 : 11, slightly wider behind than long. Ratio of eyes AME : ALE : PME : PLE = 7 : 5.5 : 8 : 6.5. AME separated from one another by 1.5 times their diameter, from ALE by a little more than twice their diameter. PME separated from one another by 1.7 times their diameter, from PLE by slightly more than this distance. Laterals separated from one another by the diameter of PLE; AME separated from PME by nearly 1.5 times as far. Height of clypeus equal to a little more than the diameter of AME.

Chelicerae. Moderately well developed, porrect and divergent; basal segment .78 mm. long and, therefore, about half as long as the carapace ; the prolateral spur is simple, rather poorly devel- oped, and not distally bifid ; the fang is moderately sinuous and distinctly bent posteriorly in distal half; the promargin of the fang groove has seven teeth but the last three are minute den- ticles and would probably be subject to much variation in a large population; the retromargin also has seven teeth with the last two very small (Pigs. 1-2).

Maxillae. Nearly parallel ; somewhat concave along outer border ; about three times as long as wide at narrowest level.

Lip. Wider at base than long in ratio of about 15 : 11 ; less than one-half as long as maxillae ; sternal suture gently pro- curved ; sternal tubercles well developed and robust at ends of sternal suture.

Sternum. As usual in the genus ; with fourth coxae separated by about one-half their width.

Legs. 1243. Width of first patella at "knee" .162 mm., tibial index of first leg 4. Width of fourth patella at "knee" .152 mm., tibial index of fourth leg 7.

BREVIORA NTO . 67

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters) 2.990 MO 3.250 .3.380 .975 11.245

2.

2.080

.552

1.755

1.885

.700

6.972

3.

1.105

.390

.617

.845

.325

3.282

4.

2.210

.390

1.625

1.950

.520

6.695

Palp

.660

.154

.176

.970

1.960

All le^s with moderately coarse spines and the normal eoatinp: of hair.

Palp. Both tibia and patella are short with the former only slightly longer than the latter; the paracymbium is very trans- parent but appears to have the shape shown in Figure 5; the cyrabium is long and slender; the conductor and embolus are also long, slender, and gently curved (Figs. 3-4).

Abdomen. Not prolonged posterior to spinnerets; only slightly swollen in anterior half; slightly notched dorsally at base; 2.73 mm. long; .95 mm. wide about one-third of its length from liase where it is widest.

Color in alcohol. Color apparently well preserved. Legs, mouth parts and cephalothorax with various shades of yellowish to light reddish brown. Abdomen with a well defined folium and many closely placed yellowish white silvery spangles on dorsum and along lateral sides. Venter plain yellowish.

There is only one specimen known at present and that was col- lected by Prof. J. Fid. Tristan of San Jose, Costa Rica, in his home city with no date recorded.

REFERENCES

Banks, Nathan

1898. Arachnida from Baja California and other parts of Mexico. Proc. California Acad. Sei., Ser. 3, Zoology. 1, (7): 205-309, 5 pis.

1909. Arachnida from Costa Rica. I'roc. Acad. Nat. Sci. Philadelphia, April, 1909: 194-234, 2 pis.

BREVIORA

Mmseiuiirii of Compsirative Zoology

Cambridge, Mass. January 31, 1957 Number 68

THE GENUS TETRAGNATHA (ARANEAE,

ARGIOPIDAE) IN JAMAICA, B.W.I., AND

OTHER NEIGHBORING ISLANDS

Arthur M. Chickering

Albion College, Albion, Michigan

For several years before her death in 1953 Miss Elizabeth B. Bryant, Museum of Comparative Zoology at Harvard College, had been engaged in a comprehensive study of a collection of spiders from Jamaica, B. W. I. This collection had come from several sources but it had been assembled largely through the interest of Mr. C. Bernard Lewis, Director and Curator, Sci- ence Museum, Institute of Jamaica, Kingston, Jamaica. After Miss Bryant's death this collection was placed in my possession for continued study. On my way to Panama in June, 1954, I was able to stop in Jamaica for a reconnaissance of the island preparatory to what may be a more or less extensive study of the spiders of that country.

As an extension of my study of the genus Tetragnatha Latreille, 1804 in Panama, I have been much interested in examining the genus in Jamaica and in comparing the species found there with the tetragnathids in several of the larger islands of that general region. This paper is a result of that study, and types of the new species named here are deposited in the Mu- seum of Comparative Zoology.

It is again a pleasure to acknowledge my indebtedness to the following persons for their continued encouragement in the pur- suit of my studies : Dr. A. S. Romer and Dr. P. J. Darlington, Jr., Director, and Curator of Insects, respectively, in the Mu- seum of Comparative Zoology at Harvard College, and Miss Nelda E. Wright, Editor of Publications in the same institu- tion. Without the privileges which have been extended to me

2 BREVIORA NO. 68

for many years in this museum the continued progress of my studies would have been much more difficult.

Genus TetraGNATHA Latreille, 1804

The genus has been well defined by Seeley (1928) and is, in general, well understood by araneologists. There are, however, certain characteristics of the genus which have in the past made it difficult to identify the species correctly and numerous errors must eventually be eliminated. It is also my opinion that suffi- cient attention has not usually been given to the question of varia- tion within species in respect to several of the most important structural features used by taxonomists for identification. F. P. Cambridge (1897-1905) emphasized the value of the character- istics of the male palp such as the form of the paracymbium, shape and course of the conductor and embolus as well as the features of the eyes, chelicerae, and legs. Petrunkevitch (1930) and Wiehle (1939) were the first to appreciate the value of the genital area, which lacks an epigynum, in identifying females which are often exceedingly difficult to place with certainty. Color has been shown to be extremely variable and nearly worth- less as a means of identification. Cheliceral teeth are often quite variable in number, degree of development, and relative posi- tion. Size, when mature, is also subject to great variation in sev- eral species. I have tried to take account of all of these salient features in making my determinations.

THE GENTJS IN JAMAICA

Only the bibliographical references considered essential are given in this paper. Extensive bibliographies may be found in several sources.

Tetragnatha antillana Simon, 1897

T. antillana Petrunkevitch, 1930

T. antillana Bryant, 1940

T. antillana Bryant, 1942

T. festina Bryant, 1945 (male only)

T. haitiensis Bryant, 1945

1956 TETRAGNATHA IN JAMAICA 3

This species appears to be common in Jamaica. It was found abundant at Mavis Bank over water by R. P. Bengry. Collection records: One male from the Blue Mts., southwest side of Main Range, between 3000-4000 ft. elevation, August, 1934 (P. J. Dar- lington, Jr.) ; both sexes from Mavis Bank, over water, March, 1953 (R. P. Bengry) ; one female from Rio Cobre, June, 1954,

Tetragnatha caudata Emebton, 1884

Eucta caudata Petrunkeviteh, 1911 T. caudata Seeley, 1928 T. caudata Bryant, 1940

Miss Bryant had a single female from Cuba. The species ap- pears but once in the collection from Jamaica placed at my dis- posal ; Papine, five miles north of Kingston, April, 1937.

Tetragnatha exigua sp. nov. (Figures 1-5)

Male holotijpe. Total length including chelicerae 2.795 mm., without chelicerae 2.34 mm. Carapace 1.04 mm. long; .67 mm. wide opposite second coxae where it is widest ; with the usual gen- eral form of the genus ; .209 mm. tall at about the middle ; nearly level from PE to posterior declivity; median depression very shallow, opposite interval between second and third coxae.

Eyes. Eight in two rows as usual; lateral ocular tubercles rather prominent ; viewed from above, both rows moderately re- curved ; viewed from in front, anterior row slightly recurved and posterior row slightly procurved, both measured by centers ; cen- tral ocular quadrangle wider behind than in front in ratio of about 4 : 3, wider behind than long in about the same ratio. Ratio of eyes AME : ALE : PME : PLE = 5.5 : 4 : 5 : 4. AME separated from one another by about 1.2 times their diameter, from ALE by about the same distance. PME sep- arated from one another by a little less than twice their diam- eter, from PLE by about two thirds as far. Laterals separated from one another by about 1.25 times their diameter. AME sep- arated from PME by a little more than the diameter of AME, hence further from one another than laterals are from one another in ratio of about 6 : 5. Height of clypeus equal to nearly 1.5 times the diameter of AME.

BREVIORA

NO. 68

Chelicerae. Well developed, moderately porrect, quite diver- gent in distal two thirds, somewhat swollen in middle ; prolateral spur a simple spine not bifid distally; fang- slender, slightly sinuate, with a blunt tubercle on inner margin about one-fifth of its length from base ; promargin of fang groove with four teeth, retromargin with four smaller teeth; with no "large tooth" on the promargin (Fig. 1).

Maxillae. Nearly parallel ; slightly concave in middle of lateral border ; somewhat more than twice as long as lip ; three times as long as wide in middle.

r

n

External Anatomy of Tetragnatha exigua sp. nov.

Fig. 1. Chelicerae of male from in front.

Fig. 2. Paracymbium of male palp.

Fig. 3. Distal end of cymbium, conductor, and embolus.

Fig. 4. Cheliceral teeth of female.

Fig. 5. Genital fold of female.

Liip. Much widened in basal third where it is wider than long in ratio of 22 : 14; sternal suture only slightly procurved; with the usual sternal tubercles well developed at ends of sternal suture.

Sternum. Quite convex ; surface finely pitted and granulated ; with the usual form ; continued laterally and posteriorly between all coxae ; only a little longer than wide ; posterior coxae sepa-

1956 TETRAGNATHA IN JAMAICA 5

rated by a little more than their width.

Legs. 1243. Width of first patella at "knee" .1083 mm., tibial index of first leg 4. Width of fourth patella at "knee" .0758 ram., tibial index of fourth leg 5.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1. 2.275 .390 2.275 1.755 .718 7.413

2. 1.625 .325 1.430 1.380 .580 5.340

3. .910 .198 .445 .550 .308 2.411

4. 1.430 .260 1.170 1.235 .455 4.550 Palp .440 .120 .176 .396 1.132

Spines. True spines appear to be entirely lacking in this species (a very unusual feature) ; hair and bristles are sparsely present. Trichobotliria are present but have not been accurately observed.

Palp. Both tibia and patella are short with tibia longer than patella in ratio of about 3 : 2. The paracymbium is unusually broad. The conductor and embolus are shaped and related essen- tially as shown in Figures 2 and 3.

Abdomen. Slender ; broadest near base and gradually tapered to a blunt point posteriorly; bluntly truncated at base which is not notched; 1.495 mm. long; longer than wide in ratio of about 23 : 9 ; not continued posterior to spinnerets. Other features as usual in the genus.

Color in alcohol. First and second femora yellowish ; all other segments of legs a dusky yellowish. Palps light yellowish except the reddish brown tarsi. Chelicerae : basal segment a deep red- dish brown; fang yellowish. Lip a deep reddish brown, lighter along distal border. Maxillae yellowish in medial third and brown elsewhere. Carapace a deep reddish brown, darker along the margins; median region with a narrow dark stripe posteri- orly and widening at the median depression and extending to PLE ; all eyes except AME surrounded by black pigment. Ster- num : a deep reddish brown. Abdomen : nearly white dorsally with a few silvery spangles ; in the posterior third there are very poorly outlined median gray spots with a series of very narrow black transverse lines; a fairly broad gray stripe extends along

6 BREVIORA NO. 68

each lateral side ; the venter is generally white with a little gray around the genital area and spinnerets.

Female allotype. Total length including nearly vertical cheli- cerae 3.12 mm. Carapace 1.28 mm. long; .715 mm. wide opposite second coxae where it is widest ; otherwise essentially as in male.

Eyes. Central ocular quadrangle wider behind than in front in ratio of 5 : 4, wider behind than long in ratio of 5 : 4. Ratio of eyes AME : ALE : PME : PLE = 6 : 4.5 : 5.5 : 5. AME separated from one another by five-sixths of their diameter, from ALE by 1.5 times their diameter. PME separated from one another by slightly more than 1.6 times their diameter, from PLE by the same distance. Laterals separated from one another by the diameter of PLE. AME separated from PME by the diameter of PLE, hence as far from one another as the laterals are from one another. Height of clypeus equal to about tw^o- thirds of the diameter of AME.

Chelicerae. Moderately well developed ; nearly vertical and parallel ; basal segment .454 mm. long and, therefore, about one- third as long as cephalothorax ; fang slender and evenly curved ; promargin of fang groove with four well-developed teeth fairly evenly spaced ; retromargin with four smaller and fairly evenly spaced teeth (delicacy of the specimen makes it difficult to ob- serve teeth accurately).

Maxillae, Lip, and Sternum. Essentially as in male.

Legs. 1243. Width of first patella at "knee" .119 mm., tibial index of first leg 5. Width of fourth patella at "knee" .097 mm., tibial index of fourth leg 7.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1.

2.210

.378

2.015

2.015

.716

7.334

2.

1.625

.330

1.170

1.430

.585

5.140

3.

.845

.200

.520

.550

.396

2.511

4.

1.495

.265

1.105

1.170

.401

4.436

Spines, hairs, and trichobothria essentially as in male.

Abdomen. 1.95 mm. long; broadest near middle where it is .910 mm. wide ; slightly notched at base ; genital area essentially as shown in Figure 5. Otherwise essentially as in male.

1956 TETRAGNATHA IN JAMAICA 7

Color in alcohol. Abdomen : tlorsally the cardiac area is nearly colorless; there are numerous silvery spangles and a vaguely outlined folium; the venter has a central slightly gray- ish stripe with a stripe on each side outlined by silvery spangles. Otherwise essentially as in male.

Type locality. Ilolotype male, allotype female, and three para- type males from Hanover, Askenish, Trail to Dolphin Head, Jamaica, June 24, 1954.

Tetragnatha PAiiLESCENS F. P. Cambridge, 1903

Eugnatha pallcscens Petrunkevitch, 1911 r. pallescens Petrunkevitch, 1930 T. pallescons Bryant, 1940 T. pallescens Bryant, 1945

Collection records: A male and a female from Ocho Rios, January 1929 (W. S. Brooks) ; several of both sexes from St. Catherine, Port Henderson, Salina, November, 1949 (Bengry, Lewis, Wiles) ; both sexes from St. Thomas, Lysson, June, 1954.

Tetragnatha tenuissima O. P. Cambridge, 1889

T. tenuissima Petrunkevitch, 1930 T. tenuissima Bryant, 1940 T. tenuissima Bryant, 1945

Only one specimen, a male, has appeared in the collection available to me; St. Elizabeth, Magotty, May, 1953 (G. R. Proc- tor).

Tetragnatha visenda sp. nov. (Figures 6-9)

3Iale holotype. Total length including chelicerae 8.58 mm. ; without chelicerae total length 7.475 mm. Carapace 2.60 mm. long; 1.495 mm. wide opposite second coxae where it is widest; with the usual general form of the genus ; .66 mm. tall opposite third coxae just anterior to posterior declivity.

Eyes. Eight in two rows as usual ; lateral ocular tubercles only moderately prominent; viewed from above, posterior row mod- erately recurved, anterior row strongly recurved; viewed from

8

BREVIORA

NO. 68

in front, anterior row moderately recurved, posterior row slightly procurved, all measured by centers; central ocular quadrangle wider behind than in front in ratio of 6 : 5, wider behind than long in ratio of 9 : 8. Katio of eyes AME : ALE : PME : PLE = 11 : 5.5 : 8 : 7.5. AME separated from one another by slightly more than their diameter, from ALE by

r

<^\

External Anatomy of Tetragnatha

Fig. 6. T. visenda sp. nov. ; clieliceral teeth of male from below.

Fig. 7. Idem; the prolateral spur of male.

Fig. 8. Idem; the male paracymbium.

Fig. 9. Idem; distal end of male tarsus.

Figs. 10-11. T. versicolor Walck.; distal ends of conductors and emboli

from Cuba and Michigan, respectively. Figs. 12-13. T. parva Bryant ; distal end of male tarsus and paracymbium,

respectively.

nearly twice their diameter. PME separated from one another by 2.5 times their diameter, from PLE by slightly more than this. Laterals separated from one another by slightly more than the diameter of PLE. AME separated from PME by slightly

1956 TETKAGNATHA IN JAMAICA 9

more tlian the diameter of AME, thus are farther from one another than laterals are from one another in ratio of about 12 : 7.5. Height of clypeus equal to a little more than two-thirds the diameter of AME.

Chelicerae. Well developed; moderately porrect; quite diver- gent in distal two-thirds of basal segment ; somewhat swollen in distal half; prolateral spur well developed and clearly bifid with the larger lobe directed inward (Fig. 6); the fang is long, slender, only slightly sinuate; the fang groove has the so-called "large tooth" with eight others on the promargin and eight on the retromargin (Fig. 6).

Maxillae. Slender; considerably divergent in distal halves; a little more than twice as long as lip ; longer than wide in middle in ratio of 4 : 1.

Lip. Only slightly wider at base than long; sternal suture clearly procurved; with the usual sternal tubercles well de- veloped at ends of sternal suture.

Sternum. Only slightly convex; somewhat swollen opposite second coxae ; with the usual general form ; longer than wide in ratio of 12 : 7 ; continued laterally and posteriorly between all coxae ; posterior coxae separated by about one-fourth their width.

Legs. 1243. Width of first patella at "knee" .395 mm., tibial index of first leg 5. Width of fourth patella at "knee" .260 mm., tibial index of fourth leg 6.

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1.

6.500

1.105

6.890

7.475

1.560

23.530

2.

4.680

.910

4.225

4.420

.975

15.210

3.

2.340

.550

1.397

1.755

.600

6.642

4.

4.940

.715

3.835

4.420

.845

14.755

Palp

1..560

.370

.520

.850

3.300

Spines. All legs with spines of moderate size and length; a sparse coating of hair is also present. Trichobothria are present on femora and probably other segments of legs but their position has not been accurately observed.

Palp. Tibia and patella both short with tibia longer than patella in ratio of about 3 : 2. The paracj^mbium is rather long

10 BREVIORA NO. 68

and slender with the chitinoiis knob about one-third of length of the structure from base (Fig. 8). The conductor terminates in a characteristic manner best shown in Figure 9.

Abdomen. Slender; only slightly concave at base; widest near middle; 5.07 mm. long and about 1.43 mm. wide in broadest region ; not continued posterior to spinnerets ; other features as usual in the genus.

Color in alcohol. All legs light yellowish brown, lighter below ; first and second somewhat dusky dorsally and dorsolaterally with occasional grayish patches ; third and fourth mostly lacking the dusky coloring and grayish patches. Chelicerae reddish brown, grayish along lateral surfaces. Lip dark brown with yellowish distal border. Maxillae yellowish in medial halves, darker along lateral halves. Carapace reddish brown with darker radiating streaks and an irregular granular border. Sternum reddish brown Avith darker irregularly grouped fine dots. Abdo- men : dorsum light yellowish because of presence of numerous irregular subchitinous yellowish white deposits making this re- gion very granular in appearance ; there are also many short irregular grayish lines which become more longitudinal in posi- tion and prominent along the lateral sides; the venter has a median grayish stripe with a broader granular j^ellowish stripe on each side together with a white spot just lateral to each an- terior spinneret and a smaller white spot just dorsal to the larger one.

Type locality. The male holotype was taken at St. Catherine, Port Henderson, June 20, 1954. One male paratype is in the collection from a house in St. Andrew, August, 1955 (G. R. Proctor).

THE GENUS IN CUBA

Tetragnatha antillana Simon, 1897

There is but one specimen, a male, in the collection in the Museum of Comparative Zoology but the species has been re- corded from several localities.

Tetragnatha caudata Emerton, 1884 Two females are in the collection in the Museum of Compara- tive Zoology, both taken at different times in Soledad gardens.

1956 TETRAGNATHA IN JAMAICA 11

Tetragnatha elongata Walckenaer, 1805

This species is well represented by both sexes in the collection and appears to be the most common of all of the eight species recorded from the island.

Tetragnatha guatemalensis 0. P. Cambridge, 1889

T. banhsi McCook, 1893

T. seneca Seeley, 1928

T. banlcsi Levi and Field, 1954

Drs. Gertsch and Levi have apparently agreed that T. seneca Seeley is the same as T. hanksl MeCook. I have made careful comparisons of the specimens from Cuba identified as T. seneca Seeley with my numerous specimens of T. guatemalensis 0. P. Cambridge from Panama and other parts of Central America with the result that I am convinced that here we have another case of synonymy. The characteristics of eyes, several features of the male palps such as vermiform distal end of the paracym- bium and course and shape of both conductor and embolus to- gether with the general characters of the chelicerae all point toward this conclusion. Some may object that I am taking too much liberty with the cheliceral teeth because the "large tooth" is absent in T. seneca Seeley. This "large tooth" is not well developed in T. guatemalc7isis 0. P. Cambridge and could very well have been reduced to the condition found in T. seneca Seeley. The specimens in the Museum of Comparative Zoology identified as T. ha^iksi McCook also agree well with T. seneca Seeley as concluded by Levi and Field (1954).

Tetragnatha orizaba Banks, 1898

There are several specimens of both sexes from several locali- ties in Cuba. Also recorded from Hispaniola but from no other place in the West Indies so far as know^n to the author of this paper.

Tetragnatha pallescens F. P. Cambridge, 1903

Both sexes are represented in the collection from Havana and Soledad.

Tetragnatha tenuissima 0. P. Cambridge, 1889 Both sexes have been recorded from several localities.

12 BREVIORA NO. 68

Tetragnatha versicolor Walckenaer, 1841

T. extensa Emerton, 1884

T. d&ntigera F. P. Cambridge, 1903

T. extensa Seeley, 1928

The specimens from Soledad identified as T. dentigera F. P. Cambridge are, I believe, correctly recognized. These are espe- cially interesting because of the fact that I have been forced to the conclusion that T. dentigera F. P. Cambridge is a synonym for T. versicolor Walck. I have examined a large number of specimens assigned to the latter species and as many as possible of the former. The cheliceral teeth, several features of the male palps (paracymbium, conductor, embolus) and the eyes all point rather decisively toward the synonymy which I have indicated. The tip of the conductor is like nothing else in the genus so far as I have seen. I have provided a drawing of the tip of the con- ductor from a specimen collected in Cuba and another from a specimen of T. versicolor taken in Michigan. There are slight differences but the basic plan is the same and the differences are well within the normal variation of a species (Figs. 10, 11).

THE GENUS IN HISPANIOLA

Tetragnatha antillana Simon,, 1897

T. festina Bryant, 1945 (male only) T. haitiensis Bryant, 1945 (females)

The males of T. festina Bryant have the bifid paracymbium, other features of the male palpi, eyes, and general cheliceral characters associated with T. antillana Simon. 2\ haitiensis Bryant has the general form, cheliceral characters, and genital area characteristic of females of T. antillana. The small differ- ences noted by Miss Bryant and myself are all, I believe, within the normal variation for a species ranging over a Avide area.

Tetragnatha confraterna Banks, 1909

T. eloufjata Bryant, 1945

One female from Puerta Plata, Dominican Republic, was identified as T. elongata Walck., 1805. I have examined this specimen very carefully and I am convinced that it has been

1956 TETRAGNATHA IN" JAMAICA 13

misidentified. Its slightly extended abdomen, cheliceral char- acters, eyes, and f^enital area seem to place it in the species 7'. confraterna Banks where it is provisionally left.

Tetragnatha ORIZABA Banks, 1898

r. Orizaba Bryant, 1945

The specimens of both sexes from the Dominican Republic seem to agree well with our current understanding of this species.

Tetragnatha pallescens F. P. Cambridge, 1903 r. pallescens Bryant, 1945

Numerous specimens of both sexes from Haiti seem to indicate that this may be the most common species in Hispaniola.

Tetragnatha tenuissima 0. P. Cambridge, 1889

T. tenuissima Bryant, 1945

T. festina Bryant, 1945 (females only)

The cheliceral teeth, eyes, general form, lack of leg spines, and the genital area all indicate that T. festina females belong with T. tenuissima 0. P. Cambridge.

THE GENUS IN PORTO RICO

I have had very little opportunity to study the species of Tetragnatha from Porto Rico. Petrunkevitch (1930) listed the following species from this island : T. antillana Simon ; T. elymiquensis Petrunkevitch; T. lahoriosa Hentz; T. pallescens F. P. Cambridge ; T. piscatoria Simon ; T. suhextensa Petrunke- vitch; T. tenuissima 0. P. Cambridge; T. vicina Simon. There appears to be considerable doubt about the occurrence of T. vicina in Porto Rico. In 1947 Miss Bryant described T. parva from the Luquillo Mountains thus bringing the total number of recorded species in this island to nine. Two figures of the tip of the conductor and closely related structures have been prepared to supplement those provided by the author of the species (Figs. 12,13).

14 BREVIORA NO. 68

THE GENUS IN THE VIRGIN ISLANDS

Very little opportunity has been afforded me to study the spiders of these small islands. I have carefully examined all specimens, however, now in the collection of the Museum of Com- parative Zoology and am prepared to summarize my findings as follows: The vial labelled T. antillana Simon contains specimens belonging to this species but it also contains two females which I am tentatively assigning to T. confraterna Banks. The same vial contains a male palp which I believe was derived from this same species. Perhaps we may at least tentatively regard this species as being in the West Indies. The single male assigned to T. piscatoria Simon is, in my judgment, not this species but a specimen belonging to T. visenda sp. nov.

GENERAL SUMMARY

At the present time it seems that either fifteen or sixteen spe- cies of Tetragnatha may be considered to be known from the West Indies under consideration here. T. elijunquensis Petrun- kevitch, T. exigua sp. nov., T. parva Bryant, T. suhextensa Petrunkevitch, and T. visenda sp. nov. are known only from the islands from which they were described. T. antillana Simon has been recorded from all islands mentioned here. T. caudata Emerton has been recorded from Cuba and Jamaica. T. con- fraterna Banks is probably to be regarded as present in both Hispaniola and the Virgin Islands. T. elongata Walck. is only certainly known from Cuba. T. guatenialensis 0. P. Cambridge is now known only from Cuba. T. lahoriosa Hentz is, apparently, present only in Porto Rico. T. orizaha Banks appears to be in Cuba and Hispaniola. T. pallescens F. P. Cambridge has been recorded from all islands considered here except the Virgin Islands. T. piscatoria Simon is known only from Porto Rico among the islands considered here but it was described from St. Vincent. T. tenuissima 0. P. Cambridge is known from all of these islands except the Virgin Islands. T. vicina Simon has been recorded from Porto Rico but its presence there appears doubtful.

1956 TETRAGNATHA IN JAMAICA 15

BIBLIOGEAPHY /

Banks, Nathan

1898. Aiaclmida from Baja California and other parts of Mexico.

Proc. California Acad. Sci., Ser. 3, Zoology, 1. (7): 205-309,

5 pis. 1909. Arachnida from Costa Rica. Proc. Acad. Nat. Sci. Philadelphia,

April, 1909: 194-234.

Bryant, Elizabeth B.

1940. Cuban spiders in the Museum of Comparative Zoology. Bull. Mus. Comp. Zool. at Harvard College, 86 (7): 249-532, 22 pis.

1942. Notes on the spiders of the Virgin Islands. Bull. Mus. Comp. Zool. at Harvard College, 89 (7): 319-363, 3 pis.

1945. The Argiopidae of Hispajiiola. Bull. Mus. Comp. Zool. at Har- vard College, 95 (4) : 359-418, 4 pis.

Cambridge, O. P. and F. P. Cambridge

1889-1905. Arachnida-Araneida. Vols. I-II. In: Biologia Centrali- Amerieana. Dulau & Co., London.

Levi, Herbert W. and Hoavard M. Field

1954. The spiders of Wisconsin. Amer. Midland Natur., 51 (2): 440- 467, 113 figs.

Petrunkevitch, Alexander

1930. The spiders of Porto Eico. Pt. 2. Trans. Connecticut Acad. Arts and Sciences, 30: 159-355, 240 figs.

Seeley, E. M.

1928. Eevision of the spider genus Tetragnatha. Bull. New York State Mus., 278: 99-150.

WiEHLE, H.

1939. Die einheimischen Tetragnatha-Arten. Nova Acta Leopoldina (Halle), N.F. 6 (41) : 363-386.

BREVIORA

Mmseiiinni of Cooipsirative Zoology

Cambridge, Mass. January 31, 1957 Number 69

A NEW ZODARIID SPIDER FROM PANAMA By Arthur M. Chickering

Albion College, Albion, Michigan

Until the discovery of the species described in this paper only one zodariid spider was known from Panama, althouoh a few others had been reported from Guatemala and Mexico. The single species hitherto known from Panama is Store7ia harroana (Chamberlin), 1925.

During the summer of 1954, while sifting decaying hay along the roadside near Corozal, Canal Zone, I discovered one im- mature and two mature males together with one mature and three immature females all belonging to what I regard as a new species referred to the genus Leprolochus Simon, 1892. So far as I know, only two species in this genus have been previously reported. The genotype of this rather remarkable genus, Lepro- lochus spinifrons Simon, Avas from Brazil and L. parahyhae Mello-Leitao, 1917 was from Venezuela. The latter is known only from females. The description of this new species of Leprolochus is given in this brief paper in accord with my usual descriptive formula, and the types have been deposited in the Museum of Comparative Zoology.

Genus LEPROLOCHUS Simon, 1892

Leprolochus decoratus sp. nov. (Figures 1-6)

Male holotype. Total length 2.21 mm., including spines pro- jecting from head region. Carapace 1.235 mm. long, .770 mm. wide opposite second coxae where it is widest; bluntly rounded in front and with seven robust and bluntly pointed black spines

External Anatomy of Leprolochtis decoratua Fig. 1. Anterior cephalic region of male from above. Pig. 2. Chelicerae, eyes, and special spines of male from in front. Figs. 3-4. Tarsus and tibia of male palp; ventral and retrolateral views,

respectively. Fig. 5. Anterior cephalic region of female from above. Fig. 6. Ventral view of epigj'num.

1957 NEW ZODARIID SPIDER FROM PANAMA 3

projecting forward somewhat below AME (Figs. 1-2) ; the para- type male has nine of these spines; with a short and shallow median longitudinal thoracic groove opposite the third coxae ; finely granulated, especially over pars thoracica; gently arched from PME to posterior border with highest part shortly behind these eyes.

Eyes. Eight in two rows; posterior row only slightly longer than anterior row; anterior medians dark, all others light. Viewed from above, posterior row strongly procurved, anterior row moderately so. Viewed from in front, both rows strongly procurved. Ratio of eyes AME : ALE : PME : PLE = 5.5 : 5.25 : 5 : 5. AME separated from one another by about 4/5 of their diameter, from ALE by about 8/5 of their diameter. PME separated from one another by about 9/5 of their diameter, from PLE by nearly twice their diameter. Laterals separated from one another by about 7/10 of the diameter of PLE. Central ocular quadrangle wider behind than in front in ratio of about 9 : 7, only slightly longer than wide behind. Height of clypeus equal to somewhat more than six times the diameter of AME.

Chelicerae. Stout, vertical, parallel ; basal segment .352 mm. long ; with moderately distinct basal boss and fine striations anterior to boss ; the fang is weak, short, and apparently evenly curved ; fang groove obscure and with no teeth observed ; the pro-margin has a fleshy ridge with a row of stiff bristles.

Maxillae. Distinctly convergent; tapered distally and nearly meeting beyond lip.

Lip. Nearly triangular; wider at base than long in ratio of about 3:2; reaches about 2/5 of the length of the maxillae. Ster- nal suture straight.

Sternum. Cordiform ; widest between first and second coxae where it is wider than long in ratio of 25 : 23 ; anterior border nearly twice as wide as base of lip ; very convex ; sharply pointed at posterior end but not extended between fourth coxae which are separated by 4/5 of their greatest width. All coxae moder- ately elongated.

Legs. 4312. Width of first patella at "knee" .108 mm., tibial index of first leg 15. Width of fourth patella at "knee" .130 mm., tibial index of fourth leg 15.

4

BREVIORA

NO. 69

Femora

Patellae Tibiae (All measurements

in

Metatarsi millimeters)

Tarsi

Totals

1.

.660

.264 .462

.660

.528

2.574

2.

.638

.250 .440

.660

.506

2.494

3.

.638

.275 .450

.748

.484

2.595

4.

.811

.275 .660

.990

.690

3.426

Palp

.330

.110 *.443

.374

1.257

* Including much extended apophysis.

Spines. All legs with inconspicuous spines difficult in some instances to distinguish from coarse bristles. First leg: femur dorsal 1-1-1-2, prolateral 1 distal, retrolateral and ventral 1 weak proximal ; patella with numerous coarse bristles but probably only prolateral 0-1-1-0 spines; tibia dorsal 1-1-1 (all weak), prolateral 0-1-0-1, retrolateral 0, ventral only 2 distal; meta- tarsus 0. Second leg : essentially as in first except metatarsus which has a tuft of ventral bristles at distal end together with four spines on ventral and lateral sides. Third leg : femur apparently only dorsal 1-0-1-2, patella dorsal 1-1, prolateral 0 1-1-0, retrolateral 0-1-0, tibia dorsal 1-1-1, prolateral appar- ently 0-1-0-0 and wdth a group of distal bristles forming a comb- like structure, retrolateral 1 distal, ventral only distal 2 ; meta- tarsus only with distal spines essentially as in second. Fourth leg : essentially as in third.

Trichohothria. Observed as follows : tarsi with a row of two or three with the most distal very long ; metatarsi the same ; tibiae apparently with two rows of two each in dorsolateral positions ; palpal tibia with two dorsal in a row with distal one very long.

Claws. Two claws throughout, each pectinate in a single row ; with no claw tufts.

Palp. Patella very short ; tibia with a short body but with a lateral apophysis which extends nearly the full length of the tarsus ; also with a pair of retrolateral spines as shown in Figures 3 and 4.

Abdomen. Regularly ovoid ; longer than wide in ratio of 17 : 13; overlaps cephalothorax only slightly; widest slightly behind

1957 NEW ZODARIID SPIDER FROM PANAMA 5

the middle ; provided with a moderately well developed scutum ; well supplied wdth short appressed and recurved stiff bristles both black and transparent ; postabdomen and anal tubercle project posterior to scutum ; venter well chitinized ; with a short, recurved lip just anterior to genital groove ; tracheal spiracle obscure but apparently just anterior to base of spinnerets and appears to be marked by a row of relatively long stiff bristles; position of probable vestigial colulus obscure. With six pairs of spinnerets partly obscured by a group of bristles.

Color in alcohol. Carapace : cephalic parts a light brown ; re- mainder a medium brown with darker streaks. Chelicerae light brown with other mouth parts and sternum yellowish brown. Legs : all coxae nearl}^ white ; femora brown, lighter beneath ; other segments light brown. Abdomen : dorsally light brown with three irregularly dark reddish brown cross bars in the posterior two thirds and an irregular central stripe reaching to base from first cross bar ; the posterior end of the abdomen behind the scutum and the postabdomen are white; the venter is yellow with varying shades.

Female allotype. Total length 3.90 mm. Carapace 1.625 mm. long; 1.105 mm. wide opposite interval between second and third coxae where it is widest ; gently arched from PME to median, short, longitudinal thoracic groove from w^liich it descends rather gradually to posterior border ; cephalic portion well separated from thoracic part ; with eleven robust black spines in the posi- tion of the seven similar spines in the male (Fig. 5).

Eyes. Ratio of eyes AME : ALE : PME : PLE = 6.5 : 5.3 : 6 : 6. AME separated from one another by about their diam- eter, from ALE by twice their diameter. PME separated from one another by 13/6 of their diameter, from PLE by 17/6 of their diameter. Laterals separated from one another by nearly the diameter of ALE. Central ocular quadrangle wider behind than in front in ratio of 4 : 3, longer than wide behind in ratio of 7 : 6. Height of clypeus equal to about eight times the diam- eter of AME.

Chelicerae, Maxillae, Lip, and Sternum. Essentially as in male.

Legs. 4321. Width of first patella at "knee" .16245 mm., tibial index of first leg 15. Width of fourth patella at "knee" .2166 mm., tibial index of fourth leg 15.

BREVIOBA NO, 69

Femora Patellae Tibiae Metatarsi Tarsi Totals

(All measurements in millimeters)

1.

.975

.390

.682

.715

.650

3.412

2.

.975

.390

.650

.975

.650

3.640

3.

1.040

.455

.780

1.235

.780

4.290

4.

1.170

.455

1.000

1.625

1.040

5.290

Spines. First leg : femur dorsal 1-0-1-2, probably prolateral and retrolateral only distal 1 or 2, ventral Ir (weak) near middle ; patella only prolateral 0-1-1-0 ; tibia dorsal 1-1-1, prolateral 1-1-1, retrolateral 0-1-0-1, ventral O-lr-0-2 ; metatarsus apparently with only two distal but with many spiniform bristles. Second leg : femur probably as in first; patella dorsal 1-1, prolateral 0-1-1-0, retrolateral 0-1-0 ; tibia probably as in first ; metatarsus as in first except with a distal ventral brush of bristles and probably a group of 3 or 4 weak spines. Third leg: femur, patella, and tibia essentially as in second ; metatarsus prolateral 0-1-0, retro- lateral 0-1-0-1, ventral with brush of bristles as in second and with five clear distal spines on ventral and both lateral surfaces. Fourth leg: essentially as in third except brush is absent but the spines are retained. Palp : with numerous spines on femur, patella, tibia, and tarsus ; tarsal claw finely pectinate with about ten slender teeth ; claw is opposed to a chitinous tubercle. Trich- obothria essentially as in male so far as observed.

Abdomen. Well rounded, oviform; 2.405 mm. long, 1.95 mm. wide near middle; without scutum such as that possessed by male; with six spinnerets, the anterior pair quite robust and somewhat the longest ; without definite colulus ; tracheal spiracle close to base of the anterior spinnerets.

Epigynum. Simple but well developed. There is a central shallow depression bordered anteriorly by a strongly chitinized recurved border. Two internal tubules occur at the posterior boundary and on each side there is a relatively large spermatheca (Fig. 6).

Color in alcohol. Carapace essentially as in male except that at the base of pars cephalica there is a dorsal, somewhat oval brown spot on each side from which a thin irregular line extends forward to PLE. The dorsal abdominal region is also very sim- ilar to that of male except that the reddish brown markings are

1957 NEW ZODARIID SPIDER FROM PANAMA 7

broader and more united. Otherwise essentially as in male.

Type locality. The male holotype, female allotype, one mature male paratype together with four immature specimens from Corozal, C. Z., July 10, 1954.

BIBLIOGRAPHY

Banks, Nathan

1929. Spiders from Panama. Bull. Mus. Comp. Zool. at Harvard Col- lege, 69:53-96, 4 pis.

Cambridge, O. P. and F. P. Cambridge

1889- Arachnida-Araneida. In: Biologia Central- Americana. Dulau 1905. & Co., London.

Chamberlin, R. V.

1925. Diagnoses of New American Arachnida. Bull. Mus. Comp. Zool. at Harvard College, 67:211-248.

Chickering, Arthur M.

1947. The Male Allotype and Female Hypotype of Storena Barroana (Chamberlin). Papers Michigan Acad. Sci., Arts, Letters, 31 133-140.

ROEWER, C. Fr.

1942. Katalog der Araneae. Vol. 1. Bremen.

Simon, Eugene

1892- Histoire Naturelle des Araignees. Deuxi^me Edition. 2 Vols. 1903. Librairie Encyclopedique de Roret, Paris.

BREVIORA

Museiiam of Comparative Zoology

Cambridge, Mass. January 31, 1957 Number 70

"ANGUIMORPH" TOOTH REPLACEMENT IN

AMPIIISBAENA ALBA LINNAEUS, 1758, AND

.1. FULIGINOSA LINNAEUS, 1758

(REPTILIA: AMPHISBAENIDAE)

By Carl Gans

In the course of an investigation into the status of the acro- (lont amphisbaenids it proved necessary to prepare the skulls of some related forms. When cleaning the mandible of a female specimen of Amphishoenn alha Linnaeus, 1758, it was noted that six to eight teeth were in the process of being replaced, while two or three others had only recently moved into position, and were as yet but imperfectly fused to the dentaries. The replace- ment teeth appeared to lie interdentally, between, rather than below^, their predecessors.

McDowell and Bogert, in their recent revision of the angui- morph lizards (1954, pp. 102, 104, tig. 30), have stated that "alternate" tooth replacement was restricted to this group, all other lizards possessing "vertical" replacement. The amphis- baenids are generally considered to be Scincomorpha (Camp, 1923, p. 296) and if the distinction between an anguimorph and iiou-anguimorph pattern is as clearcut as McDowell and Bogert suppose, the occurrence of a pattern descriptively alternate would thus be unexpected here. For this reason and because certain other aspects of dental replacement in these forms seem worthy of special attention, it appears desirable to describe the female specimen mentioned above, as well as two specimens of A. fuligmosa Linnaeus, 1758, in which a similar replacement pattern was noted, in the hope of stimulating further research into these matters.

BREVIOKA

NO. 70

The mandibular ilentition of Amphishaena alba

Figures 1 to 5 show lal)ial and lingual views of the mandil)h> of an adult (body length 440 mm.; length of mandibular ramus 15 mm.i) female specimen (MCZ 54299) from "Brazil." This had been cleaned by dissection following controlled applications of full strength commercial bleach to selected portions of the soft parts.

Fig. 1. (Upper) Amphisbaena alba. Lingual view of right mandibular ramus of MCZ 54299.

Fig. 2. (Lower) AiiiiiliLsb(n no alba. Labial view of same niamliliular ramus as Figure \.

1 The length of the mamllbulur riinius has been selected as a convenient indi- cator of the total length of the specimen and Is hence abbreviated as Imr.

1957

AMl'lllSBAENID TOOTH REPLACEMENT

The first pa it of this description covers the general aspect of the tlentitiou and ap])lies equally well to the female cited and to larger si)eeinieus listed below. The description of the replacement l)attern and of individual or possibly ontogenetic variation fol- lows upon this.

The dentition is weakly pleurodont, the height of the dorsal margin above the lingual shelf on which the teeth rest barel}^

Fig. 3. AmpMshaena alba. (MGZ 54299) Sketch of lingual view of right mandibular ramus showing location of the replacement teeth.

equaling the width of the tooth base. There are eight teeth on each side ; the third is largest, the fourth, fifth, sixth, and seventh are slightly smaller, while the second, eighth, and first are pro- gressively smaller in that order. All of the teeth are curved, this being most noticeable near the tip, and the curvature is directed medially and slightly caudad. There is a rotation in the tooth alignment so that the planes of curvature of the individual teeth do not lie parallel to one another. The teeth are of oval cross- section w4th the long axis lying in the plane of curvature.

The base of the fully formed tooth is hollow, the pulp cavity extending two-thirds of the total height of the tooth. About midway up the tooth the diameter of the cavity contracts so as to continue upwards as a thin cylindrical tube.

There is no tooth-bearing shelf as in a typical pleurodont den- tition; instead, a ridge of bone (Fig. 6) rises to the projecting dorsal margin of the dentary between each two adjacent teeth (where replacement teeth are present this ridge is hollowed out to afford lodgement for the pit of the new tooth). Consequently

UREVIORA

NO. 70

tilt' base of eaeh tootli is almost entirely surrouiulcd by bone, to ^vhicll it is ankylosed by a ring of cement. The lowest exposed ])oint lies on the lingual aspect and careful, but thorouph, prepa- ration shows here a single round foramen.' The foramen leads into the ptdp cavity and ])resumably carries its vascular and nerve siipi)ly. A line extending to the level of the tip of the interior cavity is visible on the lingual side. Inspection along the lingual aspect of even the smallest cap-shaped tooth germs re- veals a slight scalloping so that this line may be formed during

Fig. 4. AinpJtLshacna alhn. Lingual view of the left niaiulilnilar ramus of MCZ 54299.

tooth development by the fusion of the anterior and posterior portions. Several of the skulls showed longitudinal cleavage of the teeth along this line, which coincides with the long axis of the oval tooth cross-section.

\'arious stages of tooth replacement are shown in MCZ 54299. In the subsequent description of it and other specimens, L and R will stand for left and right mandibular ramus, and the number following this for the particular tooth or alveolus counting from front to back.

At L-7, R-3 and R-4 the replacement tooth is but a small hollow conical shell, thicker at the top than at the sides, its lower edge somewhat excavated on the lingual side. It lies in a small and deep depression in the deutary in line with the posterior edge

1 These foramina, as shown in Figure 6, are of a larger specimen, which could be cleaned completely without danger of dislodging the firmly fused teeth.

[9i

)*

AMPHISBAENID TOOTH KKPI.ACEMENT

of the tooth it is replaciiij?. As a result, this pit lies on the ascendino; ridoe directly between the tooth to be replaced and the one posterior to it. Even at this stage there is already some erosion into the base of the tooth beinj^- replaced.

L-4 and L-5 show a slightly more advanced stage with pro- gressive erosion into the precursor and an increase in size of the enamel cap of the new tooth.

In R-8 the replacement tooth has reached approximately one- third of its final height, though its tip is still below the level of the projecting dorsal margin of the dentary. It has, however, destroved almost half of the base of the tooth Iving above it.

8 7 6 5 4 3

Fig. 5. Amphishacna alba. (MCZ 54299) Sketch of lingual view of left iiiniidilmlar ramus showing location of the replacement teeth.

At R-6 the tooth has just moved into its final position, its predecessor having been pushed out at some prior stage. It is still only loosely held in place by struts of cement, and appears slightly smaller, and thinner-walled than its neighbors. Its dark appearance is due to the contents of the large, soft-tissue-filled pulp cavity being visible through the translucent walls. This may indicate that the buihl up of the internal dentine layers is not completed until after the tooth is finally cemented into place.

The heavier enamel and more opaque, shell-like appearance of L-3 may indicate a more advanced stage of development. It is, however, still darker than its neighbors, and its base is still far from fused to the dentary. Its slightly cocked position seems to indicate that it is not fully aligned as yet.

All of the replacement teeth except the last two lie freely in the soft tissues and show no fusion to the dentaries.

6

BREVIORA

NO. 70

Another interesting point is that the alveolus formation ap- pears to start in the interdental ridge and only begins to extend into the base of the precursor tooth as growth takes place. No pits or alveoli were found at the base of any tooth that Avas not undergoing replacement.

Though the replacement pattern is interdental it differs from that described by Camp (1923, p. 329, fig. H) for Gerrhonotus s. scincicauda (Skilton), 1849 (and from that seen in a skull of the same form in the MCZ). In this specimen there is no trace of cavity or alveolus formation and the replacenumt tooth appears

Fig. 6. Ampliishaena alha. (MCZ 32257) Dorsal view of tip of mandible, to show interdental bone ridges, basal foramina, and fusion lines.

to lie always in the layer of tissue next to the bone. Gerrhonotun .s. scincicauda is also definitely pleuro<lont and does not possess the interdental ridge that produces what is almost a sub-thecodont condition in Amphishaena alba.

The pattern in aJha is actually closest to that of the snakes ( Bogert, 1943, p. 327 ff.). Here the replacement occurs in pairs, alternate teeth being replaced, while those between them arc functional. This results in the characteristic tooth - alveolus -

1957 AMl'lllSBAENID TOOTH REPLACEMENT 7

tooth - alveolus succession of the prepared dentate bones of the snake skull. Ainphishaena alha differs from this in that at each locus only one replacement tooth appears to be present at a time, and that this erodes its own pocket into the interdental ridge.

Four Aiitphisbaena alba skulls and mandibles from museum collections were available for comparison (MCZ 4031 Brazil, linr = 23.5 mm; MCZ 32256 Surinam, Imr = 20 mm; MCZ 32257 -- Brazil, Imr = 22.5 mm ; AMNH 73233 no data, Imr ^ 17 mm). In only one of these was any soft tissue still present along- the lingual aspect of the dentary, but in this as in the otiiers there was not the slightest indication either of replace- ment teeth or of eroded areas in the interdental ridges which might have lodged replacement teeth. This lack of evidence of tooth replacement may perhaps be related to the fact that all of the skidls were larger (see Imr above), and hence probably be- longed to older specimens. The tip of one of these lower jaws is tigured (Pig. 6) and illustrates not only the basal foramina pre- viously referred to, l)ut also the interdental bone ridges and the strong cementing of llic tooth bases. The complete absence of evidence of replacement activity in these adult specimens, con- trasted Avith the large number of teeth undergoing almost simul- taneous replacement in the above described smaller specimen, may indicate that the ability to replace the teeth is lost in the adults.

It is interesting to observe that there is no evidence for the replacement of maxillary oi- pi-emaxillary teeth in any of the skulls of .1. aJha examined.

Tlie mandibular dentition of Ainphishaena fuliginosa

Ail available dried amphisbaenid skulls were examined for evidence of tooth replacement. Only two medium-sized speci- mens of Amphishaena fuliginosa (MCZ 2154 South America, lmr-8mm ; MCZ 7799 Riobamba, Ecuador, lmr-8 mm ; listed by Zangerl 1944, p. 426 as specimens A and B) demonstrated a discernible tooth replacement pattern.^

Figures 7-10 show lingual views of two of the mandibles of

1 A very clean skull of Rhinriira floridana Baird, (1S5S) (Gainesville. Florida) from the' collection of Walter Auffenberg also reveals that some sort of tooth replacement occurs in this form. Since there are many reasons such as over- cleaning', etc., which might explain the absence of tooth replacement evidence in any given specimen, no useful purpose would be served by listing the names or numbers of specimens examined with negative resuUs.

8

BREVIORA

NO. 70

A. fuliginom. The dentition is again pieurodont, with the lateral shelf slightly higher than the width of the tooth base. The teeth appear to lie in a slight trench formed between the lingual shelf and the ascending face of the dentarv. This trench is crossed by only faint and occasional ridges.

There arc seven teeth or lar<>e alveoli in each of the four

Fig. 7. (Upper) Amphishaena fnliginosa. (MCZ 2154) Lingual view of loft iii,niidit)nl;ir ramus.

Fig. 8. (Lower) Amphisbaena fuliginosa. (MCZ 7799) Lingual view of liglit ninnrliliular ramus.

\'Xu

AMI'IUSBAENID TOOTH REPLACEMENT

iiiaiHliliular i-aiui. Ilowcvcr, tlu' broken tooth spacinji' ot" the soiiu'wliat luacoratod nuuidil)l(' of one of tlic spccinicns (i\ICZ 779!)) (Figures 8. 10) leads one to suspect that Ave are dealinj; with a row of ei<rht teeth. Tlie third and fourth teeth are a<i'ain the hu'ji'est, with numbers five, six and two equal to each other and sliji'htly smaller, while seven and one are smallest. In other aspects the mandibles elosel}' resemble those of A. aiha though the cementing of the teeth is not quite as solid.

Three replacement teeth are visible in each of the mandibular rami of MCZ 2154 and two or three in each of MCZ 77!)!). These

7 6 5 4 3 2 1

Fig. 9. (Upper) Amphisbaena fuliginosa. Sketch of mandibular ramus shown in Figure 7 to show locations of replacement teeth.

Fig. 10. (Lower) Amphisbaena fuliginosa. Sketch of mandibular ramus shown in Figure 8 to show locations of replacement teeth.

10 BREVIORA NO. 70

again lie clearly interclentally and the general pattern of the dentition is very similar to that previously described for A. alha. The largest replacement tooth, the precursor of which is still in jiosition, is but slightly higher than the dorsal margin.

However, there is one item apparent in these jaws that cannot be seen in the specimen of A. alha. where the erosion always takes place on the posterior edge of the tooth undergoing replacement. In A. fuligmosa there are several instances of teeth eroded on the front only or on both front and rear. This is the ease in R-5 and L-7 of MCZ 2154, as well as in R-3 and R-4, and possibl}' L-4 of MCZ 7799. Both in R-5 of MCZ 2154 and in R-3 of MCZ 7799 the developing anterior replacement tooth is considerably larger than the tooth that is eroded. The anterior tooth has thus eroded itself an alveolus wider than that of its precursor. The general spacing indicates that what is involved here is a re- arrangement, respacing or perhaps a change in the tooth number, possibly due to the growth of the mandible.

Both sets of upper jaws show evidence of tooth replacement, with alveoli present above both maxillary and in-emaxillary teeth. As far as can be seen from the skulls, which are somewhat over- cleaned for this purpose, the replacement proceeds alternately as described above. The only complications are due to the differ- ent alignment of the various teeth which make terms like "alter- nate" and "vertical" very difficult to apply here.

Discussion and Summary

iu AnipJiisha< 11(1 alba and A. fuliginosa:

The teeth ai'c plciirodoiit in that they lie against the lingual side of the dentary on a low shelf.

A ridge rises between each two teeth. This character, which gives the dentition a sub-thecodont appearance, is most strongly expressed in older or larger individuals.

The teeth when fully grown have a pulp cavity extending through more than two-thirds of their total height, and are fused to the dentary by a heavy layer of cement around their base. Vascular supply reaches the pulp cavity by a foramen located on the ventral edge of the tooth 's lingual aspect.

Tooth replacement is alternate, with the developing tooth lying in a deep pit close to the posterior edge of its precursor. During

U),")7 A.MI'IIISBAKXII) TOOTH REPLACEMENT 11

jiiowtli it ci'odcs away the posterior aspect of its i)recursor and sometinit's the anterior face of the tooth next in line. The latter featnre may ])rovide for rearran<iement or ehan<'e in nnml)er of teeth. As the mandible lengthens and the individnal teeth in- erease in size, it becomes i)ossible for the rei)lacement teeth to realign themselves (thus extending the lengtli of the tooth row. and maintaining or increasing the interdental gap), rather than being restricted to entering the exact alveolus vacated by their precursor. Shortly after the new tooth extends higher than the dorsal margin it either displaces the tooth above it or the latter breaks away due to the dissolving of its base.

When initially entering the alveolus the new tooth is dark and lias but a thin coating of enamel. Internal dentine deposition api)ears to continue until the tooth is well cemented into place.

In AivpJiishacna alho there is evidence that tooth replacement t'ither becomes rare or stops altogether beyond a certain size. If tiie function of the replacement were related to the provision of larger teeth, this phenomenon might result from a flattening of the growth curve with age.

None of the skulls of A. alba examined showed any evidence of tooth replacement on maxillaries or premaxillaries.

Beth taxouDmieally and in a descriptive sense these results modify tlie simple j^icture presented by McDowell and Bogert. It is hoped that this brief note will stimulate supplementary investigation of this problem in every family and genus of the Squamata. This seems particularly desirable since even a cur- sory inspection of the lizard skeletons in the MCZ shows such divergence in the patterns ^ of tooth replacement that clearly they cannot be described in terms of just tw'o categories angui- morph or non-anguimorph.

1 Some of these patterns have previously been referred to in the literature (e.g. Camp 1923, p. 329, fig. H).

12 BREVIORA NO. 70

1 wish to acknowledge the aid of the following friends, who read the manuscript and commented thereon : Walter Auffen- berg, Charles ]M. Bogert, Tilly Edinger, Gordon Edmund, Rich- ard van Frank, Arthur Loveridge, Samuel B. McDowell Jr., M. Graham Netting, Neil D. Richmond, and Ernest E. Williams.

This paper was completed while working under a National Science Foundation Predoctoral Fellowship.

LITERATUEE CITED

Bogert, C. M.

1943. Dentitiouiil phenomena in cobras and other elapids witli notes on adaiitive modififotions of fangs. Bull. Anipr. Mus. Nat. Hist., vol. 81, ;irt. .S, pp. 285-360.

Camp, C. L.

1923. Classification of the lizards. Bull. Anwv. .Mas. Xat. Hist., vol. 48, art. 11, pp. 289-481.

McDowKLL, S. B., Jk. and C. M. Bogekt

19.")4. The systematic position of Laitthanotu.s and the atliaities of the anguinomorphan lizai'ds. Bull. Aniei-. Mus. Nat. Hist., vol. 10."), art. 1, pp. 1-142.

Z.\NGERL, R.

1944. Contributions to the osteology of the skull of the Anipliisbaeni- dae. Amer. Midland Nat., vol. 31, no. 2, pp. 417-454.

BREVIORA

Miasemei of Comparative Zoology

Cambridge, Mass. March 29, 1957 Number 71

TAXONOMIC NOTES ON THE NEW WORLD FORMS

OF TROGLODYTES

By Raymond A. Paynter, Jr.

While preparing a list of the Troglodytidae, for the continua- tion of the "Check-list of Birds of the World," it became evi- dent that my taxonomic treatment of the family diverged in a number of respects from the classification employed by Hellmayr (1934), the current standard reference to the New World wrens. It seems advisable, therefore, to present, in more detail than possible in the Check-list, the reasons which have instigated these changes. This paper will be concerned with the New World forms of Troglodytes.

I am greatly indebted to Dr. Dean Amadon of the American Museum of Natural History and to Dr. Herbert Friedmann of the United States National Museum for loaning me specimens under their care, and to Dr. Ira N. Gabrielson for very gen- erously allowing me to borrow critical specimens from his private collection.

TROGLODYTES TROGLODYTES

It has been nearlv 40 vears since the New World forms of Troglodytes troglodytes were last revised. At that time, Ober- holser (1919) recognized nine races, three of which he described in the course of his study. Four races have been designated subsequently. After examining nearh^ 450 specimens of the species from North America, I find that I cannot recognize five of the 13 races proposed. A synopsis of the races is presented below.

One of the puzzling features exposed in this review is that the number of male specimens outnumbers that of females by

2 BREVIORA No. 71

almost tAvo to one. An unbalanced sex ratio does not seem to occur in the wild, at least in the European races (Armstrong, 1955). I suspect that collectinf? bias is involved in the museum material, perhaps caused by the greater ease Avith Avhich the vociferous males mav be found. Bias created bv careless sexing of specimens is also a strong- possibility and suggested by the fact that the non-breeding specimens display a greater disparity in the sex ratio than birds taken during the nesting season. Were accurately sexed material available, sexual dimorphism in size might be more pronounced than now evident and some of tlie apparent interracial overlap in size might be diminished.

Troglodytes troglodytes hiemalis Vieillot

TrncilDilijIrft hicnuilis Vieillot, LS19, Xouv. Diet, d'llist. Njit., nouv. ed., 34, p. 514: Xova Scotia and New York; lestrieted to Nova Scotia by Oberholser, 1902, Auk, 19. j.. ITS. Troglodytes troglodylc.s aqi(ilonarif! Burleigh and Peters, 1948, Proc. Biol. Soc. Washington, 61. p. 116 Tompkins, Ne\s-foundland. T. t. hiemalis is the lightest colored of the continental forms. The Newfoundland population was named aquilonaris and was described as differing from hiemalis in being darker and less rufescent dorsally, and paler and more heavily squamated ven- trally characters which are notoriously variable throughout the species. A series of 36 specimens from Newfoundland, in- cluding part of the type series, has been examined. Some birds possess the characters ascribed to the race, but these individuals fall well W'ithin the range of variability of hiemalis and occur with the same frequency. T. t. aquilonaris is considered to be untenable.

The race breeds from the southern part of the District of Mackenzie and southern Newfoundland south to Pennsylvania, Minnesota, and Alberta. It winters in the southeastern half of the United States.

Troglodytes troglodytes pullus (Burleigh)

Xannu-s hiemalis pullus Burleigh, 1935, Proc. Biol. Soc. Washington, 48, p. 61 Mount Mitchell (alt. 6,500 ft.), North Carolina. This race, which breeds in the Appalachians, is barely dis- tinguishable from hiemalis, but a series of 13 breeding specimens from Tennessee, Virginia, and North Carolina shows that

1957

XEW WORLD FORMS OF TROGLODYTES

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4 BREVIORA No. 71

dorsally it is, almost constantly, slightly darker than birds of the northern race collected in the same season. A shorter wing and smaller bill were also attributed to pullus. However, 21 adult males collected between late May and early August in New- foundland, New Brunswick, Quebec, Alberta, Maine, New Hamp- shire, New York, and Michigan had a mean wing length of 49.38±:.28 mm., and 20 of the same series had a mean culmen length of 14.25±:.15 mm., dimensions which do not differ sig- nificantly from those of breeding males from the Api)alachians (Table 1).

The race breeds in the mountains from Virginia to Georgia. It is absent from its breeding grounds during the winter and presumably mingles with hiemalis in the southeastern United States. Differentiation of the two races in the winter is uncertain. The post-nuptial plumage of the species is considerably darker than the breeding plumage and although one might assume that in this dress pullus remains slightly darker than hiemalis, there is no evidence that this is true.

Troglodytes troglodytes pacipicus Baird

Troglodytes hiemalis var. pacificiis Baird, 1864, Rev. Anier. Birds, 1, (18(i-t- 1873), p. 14.5 Simiahmoo, Puget Sound, Washington.

The western continental race is much more richly colored throughout than hiemalis and pullus and somewhat more fre- (piently lacks dorsal barring. Ridgway (1904) stated that the bill of pacificus is more slender and less curved, but I cannot confirm this. The mean wing length of males of pacificus is substantially shorter than that of the males of the eastern races (Table 1), although the overlap in measurements is almost complete, rendering the character of little taxonomic value. Females of pacificus also have shorter wings than their eastern counterparts, but the difference is less pronounced than in the males.

The breeding range is from southeastern Alaska and the southern Yukon southward through the mountains to Idaho and central California. In the winter the race moves to lower elevations and occurs casually south to Arizona and New Mexico.

1957 NEW WORLD FORMS OF TROGLODYTES 5

Troglodytes troglodytes iielleri (Osgood)

Anorthura hiemalis hcUeri Osgood, 1!)01, Aiik. 18. \k ]S] Englisli B;iy,

Kodiak IsLand, Alaska.

Kodiak and Afognak Islands are occupied by a nonmigratory race which is most similar to pad fie us, but distinguished by its less rich coloration, reduced abdominal vermiculations. and slightly larger average size.

Troglodytes troglodytes semidiensis (Brooks)

Xanniis IiicmaJis semidiensis Brooks, 1915, Bull. Mus. Comp. Zool., 59. p. 400 Chowiet Island, Semidi Islands, Alaska.

It is with reluctance that this form is accepted. It appears to be distinguished from kiskcnsis only by the greater average length of the bill of the males (19. 66 ±.19 mm. vs. 18. 57 ±.14 mm. ; 0.05>P>0.02). However, there are only three males in the sample from the Semidi Islands, and their bill measurements (19.0, 20.0 and 20.0 mm.) fall within the upper range of males of kiskensis (17.0 to 20.5 mm.). Under many conditions one might ascribe the observed difference to chance and with little hesitancy consider the two populations unworthy of separate designation. However, it is obvious that the birds are markedly different from helleri of the Kodiak region, Avhich is about 100 miles away, and resemble the Aleutian form, from which they are isolated by roughly 400 miles. The extent of isolation makes it appear probable that the observed differences are indicative of two populations which are morphologically distinct. It is for this reason that it .seems best to accept the data at face value and maintain semidiensis.

Specimens have been taken on Chowiet, Aghiyuk, and North Semidi Islands.

Troglodytes troglodytes kiskensis (Oberholser)

Kannus troglodytes kiskensis Olierholser, 1919, Proc. U.S. Nat. Mus., 55, p. 228 Kiska Harbor, Kiska Island, Aleutian Islands, Alaska.

Xa7inus troglodytes tanagensis Oberholser, 1919, Proc. U.S. Nat, Mus., 55, p. 230 Tanaga Bay, Tanaga Island, Aleutian Islands, Alaska.

Kannus troglodytes petrophilus Oberholser, 1919, Proc. U.S. Nat. Mus., 55. p. 232 Unalaska, Unalaska Lsland, Aleutian Islands, Alaska.

6 BREVIORA No. 71

Nannus troglodytes stevensoni Obeiholser, 1930, Proc. Biol. Soc. Washington,

43, p. 151 Amak Island, Alaska. Troglodytes troglodytes seguamensis Gabrielson and Lincoln, 1951, Proc. Biol. Soe. Washington, 64, p. 73 Seguam Island, Aleutian Islands, Alaska.

The Avreiis of the Aleutian Islands present a number of prob- lems, the most difficult of which is the condition of the available specimens. Almost all collecting has been done during the late spring and the summer, when the birds are in their most worn plumage, or are immature. Of 110 Aleutian specimens examined, only 15 were taken between the months of October and April. The condition of wear is frequently variable ; even birds from the same island, collected on the same day, maj^ exhibit marked dif- ferences. The poor condition of the specimens seems to have been the main reason for the description of six races from the region, although only two stand up under critical examination.

On the basis of five adult males and ten young birds, the population of Amak and Amagat Islands was named "steven- soni." It was defined as differing from "petrophilus," of the Fox Islands, in being more gray in both adult and juvenal plumages, in having fewer vermiculations below in adult plum- age, and in having a slightly shorter bill in the adult. I cannot appreciate any of the supposed color characters. The difference between the mean lengtji of the bills of three males from Amak and Amagat Islands (17.50±.14 mm.) and that of nine males from the Fox Islands (18.16±.14 mm.) is without question insignificant. (P>.10).

In the original description, " petrophiliis" was compared with alascensis, of the Pribilofs, even though the describer (Ober- holser, 1919) admitted that his new race was nearest to "tana- <je)isis," a form from the Andrean Islands which he described in the same paper. The race " petrophilus" was then casually mentioned as differing from " tanacjensis" {op. cit., p. 233) in having a shorter bill, shorter wing, more rufescent dorsum, and more ochraceous ventral surface. The series available to Ober- holser, as well as material collected since that time, has been examined but not even a trend toward the characters described can be noted. Tlie series is, of course, distinct from the Pribilof j)opulation, but a sample from anywhere in the Aleutians would have shown the same thing.

1057 NEW WORLD FORMS OF TROGLODYTES 7

The most recently' named Aleutian subspecies is "segvam- ensis," which was restricted to Seguam, Amutka, and Yunaska Islands, with intermediates occun-ino- on the Islands of the Four Mountains. It was described, from badly-worn breedinp: specimens, as beinp: the palest and grayest of the Aleutian races. Reduced barring on the flanks was another supposed character and it was indirectly suggested that the race has a longer culmen than that of "petrophilus."

The type series of "scguamensis" is rather lighter, on the whole, than the material from the eastern Aleutians. However, the series of "seguamensis'' is extremely Avorn and comparable coloration maj^ be found among equally-worn specimens from anywhere in the Aleutians. The tips of the ventral feathers are abraded, resulting in the apparent reduction of barring. As may be seen from Table I, no significant difference in the length of the bill is evident.

As has been shoAvn, "tanagensis" is indistinguishable from " petrophilus." This leaves only Jtiskensis, to which all of the birds of the Aleutians, east of the Near Islands, are referred. It is a large race, with only a few of its smallest examples over- lapping the largest found on the mainland. It is rather richly colored and reminiscent of helleri, although lighter brown dor- sally. It most nearly resembles meligerus, Avhich is considered below.

The range may be defined as extending from Amak and Amagat Islands, off the western side of the tip of the Alaskan Peninsula, west through the Aleutians to Buldir Island. There are no records from the Alaskan Peninsula or Unimak Island. This may be due to the absence of collectors, since the climate (presumably the limiting factor on the mainland) at the end of the peninsula and on Unimak is probably not markedly different from that slightly farther out on the Aleutians or on Amak and Amagat.

"■fe^

Troglodytes troglodytes meligerus (Oberholser)

Anorthura mcligera OVK-rholser, 1900, Auk, 17. p. 25 Attn Island, Aleutian Islands, Alaska. 1\ t. meligerus differs from kiskensis in its generally longer wing and tail, and in having more and darker vermiculations on

8 BREVIORA No. 71

the flanks. It is a fairly well-defined race.

It has been recorded from the Near Islands (Attn and Agattu Islands),

Troglodytes troglodytes alascensis Baird

Troglodytes alascensis Baird, 1869, Trans. Chicago Acad. Sci., 1. p. 31o, pi. 30, fig. 3 Saint George Island, Pribilof Islands, Alaska.

The race differs from kiskensis and meligerus in being darker dorsally and in having lighter and fewer vermiculations on the flanks. Although the data are scant, it w'ould seem to have a somewhat shorter bill and longer wing and tail than either of these races. With the limited material at hand, the darker dorsal color appears to be the most distinctive character.

It occurs on the Pribilofs, on Saint George, Saint Paul and Otter Islands.

TROGLODYTES AEDON COMPLEX

Many taxonomists (e.g., Chapman and Griscom, 1924; Hell- mayr, 1934; Sutton, 1951; Bond, 1956) have noted the close physical and behavorial resemblances between T. aedon of the United States ^nd Canada, T. hrn.nn ei colli s of the mountains of Mexico, and T. musculns of Central and South America and the Lesser Antilles. The slight differences between the taxa involve such characters as the amount of barring on the flanks, minor variations in wing-tail proportions, and differences in the general shade of the plumage. The groups replace one another geo- graphically, witii only minor discontinuities in their ranges. Be- cause of their allopatric, orderly distribution, they would seem to have arisen as isolates of a common progenitor and later expanded their range to form the present pattern. In spite of their obvious close affinities, there has been reluctance to con- sider them as racial groups of a common species because evidence of interbreeding or of bridging forms, has been lacking. Recent studies (Marshall, 1956) however, have shown that aedon and brunneicollis do interbreed and this evidence, plus previoush' known facts, does much toward forming a convincing argument for considering musculns conspecific with the other groups.

The hrunneicollis group, consisting of montane forms which range from southernmost Arizona through Oaxaca, may be dis-

1957 NEW WORLD FORMS OF TROGLODYTES 9

tinguished from the aedon group, which occurs from southern Canada to northern Baja California, by its more heavily barred flanks and browner coloration. Ridgway (1904) chose to con- sider the more pronounced superciliary stripe an additional character, but in reality the stripe is merely accentuated by the darker color of the surrounding plumage. These characters readily differentiate the birds of southern Mexico from the aedon group, but the northern birds are not so distinct, for there is a south to north cline toward lighter color and reduced barring, or in other words an approach toward aedon, until in Sonora, for example, the brown of the ventral surface is restricted to a buffy chest band and the barring of the flanks becomes no heavier than that which is found in extreme examples within the aedon group. The converging of characters is in itself an indication that the groups are not specifically distinct, but convincing proof has been found by Marshall (1956) in southern Arizona, where both types of birds were discovered interbreeding and specimens were collected from a population which is intermediate between the already only slightly differentiated groups. Marshall also noted that the song of the more brownish form does not differ appreciably from that of the T. a. parkmanii, the house wren of the western United States. Thus, there can be no doubt that hrunneicollis should be merged with aedon.

The problem of the relationship between aedon (now includ- ing 'hrunneicollis) and musculus cannot be resolved so simply and conclusively. T. musculus is distributed from sea level to high altitudes over much of South and Central America and in the Lesser Antilles. A moderately dark race (intermedius) with barred under-tail coverts and faintly marked flanks extends north to Tabasco and to lower elevations in southeastern Oaxaea.^ T. a. hrunneicollis, a richly colored, heavily vermiculated form, is found at high elevations in Oaxaca. No indication of inter- gradation or sympatry between the groups has been noted, but the area is ornithologically poorly known and the evidence must be considered inconclusive.

1 Chapmau and Griscom (1924), on the basis of two males from Manuel and Rfo Pilon, included Tamaulipas within the ranjip. The wing-tail ratios ot these birds, which I have examined, clearly Indicate that they are wintering examples of pnrhmnnii. The confusion of the two forms emphasizes their sim- ilarity.

10 BREVIORA No. 71

Ridgway (1904) distinguished the Middle American forms of T. muscidus from T. aedon by the difference in the relative lengths of their wings and tails, the tail of muscidus being less than five-sixths (83 per cent) the length of the wing and that of aedon being five-sixths, or more, the length of the wing. The ratios have been recalculated, using more material than available to Ridgway, and the distinction between the two groups is still maintained (Table 2). But, when the races of musculus from South America (which undeniably are representatives of the

TABLE 2 Wing-tail Eatios of Males of the Northern Mainland Races of T. aedon

Ratio

82.1%

83.8

72.7

69.8

Middle American forms) are considered, the clear-cut difference between the two "species" breaks down, although the majority of races are separable. For example, the wing-tail ratios of musculus, honaire, and chUensis are roughly 84.5, 87.5 and 89.0 per cent; the remaining races range between 69.0 (albicans) and 82.0 (rex) per cent. When the brumieicollis group is in- cluded with aedon, the overlap becomes more nearlj^ complete, for these are short-tailed forms with cahooni, hrunneicollis, and compositus, for example, having wing-tail ratios of about 75, 78, and 78 per cent respectively. It is evident, therefore, that the wing-tail ratio cannot serve to distinguish the two "species."

The absence of ventral barring is useful in distinguishing about half of the musculus groups from aedon but within the remaining forms there is a range from the faintly barred crissum of musculus to the fairly heavily barred flanks and under-tail coverts of inquietus.

Race

Character

M

Om

Range

X

aedon

wing

51.74 mm.

.29 mm.

49.0—55.0 mm.

25

tail

42.48

.30

39.0—46.5

26

parTcmanii

wing tail

52.26 43.82

.21 .28

49.0—56.0 38.0—47.0

53 48

. . J. wine tail

50.64 36.83

.28 .25

43.0 55.0 31.0—42.0

55 53

inquietua

wing

55.00

.33

53.0—57.5

17

tail

38.41

.41

36.5—40.0

17

1957 NEW WORLD FORMS OF TROGLODYTES 11

Other characters, such as the dorsal and ventral coloration, might be cited, but none is more useful in distinguishing the groups than those already mentioned. Taken collectively these distinguish each taxon from ever}^ other taxon, but no single feature, or assemblage of features, can be found which will fully separate the aedon group from the muscuUis group.

Without a mutually exclusive character, or group of characters, it is difficult to justify considering musculiis and aedon distinct species. When this fact is coupled with what is known of the similarity in behavior of the two groups {vide Chapman and Griscom, 1924, in particular), with the evidence that the groups are allopatric, and with the knowledge that the generally more distinct hmnneicollis group has been found conspecific with aedon, the case for considering musculus and aedon to be conspecific becomes as strong as that which can be made for many polytypic species. Although indisputable evidence of conspecifi- eity is lacking, it seems far better to accept the evidence as it stands than to defer to conservative tradition and the philosophy that no change is better than change based on less than inviolable proof.

Twenty-nine subspecies have been recognized. This is three less than the number recognized by Ilellmayr (1934). An addi- tional form {haldwini) was described after Ilellmayr's synopsis, and this is considered to be a synonym.

T . a. haldwini was described as being darker and grayer than the nominate form, with its breeding range in the central north- ern United States and adjacent parts of southeastern Canada (Oberholser, 1934). Although a large series was examined, in- eluding many of the specimens utilized by the describer, not even a trend toward the ascril)ed characters could be discerned, it appears that the author failed to take into account the post- mortem color changes usually associated with wrens, as well as the heterogeneity^ in color found even in fresh material from a given locality. The race is synonymized with T. a. aedon.

The race intermedins is now considered to range from southern Mexico through Costa Rica, although Nelson (1901) named peninsularis from the tip of the Yucatan Peninsula, and Chap- man and Griscom (1924) named oreopoliis from the highlands of Nicaragua.

12 BREVIORA No. 71

Paynter (1955) accepted pcninsularis, with reluctance, and stated that it could be distinguished from intermedius only by its slightly longer bill. An examination of 54 males from throughout the range of intermedius shows them to have a mean bill length of 16.16±.08 mm., with a range from 15.0 to 18.0 mm. Only one specimen, a bird from Costa Rica which approaches inquietus, has a bill as long as 18.00 mm.; the remainder fall between 15.0 and 17.0 mm. Six males of 'peninsularis from the type locality at Progreso and from nearby Santa Clara, Yucatan, have a mean culmen length of 16.83±.02 mm. (not ±.20 mm. as published by Paynter, 1955), and range from 16.5 to 17.0 mm. While the difference between the two samples is "statistically significant," the overlap is much too great to warrant recognition of two races.

Hellmayr (1934) doubted the validity of oreopolus, described by Chapman and Griscom (1924) as being a montane race slightly darker than intermedius. I cannot recognize the form and consider it to be synonymous with intermedius.

T. a. clarus, of the Guianas, Trinidad, northern Brazil, and adjacent parts of Venezuela, Colombia, and Peru, was considered by Chapman and Griscom (1924) to be indistinguishable from albicans of southwestern Colombia and western Ecuador. Never- theless, Hellmayr (1934) chose to recognize the two forms, even though he admitted that individual variation frequently bridged the slight gap he believed to exist. Approximately 50 specimens from the range of each race have been examined and it is con- eluded with Chapman and Griscom (1924) that clarus cannot be separated from albicans.

TROGLODYTES SOLSTITIALIS COMPLEX

At high altitudes from southern Mexico through Central and South America, there is a series of isolated populations of short- tailed, short-billed wrens of rich, brown coloration, with broad, tawny superciliaries, taAvny auriculars, and pronounced, dark post-ocular stripes. These have been treated as three species. The first is T. rufociliatus, a form with four isolated races ranging from Chiapas, Mexico, through El Sah^aclor, which is character- ized by heavy barring on the abdomen and under-tail coverts, concealed (or obsolete) small, white spots on upper-tail coverts,

1957 NEW WORLD FORMS OF TROGLODYTES 13

and a few white spots on the wing coverts. The second species is T. solsfitialis (a group of seven or eight subspecies, most of which are isolated from one another) which is distributed from Costa Rica through the Andes to Venezuela and Argentina. Tt may be distinguished from T. rufociliatns l)y its less saturated color, lack (or nearly so) of abdominal vermiculation, and absence of spots on both the upper tail and wing coverts. The third species is T. monticola a monotj''pic form isolated near the top of the Sierra Nevada de Santa Marta, Colombia, which is much larger than the other two forms. It resembles T. solstitialis in that it is not deeply colored and has no spotting on the rump or wings, and is similar to T. rufociliatus in that it is heavily barred below.

Hellmayr (1934) saw no specimeiLs of rufociliatus, but thought that it was intermediate betAveen hrimneicollis and solstitialis, and that eventually it might seem advisable to unite the three forms under one species. T. rufociliatus and solstitialis are morphologically quite distinct from T. a. irunneicollis, with only their rich coloration a point in common. In El Salvador T. a. intermedius and rufociliatus occur sympatrically (Dickey and van Rossem, 1938) and, as it has been shown, intermedius is conspecific with hrunneicollis. Therefore, it is impossible to con- sider rufociliatus to be a member of the aedon-hrunneicolUs- musculiis complex, even if one ignores their morphological dis- tinctiveness. Although hrunneicollis is not the link between rufociliatus and solstitialis, the two groups are very similar, with only the presence or absence of abdominal barring a conspicuous difference. Even this difference is bridged, somewhat, by the nominate form of solstitialis, since mature specimens occasionally are distinctly, and quite extensively, barred on the flanks. The spotting on the wing and upper-tail coverts of rufociliatus is not a good distinguishing character either, because within a given population there are some individuals lacking these markings.

The two groups would seem to have arisen from a progenial population which at some time became divided into two isolated populations and thereupon became somewhat differentiated. Further division and isolation within the groups then gave rise to the various subspecies now recognized. It may be postulated that the primary division occurred during the Cenozoic when

14 BREVIORA No. 71

Middle America was divided by various seaways. One population (solstitialis) maj^ have been isolated in South America and southern Central America while the other population (rufocilia- tus) was isolated in northern Central America and southern Mexico. Even if the groups did not have their origin precisely in this manner, or at this period, the almost certain fact remains that they arose from a common ancestral stock which became split into two geographically isolated units.

Since the two groups are so slightly differentiated, there is little doubt that their relationship is best expressed by treating them as conspecific forms, uniting them under the older name of solstitialis.

T. monticola, the large, heavily barred form of Santa Marta, was believed by Hellmayr (1934) to be reminiscent of hrun- neicollis. He thought it a specialized offshoot of solstitialis but a possible link between solstitialis and hnmneicollis. The resem- blance between these two forms is superficial. T. monticola has the short tail, short bill, and facial pattern of solstitialis. Its large size and barred underparts are the only features which are similar to drunneicollis, and even the barred underparts are shared by some races of solstitialis. While I do not agree that monticola is a link between l>runneicollis and solstitialis, I do think that Hellmaja- (oj). cit.) was correct in believing it to be an offshoot of solstitialis. The only character which distinguishes it from all of the forms of solstitialis is its large size. Being con- fined to a small area on the top of a mountain, it is analogous to an insular population, in^ which situation a race often is larger than its congeners. It would seem best to consider monticola as merely another race of solstitialis.

LITERATURE CITED

Armstrong, Edward A.

1955. The Wren. Collins, London, viii + 312 pp.

Bond, James

1956. Check-List of Birds of the West Indies. Acad. Nat. Sci. Phila- delphia, vi + 214 pp.

Chapman, Frank M. and Ludlow Griscom

1924. The House Wrens of the Genus Troglodytes. Bull. Amer. Mus. Nat. Hist., 50: 279-304.

1957 NEW WORLD FORMS OF TROGLODYTES 15

DiCKFA-, Donald K. and A. J. van Rossem

1938. The Birds of El Salvador. Field Mus. Nat. Hist., Zool. Ser., 23. 609 pp.

Hkllmavk, Charles E.

1934. Catalogue of Birds of the Americas and the Adjacent Islands. Field Mus. Nat. Hist., Zool. Ser., 13. pt. 7, vi + .531 pp.

Marshall, Joe T.

19.")6. Summer Birds of the Rincon Mountains, Saguaro National Monument, Arizona. Condor, 58: 81-97.

Nelson, E. W.

1901. Descriptions of a New Genus and Eleven New Species and Subspecies of Birds from Mexico. Proc. Biol. Soc. Washington, 14: 169-175.

Oberholser, Harry C.

1919. Notes on the Wrens of the Genus Nannus Billberg. Proc. U.S.

Nat. Mus., 55: 223-236. 193-1. A Revision of North American House Wrens. Ohio Jour. Sci.,

34: 86-96.

Paynter, Raymond A., Jr.

1955. The Ornithogeography of the Yucatan Peninsula. Bull. Peabody Mus. Nat. Hist., Yale Univer., 9. 347 pp.

RiDGWAY, Robert

1904. The Birds of North and Middle America. Bull. U.S. Nat. Mus., 50. pt. 3, XX 4- 801 pp.

Sutton, George Miksch

1951. Mexican Birds. First Impressions. U. of Oklahoma Press, Nor- man, XV -|- 282 pp.

^^Ui)

BREVIORA

Meseeei of Cooipsirative Zoology

Cambridge, Mass. March 29, 1957 Number 72

IS THE ANT GENUS TETRAMORIUM NATIVE IN NORTH AMERICA?

By W. L. Brown, Jr.

For some years, a controversy has continued concerning the distribution in North America of the ant genus Tetramorium Mayr. On opposite sides in this argument ha.ve been Dr. M. R. Smith (1943), who believes that all five of the Tetramorium species reported from North America have been introduced by man from overseas, and Dr. W. S. Creighton (1950), who thinks that the evidence points to prehistoric endemicity in the continent for at least two of the species mentioned ; T. cacspitnm (Linnaeus) and T. riigiventris M. R. Smith. Apparently there is no serious disagreement about the origin abroad of 2\ guineense (Fabri- cius), T. simillimum (F. Smith) and T. pacificum Mayr. The first two of these are very likely African in origin, since they occur in wild parts of Africa and have their closest relatives among the species of that continent. T. pacificum is apparently from the Indo-Australian area, although its exact source has never been seriously tracked down. Certainly, it is Old World in origin.

The two species caespitiim and rugiventris are therefore the central elements in the discussion, and it seems appropriate at this time to review the important facts in connection with these two forms and to add whatever significant observations are now available.

Tetramorium caespitum

This species is widespread in the Palaearctic region and in Africa, where it is extremely variable, as attested by an almost endless list of infraspecific variants. In terms of modern tax-

2 BREVIORA NO. 72

onomic practice, many of these entities would be considered good sibling species, as indeed some of them already have been treated by various European authors. Many others are mere synonyms of caespitum individual or nest variants that do not represent natural, self-maintaining populations. No one has challenged the placement of the North American populations with the more nearly "typical" caespitum variants.

Creighton's assertion that caespitum is a native Nearctic ant rests on two principal pieces of evidence : first, the species was known at a very early date, i.e., 1895, from states as remote as Tennessee and Nebraska ; second, there is present in North America, at least in the eastern states, a workerless parasite of caespitum, the aberrant species Anergates atraiulus (Schenek), the transport and establishment of Avhich would seem to present special difficulties.

The difficulty with the first piece of evidence is that the exact situation of the collections made in Tennessee, Nebraska and elsewhere in the "interior" of North America was never specified. All of these collections may have been, and probably were, made in or near "culture areas," that is, regions strongly disturbed by the presence of man. The experience of several practiced myrmecological field workers, including that of Dr. Creighton (personal communication) and myself, indicates that so far as known, caespitum in North America is known only from rather heavily disturbed localities, such as cities, towns, road- sides, farmyards, picnic areas and the like. I have been able to gather no records at all to show the existence of the species in places remote from the works of man in North America.

This situation contrasts with that holding in the Old World, at least so far as my own personal experience goes, and judging also from what I have been able to glean from various publica- tions dealing with the species, and from personal communications with European myrmecologists. To sum up this information, it can be stated that T. caespitum in Europe and (although given various infraspecific names) in China is often abundant in and around human habitations, just as in North America. However, it is also to be found, often in abundance, in localities that show little or no trace of human disturbance, and that are far from the nearest humanly-occupied places. The contrast is very marked if one collects, as I have, at intermediate altitudes in

1957 TETRAMORIUM IN NORTH AMERICA 3

West China, in pine-oaJc forest, and then compares the collec- tions of Tetramorium obtained with the results of a deliberate search for Tetramorimn in similar vegetational zones in Pennsvl- vania, New Jersey or Massachusetts. In West China, T. caespi- ium tends to occur uniformly throughout the pine-oak forest, regardless of roads, villages, etc., which are very sparsely dis- tributed in the areas I am recalling. In ecologically equivalent areas in the eastern United States, I have found the same species established only on or near the sites of more or less actively maintained human works. Clearly, the density of nests and individuals seen in North America is in large part proportional to the degree of urbanization of the area occupied, although the real extremes of urbanization, where almost all space is covered by concrete or asphalt, are certainly not favorable locales for colonization by this or any other ant species that lives largely in the open. In my opinion, the local, detailed distribution of T. caespitum in North America is that expected of an historically introduced, not a native ant.

The second point of evidence, that concerning the presence of tlie workerless, and therefore obligatory parasite, Anergates atra- iulus, certainly seems on the face of it a real sign of long occupancy of North America by both host and parasite. Creigh- ton emphasizes the difficulties facing trial colonists of the parasite species : first, the species seems to be relatively rare in Europe ; second, it is unlikely that the parasite female could survive a long trip; third, a female arriving in North America would be hard put to find a suitable nest of the host species to enter. To take these difficulties one at a time, we should first recognize that, while Aner gates is not the commonest of ants in collections, it is nevertheless likely to be locally very common in restricted localities. Even in Europe, host populations are normally concentrated in gardens and waste places within towns and cities, including seaports, and these are accordingly \evy likely places for Anergates to occur unnoticed by primarily country-searching myrmecologists. In the United States, Aner- gates is known chiefly from East Coast localities in urban areas near the sea exactly the kind of place from w'hicli colonists might be expected to be exported most easily. It should be added that Anergates females may be produced in very large numbers

4 BREVIORA NO. 72

from a single nest, so that a given locality may be heavily saturated with them during the period of nuptial flights.

The difficulty of transport of live Aner gates propagules is real, but far from insurmountable. Females can be carried either as individuals carrying the necessary sperm, or as estab- lished inquilines in a Tetramorium colony. There is no reason why such a voyage might not be successfully made by a parasite queen, especially when one considers the evidence of Lindroth (ms., personal communication) for transport of faunal frag- ments to North America in ballast originating in Europe. Fur- thermore, there is no reason to believe that Tetramorium nests, with or without Anergates, cannot flourish on shipboard for at least the normal span of these species as colonies, a span which seems to be sufficient even for a long voyage under sail.

The third objection is the least difficult one, for there ap- parently has been no shortage of suitable host nests at close proximity to the waterfront in at least some of the major western Atlantic ports, perhaps as far back as colonial times. If a colony of the host parasitized by Anergates arrived in ballast or other- wise stowed-away, it had only to release its flight of fertile female imagines on the new shore to create a high probability of successful establishment.

A similar series of events may have led to the establishment of the workerless parasite Xenometra moriilicornis Emery in the West Indies, together with or following the establishment of its host, Cardiocondyla cmeryi Forel. A Xenometra of the same or a very closely allied species lives with C. clegans Emery in Italy ; Menozzi (1919) thought this was the male of elegans, but specimens from his collection indicate instead its affinity with X. monilicontis. Cardiocondyla is a primarily littoral and riparian genus from the warmer parts of the Old World ; records of several species from the New World seem to indicate rather clearly that it has been introduced by man on many separate occasions (M. R. Smith, 1944).

To conclude the discussion of the bearing of parasites on the distribution of Tetramorium, 1 think we may safely consider that introduction of an obligatory parasite, while less probable than the establishment of the host, is nonetheless entirely possible if the opportunities exist for a long enough time, and if a dense host population is available to the immigrant parasite.

1957 TETKAMOHir.AI IX NORTH AMERICA 5

TeTKAMORIUM RUGIYENTRIS

The type series and onl.y recorded sample oC T. rugiventris was obtained from an upland ponderosa jiine stand about ten miles south of Prescott, Arizona, and about one mile off the liijihAvay. In empliasiziny his difficulty in accepting Smith's hypothesis, namely, that the ant was introduced with camel food or stores at the time Avhen camels were imported from North Africa during the last century, Creighton wrote : ' ' Ento- mologists frequently strain at gnats but it is seldom that they are asked to swallow a camel."

I have checked with care a syntype of T. rugiventris kindly sent by Dr. Smith. As a result of this examination, I can agree with Dr. Creighton that the ant in cpiestion is almost unques- tionably endemic to the locality where it Avas found. However, I cannot agree with either Smith or Creighton that the species rKgivcufris belongs to the genus Tctrcnnorium as it is now con- stituted. Instead, the type I have examined seems to me to be a cleareut, if somewhat aberrant, member of the genus Myrmica, closely allied to M. striolagaster Cole. M. striolagaster is re- corded from several localities in Arizona and New Mexico, and I have specimens collected by E. 0. Wilson at or near the type localit}- of rugiventris, in the vicinity of Prescott. Although the two species are separated hy the extent and strength of the gastric sculpturing and by other characters as well, it seems clear that they are congeneric, and also that they run rather close to the M. pnnctiventris group of Myrmica.

The rugiventris type actually possesses minute barbulation on the posterior tibial spurs, as can be seen at magnifications of 90 X and better, so that in this character, the species would key to Myrmica, rather than Tetramorium, in the standard keys to tribes and genera of Formicidae. Actually, however, this spur barbulation is not worth much as a tribal character within the IMyrmicinae, despite the faith that key-makers have placed in it. The false distinction between Hylomyrma Forel (tribe Myrmicini) and Lundella (tribe Tetramoriini) appears to have been based partly on this character (Brown, 1953), and the same may hold true of the supposed difference between Crato- myrmex Santschi, purportedly a member of tribe Myrmicini, and Messor Forel, of tribe Pheidolini.

6 BREVIOBA NO. 72

Dr. Smith is fortified in his opinion that rugivcntris belongs to Tetramorium by the presence in his species of raised carini- form lateral wings of the clj^peus that border the antennal fossae in front, as in Tetramorium; but this character is shown with varying degrees of clearness in other, undoubted Myrmka species, both in North America and in southern Asia, so that it cannot be used as a point of separation between the two genera. The question comes to mind, of course, as to whether Myrmica and Tetramorium. really are separate genera after all, and this is precisely the kind of question that most needs asking in ant taxonomy these days. In deciding this particular question, fur- ther study must l)e made of the males, since tetramoriine males (with the exception of a couple of African forms that require closer study) have several funicular segments fused in such a way as to reduce the numl)er of antennal segments to ten in this sex.

Unfortunately, the males of rugiventris remain unknown at present, so that it is not known whether they meet the strong criteria of this caste. From the habitus and lesser details of the worker, however, I consider the relationship with Myrmica is close enough to call for a new combination: Myrmica rugiventris. Whether or not this combination finally i)roves to be the valid one, it at least helps to establish strong doubts as to the pre- Columbian existence of Tetramorium in North America.

In fact, were it not for Tetramorium lucayanum Forel and the Xiphomyrmex spinosus complex, the New World could h( considered free of endemic members of tribe Tetramoriini. With the synonymy of Lundella under Hylomyrma (Brown, 1953), the New World lost its one endemic tetramoriine genus. T. lucayanum presents no special difficulty, because its distribu- tion (Bahamas, Puerto Rico, etc.) is highly suggestive of intro- duced status. In its morphological characteristics, lucayannm seems closest to an African group of species, but the species itself has not l^een identified with any particular continental African population. However, our knowledge of African Tet- ramorium is in a very imperfect state, and it seems to me likely that lucayanum must have come from the Dark Continent, even though it may be rare there. A parallel case involving Strumi- genys rogeri Emery has turned out to follow exactly this pattern (BroAvn, 1954).

1957 TETRAMOKIUM IN NORTH AMERICA 7

The Xiphomyrmex sinnosus complex (which may represent a single variable species) is the one example that cannot be ex- l^lained away, and it is all the more remarkable, considering its isolated position in southwestern United States and through much of Mexico, far away from the remainder of the generic range, which is entirel}' Old World tropical and warm-temperate. Pre- limina.ry examination reveals no reason to consider this complex as other than bona fide Xiphomyrmex, and its distribution is almost certainly that of a long-established endemic group of populations, surely pre-Columbian in North America. Xipho- inyrmex is separated from Tetramorium by a very minor charac- ter, 11 antennal segments in place of the 12 of Tetramorium. Future revisers could well fail to be impressed by the soundness of the generic split based on this difference, so that we may eventually see a systematic technicality bring back Tetramorium as a native American genus.

SUMMARY

Of the five species of Tetramorium so far reported as occurring in North America, only two are under dispute as possibly having existed on this continent prior to the advent of European colon- ists ; these are the species heretofore known as T. caespitum and T. rugiventris. Evidence is presented to show that T. caespitum almost certainly was introduced by man from Europe, this evi- dence consisting primarily of the demonstration that T. caespi- tum in North America, unlike the Eurasian populations, is distributed exclusively in the manner of a man-accompanying "tramp" species. The species rugiventris, on the other hand, is removed from Tetramorium, where it does not fit well, and is transferred to Myrmica. The number of Tetramorium species occurring in North America is thus reduced to four, all of them likel}' introductions from the Old "World w^ithin historical times. The only member of tribe Tetramoriini that can safely be con- sidered as endemic to the New World at the present time is the Xiphomyrmex sj^inosus complex, widespread in southwestern U. S. and Mexico.

8 15IJEVIOKA NO. 72

KEFEKENCES CITED

r>iio\v.\, \V. L., Jk.

19.").'}. C'haraetois aiul aynoiiymics among the genera of ants. Part II. Brcvioia, Mus. Comp. Zool., 18: 1-8.

1!)'>4. The ant genus Strumigcnys Fred. Smith in the Ethiopian and Malagasy regions. Bull. Mus. Comp. Zool., 112: 1-34, cf. pp. 4-7.

CUEIGIITON, W. S.

1950. The ants of North America. Bull. Mus. Comp. Zool., 104: cf, pp. 241-245, 286-294.

MiNCZZi (Mexozzi), C.

1919 (1918). Primo contribute alhi couoscenza della fauna mirmecologica

del Modenese. Atti Soc. Nat. Mat. Modena, (5) 4: 81-88, cf.

p]). 83-84.

S.\iiTii, ^r. E.

1943. Ants of the genus Tetramorium in the United States with the description of a new si)eeies. Proc. Ent. Soc. Washington, 45: 1-5.

1944. Ants of the genus Cardiocondyla Emery in the United States. Proc. Ent. Soc. Washington, 46: 30-41.

BREVIORA

Mmseiuiini of Comparsitive Zoology

Cambridge, Mass. Mak( h 29, 1957 Number 78

ADDITIONS TO THE MAMMALIAN FAUNA OF PERU AND NOTES ON SOME OTHER PERUVIAN

MAMMALS

By Oliver P. Pearson

Museum of Vortoljiate Zoology, Berkeley, California

All earlier report discussed the mammals of the highlands, or altiplaiio, of southern Peru (Pearson, 1951). Subsequent col- lecting has revealed important additions to the fauna of this area as well as considerable extensions of the range of certain species. 1 am indebted to Dr. Carl P). Koford, who collected several of the specimens mentioned, for permission to report upon them and to draw upon information in his field notes. All specimens are in the Museum of Vertebrate Zoology, Berkeley.

Hesperomys sorella (Thomas). This species was not listed in the previous report on mammals of the altiplano. Eleven speci- mens have now been taken in the Department of Puno in bunch- grass terrain at 3 mi. NE Arapa, 12,600 ft., 5 mi. S Asillo. 13,000 ft. and at Hacienda Calacala, 13,000 ft., which is 7 mi. SW Putina. They are longer-tailed, longer-eared, and tawnier than H. lepidus ducilla, which also lives in this region, and have shorter, more slanting zygomatic plates. The range of measure- ments (in mm.) is: total length, 128-147; tail, 60-71; hind foot, 17-19 ; ear from notch, 18-19 ; and greatest length of skull, 21.5- 23.2. They do not match the type of sorella perfectly but seem to be more closely related to that form than to coJlosus, carillus, or frida.

Since publishing the earlier report in which I used the name Hesperomys ducilla for the short-tailed species in southern Peru, I have examined the types of H. lepidus (Thomas) and H. ducilla (Thomas) and find that despite great difference in age of the in-

2 BREVIORA NO. 73

(li\i(liials iliey are (luite .siiiiilai- and can easily l)e included in the same species, characterized by short tails and tall zygomatic plates. Saiiliorn (1950^ also considered (liirtlJa to lie a subspecies of Jepidus.

Klifjmodoiil l(( iJiKriilus piicruhis (Philippi). Four specimens of this desert mouse have l)een reported from Peru (Pearson, 1951), all from Santa Rosa de -luli, Department of Puno. and were listed as E. p. hirtipcs, the type locality of ^vhich is Lake Poopo in Bolivia. Our recent collections included 11 specimens taken in Peru between 13,000 and 15,300 ft. in the Departments of Moquegua, Puno, and Tacna. These specimens have now been compared Avith topotypes, in the Chicago Natural History Museum, of E. p. puendus from northern Chile and are iudis- tinguishal)le. E. p. iarnpacensis Mann has also been described from northern Chile, but the new material reveals that the shape of the zygomatic ])late and the bicolored nature of the tail, features on which farapacensis was based primarily, are un- reliable. The Peruvian Elifpiindoiil ia therefore should l)e E. p. pKcridus.

FhyUotis [Galoionnjs) garlcppi Thomas. Two specimens of this rare mouse were taken in southern Peru, one at Pichupichuni. 12,600 ft., 5 mi. NW Iluacullani, and another at Pampa de Anco- marca, 13,700 ft., 76 mi. S Have, both in the Department of Puno. These are the first records of this species from Peru. Since no field measurements have been i)ublished and the type specimen has a smashed skull, we record measurements (in mm.) of our two specimens and photographs of one of the skulls (Fig. 1).

Total length

Tail

Hind foot

Ear

Skull greatest length

Zygomatic breadth

Width of braincase

jMaxillary tootlnow

Compared with the type and another Bolivian specimen in the British Museum of Natural History, and one in the Chicago Natural History Museum, the Peruvian specimens are smaller,

MVZ 115903

MVZ 11590

129

132

32

30

23

24

19

19

26.7

27.0

lo.O

15.0

12.8

13.0

5.3

5.2

li);")?

PERUVIAN MAMMALS

3

have iimcli sliorlcf I'lir, color iniicli less l)ri<ilit, and slioiici' cars, riit'oi'tuiialcly both I'cniviaii specimens arc ^ouniicr, although one w;is oh] enouji'h lo l)e pi-euiiant (April 10) and llie ollici- estrons ( F(>l)ruary 1). Tlie Peruvian s])ecinieiis auree with the

Fis. 1 (;iliove;. I'liylloti.s (Galcnoviys) (jarleppi ; I'.o x. Fig. L' (below ). Fr'li'< jdcohlfd ; ().(> x.

tyj)e in these <lia^nostic i'eatni'es: incisors slender and nia.i-kedly ])r()odont. antci'ior hoiv'er of zyg'oniatie plate distinctly convex and extendin^.^' almost to the top of the rostrum, and dorsal pro- file of ihe skull strons'lv convex.

4 RREVTORA XO. 7.'^

One of the specimens Avas caught in front of a l)ui'i'ow of Ctenomys pernayius on a hea.vily "razed pampa of dwarf grass and prostrate forbs. The other was taken near a stone wall on a similar, grazed pampa on which were also growing scattered thornbnshes about one foot tall. Cienomys (yphnus was liviiig about TOO yards away. Other mammalian associates were (/(ilea niusteloidcs, Alxodon jelskii, PhyUoih darwini, Ph. siihlimis, and Hesperomys Jepidus dnciUa. The last two of these are sur- prisingly similar in appearance to (/arlcppi. Ph. garlcppi is larger than dmiUa (total length of garlcppi more than 120 mm., foot more than 20 mm.) and smaller than most subliniis with tail usually shorter (less than 46 nun.) and not bicolor, soles of hind feet more hairy than in .'>i(bli'}iiis. Using Pearson's (1951) key to the rodents of the altiplano west of Lake Titicaca, Galeno- mys keys out as either Phyllotis osilae, Ph. darivini, or Akodon aruocinis. All of these have much longer tails than does garlcppi.

Punomys lew minus Osgood. The range of this rare mouse has been extended considerably by its capture 55 mi. ENE Arequipa, 15,300 ft., Department of Are(|uipa, and 12 mi. NE Tarata. 14,600 ft., Department of Tacna. These new specimens agree well with those from Caccachara (Pearson, 1951) and with the lype.

In addition, nine specimens were taken 8 mi. 8SW Limbani, 15,000 ft., Department of Puno. This is 130 miles north of the type locality and separated from it by the grassland of the Titicaca basin. A circuitous strip of more favorable habitat may connect the two regions by way of the mountains to the west and northwest, but no specimens have been taken there. The specimens from near Limbani, like so many forms from this more humid region of the Andes, are distinctly and consistently darker than those from farther south or west. Compared wiih the type and with the sj^ecimens mentioned in the preceding paragraph, the Limbani Punomys are greyer and darker on the back, the feet and hands duskier al)ove, ears darker, tail less distinctly bicolor, and belly considerably darker grey with a distinct bult'y wash.

The Limbani specimens, as well as those from near Are(iuipa and Tarata, were captured in barren, broken I'ock areas and, as at Caccaciuira, were near fleshy-leaved, luuigent Soiccio plants or piles of Scnecio cuttings.

l!);")? PERU VI AX MAMMALS 5

Covin tscJnuh'i osgoodi Sanborn. Two s])CM'i('s of guinea piprs live on the altiplano of Peru and at some localities ])robably oeciir together. Cavia (Galea) mustcloicles is diurnal (Pearson, 1951) and prefers i-ather open habitat with jiood visibility. At Ilaeienda Calaeala the mueh darker colored Cavia (Cavia) iscliudii lived in thick grass where it made distinct nuiways and was crepuscular. 8teel traps set in the runways failed to catch any during the day, but numerous individuals were seen and collected in the evening after sunset and in the early morning.

Lagidinui poKanuDt Eleven. In Peru mountain viscachas have been considered to live only at high altitude. It was a surjirise, therefore, to find a small population living in the fog belt, or loma.-;, at only 2200 ft. at Nana, Department of Lima. The hilltops at Nana support scattered clumps of fog-nourished TiUandsia (Bromeliaceae). On the lower slopes there is no vege- tation, but thei-e are irrigated fields on the floor of the valle3^ Among rocks on one of the hilltops were many viscacha scats, which probably accumidate for years in the absence of rain, a few viscacha bones, and at least one living viscacha. This individual was seen at a distance of oidy 6 feet and appeared to be similar to the mountain viscacha,-; of southern Peru but with more than average buffy color. A maxillary toothrow picked up nearby agrees well with specimens of Lagidiuni pcruanum.

This population living more than 6000 ft. lower than any other viscacha population known to me in Peru is isolated by several thousand feet of brushy and weedy terrain unsuitable for viscachas.

Mustehi fytnaia Lichtenstein. Weasels have long been thought to live on the altiplano of Peru but have been inadequately rep- i-esented by specimens. A mounted skin is now available, taken in the spring of 1951 at Hacienda CaJ.acala, 13,000 ft., 7 mi. ^S^Y Putina, Department of Pnno.

Orison (Gri^onella) cuja (Molina). The presence of this nuistelid on the altiplano of Peru has heretofore been inade- (juately documented. We now have the skin and skull of an adult male from Ilaeienda Pairumani, 13.000 ft., 24 mi. 8 Have, De- partment of Puno. JNIeasurements (in mm.) are: total length, 570; tail, 145; foot, 70; weight, 1700 grams. This specimen was shot at 4 p.m. with its stomach crammed with at least three mice and a lizard, indicating diurnal feeding.

6 BRFA'IORA XO. 78

Felis jacobita Cornalia. The complex taxoiiomie history of this wildcat (see Osgood, 1943) has been based on a half-dozen skins, many of them without adecinate locality information, and on drawings of a single skull (Pliilii)pi, 1873). Xo previous specimens are known from Peru. AVe now have a skin and skull of a male, trapped March 30, 1952, among rock outcrops at 15,500 ft., 57 mi. ENE Arequipa, Department of Arequipa. This is a barren region of rocks and bare ground with scattered c]um])s of buncligrass [Fcsiuca orihophxiUti) and small tola bushes. Other steel ti*aps nearby caught a. fox [Diisicifov ciilpdCHs) and a mountain caracara (PlialcohooiKs uicgaJopterus) . A mountain lion passed close by several times. Probable prey items in addi- tion to small rodents and small birds were mountain viscachas {Lagidiuni), tinamous (Tinamotis), and seed snipe {Attagis. Thinocorus) . Vicunas, the young of which might be killed by jncohito, were abundant.

The skin matches well the vai-ious published descriptions, but several features of the skull, some of them previously considered to be diagnostic, do not matcii the illustrations in Philippi's rei)ort. The audital bullae of our specimen are not bisected by a deep sulcus; in fact, scarcely a trace of the sulcus shows. Com- pared to four Felis pajeros from southern Peru and northern Chile, the nasals of our jacobita are only slightly larger and the incisors not more jn-oodont. The skidl of our jacobita is slightly flatter and more i-obust than in pajvros, and has lai-ger teeth, but skulls of the two s])ecies are not as different in ai)pearance as are those shown in Philipi)i's illustrations. A possible diag- nostic cliaracter is the orientation of the upper premolars, in pajeros these two teeth on each side, when vie\\t'(l from l)elow. lie in a straight line. In jacobita the anterior premolar toes in abruptly in front. Since no other skulls seem to bi' available and since our specimen differs considerably from the one seen by Philippi, two photographs are presented in Figure 2. Measui-c- ments (in mm.) are: total length, 990; tail, 413; foot. 133; ear, (i3 ; Wright, 4.0 kg. ; condylobasal length, 9(i ; greatest length of skull, 100; zygomatic lu-eadth, (i!>.5 : greatest length of upi)er carnassial, 14.0.

lO;")? I'KRirVIAX MAMMAliS 7

'I'lu' ]»t'la;:t' is Hut'tici- and jLii-cyci- than that of /•'. pujcros and is without a spinal crest of loiij^- hairs; tlic ears are rounded without tufts, and the tail is non-taperino-, much louf>er, an<i more eonspicuously ringed than in pajeros.

In southern i'ei-u Fclis jacohifa seems to be mueh less ahuiul- ant than F. pajeros and probably prefers higher elevations.

BIBLIOGRAPHY

( )S(H>OI>. \V. il.

1943. The iiwmminls of Cliilc. Fiold Mus. Nat. Hist., Zool. Ser., 30: 1-268.

I'EARSOX, U. P.

li>.il. ^[;llnln;lls in tin- liigliliiTxls of southern Pern. Bull. Mus. Comp. Zool., 106: 117-174.

rmi.ippi, R. A.

1S7.S. Ueber Felis Ciuina Molina and iiber die iScliJidelbildung bei Felis Pa.ieios und Felis Oolocolo. Areh. Xaturg., 39 (1): 8-15, pis. 2-3.

Sanhokx. 0. C.

1 !».■)(•. Small rodents from Peru and Bolivia. Pul). Mus. Hist. Nat. "Javier Prado," Ser. A. Zool., No. 5: 1-16.

BREVIORA

MeseiJim of Comparative Zoology

Cambridge, Mass. May 1, 1957 Number 74

THE DISCOVERY OF CERAPACHYINE ANTS ON NEW

CALEDONIA, WITH THE DESCRIPTION OF NEW SPECIES OF PHYRACACES AND 8PHINCT0MYRMEX

By E. 0. Wilson

Biological Laboratories, Harvard University

During December, 1954, and January, 1955, the author was fortunate in being able to spend an uninterrupted five-week period on New Caledonia, studying the ant fauna of the island.^ During this time three species of Phyracaces and Sphinctomyr- mex were collected, all undescribed and representing the first cerapachyine ants ever found on New Caledonia. Their presence is of some zoogeographic importance, in that they appear to ally the ant fauna of New Caledonia more closely with that of eastern Australia, as opposed to the remainder of Melanesia. Phyracaces and Sphinctomynnex are strongly developed both in species and in individual numbers in Australia, but are known from only three rather rare species (two Phyracaces and one Sphincto- myrmex) on New Guinea, and are unknown from the rest of northern and central Melanesia. In an obverse relationship, Cerapachys {s. sir.), is the predominant cerapachyine genus on New Guinea and the Fijis, but has never been collected in either Australia or New Caledonia. Finally, the New Caledonian Phyra- caces and Sphinctomyrmex are most closely allied to eastern Australian species, as indicated in the descriptions to follow.

Phyracaces cohici Wilson, new species

Diagnusis. A shining, black, medium-sized species closely re- lated to the turneri group of species of eastern Australia. P.

1 Field research was supported by grants from the Society of Fellows, Harvard University, and the Museum of Comparative Zoology.

2 BREVIORA No. 74

cohici can be easily distinguished from the latter group, which includes turneri Forel, adamus Forel, and larvatus Wheeler, by its more obtuse and rounded dorsal propodeal corners. In cohici these corners form an angle of more than 110° when viewed from the side, while in the turneri group of species they form an angle of 90° or less. P. cohici also bears a fair resemblance to the Australian P. senescens Wheeler, but can be separated from this species by its longer, flatter petiolar node and straighter and more horizontally aligned posterior petiolar teeth.

Holotype worker. Head width ^ 0.99 mm, head length 1.05 mm, scape length 0.63 mm, cephalic index 94, scape index 64, exposed length of mandibles 0.21 mm, eye length 0.26 mm, pronotal width 0.79 mm, petiole width 0.83 mm, petiole length (measured from the midpoint of the anterior border of the node to the midpoint of the posterior border of the posterior peduncle) 0.63 mm, postpetiole width 0.83 mm, postpetiole length 0.67 mm, width of next gastric segment 1.00 mm. Occipital border very feebly convex in full-face view. Alitrunk viewed from above moderately constricted medially, marginate along the entire dorsolateral border except for an interval of about 0.20 mm in the region of the mesothorax. Petiolar node viewed from directly above with strongly concave anterior border, and evenly convex lateral borders, its widest point being about in the middle (see accompanying figure). In the same view the posterolateral teeth extend well beyond the posteriormost point of the posterior node border. Seen from directly above, the postpetiole is widest in the anterior half and is laterally marginate only in the anterior half.

Entire body covered by scattered piligerous punctures spaced on an average of 0.03 to 0.06 mm apart, the interspaces com- pletely smooth and shining. On the sides of the alitrunk the punctures are unusually small, being barely visible at 40X magnification.

1 Head width and other body measurements were made in the standard fashion employed by W. L. I'.rown in his published serial studies of the daeetine ants and by myself in my recent revision of the ant genus Lasiug (1955, Bull. Mus. Comp. Zool. Ilarv., 113: 1-200). In the past, Brown and I have differed slightly in our cleflnitiuus of head length, but in the present paper I have decided for the purposes of uniformity to follow Brown's original definition, which was given as follows : "critical maximum length of the head, measuring from a transverse through the posteriormost point or points along the occipital border to a trans- verse through the anteriormost point or points on the anterior clypeal border" (1953, Amer. Midi. Nat., 50: 11). With this change I believe that our standard- ized measurements now all agree in every detail.

1957 CERAPACllYINE ANTS OF NEW CALEDONIA 3

Entire body deep blackish browu to jet black, except for the mandibles and gastric apex, which are dark reddish brown. Ap- })endag'es variably dark reddish brown.

Worker variation. Maximum head width of tj^pe series (all shown by a single colony, ace. no. 190) : 0.88-1.01 mm. The worker tj^pe series shows very little variation in other external characters.

MaJe. Head width (across and including compound eyes) 1.10-1.14 mm. Antenna 13-segmented. Mandible well developed, narrowly subtriangular, its masticatory border bearing a large, blunt apical tooth followed basally by an indeterminate num- ber of serial denticulae ; the masticatory and basal borders join- ing through an even, convex curve. Clypeus narrow, its dorsal surface gently convex, its anterior border seen from directly above moderately and evenly convex.

Parapsidal furrows well developed, parallel with each other, and extending anteriorly for about half the length of the scutum. Notaulices absent. Median notal suture present and extending posteriorly for slightly more than one-third the length of the scutum. Wing venation generalized, essentially similar to that of Cerapacliys (see Brown and Nutting, 1950, Trans. Amer. Ent. Soc, 75: 132, pi. VIII), differing primarily as follows: Rsf2'3 is lacking, and the second radial crossvein and Bsf5 form a single unit coming off the stigma. The crossvein " x" indicated by Brown and Nutting in the basal cubital-anal region of Cerapacliys manni is missing in Fhyracaces cohici.

Petiole completeh^ lacking the lateral margination that char- acterizes the Fhyracaces worker caste. Seen from above, the node is widest in its anterior half, and its anterior and posterior corners are gently rounded ; seen from the side, the node forms a single, even, strong convexity only weakly demarcated from the peduncles. Genitalia completely retractile. Subgenital plate relatively large (exposed length about 0.35 mm), sclerotized, tapering posteriorly to end in a pair of sharp, upward curving hooks each about 0.06 mm in straight length. Parameres short, broadly rounded apically.

Pilosity, sculpturing, and color essentially as in the worker.

Types. Described from a long series of workers and four males collected by the author at Ciu, near Canala, at 300 meters, and from two kilometers southwest of Ciu, at 500 meters, New Cale-

4 BREVIORA No. 74

donia. The following accessions are included : 190, 246, 263 (holotype nest series), 267, 275, 278, 298. Each represents a separate nest series with the exception of numbers 246 and 275, which are from the same nest. All of the collections were made in the period from December 21, 1954, to January 2, 1955.

This species is named for Mr. Francois Cohic, of the Institut Frangais d 'Oceanic, an able and enthusiastic student of New Caledonian entomology.

Ecological notes. All of the collections were made in the rich subtropical evergreen forest clothing the hills that extend from the south bank of the Canala River in the vicinity of the Ciu Falls. The holotype colony was found under a rock in a densely shaded part of the forest, and was occupying a single cavity and adjacent short vertical gallery in the soil. The males were very active and attempted to fly when the nest was exposed. Another colony (ace. no. 246-275) was nesting in open soil at the side of one of the forest trails. The nest entrance consisted of a single five-millimeter-wide opening surrounded by a low, indistinct turret of excavated earth. Lateral excavation revealed three or four galleries leading down from the entrance hole and into soil packed between several buried rocks. At about ten centimeters down two small adjacent chambers had been excavated in the soil against the vertical face of one of the rocks. In these were massed all of the brood and most of the workers. The following rough population estimate was made at the time of collection : 80-100 workers, 40 pupae (in cocoons), 30-40 half -grown to fully grown larvae, and 30 eggs. The reproductive of this colony was unfortunately not found.

Workers belonging to colony no. 246-275 and other colonies were encountered on several occasions foraging during the day, always in file, over the surface of the ground, and on one occa- sion (no. 190) workers were discovered in the upper layers of a moist rotting log. Twice, workers were observed in the act of raiding colonies of the ant genus Pheidole. The foraging and raiding behavior of this species will be described in greater detail in a later paper on the general subject of cerapachyine behavior.

1957 CERAPACIIYINE ANTS OF NEW CALEDONIA 5

Phyracaces dumbletoni Wilson, new species

Diagnosis. Distinguished from all other species of the genns known to me by the following combination of characters : mod- erately large size (head width of type series 1.16-1.25 mm), antennal scapes unusually long, sides of alitrunk non-marginate, body surface smooth and shining to shagreened and subopaque, body color deep blackish brown to jet black.

The only other Indo- Australian Phyracaces lacking margina- tions along the sides of the alitrunk are aherrans Clark and pygmaeus Clark of northern Queensland and hewitti "Wheeler of Borneo. These species are easily distinguished from dumhletoni by their much shorter scapes, which reach only to about the posterior margins of the compound eyes when the head is viewed in full face in dumbletoni the scapes extend beyond the eyes by about their own maximum width. Of the three species, dumbletoni most resembles aberrans Clark, being very close in both size and sculpturing.

Holotype ivorker. Head width 1.17 mm, head length 1.30 mm, scape length 1.08 mm, cephalic index 90, scape index 92, exposed length of mandibles 0.29 mm, eye length 0.33 mm, pronotal width 1.03 mm, petiole width 0.95 mm, petiole length 0.76 mm, post- petiole width 1.09 mm, width of next gastric segment 1.26 mm. Occipital border in full-face view very feebly convex. Alitrunk viewed from above only feebly constricted medially, its dorso- lateral area evenly rounded and lacking any trace of margina- tion. Petiolar node seen from directly above with moderately concave anterior border and evenly convex lateral borders, its greatest width located in the anterior half.

Puncturation as described for P. cohici, except that anterior to the postpetiole the punctures are more scattered, the majority being 0.09 to 0.12 mm apart. On the postpetiole and anterior to this segment the interspaces are for the most part "shag- reened," the shagreening in this case in reality consisting of regular, minute, contiguous foveolae each about 0.01 mm in diameter. The foveolae are deepest on the dorsal surface of the head and alitrunk, and render most of the cuticular surface there subopaque. They are absent posterior to the postpetiole.

Color as described for P. cohici.

BREVIORA

No. 74

Worker variation. Maximum head width range, Chapeaii Gendarme and Mt. Mou series: 1.16-1.25 mm (all shown by ace. no. 65, the holotype nest series). Maximum head width range, Ciu series: 1.11-1.20 mm (all shown by ace. no. 245). The Ciu workers differ significantly from the Chapeau Gendarme and Mt. Mou workers in the intensity of the f oveolar ' ' shagreening. ' ' The latter approach the condition described in the holotype, with little deviation. The Ciu specimens have the same basic form of sculpturing, but on the head and alitruncal dorsum the foveolae are much shallower, so that the surface is strongly shining under ordinary reflected light.

cohici

dumbletoni

Dorsal view of the petioles of the two New Caledonian species of Phyracaccs ; outlines drawn to scale from the holotyjoes.

Ergatoyyne. Head width 1.14 mm, head length 1.26 mm, scape length 0.76 mm, cephalic index 90, scape index 67, exposed length of mandibles 0.22 mm, eye length 0.30 mm, pronotal width 1.00 mm, petiole width 0.95 mm, petiole length 0.62 mm, postpetiole width 1.32 mm, width of next gastric segment 1.55 mm. Ocelli lacking. More similar to the worker caste than to the queen caste of other Phyracaces species, differing primarily in the following external features. (1) The alitrunk is very worker-like, apparently differing only in the somewhat stronger pleural suturation. The posterior metapleural suture, which is absent in the dumbletoni Avorker, is present although feebly de- veloped in the ergatogyne. (2) The petiole is relatively shorter

1957 CERAPACHYINE ANTS OF NEW CALEDONIA 7

in the ergatogyne. (3) The postpetiole and gaster are much more massive and more poorly demarcated from each other than are the same structures in the worker.

Types. Described from a long series of workers from Chapeau Gendarme (Yahoue), Mt. Mou, and Ciu, and a single ergatogyne from Ciu. The following accessions are included, each repre- senting a separate nest series: Chapeau Gendarme, no. 65 (holo- tj'pe nest series); Mt. Mou, single worker; Ciu, no. 245 and ' ' observation colony. ' '

This species is named after Mr. L. J. Dumbleton, formerly economic entomologist for the South Pacific Commission, and an expert student of the New Caledonian insect fauna.

Ecological notes. P. dumhletoni was collected at Chapeau Gendarme and Mt. Mou in dry, semi-deciduous "valley-pocket" forest and at Ciu in moist broadleaf evergreen forest. In all three localities it was limited primarily to the least disturbed portions of the forest, and was never encountered in the adjacent open Melaleuca woodland.

At Chapeau Gendarme a large colony (no. 65) was found nesting in several spacious galleries and chambers in the upper layers of a large, moist, fern-covered log. It contained at least 200 workers, a single ergatogyne (later lost), over 100 cocoons and larger larvae, and an undetermined number of eggs. Among the brood were found the hollowed-out propodeum of a worker of an undescribed species of Lordomyrma and the mangled remains of the larva of an undetermined ant genus. These insects ap- peared to be the prey of the Phyracaces, but of course this can- not be proven.

When first disturbed, many of the Phyracaces workers sallied out with a display of aggressiveness unusual for cerapachyines, and one succeeded in stinging me on the forearm. I think it is worth mentioning that this is the only time I have ever been stung by a cerapachyine ant, despite the fact that I have ex- cavated many nests of Phyracaces and other genera without mak- ing any effort to protect myself from the workers. The sting caused a prominent welt about six millimeters wide surrounded by an erythema about twenty-five millimeters wide. There was a persistent, dull, throbbing pain of the sort commonly resulting from the stings of ponerine ants.

8 BREVIOBA No. 74

Sphinctomyrmex caledonicus Wilson, new species

Diagnosis. Closely resembling S. steinheili Forel of eastern Australia, from which it can be distinguished by the following two characters. (1) Sculpturing, which is dominated by punc- turation, is overall denser in caledonicus. In steinheili, the inter- space distances over the genal surface are mainly as great as the adjacent puncture diameters or greater, while in caledonicus the genal punctures are contiguous and their borders form an even rugoreticulum. In steinheili, the lateral surfaces of the alitrunk are in large part feebly shining, while in caledonicus these surfaces are entirely opaque. (2) In side view, the dorsal posterior corners of the propodeum form a distinct angle of about 110° in steinheili, but are evenly rounded in caledonicus.

Holotype worker. Head width 0.56 mm, head length 0.71 mm, scape length 0.41 mm, cephalic index 79, scape index 73, exposed length of mandibles 0.06 mm, pronotal width 0.42 mm, petiole width 0.35 mm, petiole length (including peduncles) 0.38 mm, postpetiole width 0.47 mm, w^idth of next gastric segment 0.60 mm, length of gaster posterior to postpetiole (measured in a straight line) 1.35 mm.

Worker variation. Maximum head width range (internidal) 0.50-0.57 mm; (intranidal; aec. no. 195) 0.50-0.56 mm. In size as well as other external characters the worker type series is remarkably uniform.

Ergatogyne. Head width 0.62 mm, head length 0.76 mm, scape length 0.42 mm, cephalic index 82, scape index 68, exposed length of mandibles 0.09 mm, pronotal width 0.44 mm, petiole width 0.42 mm, petiole length 0.41 mm, postpetiole width 0.60 mm, length of gaster posterior to postpetiole 1.96 mm. This specimen is very worker-like, and can be distinguished only by its slightly larger size, proportionately shorter head and scapes, broader petiole, and larger postpetiole and gaster. In addition the postpetiolar-gastric constriction is somewhat weaker than in the Avorker. Compound eyes and ocelli are completely lacking, as in the worker, and the structure of the alitrunk is essentiallj- the same.

Types. Described from a long series of workers and a single ergatogyne from Ciu, 300 meters, New Caledonia. The following two accessions, representing separate nest series, are included :

1957 CERAPACHYINE ANTS OF NEW CALEDONIA 9

no. 195, no. 225. The holotype was chosen from no. 225.

Ecological notes. The two type colonies were found in a small, isolated patch of broadleaf evergreen forest on the farm of Mr. D. Fere, about half a kilometer northwest of the Ciu Falls. This little woodlot did not exceed two acres in extent, and its floor had been badly disturbed by cattle. Most of the ants present, including the Sphinctomyrmex, were found underneath rocks embedded in the soil. It is curious that the Sphinctomyr- mex was not found in the undisturbed forest on the south bank of the Canala River a short distance away, despite intensive collecting there.

Both colonies were quite large, one containing over 500 work- ers and the other over a thousand. That the colonies may have been in migration is suggested by the fact that they occupied poorly defined galleries in the soil which bore no sign of lengthy occupation. The brood of colony no. 195, collected on December 21, consisted of large numbers of mature larvae ; two days later, about three-quarters of a sample of these larvae kept alive in a bottle had spun cocoons. The brood of colony no. 229, collected on December 31, consisted of large numbers of cocoons, all of which contained prepupae of indeterminate caste. These data suggest a high degree of synchronization of brood development, a condition usually associated in ants with a nomadic way of life. An account of the behavior of the w^orkers of this species will be given in a later paper.

BREVIORA

Museiuoi of Comparsitive Zoology

Cambridge, Mass. May 1, 1957 Number 75

ON A NEW OCTOCHAETINE EARTHWORM SUPPOSEDLY FROM GUATEMALA

By G. E. Gates

INTRODUCTION

Tweuty-four species of earthworms have beeu reported from Guatemala. One of them is widely distributed in North America where it presumably originated. Another, "pantropical," may have come from the south. Four are obviously exotic, probably introduced accidentally by man since 1500 A.D. The eighteen supposed endemics are known only from the original material.

As a result of one of those accidents that occasionally advance our knowledge of an infrequently collected group of animals, another presumed endemic now can be added to the list of species known only from a single specimen.

The author's thanks are extended to Dr. P. W. Oman for providing the present specimen and for assistance rendered in connection with problems arising from the study of various other species.

Family MEGASCOLECIDAE Subfamily OCTOCHAETINAE

Genus RamiELLA Stephenson 1921

RaMIELLA AMERICANA n. Sp.

Guatemala. In soil with Ceiba pentandra in cargo on plane arriving at San Pedro, November 25, 1955, 0-0-1. U. S. Bureau of Plant Quarantine.

2 BREVioRA No. 75

External characteristics. Length, 133 mm. Diameter, 8 mm. (through clitellum). Segments, 217 (+? possibly a recent posterior amputee). Body slightly compressed dorsoventrally and transversely elliptical in cross section behind clitellum. Un- pigmented (formalin preservation). Prostomium, prolobous. First two segments with numerous fine longitudinal grooves in both ventrum and dorsum. Secondary annulation, one presetal furrow each on iv-xii, one postsetal each on vi-xii, first presetal furrows deep but from ix less marked and like the postsetal; secondary furrows lacking behind the clitellum. Setae small, closely paired, only very slightly or not at all protuberant from epidermis, unrecognizable (lacking?) anteriorlj^; ventral couples first visible on vi, the lateral on xi, CD<^AB, J-A<BC, DD ca.=|C, both a and b present on left side of xrsdii, h only on right side, ventral couples of xvii and xix penial. Follicles of ventral couples larger than the others, ventral setae probably also larger. Nephropores unrecognizable. First dorsal pore on 11/12.

Clitellum reddish brown, saddle-shaped, reaching ventrally about to B, intersegmental furrows obliterated, dorsal pores occluded, setae retained, on xiii-xx. Epidermis 3-4 times as thick as rest of the body wall, cracking into blocks on slight tension.

Spermathecal pores transverse slits of about the same width as AB, centered at A, on 7/8-8/9. Female pore in a transverse groove in A A just in front of eq/xiv. Male pores minute, about half way between B and a furrow circumscribing a male field, at 17/18. Prostatic pores minute, on lateral wall of a common lumen containing both a and h penial setae, on xvii and xix.

Male field longitudinally and broadly elliptical, demarcated by a distinct circumferential furrow somewhat lateral to B and just behind eq/xvi but less obvious in front of eq/xx. Two small but conspicuous conical protuberances on each of xvii and xix, about in region of AB, might be either localized temporary eleva- tions or eversions of slight parietal invaginations. The large aperture at the tip of each is filled by two penial setae. No other genital markings.

Internal anatomy. Septum 4/5 membranous and transparent: 5/6-9/10 thickly nuiscular, funnel-shaped, posteriorly directed; 10/11 and 11/12 much thinner though slightly strengthened, ap- posed at parietes laterally and ventrally but easily separable till

1957 NEW OCTOCHAETINE EARTHWORM 3

just lateral to hearts of xi and there strongly adherent, ap- parently united dorsally and without insertion on body wall so that gut, dorsal trunk and hearts of xi are invisible at first. Septum 12/13 and those following all membranous.

Progizzard portion of gut ca. 10 mm. long, the thin-walled oesophagus sigmoid. Gizzard large, in v. Gut in vi-xiv slender; caleiferous glands and lamellae unrecognizable. Inte.stinal origin in XV. Caeca vertically placed, 2.5-3 mm. long (high), slightly narrower dorsally, protuberant above dorsal face of gut so that the large dorsal trunk at first glance appears to be within a depression of intestinal roof, nine pairs in xxi-xxix. Typhlo- sole begins in xix-xx though a nuu'h lower ridge extends forward into xvi, reaching a height of 2 mm., lamelliform, anteriorly for a short distance Avith vertically lamellar lateral protuberances, ending abruptly in clxii. A fairly high and rather thick longi- tudinal ridge on each side in xxi-xxviii, between apertures into caeca and the median typhlosole, appears to be a lateral typhlo- sole. It probably does not function as a valve to close off lumen of gut from the caeca as the latter are filled with soil. (No supra-intestinal glands.)

Dorsal blood vessel single throughout, large, gradually dwin- dling anteriorly and disappearing into tissues of pharyngeal bulb. Supra-oesophageal vessel recognizable only in xi-xii. Ven- tral trunk unrecognizable anterior to viii. Subneural unrecog- nizable (lacking?). Extra-oesophageals or other longitudinal vessels not found. Hearts of xi-xii large, latero-oesophageal though presumed bifurcations to dorsal trunk have no blood. Hearts of x somewhat narrower and lateral, those of ix-viii still slenderer and also lateral. Paired vessels from dorsal trunk in vii-v cannot be traced to the ventral trunk. ("Lymph glands" lacking.)

Nephridia of iv represented by a large cluster of looped slender tubules on each side of anterior face of 4/5 (ducts not found nor any funnels, possibly closed, enterouephric and open- ing into pharynx). Paired, smaller clusters of similar slender tubules present on anterior faces of septa ventrally in v-xi, in viii-ix also attached to posterior faces of spermathecal ducts (ducts and funnels not found). Nephridia, from xii posteriorly on the parietes, on each side one transverse row of short loops from A nearly to mD. Nephridial tubules are as slender as

4 BREVioRA No. 75

anteriorly in xii-xvi but from xvii much thickened and flattened without marked increase in length. The thickening appears to be due to presence of the same sort of tissue as conceals the slender tubules of the micronephridia in certain species of Dichogaster. Individual loops, 9-10 on each side in each segment, are con- nected by a slender transverse cord (blood vessel? or?) but similar cords may be recognizable passing from the loop into the parietes or to the gut. Nephrostomes apparently are lacking until well posteriorly. From the medianmost loop on each side, beginning about at cxxv, a slender cord passes anteromesialh^ to and through the anterior septum, the anterior free end median to A very slightly widened. This cord and its slight terminal thickening presumably are neck and funnel respectively but even under highest power of the binocular definite funnel char- acters cannot be distinguished. Posteriorly the medianmost component of each transverse series becomes somewhat larger and of more than a single loop, and the larger widening of the free anterior end of the cord (neck) becomes definitely funnel- like. Typical, stomate exonephric holonephridia certainly are lacking throughout the entire length. Brain and circumpharyn- geal connectives well anteriorly in iii.

Metandrie (no funnels in x). Male funnels large, plicate. Male deferent ducts large (with muscular sheath?), with large lumen, on parietes in BC, passing into body wall about at level of 17/18, slightly narrowed in xvii and still more so within the body wall. Seminal vesicles, in xii, medium-sized, cut up into very many fine lobes. Prostates tubular, looped and coiled, ca. 20 mm. long, lumen central and recognizable clear to ental end, in xvii-xviii and xix-xx. Ducts 3-|- mm. long, slightly narrower than gland, with smooth surface, muscular sheen, slit-shaped lumen, muscular layer much thicker than the lining epithelium. Penisetal follicles at first separated from prostatic duct in body wall by a thin vertical sheet of longitudinal muscle, united more deeply.

Penial setae have been studied by Mrs. Dorothy McKey- Fender. Her description is as follows :

"Penial setae (Fig. A) curved, especially proximallj^ and distally, flattened in the plane of curvature and somewhat twisted, the apical, more strongly curved portion somewhat widened, forming an asymmetrical, spoon-shaped blade (Fig. C)

1957

NEW OCTOCHAETINE EARTHWORM

with attenuate, often spirally twisted, tip. Distal seventh of shaft, including tip, sculptured on sides and both faces. Sculp- ture (Fig. B), of interrupted rows of fine teeth that become slightly longer and fewer per group toward the apex of the shaft. Setae a and h of both xvii and xix alike, except that the tip of the & setae may be more strongly twisted. An apparently mature, unworn seta is 3.2 mm. long (disregarding curvature), 0.1 mm. wide and (being rectangular in section) 0.05 mm. thick."

Penial setae of Bamiella americana.

A. Camera lucida drawing of an entire seta.

Arrow indicates location of detail shown in B.

B. Detail of edge of blade of same seta, showing character of sculpture

(More highly magnified.)

C. Face view of distal portion of a fc seta from segment xix (sculpture

omitted). Drawings by Mrs. Dorothy McKey-Fender.

Spermathecae adiverticulate, ca. 5 mm. long, a band of con- nective tissue passing from each duct to the septum just in front. Ampulla narrowing somewhat entally, with several annu- lar constrictions externally, ridging of inner wall and large lumen. Duct ca. li mm. long, thicker than ampulla but asym- metrical, somewhat and gradually narrowed in parietes, lumen slit-like, wall thick, muscular. Seminal chambers, seven or eight on the bulged side, one or two smaller on the other side, com-

6 BEEVIORA No. 75

municate narrowly with duct lumen. Ovaries very small. (No ovisacs found.)

Reproduction. Testis sac with little coagulum and that within a forward pocket-like, median bulge of 10/11. Male funnels with iridescence recognizable only at margins of the folds. Flecks of red iridescence recognizable in each lobule of the seminal vesicles (sheen of muscles in connective tissue binding lobules together, green-red-orange). Clitellum probably at or near maximal tu- mescence. However, seminal chambers in spermathecal duct and the ampullae are empty. Iridescence is unrecognizable ter- minally^ in male deferent ducts. Obviously copulation had not taken place. Accumulation of mature sperm on male funnels presumably had begun only fairly recently though the clitellum appears to be already at maximal development. Sexual, and bi- parental, reproduction is anticipated.

Remarks. The gut, through xiii-xiv where coelomic cavities were filled with coagulum, is macerated but probably not enough so that intramural calciferous lamellae would have become un- recognizable. Extramural glands, since the intestine begins in XV, would have to be in xiii-xiv. In those segments, small slenderly-stalked glands, judging from experience with poorly preserved material of other species, could have been lost in wash- ing out the coagulum, if disintegration had not been previously completed. Site of stalk-gut junction, in either case, probably would be indistinguishable. As there is no reason for believing that extramural glands had been present in the holotype, absence in the species must, for the present, be assumed.

The excretory system obviously is meronephric.

DISCUSSION

Absence of calciferous glands in ix-x, location of the male pores on xviii at some distance from external apertures of all prostatic ducts, and the meronephry require (cf. key in Pick- ford, 1937, p. 98) the Guatemalan species to go into the Octo- chaetinae. In that subfamily, only two genera now appear to require consideration.

The genus Ramiella, hitherto recorded only from India and Burma, is defined by its author (Stephenson, 1930, p. 845) as follows: "Setal arrangement lumbricine. All septa present (be-

1957 NEW OCTOCHAETINE EARTHWORM 7

hind their commencement). One oesophageal gizzard in one simple segment. No calciferous glands. Excretory system purely micronephridial, the micronephridia relatively large and few in number, from seven to one pair per segment. Sexual apparatus purely acanthodriline. ' '

The phrase "purely micronephridial" was intended to convey the idea that holonephridia are lacking from one end of the body to the other. One word, meronephric, is sufficient for that pur- pose. Further characterization of the excretory system is not warranted at present^ any more than previously, the supposed reduction of meronephry to a single pair of nephridia per seg- ment being based, in one species at least, on the number of opaque white masses that were externally recognizable through a semi-transparent body wall. The definition accordingly should read, in part, only as follows: Excretory system meronephric.

Ramiellona Michaelsen 1935, also meronephric, differs from Ramiella only in having calciferous glands. The latter are characterized (p. 53) as follows: "segmental Erweiterungen des Osophagus mit blutreichen Falten der Wandung im 12 Segment und einigen vorhergehenden." The small vascular folds on the inner wall of the oesophagus in Ramiellona stadelmanni Michael- sen 1935 (the only species) are not known to differ from similarly located ridges in Ramiella hishamhai'i (Stephenson, 1914) and R. nainiana Gates 1945. As Ramiellona is not dis- tinguishable morphologically, the Honduran stadelmanni, as well as the Guatemalan americana, must go into Ramiella.

The huge distributional discontinuity thus established is not exceptional in a system with geographical ranges such as the following: South India-Ceylon and Australia-New Zealand (Notoscolex and Megascolex), Australia-New Zealand and the Pacific coastal strip of the United States {Plutellus and Mega- scolides), temperate-zone North America and peninsular India (Diplocardia) .

The two Central American species of Ramiella do nut appear to be more closely related to each other, than to any of their

1 Specimens of Ramiella nainiana Gates 1945, tixed aud preserved in accordance with his instructions, were forwarded, along with other material, to K. N. Bahl for study of the nephridia. If his failure to publish thereon had been anticipated, :jome data would have been included in the original description while the worms were still available. Ever since, live individuals of any species of Ramiella have been unavailable.

8 BREVIORA No. 75

oriental congeners. The anatomy, as known, lacks the uniformity one should expect in a good genus. Ramiella thus provides one more illustration of the inadequacy of the classical system and the futility of patchwork changes in generic definitions .

EEFEBENCES

Gates, G. E.

1945. On some Indian earthworms. Proc. Indian Ac. Sci., 21: 208-258.

MiCHAELSEN, W.

1935. Die opisthoporeu Oligochaten Westindiens. Mitt. Mus. Hamburg, 45: 51-64.

PiCKFORD, G. E.

1937. A monograph of the acanthodriline earthworms of South Africa. Cambridge, England, 612 pp.

Stephenson, J.

1930. The Oligochaeta. Oxford, 978 pp.

BREVIORA

Museniim of Coimparative Zoology

Cambridge, Mass. June IS. U).")? Number 76

TWO NEW LAXI) AXl) FKESllWATEK MOLLl'SKy FROM NEW GUINEA

By AViLLiA.M -I. Clench

The tirst species was received from Mr. James Poling of New York City, the second from ]Mrs. Marjorie Kleckham of Darn, l'ai)na. New Gninea.

]\Ir. Poling pnrchased all of the remaining stock of the land and marine mollusks from Ward's Natural Science Establish- ment in Rochester, New York, after the death of Mr. F. H. Ward. Contained in this miscellaneous material were a few lots of land shells, originally obtained from some collector in New Guinea. Subsequent correspondence by ^Ir. Poling with Ward's has failed to add to the meager data accompanying the specimens. Just wlio the collector was or the exact locality in New Guinea still remains unknown. The locality is somewhere in western New Guinea, to judgi^ by th(» associated specimens from the same ('()ll(H'tor.

VeR1)ICHL01[1T1S new subgenus

Tliis subgenus differs from all other species in the subfamily Chloritinae by being a dull jade green. All other species, now inunbering over 200, are uniforml}' brown, red-brown, or banded with some shade of broAvn or red-brown. The usual arrangement of the periostracal "hairs" is in staggered or offset rows; in this subgenus the rows are regular.

T]ipr aperies. Eusfrnxopsis (VerclicJdo)-)fis) polijufi Clench.

EUf^TOMOPSIS ( \'e1;I)U IILOKITISJ POLINGI. uew S])ecies

Figure 1 1)( scnptioii. Shell small. i-cMching 1") mm. in gi-eatest diameter,

BREVIOKA

NO. 76

plauorboid in shape, thin, fragile and minutely hirsute. Color a uniform and dull jade green. Whorls 31/2, convex, and coiled in a single plane. Spire depressed below the body Avhorl. Aper- ture subcircular. Outer lip thin and very nan-owly reflected.

Fig. 1. Kiistoniopsifi (I'crdichlorili.s) poivuiii Clciu-li (llolotyi)e, 4X). Fig. '1. ll'e.sfralunio <ill)r)-ti.'<i Clcncli (Holot.vpc, 1.5X ).

Parietal area covered with a vei-y thin glaze. Scul])turc consist- ing of numerous rows of very short, gold-colored periostracal "hairs" which arc arranged axially and slightly diagonally.

1957 TWO NEW 1,A.\I) AM) KKKSI 1 WATEIi .MOIJ.ISKS 3

i'ollowiu<i' the liiu's of <ir()\vtli. These rows nvv I'ejiular aiul not staggered or offset as in other species of Eustomopsis. UmbiliciLs narrow but deep, the nueleai' whorl being visil)h'. Periostraeiim jad(^ green overlaying a nearly glass-like shell.

Height (it. (liametor Less, diameter

7.5 mm. 15 mm. 11.5 mm. Ilolotype

Type. The holotype is in tlie ]\Iuseum of Comparative Zool- ogy, No. 212319, from western New Guinea.

Neinarks. This new species is (piite remarkable. The dull jade green color sets it well apart from all other species in the sub- family Chloritinae. So far as I am aware, all other species in the Chloritinae are uniform brown, red-brown, chocolate brown, or light brown to yellowish and banded witii brown. Many species are hirsute, that is, the periostracum forms little "hairs" in regular or offset rows, such "hairs" emanating from small pits impressed in the calcium of the shell.

It is quite possible that this species is arboreal and, as in many other groups, the species which become arboreal also become somewhat brightly colored, as, for example, species in Licjims, PoJijmiia, Placostylus, Helicostyla, Amphidromus and many other genera.

A factor of considerable interest is the green color, a rare color even in arboreal mollusks. A few species are banded with green, such as Liguus and Amphidromus but nearly solid green is exceedingly rare. Helicina viridis Lamarck and H. castilloi Clench from Hispaniola, Papustyla p^ilcherrima Rensch from the Admiralty Islands, two or three species of Helicostyla and Ckloraea from the Philipiiines are about all the known species to po.ssess this color predominantly.

AVestralunio albertisi, new species Figure 2

'^p "

Description. Shell inequilateral, subcircular in outline, rounded anteriorly, subtruncate posteriorly and reaching 53 mm. in length. Umbos rather small, slightly elevated above the dorsal margin and anterior to the center. Shell moderately compressed and rather light in structure. Color a dark blackish brown. Posterior slope fairly well marked and slightly concave. Po.s-

4 BREVIORA XO. 76

tcrioi- i-i(l<z'e not well defined. Lio-ament lon^' and very narrow. I'eriostracnm shining- on tlie disc Init somewhat scaly and dull on the posterior slope.

Nacre white, shining and slightly iridescent ]X)steriorly. ^luscle scars well outlined. Hinge plate long- and narrow, liight valve with two pseudocardinal teeth, the innermost being the larger. In addition, there is a single and long, narrow, lateral tooth. Left valve with a single pseudocardinal tootli and two long: and narrow lateral teeth.

Length

Height

Width

53 mm.

42 mm.

20 nnn.

Ilolotype

51

44.5

22.5

Paratype

57

50.0

25

Paratype, Lake ^lurray, Flv River

Types. The holotype is in the ^luseum of Comparative Zool- ogy, Xo. 212!)08, inland from Darn, Western Division, Papua. New Guinea. A single paratj^pe from the same locality is in the Australian Museum, Sydney. An additional paratype is from Lake JMurray, Herbert River, Fly River System. All specimens were collected by ^Ir. Kleckham of the Department of Agricul- ture, Papua.

This new species is related to Wcstralioiio tl!i( )isis Tapparone Canefri from the upper Fly River, Papua. It differs from W. ffyensis by being far more circular in outline and in being i)ro- portionately somewhat greater in width. In addition, the ventral margin is rounded, the posterior portion of the dorsal margin is somewhat wing-like and the posterior ridge is well defined.

This species is named for L. M. d'Albertis. a naturalist-ex- plorer who was in New (iuinea in 1872-73 and again in 1875. Much of the mollusk work accomplished l)y Tapinu-one Canefi-i was hascd upon iiuitei-ifd collected by d'Albei-tis.

REFEBENCES

Haas, F.

1924. Uiiserc hishciigcii Kenntnisso der X;i,);Ml('nf;iini;i Xcu-diiiiicns. Xnvn (iiiincii, 15: 65-7(), 1 i)l:ite.

Mc.Mkuaki., I). F.

19.16. Note.s on the t'lcsh watei- imissel.s of New (Uiinen. Xautihis. 70: 38-48, 1 plate.

n

BREVIORA

m of Comparative Zoology

Cambridge, Mass. June 21, 1957 Number 77

DACETINOPS, A NEW ANT GENUS FROM NEW GUINEA

By W. L. Brown, Jr.

and

E. 0. Wilson

In the course of an extended field trip to the southwestern Pacific area during 1954-1955 (supported by the Museum of Comparative Zoology and the Society of Fellows of Harvard University), one of us (Wilson) was able to collect in the rich tract of rain forest at the Busu River, near Lae, Australian Mandated Territory of New Guinea. Among- the samples ob- tained here were several of a small myrmicine ant with the habitus, both in life and in preservation, of certain short-mandib- ulate members of tribe Dacetini. However, further examination revealed that this species could not be a member of the Dacetini, even though it possessed certain features, especially the spongi- form appendages of the petiole, postpetiole and gaster, not otherwise known among ants outside the Dacetini. This remark- able pseudo-dacetine is described below and its affinities dis- cussed.

DaCETINOPS gen. nov.

Diagnosis, worker. Small myrmicine ant, apparently closest to Lachnomyrmex Wheeler of the American tropics. Body com- pact, integument hard and thick. Head subtriangular (with mandibles triangular), widest across occipital region, tapering toward mandibles. Antennal scrobes present, deep and distinctly

2 BREVIORA XO. 77

bounded; compound eyes moderate in size, situated ventrad of scrobes near midlenp^th of head. Antennae 11-segmented, with a fairly distinct 8-segmented club. Both sets of palpi 2-se<i- mented. Clypeus convex, obscurely bicariuate, its anterior border forming- a broadly rounded translucent apron. Mandibles triangular, broad, with straight, crenulate masticatory margins. Alitrunk compact, arched and sutureless dorsally ; propodeal dorsum and declivity almost continuous; a pair of small but distinct propodeal teeth present. Petiolar node sessile (a small condylar extension is visible from above), rounded above, and sloping from a median summit ; postpetiole transversely ellipsoi- dal, rounded above. Gaster broad and somewhat depressed ; first segment taking up nearly all the length, nearly circular in out- line as seen from above, bluntly margined along the sides in front; remaining segments small, retracted. Sting sclerotized, acute, exsertile. The lower halves of the petiole, postpetiole and anterior part of the first gastric segment bearing and covered i;y bulky festoons of whitish spongiform tissue. Sculpture con- sisting predominantly of very coarse longitudinal costation, be- coming more reticulate on head, nodes and sides of alitrunk : posterior half of gaster smooth and shining, as is also the pro- podeal declivity. Mandibles striate. Legs and antennal scrobes finely and densely punctulate. Pilosity consisting of sparse, long, flexuous tapered hairs, mostly situated at definite bilateral positions over the body. Pubescence forming a thin cover mostly only on appendages. Color deep brownish red.

Female. Dealate, but with thoracic flight segments well de- veloped, as usual for winged myrmicines of small size ; wing stumps present. Size only slightly larger than worker ; form, except as mentioned, similar to that of worker, with only the usual slight dilTerences of caste. The compound eyes are larger than in the worker, but are still modest in size. Scutum and scutellum forming together a nearly plane platform, crossed by the distinct transseutal suture ; prescutellum reduced to two small lobes, one on each side of the suture, so that the scutum and scutellum meet at the suture. Scuto-scutellar platform evenly costate longitudinally.

Type of genus : Dacetinops cibdela sp. nov.

1957

DACETINOPS, A NEW ANT GENUS

LAB

Dacetinops cihdeln, geu. et sp. uov., worker. Fig. 1, holotype, head in full-face view; most of sculpture omitted, and only the hairs nearest the periphery shown. Fig. 2, holotype, body in lateral view; hairs and all sculpture except the prominent dorsal costation omitted. Fig. 3, paratype,. detail of maxillary and labial palpi. Drawings by Nancy BufiBer.

4 BREVIORA NO. 77

Dacetinops cibdela sp. nov.

Holotype worker. TL 2.7, HL 0.64, HW 0.59 (CI 91), ML 0.16 (M2i5), WL 0.76, petiole L 0.22 mm. Index of cephalic depression (ICD) is about 62. Measurements and indices are the ones used in recent works on Dacetini by Brown (see especially Brown, 1953, Amer. Midi. Nat., 50: 7-15).

Form of head and mandibles as shown in Figures 1 and 2. Scrobes sharply defined, each divided partially by a fine anterior carina, and able to receive the entire antenna except the apical part of the club ; upper scrobe margins narrowly lamellate. Pronotum margined in front, submarginate along the sides ; liumeral angles obtuse, but distinct, subtuberculate. Propodeal region of alitrunk short, tapering rapidly liehind promesonotum ; propodeal declivity weakly concave from side to side, bounded above by an ogival margin, and marginate laterally. Propodeal teeth acute, feebly downcurved toward their apices, their bases separated by about 4 times their length. Propodeal spiracles not far under the teeth on the lateral margins of the declivity, facing posterolaterad. Petiolar node seen from above approxi- mately as broad as long, with strongly rounded sides, widest near midlength. Postpetiole about half again as broad as long, ellipsoidal or subreniform as seen from above, the more strongly rounded margin posterior. Gaster with a short, straight anterior margin opposite the postpetiole, the sides rounded away on each side.

Spongiform appendages of petiole, postpetiole and gaster finely areolate, apparently composed of cuticular material, not ar- ranged in distinct lobes like those of the dacetines, but narrowed medially below, where masses of opposite sides are joined (Fig. 2).

About 10 coarse, fairly distinct longitudinal costae across the head between the ends of the scrobes, these converging and anastomosing anteriad between the frontal lobes, and joined by short transverse ridges to form a coarse reticulum (not shown in the figures). The bottoms of the large pits or foveolae thus enclosed are sculptured finely and are subopaque. Disposition of costulae on clypeus shown in Figure 1 ; interspaces here finely sculptured and subopaque. Underside of head coarsely rugo- reticulate. Mandibles coarsely striate at base, becoming smooth

lf)57 DACETINOPS, A NEW ANT GENUS 5

and shining apiead, with scattered punctures. Dorsum of ali- trunk with smooth, straight, heavy longitudinal costae running from anterior pronotal margin to the beginning of the propodeal declivity (10 costae across pronotal dorsum), shining and with more or less shining interspaces. Sides of alitrunk and dorsal surface of petiolar node coarsely rugo-reticulate. Basal half of postpetiole and of gaster with coarse longitudinal costae (9-10 across gastric base), remainder of gaster, postpetiole, declivity and concave lateral faces of propodeum smooth and shining, often with scattered punctures. Legs, antennae, scrobes and a band along each side of the first gastric sternum densely punc- tulate, opaque.

Long, fine, tapered hairs scattered over dorsum of head (those nearest the cephalic borders are shown in Figure 1), alitrunk, both nodes and gaster as well as a few on the gular surface of the head, on the coxae and on the underside of the femora ; length ranging about 0.12-0.19 mm. Short, fine, appressed to decumbent hairs forming a pubescence on the legs, antennae and sternum and apex of gaster; similar hairs are sparsely dis- tributed over both surfaces of head, mostly one hair to a foveola, and over mandibles.

Basic body color deep brownish red ; mandibles, legs and antennae, especially the club and the first funicular segment, more yellowish.

Holotype worker (deposited in the Museum of Comparative Zoology) taken in rain forest at the lower Busu River, near Lae, New Guinea (E. 0. Wilson leg., No. 1058), May 17, 1955, either as a stray or in berlesate from under bark of a large Zoraptera- stage log. Three additional workers (paratypes) were taken in this collection and bear the same number.

Paratypes, workers. 9 mounted dry, 3 in alcohol, all collected in the same tract of forest as the holotype (Wilson Nos. 899, 942, 978, 1052, 1058, 1113). Deposited in Museum of Comparative Zoology, U. S. National Museum, Coll. 6. C. Wheeler, and one or more each in Australian and European collections as yet un- selected. Total maximum variation for all series: TL 2.2-2.6, HL 0.54-0.63, HW 0.50-0.57 (CI 91-93), ML 0.15-0.16 (MI 24-27), WL 0.65-0.76, petiole L 0.18-0.22 mm. Greatest intranidal varia- tion occurs in series No. 1052: TL 2.2-2.5, HL 0.54-0.62, HW

6 BREVIORA NO. 77

0.50-0.56 (CI 91-92), ML 0.15 (MI 25-27), WL 0.65-0.75 mm. Very little variation among the workers of these lots. Sculpture, particularly of cephalic dorsum, varies in minor details ; spongi- form appendages a trifle more voluminous in some specimens than in others.

Paratypes, dealate females. 3 specimens, taken with workers (Nos. 942, 1052, 1113, the last in alcohol, not measured). De- posited in the Museum of Comparative Zoology and elsewhere with the workers. Queen variation (Nos. 942 and 1052) : TL 2.6-2.8, HL 0.58-0.62, HW 0.56-0.60 (CI 96-97), ML 0.17-0.19 (MI 29-30), WL 0.79-0.85 mm.

Larva. A single medium larva was preserved in alcohol. This specimen is short and thick, with head turned ventrad. Without proper preparation, none of the details of this larva can be made out, except that the hairs are varied in length, with some of the longer dorsal ones anchor-tipped. This larva has been sent to Dr. G. C. Wheeler for expert study (See Breviora No. 78).

Biology. The six collections were all made on the forest floor in heavy to medium rain forest. Nos. 978 and 1052 were strays taken from soil-leaf litter berlesates. No. 899 was a stray sample from the forest floor, beneath a log. No. 942 was a small nest in a cavity under the bark of a Zoraptera-stage branch about 6 cm. in diameter, buried in the leaf litter ; the queen and two workers were taken. No. 1113 is a queen with one worker, originally taken with a few larvae and two eggs in a small cavity in the middle of a small piece of rotten wood buried in the leaf litter. This apparently incipient colony was kept in an artificial nest for 10 days, but showed no signs of predatory or nest-founding behavior during that time. All of the collections were made during the first three weeks in May, 1955.

From these observations, we may perhaps conclude that D. cibdela is normally a dweller in the leaf litter of the rain forest floor, where it frequently, perhaps usually, nests in or beneath the bark of rotting logs or other large or small masses of rotting wood. Apparently the nests are small in volume and in popula- tion, like those of other specialized small myrmicines with simi- larly slight differences separating the female and worker castes. Wilson noted that the workers of this species walk in a slow, deliberate fashion reminiscent of dacetines and basicerotines.

1957 DACETINOPS, A NEW ANT GENUS 7

Relationships. Although from its general habitus this species seems to belong to the Dacetini, a closer look does not bear out placement with that tribe. In the first place, its resemblance is general, and not particular to any one dacetine genus or sub- tribe. The shape of the head and mandibles are fundamentally different from those of any dacetine, as is also the arrange- ment of the spongiform appendages, which at first sight are so dacetiue-like. The antennae are 11-segmented, like those of the primitive dacetines, but the distinct club is not a dacetine fea- ture; the palpi of Dacetinops are segmented 2, 2, against 5, 3 for the primitive dacetines and 1, 1 for the higher dacetines. The details of structure of the alitrunk and petiolar node are also not like those of any dacetine, and the sculpture recalls that of the dacetines only at the gastric base, but even here, the costulae are of a basically different type. Ties with the Basicerotini are even fewer, the position of the eyes being one important difference ; there seems to be no need to compare further with basicerotines.

As already mentioned, the closest similarity holds with the neotropical genus Lachnomyrmex Wheeler (with three described species). The best account of Lachnomyrmex is that of M. R. Smith (1944, Proc. Ent. Soc. Washington, 46:225), which gives excellent figures of the genus. Lachnomyrmex also has an 11- segmented antenna, but the club is 2-segmented. This difference in club segmentation is really not a very important character, since only a slight change in size of the antepenultimate seg- ment would be needed to make the Lachnomyrmex club 3-seg- mented. Lachyiomyrmex has no posterior expansion of the head, as in Dacetinops, and the form of the node is somewhat dif- ferent, as well as the proportions of the parts of the alitrunk. but the differences here are not as serious as those between Dacetinops and the dacetines. Dacetinops appears to belong, with Lachnomyrmex, to a group of small myrmicines that in- cludes also the New World Kogeria Emery (the so-called Rogeria from the Melanesian area are not true members of the genus, but are closer to Lordomyrnia), Apsychomyrmex Wheeler, and Adelomyrmex Emery, as well as the primitive Agroecomyrmex Wheeler of the Baltic Amber. Apparently these are specialized relicts of an ancient and widespread myrmicine fauna that still retain some marks of their eetatommine ancestry.

BREVIORA

Mmseiiim of Cooiparsitive Zoology

Cambridge, Mass. June 21, 1957 Number 78

THE LARVA OF THE ANT GENUS DACETINOPS BROWN AND WILSON

By George C. Wheeler aud Jeanette Wheeler

Department of Biology University of North Dakota

Genus DaCETINOPS Brown and Wilson

LARVA

Body hairs sparse, of two types; (1) with short-bifid tip and (2) anchor-tipped, with sinuate shaft. Cranium subcordate. Antennae minute. Head hairs few (about 26), with short-bifid tip. Posterior surface of labrum with numerous spinules. Man- dibles long and slender; apex forming a long sharp-pointed tooth which is curved medially ; with a narrow blade projecting medially from the anterior surface and bearing two long coarse medial teeth. Maxillae small, with the apex paraboloidal ; palp a stout peg; galea digitiforra. Anterior surface of labium with numerous spinules. Hypopharynx spinulose and with numerous longitudinal ridges near the pharynx.

Dacetinops cibdela Brown and Wilson

(Text figure 1)

Immature Larva. Length through spiracles about 1.25 mm. Short and stout; prothorax directed ventrally; head ventral; anterior end formed from the dorsum of the mesothorax; dorsal profile C-shaped; abdominal somites II and III produced ven-

BREVIORA

NO. 78

trally into rounded transverse welts. Anus ventral, with a small posterior lip. Spiracles small, mesothoracic slightly larger than the rest. Integument apparently without spinules. Body hairs sparse and uniformly distributed, of two types: (1) 0.036-0.15 mm. long, slightly curved, with short-bifid tip, some on each somite, longest on the thorax and ventral surface; (2) anchor- tipped, with sinuate shaft, about 0.16 mm. long, four in a row across the dorsum of each abdominal somite II-V. Cranium subcordate in anterior view, clypeus bulging. Antennae minute,

Text figure 1. Dacetinops cibdela Brown and Wilson, larva. A, head in anterior view, X93 ; B, left mandible in anterior view (stippled to show thickness), X271 ; C, left mandible in posterior view (shaded to show con- tours), X271 ; D, immature larva in side view, X54; E, labrum (left half of drawing in posterior view, right half in anterior view), X271; F, hypopharynx in anterior view, X271 ; G, anchor-tipped body hair, X185; IT ;ind T, two bifid-tipped body hairs, X185.

1957 LARVA OF THE ANT GENUS DACETINOPS 3

each with three sensilla, each of which bears a minute spinule. Head hairs few, 0.036-0.09 mm. long, slightly curved, with short-bifid tip. Labrum short, wide and bilobed ; each lobe with about 13 sensilla on and near the ventral border; posterior sur- face of each lobe with a central cluster of 4 or 5 sensilla, lateral to which the spinules are large, isolated and sparse, but medially and dorsally the spinules are much more numerous, smaller and arranged in short transverse rows which are grouped into longer rows. Mandibles heavily sclerotized, rather narrow and elongate, subtriangular in anterior view; with one large apical tooth which is curved medially and posteriorly; with a median blade arising from the anterior surface and bearing two large medial teeth. Maxillae small, apex with a few short encircling rows of minute spinules; palp a stout peg with five apical sensilla (three minute and bearing a spinule each, two larger and en- capsulated) ; galea digitiform, with two apical sensilla. Anterior surface of labium spinulose, the spinules numerous and in short arcuate rows, the rows arranged in a longer subtransverse pat- tern, the spinules longer ventrally ; palp a low knob with five apical sensilla (three minute and bearing a spinule each, two larger and encapsulated) ; opening of sericteries in a wide de- pression on the anteroventral surface of the labium. Hypo- pharynx spinulose, the spinules numerous, minute and arranged in subtransverse rows; numerous longitudinal ridges near the pharynx. (Material studied : one damaged larva from New Guinea, collected by E. 0. Wilson; courtesy of W. L. Brown.)

A single damaged immature larva is a frail foundation on which to base the taxonomic relationships of a genus. Neverthe- less we must hazard a few guesses, for that is all the material we have. Such guesses are the more difficult because of the nearly diagrammatic structure of this larva. If only it had a few (or even one) striking peculiarity something overdeveloped or something lacking. If we extrapolate the mature body shape, enlarge the antennae to normal size, inflate the bases of the mandibles a little and reduce somewhat the size of the medial mandibular teeth, the result might be considered the synthetic type for the subfamily Myrmicinae.

AVe are now in the process of synthesizing our studies of the larvae of the Myrmicinae. Consequently we do not yet have clear

4 BREVIORA NO. 78

ideas as to what characters are phylogenetically basic. However, we do feel that a few provisional opinions about the relation- ships are not out of order.

The larva of Dacetmops resembles the larva of Rog&ria so closely that the same generic description would apply to either except for the hairs : Rogeria lacks the anchor-tipped hairs which we suspect will prove to be of basic phylogenetic importance. The larva of Dacetmops also resembles the larvae of the tribe Basicerotini but differs in having anchor-tipped hairs and by lacking spinules on the mandibles. The worker of Dacetinops is convergently similar to the Dacetini, but its larva is not. In fact, the larva of Dacetinops shows no close affinity with any known genus.

BREVIORA

Muiseimm of Coimparative Zoology

Cambridge, Mass. August 9, 1957 Number 79

DASYPELTIS MEDICI LAMUENSIS, A NEW RACE OF

EGG-EATING SNAKE (OPHIDIA, REPTILIA) FROM

COASTAL EAST AFRICA

By Carl Gans

Museum of Couii);iiativc Zoology and Carnegie Museum, Pittsburgh, Pa.

Some time ago an analysis of the African egg-eating snakes of the genus Dosypeltis indicated the presence of an undescribed race from coastal East Africa. However, as the entire revision of this genus will not be published for several months, I am descril)ing the new' race here in order that Mr. Arthur Loveridge may include it in his checklist of the reptiles and amphibians of East Africa, which is now in press.

In 1942, Loveridge revived Dasypelfis medici as a subspecies of the wide-ranging D. scahra. Examination of additional speci- mens from supplementary localities convinced me that two forms are involved, and furthermore that D. scahiri and medici, though sympatric in many areas, are distinct species which, besides their color patterns, show differences in a number of other characters. Among these are ventral and caudal counts of both male and female species, body proportions, etc.

In his 1942 paper Loveridge mentioned in passing that north- ern specimens of medici were unicolored and did not possess the characteristic color pattern upon the basis of which this species was initially described by Bianeoni. The ''uniform" coloration has been found to be associated with significantly lower ventral counts and the northern specimens are here recognized as a distinct race.

The detailed acknowledgments to the many who helped with the analysis Avill be given in the main paper. Here I restrict myself to thanking Ernest E. Williams and Arthur Loveridge

BREVIORA

NO. 79

of the Museum of Comparative Zoology (MCZ) for checking this paper, the John Simon Guggenheim and National Science Foundations for supporting the research upon wliicli this paper

Figure 1. Dasypeltis medici. Dorsal views showing pattern atop head. Top. D. m. medici BM 12-1-30-6 from Matemo, Mozambique. Bottovi. I), m. lamuensis BM 51-1-3-73 from Kilifi, Kenya. Note that the pattern is almost completely faded out on the parietal scales of lamuensis, and has disappeared in the nuchal region. The pustulosity or pitting on the head shields and the dark pigmentation of the pits is clearly apparent.

I9;i7 DASYPELTIS MKDKI LAMUENSIS 3

is based, and i\lr. J. C. Battersby of the British Museum (Natu- ral History). London (BM), as well as Dr. Heinz Wermuth of the Zoolooisehen Museums der Universitat, Berlin (ZMTH, for

the loan of paratype material.

DASYPELTIS MEDICI (Bianeoni), 1859

1. rattern, (>onsisting of narrow lateral liars (of which at least the first liirce to eight fuse to form narrow Vs on the nape), extends the entire length of snake from the head on to the tail; ventrals more than 229

in males, more than 236 in females D. m. medici.

2. Pattern, if present, restricted to top of head (see Fig. 1) ; ventrals 229 or less in males, less than 233 in females D. m. lamucnsis.

Dasypeltis MEDICI MEDICI (Bianconl), 1859

Dip.su.s Medici Bianeoni, 1859, p. 277. No locality designated, luit

Mozambique b.v inference. Type in Bologna Museum. Dasiipeltis scaher var. fasciolata Peters, 1868, p. 451. Type (ZMU 5737)

from " Angeblich aus Zanzibar". Dasypeltis elongata Moequard, 1888, p. 131. Type locality: Zanzibar. Type in Museum d 'Histoire Naturelle, Paris. Diagnosis. Three to eight narrow Vs commencino- on the nape, followed by a series of narrow, lateral, dark red-brown bars. These bars wall encircle a pink to silvery-white vertebral dot, if they coalesce with their fellows from the opposite flank. In some specimens the posterior ])ands are situated betw^een more or less clearly expressed oval dorsal saddles, while various intermediate patterns (see Fig-. 2) have also been observed. The dorsum is a light reddish-brown, which shows a considerable amount of mottling under the binocular microscope. The apical scale pits, as well as those of the head region, are a dark brow-n, this pig- mentation being found on all scales and very sharply set off. The ventrum is a clear pink, more or less regularly stippled with grey. Beneath the tail of many specimens, particularly^ in males, this stippling is arranged in two to four lines. In other individuals, hoAvever, the distribution of the stippling appears to be haphazard. Three lateral scale rows are definitely reduced and inclined, and they, as well as some of the adjacent rows, generally have strongly serrated keels. The frontals shoAv pitting or pustulosity over their entire surface as do most of the other cephalic scales (see Fig. 1). The inter-prefrontal suture is not

BREVIORA

NO. 79

depressed. Veiitrals in males 235-253, in females 237-259 ; eau- dals 82-109 and 71-80, respectively.

Range. Coastal East Africa. Extreme southern Kenya, Tan- ganyika, northern and central INIozambiqnc ; inland to Xyasaland.

Figure 2. Dasypeltis m. medici. Dorsal views at midbody showing color pattern variants of this race. Top. BM 12-1-30-6 from Matemo, Mozam- bique. Bottom. BM 97-6-9106 from between Nkata Bay and Ruarwe, Xyasaland. Note lighter middorsal spots between the saddles, the regular pigmentation of the apical scale pits, and the general speckling of the ground color.

1 ;).■)( DASYFELTIS MEDICI LAMUENS18 5

Disfribidio)! records. (Maj) 1 shows Die relation of these localities to eaeh other. Records from the literature are starred.) KENYA: Takannon. Between Voi and Ndi, Taita. TANGAN- YIKA: Zanzibar Coast. Kibonp:oto' Kilimanjaro ( Intergrade"? specimen^. Amani, Usamljara Mts. Usambara. Makindn Kiver. Morogoro. Nyange, lUugurn Mts. Northern Ukutu (Kuthu) Steppe. Tendaguru. Mikindani. Nchingidi. Liwale. ZANZI- BAR. JMAFIA. MOZAMBIQUE: Matemo. Ribaue, Nyassa Prov. NYA8ALAND: Between Nkata Bay and Rnarwe. Zomba. Cholo Mtn.

DaSYPELTIS MEDICI LAMUENSIS Subsp. IIOV.

Uasyiniiis pahnarum I'etors (not of Loach), 1878, p. 20G. 1 ex. Taita,

Kenya (J. M. Hildebrandt) : ZMU. DasypcUis scahra var. F (part), Boulenger, 189-4, p. 356. 1 ex. Mt. Kili- manjaro, Tanganyika (F. J. Jackson) : BM. Dasyprltis scahrr Uthnioller (part), 1934, p. 113. 1 ex. nr. Goniberi, Kili- manjaro, Tanganyika (Uthmoller) : Zool. Staatssanimlung, Miinchen. DasypcUis scahcr Loveridge (part), 1936, p. 256. 1 ex. Mt. Mbololo; 2 ex.

Lamu Island, Kenya (Loveridge) : MCZ. DasypcUis scahcr Scortecci (part), 1939, p. 276. 1 ex. Belet Aniin, Ital. Somaliland = Somalia (S. Patrizi) : Genoa Civ. Mus. Stated to be nnifonn grey dorsally. DasypcUis scahcr mcdici Loveridge (part), 1942, p. 283. Mention of uni- formly colored northeastern material : MCZ. Diagnosis. Uniform reddish-brown dorsally, fading to butf in some specimens after preservation (Loveridge). Pink ventrally with a fine speckle of a slightly darker pinkish-brown denser on the sides (Fig. 4, bottom). Some specimens are a uniform olive grey, fading to a plumbeous grey after preservation. These have a light grey ventrum, minutely flecked with white. A vague or distinct mottling of the ground color may be seen under the binocular microscope (see Fig. 4, top). Apical pits of body and marginal pits of head scales distinctly pigmented with a darker reddish-brown (see Fig. 1). With proper illumination faint to clear V-shaped markings can be discerned on the parietal scales of some specimens. Three lateral scale rows are distinctly reduced and inclined, and they, as well as several of the adjacent rows,

1 The status of this specimen is discussed helow and its data are not included in the above ranges tor tliis subspecies.

BREVIORA

NO. 79

Figure 3

1957 i\\s\i'i;i/ris medici hAMUENSis 7

generally have strongly serrated keels. Each frontal scale shows pnstulosity or pitting aroniul tlie ])(M'ipluM'v and in the center. The suture lietwcen the pi'efroutals is not depressed. V^nitrals in males 226-22!), in females 22G-232; caudals 84-94 and 72-84, respectively.

Holoti/pc. jMuseuni of Comparative Zoology No. 40582, an adult male from Lanui Island, Ken\a, collected by Arthur Lovei-- idge 12 May 1934.

Allotype. Museum of Comparative Zoology No. 40588, an adult female collected with the type.

Paratypes. British Museum (Natural History) Nos. 51-1-3-72 and 51-1-3-73 from Kilifi, north of Mombasa, Kenya. Also Brit- ish Museum (Natural History) No. 98-1-8-15 from Maungu, near Voi, Kenya ; iMuseum of Comparative Zoology No. 40580 from Mt. Mbololo, Kenya ; Zoologisches Museum der Universitat (Berlin) No. 9244 from Taita (region), Kenya; and British Museum (Natural History) No. 87-11-3-31 from Mt. Kilimanjaro, Tanganyika.

Description. Both the Lamu holotype and allotype are a uni- form brownish-red dorsall}^ and a speckled pink on the venter. Under proper illumination a very faint, posteriorly-directed V may be discerned on the parietal scales (see Fig. 3, top). Scale pits on head and body with sharply defined darker brown pig- mentation, though this is only slightly darker than the dorsal color (Fig. 3, bottom) and not as clearly visible as on lighter specimens (Fig. 4, middle). The frontals are pitted around their margins and pustules are also present in their centers, with the other head shields showing a similar pattern. The suture between the prefrontals is not depressed, the ocular-temporal formula is l+2-(-2-|-3, and the upper labials and eye contact are 7(34). Counts of ventrals are 226 for both type and allotype, caudals are 94 and 84, dorsals 23 and 24 at midbody respectively. The third, fourth and fifth rows of dorsal scales from each side are reduced, inclined and serrated, and the keels of the dorsal

Figure 3. Dasypellis m. hnniiensis. Views of the holotype (MCZ 40582). Top. Dorsal view of head. Middle. Lateral view of head. Bottom. Dorso- lateral view of specimen in the midbody region. Note the very faint pattern on the parietals, and the fact that the prefrontals, oculars, anterior labials, etc. are covered with pigmented pits.

8

BREVIORA

NO. 79

Figiiie 4

1957 DASYPELTIS MEDICI LAMUENSIS 9

scales are serrated around the body in the anal ref^ion of the type, hut not of the pai-atype. Body and tail lengths are 498+ 132 nun. and 570-|-128 nnn. respectively .

Range. Coastal East Africa from Somalia soutli to the Kenya- Tanganyika frontier.

Disiribufion records. SOMALIA: Belet Amin (Scortecci, hm). KENYA: Lamu Island (Loveridge, WSd) ; MCZ 40582, 40583. Kilifi, north of Mombasa. BM 51-1-3-72, 51-1-3-73. Mauno-u near Voi. BM 98-1-8-15. Mt. Mbololo (Loveridge, 1936); MCZ 40580. Taita (Peters, 1878); ZMU 9244. TAN- GANYIKA: Mt. Kilimanjaro (Bouleno-er, 1894) ; BM 87-11-3-31. Near Gomberi, Mt. Kilimanjaro (Uthmoller, 1934).

Discussio)t. The map shows the relative position of collectino' localities for both patterned and unicolored forms of D. medici. The species appears to be restricted to the coastal regions below 1000 meters, generally characterized by their reddish laterite soils (Loveridge, 1942) . As may be seen from the map, all but two of the Kenya specimens are definitely unicolored. Pattern is present in the Voi-Ndi juvenile, but the Takaungu individual is an almost completely faded adult, which lacks even the pigmenta- tion of the apical pits so characteristic of the species. Two of the three specimens from the vicinity of Mt. Kilimanjaro (seen, or reported on in the literature) were unicolored, while the dorsal pattern of the third is faded out posteriorly. All other speci- mens and records from Tanganyika have, or are said to have, a color pattern.

Of the other characters examined only ventral counts show elinal variation. Figure 5 demonstrates these counts for male and female specimens of the two color phases, showing the sharply

Figure 4. Dasypeltis ?h. lamucnsis. Views of the midbody region of different specimens to demonstrate certain characteristics of the pigmenta- tion. Top. BM 51-1-3-73 from Kilifi, Kenya. Lateral view of dark grey specimen, showing the extensive speckling of the ground color, which almost masks the pigmentation of the apical pits. Middle. BM 98-1 8-15 from Maungu, Kenya. Lateral view of light colored specimen which clearly shows both pigmentation of apical pits and lateral scale arrangement. Note that the ground color is still speckled. Bottom. BM 51-1-3-72 from Kilifi, Kenya. Ventrolateral view of specimen showing color invasion and speckling of light colored ventrum.

10

BREVIORA

xo. 7!)

LAMU ISLAND

MT. KILIMANJARO Q^ CY^ <^

I ® Okilifi VaitaO Y"^ka"ngu

USAMBARA ^ AMANI

MAKLNDU^

MOROGORO 0

NYANGE0 UKUTU*

ZANZIBAR ISLAND

MAFIA ISLAND

KILWA ( TENDAGURU(

LI WALE #

NCHINGIDI V •MIKINDANI

Jjasi/pt'ltis medici. Map showing loc-alities for all specimens acliiaUy examine.! as well as for the literature record from Belet Amin. Solid dots stand for records of m. medici, open circles for records of m. lamuoi.si.s. Divided circles refer to the paiis of immediately adjacent records discussed in the text.

195';

DASYPELTIS MEDICI LAMUENSIS

11

distinct ranfjcs. The only exception is furnished by the faintly patterned specimen from Kibonjroto' Kilimanjaro. This has a ventral count of 218, i.e. 8 less than the lowest count for a unicolored specimen and 19 less than the lowest count for a patterned specimen.

In view of the fact that Ihe two populations replace each other jreographically, that they ditfer in two characters which break in the same zone, and that there exists a possibly intermediate speci- men from a geographically intermediate region, they are here considered to be subspecifically distinct. This decision has been

MALES

lamuensis

me

FEMALES

lamuensis

medic

220 230 240 260

V E N T R A L S

Figure 5. Dasypcltis vtcdici. Graph showing ventral counts of all speci- mens of the two races actually examined. Note the anomalous count of the Kibongoto specimen.

12 BREVIORA NO. 79

consciously influenced by a desire to focus attention upon this population in the hope that this will permit re-examination of the matter on the basis of more nearly adequate collections.

It has been suf>'0'ested that this situation represents two adja- cent species and that the patterned Kilimanjaro specimen is a hybrid. While the material is insufficient to decide the point, there is some evidence against this idea. Thus both of these color patterns feature a sharply defined pigmentation of the apical scale pits and are the only forms within the genus that have this characteristic. Besides which, 45 out of 46 specimens of the combined sample have frontals that are entirely, rather than marginally, pustulated or pitted, a frequency not approached by any other population, with the exception of Dasypcltis scahra from South Africa. In view of these facts and in the absence of more detailed data, the two forms are considered to belong to a single polytypic species.

LITERATURE CITED

BiANCONi, J. Joseph

1850- Specimma Zoologica Mosambicana. Bononiae, fasc. 1-1(5, ii 1862. +406(?) pp., (fasc. 14 (1859), pp. 277-278, tab. 14). First

published in parts in Mem. R. Accad. Sci. Inst. Bologna. (Vol.

10, pp. 501-2, pi. 26.)

BOULENGEK, GEORGE ALBERT

1894. Catalogue of the snakes in the British Museum (Natural His- tory). Volume II, containing the conclusion of the Colubridae Aglyphae. London, xi -[-382 pp. (353-357).

LovERiDGE, Arthur

1936. Scientific results of an expedition to rain forest regions in

Eastern Africa. V. Reptiles. Bull. Mus. Conip. Zool., vol. 79,

no. 5, pp. 209-337 (256-57, pi. 4, fig. 1). 1942. Scientific results of a fourth expedition to forested areas in

East and Central Africa. IV. Reptiles. P>ull. Mus. Com]). Zool.,

vol. 91, no. 4, pp. 237-373 (282-86).

MocQu.vRD, Francois

1888. Sur une collection de reptiles et de batraciens rapportes des Pays Comalis et de Zanzibar par M. G. Revoil. Mem. pui)!. Soc. Philomat. Occas. Centcnnaire Foundation, i)p. 109-134 (131, plate XII, figs. 2, 2a, 2b, 2e).

1957 DASYPELTIS MEDICI LAMUENSIS 13

Peters, Wilhelm Carl Hartwig

1868. liber eine neuc Nagergattung, Chiropoilowi/.s prninllatus, so

wie iiber ciiiigo neuc odcr weniger bekannto Ainphibien und

Fisehe. Monatsbcr. kgl. Akad. Wiss. Berlin, pp. 448-461 (451). 1878. Ubcr die von Hrn. J. M. Hildebrandt wiihrend seiner letzten

ostafrikaniscluMi Reise gesanimelten Saugetliiere und Aniphibien.

Mouatsber. konig. Preuss. Akad. Wiss. Berlin 1878, pp. 194-209

(206).

ScoKTEoci, Giuseppe