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Full text of "Bulletin of the Museum of Comparative Zoology at Harvard College"

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HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



(US ISSN 0027-4100) 



auUetin of the 

Museum of 

Comparative 

Zoology 



^« / 2 1995 

Types of Shelled Indo-PaGffj^'iVibtlusks 
Described by W. H. Pease 



RICHARD i. JOHNSON 



HARVARD UNIVERSITY 

CAMBRIDGE, MASSACHUSETTS, U.S.A. 



VOLUME 154, NUMBER 1 
/DECEMBER 1994 



(US ISSN 0027-4100) 



PUBLICATIONS ISSUED 

OR DISTRIBUTED BY THE 

MUSEUM OF COMPARATIVE ZOOLOGY 

HARVARD UNIVERSITY 



Breviora 1952- 

BULLETIN 1863- 

Memoirs 1865-1938 

JOHNSONIA, Department of Mollusks, 1941-1974 

Occasional Papers on Mollusks, 1945- 

SPECIAL PUBLICATIONS. 

1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963. Phylogeny and 
Evolution of Crustacea. 192 pp. 

2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Tere- 
dinidae (Mollusca: Bivalvia). 265 pp. 

3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- 
derms. 284 pp. 

4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the 
Present Day. 236 pp. 

5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology 
and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 
725 pp. 

6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. 

Other Publications. 

Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. 
"Reprinted 1964. 

Brues, C. T, A. L. Melander, and F. M. Carpenter, 1954. Classification of 
Insects. {Bulletin of the M.C.Z., Vol. 108.) Reprinted 1971. 

Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. 

Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First Inter- 
national Symposium on Natural Mammalian Hibernation. {Bulletin of 
the M.C.Z.,Wol 124.) 

Ornithological Gazetteers of the Neotropics (1975-). 

Peters' Check-list of Birds of the World, vols. 1-16. 

Proceedings of the New England Zoological Club 1899-1947. (Complete 
sets only.) 

Proceedings of the Boston Society of Natural History. 

Price list and catalog of MCZ publications may be obtained from Publications 
Office, Museum of Comparative Zoology, Har\'ard Universitv, Cambridge, 
Massachusetts 02138, U.S.A. 

This piil)licati<)n lias been printed on acid-free pernianent paper stock. 
© The President and Fellows of Harvard College 1 994. 



I 



MCZ 
LIBRARY 

JAN 1 '^ 1995 

HARVARD 
UNIVERSITY 

TYPES OF SHELLED INDO-PACIFIC MOLLUSKS DESCRIBED BY 
WILLIAM HARPER PEASE (1824-71) 



RICHARD I. JOHNSON^ 
TABLE OF CONTENTS 

Abstract 1 

Introduction 1 

Remarks 4 

Acknowledgments 5 

Abbreviations 5 

List of the Taxa of Shelled Mollusks Described 

b\ Pease 5 

Additional or Corrected Locality Data 29 

Literature Cited 29 

Index 34 

Plates 42 

Abstract. Pease described some 500 species of 
mollusks from the Indo-Pacific region. Of these, some 
380 were shell-bearing. Type material was located 
for all but 42, consisting of syntypes, some previously 
figured or measured, which have hitherto been re- 
garded as holotypes or selected as lectotypes. Many 
lectotypes are chosen here and illustrated for the first 
time. All of the holotypes and lectotypes in the Mu- 
seum of Comparative Zoology are figured. 



INTRODUCTION 

The original William Harper Pease col- 
lection is in the Museum of Comparative 
Zoology, Harvard University. After Pease 
died in 1871, his collection was sent to 
Boston to be sold "entire." It was offered 
to a Mrs. Witthaus of New York for $3,000. 
In a letter to John Gould Anthony, Curator 
of Mollusks at the Museum of Compara- 
tive Zoology, she said that she had seen 
the collection and found it to be damaged 
by careless packing. She arranged to pur- 
chase only the specimens she wanted, the 
larger pretty shells (R. D. Turner, personal 



' Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge, Massachusetts 02L38. 



communication). The remaining collec- 
tion was sold to Louis Agassiz, director of 
the Museum of Comparative Zoology. Most 
of the types and smaller specimens were 
carefully glued on pieces of glass or slate 
for exhibition by curator Anthony. This 
process, as well as the formula for the glue, 
is included in Turner's (1946) definitive 
biography of Anthony. In this century, the 
shells were removed from the plaques by 
William J. Clench, cataloged, and num- 
bered. Each lot is usually accompanied by 
labels in Anthony's distinctive and accu- 
rate calligraphy as well as one penciled on 
a scrap of paper by Pease. There is a type- 
written list of the collection in the De- 
partment of Mollusks compiled by Wil- 
liam F. Clapp (1923, Dept. Library, no. 
5912), "copied from Pease catalogue writ- 
ten on loose sheets of stationery." Aside 
from the type material, and that collected 
by Andrew Garrett, most of the lots have 
locality data that are so general as to ren- 
der the specimens useless for study. The 
Witthaus material became the property of 
Vassar College, Poughkeepsie, New York, 
and was presented to the Museum of Com- 
parative Zoology in 1944. 

Solem (1976: 9) took such a dim view 
of the Pease collection that he virtually 
ignored it when monographing the En- 
dodontoid Land Snails from Pacific Is- 
lands, preferring to select lectotypes from 
any other collection that had Pease ma- 
terial, such as the National Museum of 
Natural History, Washington, D.C., or the 
Bernice Pauahi Bishop Museum, Honolu- 



Bull. Mus. Comp. Zool., 154(1): 1-61, December, 1994 1 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



lu, even when the material had been orig- 
inally received from the Museum of Com- 
parative Zoology as duplicates. He claimed 
that when the collection was shipped "from 
England [sic] to Harvard University, ap- 
parently there was extensive mixing of sets 
when cabinets were tilted and handled. In 
subsequent years (obviously referring to 
duplicates sent out from the Museum of 
Comparative Zoology rather than by 
Pease), these shells have been traded wide- 
ly to other museums and amateur collec- 
tors. Virtually all traded Pease material 
that I examined contained more than one 
species, living on different islands" (ibid.). 
Solem further declared that "Pease him- 
self was careless in handling his collection 
since in a letter to a correspondent [An- 
drew Garrett] wrote that his small daugh- 
ter delighted in playing with shells in the 
cabinets" (Alison Kay, personal commu- 
nication). Kay (1975: 18) had actually 
quoted Pease as having written, "Our little 
girl is growing and troubles me in my col- 
lections, by assorting my duplicates and 
arranging them in boxes." 

Pease began describing shells in the Pro- 
ceedings of the Zoological Society of Lon- 
don, often remarking that examples were 
in the collection of Hugh Cuming, who 
had died in 1865. The Cuming collection 
was acquired by the British Museum (Nat- 
ural History). Kay (1965) located all of the 
available types of the marine species, re- 
described them, selected lectotypes when 
necessary, and figured them, most for the 
first time. Many of those species not found 
by Kay were found in the Pease collection 
and are figured here. Syntypes of all but 
14 of the land and freshwater species de- 
scribed in the Proceedings were located in 
the Museum of Comparative Zoology. 
None of those missing could be located in 
the British Museum (Natural History) (P. 
B. Mordan, in litt.). Clench (1975) listed 
all of the Pease taxa but made no effort to 
supply any information as to the location 
of any of the type specimens, neither those 
in the British Museum (Natural History) 
nor those described in the Journal de Con- 



chyliologie, which were sent to H. Crosse, 
who translated the papers into French. Fi- 
scher-Piette (1950) listed all of the types 
that had been in the collection of the Jour- 
nal, and later presented them to the Mu- 
seum National d'Histoire Naturelle, Paris, 
figuring those not done so previously. 
Clench also did not notice that Baker (1963, 
1964), who listed all of the land and fresh- 
water types in the Academy of Natural 
Sciences of Philadelphia, included those 
by Pease described in the American Jour- 
nal of Conchology . 

After the death of Cuming, Pease chose 
P. P. Carpenter to sponsor his publications 
in the Proceedings of the Zoological So- 
ciety, but Carpenter (in Pease, 1865a: 675- 
676) included a list of synonyms based on 
specimens Pease sent to Cuming deter- 
mined by Cuming and H. Adams. Later 
(in Pease, 1865d: 516-517), Carpenter 
published a list of synonyms based on his 
own observations. This ended their rela- 
tionship. It was not until 1872 that another 
paper by Pease appeared in the Proceed- 
ings. In it were described 14 new species 
of Triphoris, none of which were sent to 
the British Museum (Natural History). 
Twelve of them were found in Pease's col- 
lection and are figured here for the first 
time. While the opisthobranch mollusks 
that are shell-less are not included in the 
following list, those that were studied by 
Pruvot-Fol (1947) have been noted in the 
accompanying index of all of the Pease 
taxa. 

William Harper Pease was born in 
Brooklyn, New York, in January 1824. He 
joined the Lyceum of Natural History of 
New York in 1841; traveled with General 
Winfield Scott to Mexico in 1848, his as- 
signment a matter of conjecture; arrived 
in Honolulu on December 11, 1849; and 
in 1850 applied for citizenship in the Ha- 
waiian Kingdom, purchased land, and be- 
came Local Agent of the Government on 
the island of Kauai. Little else is known 
about him to this time. Returning to Ho- 
nolulu about 1856, Pease became Com- 
missioner of Water Rights and Rights of 



MoLLUSKS Described by W. H. Pease • Johnson 



Way as well as Assessor of the City of Ho- 
nolulu, though Solem (1976-77) thought 
he was a merchant. His main interest was 
natural history, especially the shells of the 
Pacific. In a letter of 1865, quoted by Kay 
(1975: 6), he wrote, "That is all I think or 
care about." Prominent among his friends 
was Levi Haalelea, Chamberlain to the 
Court. He kept his shell collection, library, 
and an office in Haalelea's home opposite 
the Palace. When Haalelea died in 1865, 
Pease moved the library to a small building 
adjoining his own house at 25 Liliha Street. 

Most of the material that Pease de- 
scribed after 1860 was collected by An- 
drew Garrett (1823-87), whom Solem 
(1976: 7) mistakenly thought was a mis- 
sionary. Garrett was actually a self-trained 
naturalist, who had spent some of his life 
as a sailor before becoming a professional 
collector of shells, fishes, and other natural 
history objects of the South Pacific Islands. 
He arrived in Honolulu in 1852 and later 
moved to the island of Hawaii, where he 
remained, except for collecting trips to 
other Pacific Islands, until 1863 when he 
settled permanently in the Society Islands. 
While in the Hawaiian Islands he became 
a close friend and employee of Pease, and 
the two carried on an active correspon- 
dence until the death of the latter in 1871. 

Garrett collected for Louis Agassiz of 
the Museum of Comparative Zoology, for 
the Museum Godeffroy, Hamburg, and for 
Pease, who sponsored his trips in part and 
acted as his agent in Honolulu. He sent 
Pease descriptions of shells and nudi- 
branchs, which the latter used in his pub- 
lications. 



The relations of these two men was friendly, 
although Pease took almost a paternalistic attitude 
toward Garrett. There was a difficult side for Gar- 
rett. It is true that, in a sense, he was an employee 
of the other man, but in some ways Pease seems to 
have taken advantage of Garrett's abilities and ef- 
forts. The latter worked hard and assiduously. He 
collected amid the most trying and dangerous con- 
ditions. He scrupulously described and drew pic- 
tures of many of the specimens gathered, partic- 
ularly the fishes. In the case of fishes and shells he 
suggested the scientific names. But despite all this, 



he lived in the shadow of Pease, while the more 
articulate writer and more assertive man earned 
the glory. (Thomas, 1979: 20) 

After the death of Pease, beginning in 
1872 until his death in 1887, Garrett began 
publishing his own descriptions of new 
species and monographs of various groups 
of mollusks, which included much more 
accurate locality data than those given by 
Pease. 

Few details of the life of Pease are re- 
peated here. His biographer, Karl W. 
Greene (1960), based his account on the 
letters Pease wrote to Garrett, which, along 
with the latter's shell collection, are now 
in the Bernice Pauahi Bishop Museum, 
Honolulu. While details of the purchase of 
the Garrett collection are lacking, it was 
in the museum by 1899. Kay (1975) also 
wrote a biography of Pease. Although 
Greene (1960 (8): 12) mentioned Kay's ar- 
rival in Honolulu, and while she must have 
been aware of his articles in Hawaiian Shell 
News, Kay did not allude to Greene's work. 

The life and travels of Andrew Garrett 
are treated definitively by W. Stephen 
Thomas (1979), who devoted many years 
to the study. Thomas' interest in Garrett 
was inspired by his obtaining from some 
of his cousins over 250 drawings of fishes 
and shells that Garrett had made for Cap- 
tain John W. Leonard of the whaleship 
Lydia. Clench (1979) listed all of the mol- 
lusks described by Garrett but again did 
not make any effort to locate his type spec- 
imens, many of which are in the Academy 
of Natural Sciences of Philadelphia, the 
Museum of Comparative Zoology, and, of 
course, the Bishop Museum. 

Kay (1975: 18) mentioned that neither 
portrait nor photograph of Pease had ever 
been found, but that Lady Franklin, the 
widow of Sir John Franklin, the Arctic ex- 
plorer, met Pease at the plantation of Rob- 
ert G. Wyllie, a Scotsman who was Ka- 
mehameha IV's minister of foreign affairs. 
Lady Franklin wrote, 

I find it so difficult to make out what he says that 
much is lost to me — this proceeds partly perhaps 
from a want of some teeth in front of his mouth, 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



but chiefly from his holding and chewing tobacco 
which not only thickens his speech, but causes him 
to be constantly spitting. (Korn, 1958) 

Perhaps, it is just as well that no portrait 
was found. 

In spite of his appearance, Pease seems 
to have moved in the best circle of Ha- 
waiian society of the time. As already men- 
tioned, he was an intimate of Levi Haa- 
lelea, an acquaintance of Robert G. Wyllie, 
Minister of Foreign Affairs, and James 
Walker Austin, sometime Associate Justice 
of the Supreme Court of the Kingdom of 
Hawaii. Austin, a relative of my children, 
was executor of all of the preceding estates 
including that of Pease (Austin, 1921: 64). 

REMARKS 

For convenience the following list of taxa 
is arranged alphabetically by species name. 
It contains references only to those that 
are shelled: a complete list of molluscan 
names introduced by Pease was compiled 
by Clench (1975) and may be found here 
in the Index alphabetically by genus with 
supplemental data. In addition to the orig- 
inal reference, subsequent ones are also 
cited if they include the first figure of a 
type or other relevant information. More 
detailed locality information is included 
from subsequent references by both Pease 
and Garrett, the latter having collected 
most of the material. 

Pease put a question mark before some 
genera when he was not sure where the 
species belonged. A generic name in pa- 
rentheses is a second citation indicating 
that he later changed the generic place- 
ment. Data in brackets have been found 
on original labels, are additions or correc- 
tions from recent maps, or are comments 
by this author. No attempt has been made 
to discuss the present status of any of the 
taxa. This is a task for individual revisers. 

In the past, if Kay (1965), Fischer-Piette 
(1950), Baker (1963, 1964), Houbrick 
(1992), or the present author was able to 
locate the single figured or measured syn- 
type, it was usually regarded as the holo- 
type. Most of these designations, included 



in this paper, were made long before the 
promulgation of the third edition of the 
International Code of Zoological Nomen- 
clature (1985), which invalidates this prac- 
tice under Article 74 (b): ". . . should an- 
other syntype or syntypes be discovered 
[the first subsequent author is to be re- 
garded] to have chosen a lectotype." If this 
change is to be made it is left here, in these 
instances, to the first subsequent reviser. 

Baker (1963, 1964) listed all of the types 
of land snails in the Academy of Natural 
Sciences of Philadelphia and used his own 
system of abbreviations for type designa- 
tions, which have been interpreted here as 
follows: 

TOD type by original designation [ho- 
lotype] 

TOM type because only one example 
was included in the original de- 
scription, or was indicated by only 
one set of dimensions . . . [holo- 
type] 

TSD type by subsequent selection, fol- 
lowed by "now" if apparently first 
designated in these lists . . . [lec- 
totype] 

At the request of Dr. Gary Rosenberg, 
of the Academy of Natural Sciences of 
Philadelphia, all of the types, either fig- 
ured for the first time or refigured, are 
regarded as lectotypes in compliance of 
ICZN 74 (b). 

Unless previously selected as lectotypes 
by someone else, the types of the species 
of genus Parttila from the Society Islands 
have been included only as syntypes, be- 
cause so many were regarded as mere color 
forms by Crampton (1916, 1932) and be- 
cause most of these are now extinct (Tudge, 
1992). 

Pease introduced a number of names in 
the literature that are believed to be nude. 
When these were represented by speci- 
mens, the number of the lot is included. 
These specimens are of no significance un- 
less, peradventure, the taxa were subse- 
quently validated by another author and 
overlooked by the present one. Several of 



MoLLUSKS Described by W. H. Pease • Johnson 



the land shells from the Society Islands are 
known only from the specimens figured 
by Garrett (1884) and are regarded as lec- 
totypes. Examples of some of the described 
taxa that are missing may occur in the 
Garrett collection in the Bernice Pauahi 
Bishop Museum under the names of Pease 
and, while not exactly strictly speaking 
types, might serve as neotypes. 

ACKNOWLEDGMENTS 

It gives me great pleasure to thank those 
people whose cooperation eased the task 
of gathering information. Peter Mordan, 
of the Mollusca Section, British Museum 
(Natural History), was able to inform me 
that certain of the 14 non-marine mollusks 
not in the Pease collection were repre- 
sented in that collection and Ms. Kathie 
Way supplied other data. Drs. George M. 
Davis and Gary Rosenberg of the Acade- 
my of Natural Sciences of Philadelphia an- 
swered every request for material and data, 
and there were many; Ms. Doree Bardes 
also helped. Dr. Alison Kay left a number 
of helpful notes in many lots of the marine 
types in the Museum of Comparative Zo- 
ology in conjunction with her work (1975). 
Mr. Tan Koh Siang identified the types of 
Sistrum triangulatum, which otherwise 
would have been overlooked. With the ex- 
ception of two photographs by Dr. James 
H. McLean, Los Angeles County Museum 
of Natural History, and two others by Dr. 
Kenneth J. Boss, all of the photographs and 
measurements were made by Dr. Silvard 
P. Kool. Dr. Kenneth J. Boss made many 
helpful suggestions. Finally, thanks are ex- 
tended to Mrs. Marion D. Britz, who cop- 
ied and recopied as many corrected copies 
of this manuscript as did Sonia Tolstoy of 
Anna Karenina. Drs. Alan R. Kabat and 
Gary Rosenberg kindly reviewed the 
manuscript and made a number of frank 
criticisms that saved this author from some 
really egregious errors. I would also like to 
thank Drs. C. C. Wang, William U. Shi- 
pley, Merrill Goldstein, Mandel E. Cohen, 
and Blair Ardman. Publication costs of this 



study were covered in part by the Wet- 
more Colles Fund. 

ABBREVIATIONS 

In general, this author disdains refer- 
ences that merely give a date, forcing the 
use of the bibliography over and over. To 
partially alleviate this problem, the prin- 
cipal references are cited in the catalog by 
the following abbreviations. 

ACM Annals of the Carnegie Museum 
AJC American Journal of Conchology 
ANSP Academy of Natural Sciences of 
Philadelphia, Philadelphia, Penn- 
sylvania 
BMCZ Bulletin of the Museum of Com- 
parative Zoology, Cambridge, 
Massachusetts 
BMNH British Museum (Natural History) 
BPBM Bernice Pauahi Bishop Museum, 

Honolulu, Hawaii 
CM Carnegie Museum, Pittsburgh, 

Pennsylvania 
JANSP Journal of the Academy of Nat- 
ural Sciences of Philadelphia 
JdeC Journal de Conchyliologie 
MCZ Museum of Comparative Zoology, 

Cambridge, Massachusetts 
MNHN Museum National d'Histoire Na- 

turelle, Paris, France 
MofC Manual of Conchology 
PZS Proceedings of the Zoological So- 
ciety of London 
USNM National Museum of Natural His- 
tory, Washington, D.C. 

Titles of other journals cited in the catalog 
are usually given in full. 



LIST OF THE TAXA OF SHELLED 
MOLLUSKS DESCRIBED BY PEASE 

abbreviata, Partula 

1865a, PZS for 1864: 675 [>2omen nudum]. MCZ 2505. 

abbreviata, Planaxis Plate 9, Figure 2 

1865d, PZS for 1865: 515 (Islands of the Central 
Pacific); 1868g, AJC 4: 101, pi. 12, fig. 16 (Tahiti). 
Lectotype, here selected, ANSP 18261; paralecto- 
types MCZ 187833 (Plate 8, Figure 13) and 187834; 
not located in BMNH by Kay (1965: 86). 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



abbreviata, Realia 

1865a, PZS for 1864: 674 (Islands of the Central 
Pacific); 1869a, JdeC 17: 155, pi. 7, fig. 5, as Om- 
phalotropis (Tahiti). Holotype in MNHN, teste Fi- 
scher-Piette (1950: 72); paratypes MCZ 74936 and 
187842. 

acetabulum, Helix 

1861d, PZS for 1861: 242 (Tahiti). Holotype ANSP 
47844a, teste Baker (1963: 231) selected as lectotype 
by Solem (1976: 357, figs. 156d-f); paralectotypes 
MCZ 17248 and 176567. 

aciculata, Eulima 

1861b, PZS 28: 438 (Sandwich Islands). Lectotype 
BMNH 1962839 selected by Kay (1965: 66, pl. 9, fig. 
2); paralectotype MCZ 31705 figured by Pilsbry 
(1917: 222, figs. 11a, b); paralectotype MCZ 187747. 

aculeata, Scutellina Plate 1, Figure 9 

1868g, AJC 4: 100 (Hawaii [Island]). Lectotype, here 
selected, MCZ 83720 labeled as holotype. 

affine, Sistrum 

1863b, PZS for 1862: 244 (Kingsmill Islands); 1868d, 
AJC 3: 277, pl. 23, fig. 13. Lectotype BMNH 1964304 
selected by Kay (1965: 79, pl. 13, figs. 13, 14). 

affinis, Partula 

1865a, PZS for 1864: 675 [nometi nudum]; 1868b, AJC 
3: 224 (Tahiti). Holotype ANSP 59560a, teste Baker 
(1963: 204); paratypes MCZ 25313. 

affinis, Realia 

1865e, AJC 1: 288 (Polynesia); Tryon, 1866a, AJC 2: 
82, pl. 5, fig. 4 (Hervey Isles); Pease, 1869a, JdeC 
17: 152 as Omphalotropis (Aitutake, Hervey Isles). 
Holotype ANSP 13346a, teste Baker (1964: 178); 
paratypes MCZ 187840 and 187841. 

affinis, Triphoris 

1861b, PZS 28: 434 ([Kauai] Sandwich Islands). Ho- 
lotype BMNH 1962808 figured by Kay (1965: 51, pi. 
10, fig. 1); paratypes MCZ 50074. 

alba, Thala Plate 9, Figure 20 

1868a, AJC 3: 215, pl. 15, fig. 8 (Paumotus). Lecto- 
type ANSP 28755 selected by Cernohorsky (1976: 
501, pl. 450, fig. 6); three paralectotypes ANSP 
325385; paralectotypes MCZ 260612. 

albinea, Helicina maugeriane 

1871d, PZS for 1871: 466 (Raiatea [restricted to a 
single valley on the east side of Tahaa, teste Garrett, 
1884: 101, pl. 3, fig. 64]). Lectotype, here selected, 
ANSP 14538 is the specimen figured by Garrett; 
five paralectotypes MCZ 202599 from Garrett; not 
located in BMNH, teste Mordan, personal com- 
munication. 

albocincta, Engina 

1860b, PZS 28: 142 (Sandwich Islands). Lectotype 
BMNH 1961454 selected by Kay (1965: 16, pl. 2, 
figs. 9, 10); paralectotypes MCZ 297948. 

alta. Helix 

1868), AJC 4: 153, pl. 12, fig. 1 (Ponape). Holotype 
ANSP 49016, teste Baker (1963: 233); paratypes MCZ 
94770. 

alternata 'Pease' H. H. Smith, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1902, ACM 
1: 447 (Moorea). Syntypes MCZ 25069. 



alternata, Triphoris 

1861b, PZS 28: 434 ([Kauai] Sandwich Islands); 
1868h, AJC 4: 127 non Adams 1852. Changed to: 
Triphoris btcolor. Lectotype BMNH 1962815 selected 
by Kay (1965: 54, pl. 10, fig. 4); paralectotypes MCZ 
50057 and 73735. 
ambigua, Limnaea 

1870b, AJC 6: 6, pl. 3, fig. 5 (Hawaiian Islands). 
Holotype ANSP 21240a, teste Baker (1964: 152); 
paratypes MCZ 73469. 
anibusta, Auriculella Plate 2, Figure 10 

1868f, JdeC 16: 345 ([Waianae Mountains, Oahu]). 
Lectotype, here selected, MCZ 45152; paralecto- 
types MCZ 298488. 
analogica, Pithys 

1870c, JdeC 18: 396 (Marquesas [Islands]); 1871b, 
PZS for 1871: 454. Lectotype BPBM 115307 from 
MCZ 17260 selected by Solem (1976: 328, figs. 143a- 
d); paralectotypes MCZ 17260. 
angasii, Rissoina 

1871e, AJC 7: 20. New name for turricula Angas 1867 
non Pease 1860. 
angicostata, Hindsia 

1860b, PZS 28: 142 (Sandwich Islands). Lectotype 
BMNH 1961159 selected by Kay (1965: 16, pl. 1, 
figs. 15, 16). 
angiostoma, Cythara 

1868h, AJC 4: 105. New name for Pleurotoma triticea 
Reeve 1843 non Kiener 1839. 
angiostoma, Thala Plate 9, Figure 22 

1868a, AJC 3: 216, pl. 15, fig. 9 (Paumotus). Lecto- 
type, here selected, ANSP 28754; paralectotypes 
MCZ 256500. 
angulata, Carelia adusta 
\870c, JdeC 18: 403 (Kauai). Lectotype ANSP 23434 
selected by Baker (1963: 197) figured by Hyatt and 
Pilsbry (1911: 116, pl. 20, fig. 16); paralectotypes 
MCZ 45167. 
angulatus, Euchelus Plate 7, Figure 13 

1868d, AJC 3: 283, pl. 23, fig. 27 (Annaa Island). 
Lectotype, here selected, ANSP 40671 is the figured 
type; two paralectotypes ANSP 391031; paralecto- 
types MCZ 89796. 
annectens, Bulimus [Partula] 

1865a, PZS for 1864: 671 (Islands of the Central 
Pacific [two valleys on the west coast of Huaheine, 
teste Garrett, 1884: 66]). Syntypes MCZ 26352. 
antiqua, Leptachatma 

1870a, JdeC 18: 94 (Kauai); Crosse, 1876, JdeC 24: 
97, pl. 3, fig. 6. Holotype in MNHN, teste Fischer- 
Piette (1950: 149). 
aperta, Haminea Plate 4, Figure 17 

I868e, AJC 4: 72 (Tahiti). Lectotype, here selected, 
ANSP 57575. 
aperta, Tornatellina Plate 2, Figure 18 

1865b, PZS for 1864: 673 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 473 (Tahiti). Lecto- 
type, here selected, MCZ 175722; paralectotypes 
MCZ 298457. 
approximala, Nassa Plate 8, Figure 4 

1868d, AJC 3: 272, pl. 23, fig. 3 ([Ascension Island]). 



MoLLUSKS Described by W. H. Pease • Johnson 



Lectotype MCZ 151800 selected by Cernohorsky 
(1984: 128, pi. 23, fig. 13); paralectotype MCZ 303191. 

approximata 'Pease' Garrett, Partula 

1884, JANSP (2) 9: 75 (several small valleys on the 
south west part of Raiatea). Lectotype ANSP 59451a 
selected by Baker (1964; 204) figured by Pilsbry 
(1909: 243, pi. 17, fig. 15); paralectotypes MCZ 50851, 
24927, and 25197. 

approximata, Turricula 

1860b, PZS 28: 146 (Sandwich Islands). Lectotype 
BMNH 1961192 selected by Kay (1965: 27, pi. 3, 
figs. 1, 2). 

argutus, Bulimus [Partula] 
'l865b, PZS for 1864: 670 (Islands of the Central 
Pacific [Huaheine, teste Garrett, 1884, JANSP (2) 9: 
62, pi. 3, fig. 57]). Syntypes MCZ 26353. 

armata, Vertigo 

1871d, PZS for 1871: 461 (Bolabola [Borabora Island, 
Society Islands]). Holotype MCZ 48315 figured by 
Pilsbry and Cooke (1920: 327, pi. 30, figs. 12, 13); 
paratypes MCZ 31398. 

arwatus, "7" Acanthochites Plate 1, Figure 1 

1872a, AJC 7: 195 ([Pauloa] Oahu). Holotype MCZ 
73452, only specimen. 

asperum, Cerithium 

1861b, PZS 28: 433 (Sandwich Islands). Holotype 
BMNH 1961203 figured by Kay (1965: 47, pi. 5, figs. 
11, 12); paratype MCZ 239650. 

assimilis, Mitra Plate 7, Figure 20 

1868a, AJC 3: 211, pi. 15, fig. 1 ([Oahu] Polynesia). 
Lectotype ANSP 28718 selected by Cernohorsky 
(1976: 487, pi. 436, fig. 1) and the type locality re- 
stricted to: Huahine [sic] Island, Society Islands; 
paralectotypes MCZ 260601 and 260603; paralec- 
totype ANSP 391048. 

assimilis, Partula 

1868b, AJC 3: 230, pi. 15, figs. 28, 29 [both as 28] 
(Roratonga); 1870c, JdeC 18: 401. Syntypes MCZ 
24906 and 25137; not located in ANSP by Baker 
(1963: 204). 

assimilis, Terebra 

1869d, AJC 5: 67 (Oahu); 1871e, AJC 7: 20 is Terebra 
contigua Pease 1871, new name for T. assimilis non 
Angus 1867. Not located in ANSP. 

aliensis, Pithys Plate 2, Figure 8 

1870c, JdeC 18: 394 (Atiu [Island, Cook Islands]); 
1871d, PZS for 1871: 453. Lectotype, here selected, 
MCZ 17335; paralectotypes MCZ 17336. 

atra, Planaxis Plate 9, Figure 1 

1869d, AJC 5: 72, pi. 6, fig. 4 (Marquesas [Islands]). 
Lectotype, here selected, ANSP 18282; three para- 
lectotypes ANSP 391038; paralectotypes MCZ 
187836 and 187837. 

attenuala, Cirsotrema. 

1861a, PZS 28: 400 (Sandwich Islands). Lectotype 
BMNH 1962796 selected by Kay (1965: 43, pi. 10, 
figs. 9, 10). 

attenuata, Partula 

1865a, PZS for 1864: 672 (Islands of the Central 
Pacific [Tahiti]). Syntypes MCZ 24901 and 25047. 



aurantmm, Operculatum 

1868d, AJC 3: 287 (Hawaii [Island]). Not located in 
ANSP. 

australis, Tritonidea 

1871e, AJC 7: 21 (Australia). New name for Triton- 
idea assimilis Angas 1867 non Buccitium assimile Reeve 
1846. 

bacca. Vertigo 

187 Id, PZS for 1871: 462 (Kalapana [Puna District] 
Hawaii Island); specimens lost, teste Pease. 

baetica 'Pease' Paetel, Lampania 

1887, Catalog der Conchylten-Sammlung (Berlin) 1: 351 
[nomen tiudum]. 

balteata, Clathurella 

1860b, PZS 28: 143 (Sandwich Islands). Lectotype 
BMNH 1962786 selected by Kay (1965: 34, pi. 5, 
figs. 3, 4); paralectotypes MCZ 50004. 

balteata, Leptachatina 

1870a, JdeC 18: 91 ([Waimea] Kauai); Crosse, 1876, 
JdeC 24: 97, pi. 4, fig. 4. Holotype in MNHN, teste 
Fischer-Piette (1950: 149); paratypes MCZ 142986. 

balteata, Nassa 

1869d, AJC 5: 71, pi. 8, fig. 4 (Ebon Island). Lecto- 
type, here selected, ANSP 16366 is the figured type. 

balteata, Rissoina 

1869d, AJC 5: 72 (Hawaii [Island]). Not located in 
ANSP. 

bell a, Daphnella 

1860b, PZS 28: 147 (Sandwich Islands). Lectotype 
BMNH 1962786 selected by Kay (1965: 34, pi. 5, 
figs. 3, 4); paralectotypes MCZ 50013 and 50014. 

bella, Helicitia 

1865a, PZS for 1864: 676 [numen nudum]. MCZ 297930. 

bella 'Pease' Hartman, Partula 

1871a, PZS for 1871: 473 [nomen nudum]; 1881, BMCZ 
9: 193 ([Raiatea]). Syntypes MCZ 3639. 

bella, Turricula 

1860b, PZS 28: 145 (Sandwich Islands). Lectotype 
BMNH 1961204 selected by Kay (1965: 26, pi. 3, 
figs. 12, 14); paralectotypes MCZ 50017. 

biangulatum, Cyclostoma Plate 2, Figure 13 

1865a, PZS for 1864: 674 (Islands of the Central 
Pacific [Aitutaki, Cook Islands, teste Garrett (1881: 
404) who collected the specimens]). Lectotype, here 
selected, MCZ 141031; paralectotypes MCZ 141032. 

bicarinata, Amathina 

1861a, PZS 28: 399 (Sandwich Islands). Lectotype 
BMNH 1962792 selected by Kay (1965: 40, pi. 6, 
figs. 3, 4). 

bicarinata, Clathurella Plate 6, Figure 17 

1863c, PZS for 1862: 243 (KingsmiU Islands); 1868a, 
AJC 3: 222, pi. 15, fig. 23. Lectotype, here selected, 
ANSP 15806; not located in BMNH by Kay (1965: 
86). 

bicolor, Partula 

1871f, AJC 7: 26, pi. 9, fig. 4 (Guam). Syntype MCZ 
25345; not located in ANSP by Baker (1963: 204). 

bicolor, Triphoris 

1868h, AJC 4: 127. New name for Triphoris alternata 



8 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



Pease 1860 non C. B. Adams 1852. See under: alter- 
tiata, Triphoris. 

bifasciata, Borsonia 

1860b, PZS 28: 143 (Sandwich Islands). Holotype 
BMNH 1962758, teste Kay (1965; 18, pi. 8, figs. 4, 
5). 

bilineata, Partula 

1865b, PZS for 1864: 675 [tiomen nudum]; 1866a, AJC 
2: 201 (Tahaa [confined to Faa-apa Valley on the 
east coast, teste Garrett, 1884: 62]); 1867a, AJC 3: 81, 
pi. 1, fig. 10; 1871b, PZS for 1871: 473 (Tahiti). Lec- 
totype ANSP 59438a selected by Baker (1963: 204) 
is the figured type; paralectotypes MCZ 24903. 

hipes, Syphonota 

1860a, PZS 28: 23 (Sandwich Islands). Syntype MCZ 
297867; not located in BMNH by Kay (1965: 84). 

boeticum, Centhmm 

1861b, PZS 28: 433 (Sandwich Islands). Lectotype 
BMNH 1962802 selected by Kay (1965: 48, pl. 10, 
fig. 8). 

brazieri, Helicina Plate 5, Figure 11 

1870c, JdeC 18: 397 (Niue [Island]). Lectotype, here 
selected, MCZ 74947; paralectotypes MCZ 298500. 

brazieri, Partula 

18711, AJC 7: 27, pl. 9, fig. 5 (Tutuila). Holotype 
ANSP 59846, teste Baker (1963: 204). 

brevicula, Leptachatina 

1869c, JdeC 17: 169 (Kauai). Measured holotype and 
paratype in MNHN, teste Fischer-Piette (1950: 72); 
paratype ANSP 57802 figured by Cooke (1910: 24, 
pl. 8, fig. 54); paratype MCZ 45195. 

brevicula 'Pease' Garrett, Partula 

1884, JANSP (2) 9: 48 (Faahuaite Valley, Tahiti). 
Lectotype ANSP 59558a selected by Baker (1963: 
204) figured by Pilsbry (1909: 191, pl. 26, fig. 12); 
paralectotypes MCZ 24997. 

brevis, Cithara [Cythara] Plate 6, Figure 10 

1868a, AJC 3: 217, pl. 15, fig. 11 ([Anaa Island] Pau- 
motus). Lectotype, here selected, ANSP 15652 is the 
figured type; paralectotype ANSP 391038; para- 
types MCZ 49988 and 49989. 

brunnea, Avicula Plate 1, Figure 3 

1863c, PZS for 1862: 244 ([Molokai] Sandwich Is- 
lands). Lectotype, here selected, MCZ 298465; para- 
lectotypes MCZ 297881; not located in BMNH by 
Kay (1965: 86). 

brunnea, Clatliurella 

1860b, PZS 28: 143 (Sandwich Islands). Lectotype 
BMNH 1962763 selected by Kay (1965: 19, pl. 8, 
figs. 1-3); paralectotypes MCZ 50006 and 50007. 

brunnea, Strigatella Plate 9, Figure 10 

1868a, AJC 3: 215, pl. 15, fig. 7 ([Paumotus] Polyne- 
sia). Lectotype, here selected, ANSP 29722 is the 
figured type; five paralectotypes ANSP 391049; 
paralectotype MCZ 260610. 

brunneus, Triphoris Plate 10, Figure 6 

1871a, PZS for 1870: 777 (Apaiang [Abaiang Is- 
land]). Lectotype, here selected, MCZ 73922. 

buccmoides, Clatliurella 

1860d, PZS 28: 144 (Sandwich Islands). Holotype 
BMNH 1961156, teste Kay (1965: 23, pl. 2, figs. 1, 
2). 



calliostoma, Helicina Plate 5, Figure 5 

1871d, PZS for 1871: 466 (Marquesas [Islands]). Lec- 
totype, here selected, MCZ 74950; paralectotypes 
MCZ 298501. 

canaliculata, Clatliurella Plate 6, Figure 16 

1868a, AJC 3: 219, pl. 15, fig. 17 (Paumotus). Lec- 
totype, here selected, MCZ 231978; paralectotypes 
MCZ 303197; not located in ANSP. 

cancellata, Coralliobia 

1861a, PZS 28: 399 (Sandwich Islands), "only a sin- 
gle dead specimen found." Not located in BMNH 
by Kay (1965: 86). 

cancellata, Scutellina 

1861b, PZS 28: 437 (Sandwich Islands). Lectotype 
BMNH 1962833 selected by Kay (1965: 64, pl. 11, 
figs. 8, 9). 

cancellatus, Strombus 

1861a, PZS 28: 398 (Sandwich Islands). Lectotype 
BMNH 1961180 selected by Kay (1965: 40, pl. 4, 
figs. 8, 9). 

Candida, "?" Collonia 

1861b, PZS 28: 436 (Sandwich Islands). Lectotype 
BMNH 1962826 selected by Kay (1965: 61, pl. 9, fig. 
8); paralectotypes MCZ 245260. 

Candida, Cypraea 

1865d, PZS for 1865: 515 (Islands of the Central 
Pacific); 1868g, AJC 4: 95, pl. 11, figs. 12, 13 (Apaian 
Island). Lectotype BMNH 1964318 selected by Kay 
(1965: 83, pl. 14, figs. 3, 4); paralectotypes MCZ 
297949. 

Candida, Volvatella 

1868e, AJC 4: 73, pl. 7, fig. 6 ([Tahiti] Polynesia). 
Syntypes MCZ 297940, mostly fragments. 

capillata. Helix Plate 2, Figure 1 

1866b, AJC 2: 292 (Sandwich Islands); 1871b, PZS 
for 1871: 474 as Pitys (Kauai). Lectotype ANSP 1975a 
(now 1975) selected by Solem (1976: 368 [not fig- 
ured]) is the measured type, teste Baker (1963: 232); 
paralectotypes MCZ 17585. 

castanea 'Pease' Garrett, Partula 

1884, JANSP (2) 9: 75 (Faaloa, on the east coast [of 
Raiatea]). Lectotype ANSP 59977a selected by Baker 
(1963: 204) figured by Pilsbry (1909: 244, pl. 17, fig. 
2); paralectotypes MCZ 297856. 

celsa, "?" Pitliys 

1870c, JdeC 18: 396 (Raiatea); 1871b, PZS for 1871: 
445 as Endodonta. Lectotype BPBM 3484 selected by 
Solem (1976: 358, figs. 158a-c); paralectotypes MCZ 
17243 and 17249. 

cerithiopsis, Rissoina 

1862b, JdeC 10: 382 [nomen nudum]. 

cmcta, Helicina 

1865a, PZS for 1864: 676 [nomen nudum]. 

cmctus, Melampus 

1865a, PZS for 1864; 676 [nomen nudum]. 

cinerea, Littorina 

1869d, AJC 5: 78, pl. 8, fig. 14 (Marquesas [Islands]). 
Lectotype, here selected, ANSP 18811 is the figured 
type. 

cmgulifera, Triphoris 

i861b, PZS 28: 434 ([Haena Point, Kauai] Sandwich 
Islands). Lectotype BMNH 1962812 selected by Kay 



MoLLUSKs Described by W. H. Pease • Johnson 



(1965: 52, pi. 6, figs. 9, 10); paralectotypes MCZ 
50056, 50076, and 73737. 

citrina, Partula 

1865b, PZS for 1864: 675 [nomcn nudum] 1866a, AJC 
2: 195 (Tahitian Archipelago [restricted to a single 
valley called Uparu on the west coast of Raiatea, 
teste Garrett, 1884: 64]). Syntypes MCZ 24885 and 
25027. 

clara, Partula 

1865b, PZS for 1864: 671 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 473 (Tahiti). Syntypes 
MCZ 24931 and 25141. 

clathrata, Emarginula 

1863b, PZS 'for 1862: 241 (Pacific Islands); 1868g, 
AJC 4: 99, pi. 1 1, fig. 24 (Rowland Island). Lectotype 
BMNH 1964290 selected by Kay (1965: 74, pi. 12, 
fig. 11); paralectotypes MCZ 179161. 

clavata, Triphons 

1861b, PZS 28: 434 (Sandwich Islands). Lectotype 
BMNH 1962814 selected by Kay (1965: 53, pi. 10, 
fig. 2); paralectotype MCZ 50060. 

cognata 'Pease' Garrett, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1884, JANSP 
(2) 9: 67, 68 (Faahiti Valley, Huaheine). Specimens 
figured by Pilsbry (1909: 254, pi. 22, figs. 12, 17; pi. 
33, fig. 3). Lectotype ANSP 59991a selected by Baker 
(1963: 204); paralectotypes MCZ 24817 and 25117. 

colorata, Helicina 

1868), AJC 4: 156, pi. 12, fig. 9 (Annaa [Island]). 
Holotype ANSP 145541a, teste Baker (1964: 101); 
paratypes MCZ 297929. 

compacta, Labiella Plate 4, Figure 5 

1869c, JdeC 17: 172 ([Palauea] Maui; Coll. [of] Pease). 
Lectotype, here selected, MCZ 45196; paralectotype 
MCZ 298489. 

compacta, Limnaea 

1870b, AJC 6: 6, pi. 3, fig. 4 (Oahu). Syntypes MCZ 
302381; not located in ANSP by Baker (1964: 151). 
Is Physa mexicana Philippi [introduced], teste D. W. 
Taylor, personal communication. 

compacta, Partula 

1865b, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 200 (Raiatea [Hamoa Valley on the east coast of 
Raiatea, teste Garrett, 1884: 55]); 1867a, AJC 3: 81, 
pi. 1, fig. 9. Lectotype ANSP 59983 selected by Baker 
(1963: 204) is the figured type; paralectotypes MCZ 
24993. 

complanalum, Registoma Plate 4, Figure 3 

1861c, PZS 28: 440 (Ebon Island). Lectotype, here 
selected, MCZ 141018; paralectotypes MCZ 141019. 

compressa, Scutellina Plate 1, Figure 10 

1868g, AJC 4: 99, pi. 11, figs. 25, 27 (Tahiti). Lec- 
totype, here selected, MCZ 302552. 

compta, Cypraea 

1860c, PZS 28: 189 (Jarvis Island). Holotype BMNH 
1964276, teste Kay (1965: 36, pi. 12, figs. 1, 2). 

compta, Partulma 

1869c, JdeC 17: 175 (Molokai). Holotype in MNHN 
figured by Fischer-Piette (1950: 73, fig. 54); para- 
types MCZ 25826 and 25828. 

concinna, Partula 

1872a, AJC 7: 196 (New Hebrides). Though not a 



type Carnegie Museum 4244, "Tanna Island, New 
Hebrides [Vanuatu] (Cox)" was figured by Hartman 
(1886: 35, pi. 2, fig. 16) as this species and by Pilsbry 
(1909: 288, pi. 36, figs. 9, 12); not located in ANSP 
by Baker (1963: 204). 

concinna, Truncatella Plate 10, Figure 18 

1871d, PZS for 1871: 468 (Apaiang [Abaiang Island, 
Kingsmill Islands]). Lectotype, here selected, MCZ 
178650; paralectotypes MCZ 298459. 

congrua. Helix Plate 2, Figure 4 

1868), AJC 4: 154, pi. 12, figs. 3, 4 (Ponape); 1871b, 
PZS for 1871: 457 as Trochomorpha contigua to replace 
H. congrua 1868 non Pfeiffer 1858. Lectotype, here 
selected, MCZ 12161; paralectotypes MCZ 12159 
and 297861; not located in ANSP by Baker (1963). 

conica, Laimodonta 

1863b, PZS for 1862: 242 (Pacific Islands); 1868g, 
AJC 4: 101, pi. 12, fig. 15 (Paumotus); 1871b, JdeC 
19: 94 (Annaa). Lectotype ANSP 22610a selected by 
Baker (1964: 151) is the figured type; a lectotype 
BMNH 1964292 was also subsequently selected by 
Kay (1965: 75, pi. 13, figs. 7, 8). 

conica, Tormia 

1865d, PZS for 1865: 514 (Islands of the Central 
Pacific). Not located in BMNH by Kay (1965: 86). 

conoidalis, Tectura Plate 1, Figure 6 

1868g, AJC 4: 98, pi. 11, fig. 22 (Roratonga [Island]). 
Lectotype, here selected, MCZ 302558; paralecto- 
types MCZ 298470; specimens identified by Pease 
from a different locality [idiotypes] ANSP 40993. 

conoidalis, Trochus Plate 9, Figure 26 

1868d, AJC 3: 287, pi. 24, fig. 8 (Paumotus). Lecto- 
type, here selected, ANSP 18868 is the figured type; 
three paralectotypes ANSP 267209; paralectotypes 
MCZ 104618, 150597, and 298236. 

consimilis. Helix 

1865b, PZS for 1864: 675 [nomen nudum]; 1868b, AJC 
3: 227 (Tahiti [error for Raiatea]). Lectotype BMNH 
71 .1.5.28 selected by Solem (1976: 174 [not figured]); 
paralectotypes MCZ 17262 and 297952. 

conspersa. Bulla 

1869d, AJC 5: 72, pi. 8, fig. 9 (Marquesas [Islands]). 
Lectotype, here selected, ANSP 57505 is the figured 
type; paralectotypes MCZ 297880 and 298464. 

contigua, Melania Plate 4, Figure 19 

1870b, AJC 6: 7 (Kauai). Lectotype, here selected, 
MCZ 74887; paralectotypes MCZ 298908; not lo- 
cated in ANSP by Baker (1964: 181). 

contigua, Terebra 

1871e, AJC 7: 20. New name for Terebra assimilis 
Pease 1869 non Angas 1867. See under: assimilis, 
Terebra. 

contigua, Trochomorpha 

1871d, PZS for 1871: 457. New name for Helix con- 
grua Pease 1868 non Pfeiffer 1858. See under: con- 
grua. Helix. 

conula. Helix Plate 3, Figure 6 

1861d, PZS for 1861: 243 (Tahiti). Lectotype, here 
selected, MCZ 297945; paralectotypes MCZ 298469. 

coreensis. Turcica Plate 10, Figure 23 

1860c, PZS 28: 189, pi. 51, fig. 2 (Corea [Korea] Sea). 



10 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



Lectotype, here selected, MCZ 104609 is the figured 
type; not located in BMNH by Kay (1965: 85). 

corrugata 'Pease' Tryon, Borsonia 

1884, MofC (1)6: 228 [nomen nudum]. 

corrugata, Helkina Plate 5, Figure 4 

1865a, PZS for 1864: 673 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 476 (Raiatea). Lecto- 
type, here selected, MCZ 297925; paralectotypes 
MCZ 298904. 

corrugata, Trivia Plate 10, Figure 25 

1868g, AJC 4: 95, pi. 11, figs. 14, 15 (Paumotus). 
Lectotype, here selected, MCZ 303451; paralecto- 
types MCZ 303197; three specimens identified by 
Pease [idiotypes] ANSP 39703 labeled as from the 
Sandwich Islands. 

corrugatus, Euchelus 

1861b, PZS 28: 435 (Sandwich Islands). Lectotype 
BMNH 1962821 selected by Kay (1965: 58, pi. 8, 
figs. 12, 13); paralectotypes MCZ 89897. 

costata, Engina 

1860b, PZS 28: 142 (Sandwich Islands). Lectotype 
BMNH 1961163 selected by Kay (1965: 14, pi. 1, 
figs. 17, 18). 

costata, Hydrocaena 

1865b, PZS for 1864: 676 [nomen nudum]. 

costata, Leptothyra Plate 7, Figure 19 

1869d, AJC 5: 70 (Maui). Lectotype, here selected, 
MCZ 245261; paralectotypes MCZ 298461. 

costata, Realia 

1868b, AJC 3: 225 (Tahaa); 1869a, JdeC 17: 158, pi. 
7, fig. 2 as Scalinella. Holotype and paratype in 
MNHN, teste Fischer-Piette (1950: 72); Baker (1964: 
178) claimed the holotype is ANSP 13359a; para- 
types MCZ 74939, 74951, and 187864. 

costata. Vertigo Plate 2, Figure 20 

1871d, PZS for 1871: 461 ([Kona] Hawaii [Island]). 
Listed as an undetermined species by Pilsbry and 
Cooke (1920: 272). Holotype MCZ 45238, teste Cooke 
on label; not MCZ 45327 which is a Goniobasis as 
mentioned by Pilsbry and Cooke (1926: 224). 

cost at us, Triphoris Plate 10, Figure 1 

1871a, PZS for 1870: 775 (Annaa [Island]). Lecto- 
type, here selected, MCZ 273206; paralectotypes 
MCZ 298481. 

costellifera, Anachis 

1863d, PZS for 1862: 279 (Pacific Islands). Not lo- 
cated in BMNH by Kay (1965: 86). 

costellifera, Terebra Plate 9, Figure 19 

1869d, AJC 5: 66 ([Honolulu] Oahu). Lectotype MCZ 
248800 selected by Bratcher and Cernohorsky (1987: 
207, pi. 64, fig. 252b). 

costellifera. Truncal ella 

1871d, PZS for 1871: 468 (Vavau Island [Tonga Is- 
lands]). Not located in BMNH, teste Mordan (per- 
sonal communication). "Mr. Pease's T. costellifera, 
which Mr. Brazier obtained at Vavau, Tonga Islands 
is undoubtedly the same as T. rustica (Mousson)." 
(Garrett, 1887: 300). 

costulata, Rissoina Plate 9, Figure 6 

1868d, AJC 3: 295, pi. 24, fig. 28 (Paumotus). Lec- 
totype, here selected, ANSP 19241 is the figured 



type; three paralectotypes ANSP 391036; paralec- 
totypes MCZ 178856. 

costulosa, Atys Plate 6, Figure 5 

1869d, AJC 5: 73 ([Waimalu] Oahu). Holotype MCZ 
31714, only specimen, figured by Pilsbry (1917: 218, 
fig. 6). 

costulosa, Leptachatina 

1870a, JdeC 18: 90 (no locality); Crosse, 1876, JdeC 
24: 97, pi. 3, fig. 4 (Kauai). Holotype in MNHN, 
teste Fischer-Piette (1950: 149); paratype MCZ 45191. 

costulosa, Succinea Plate 4, Figure 15 

1865b, PZS for 1864: 677 (Tahitian Archipelago); 
1871d, PZS for 1871: 472 (Tahiti). Lectotype, here 
selected, MCZ 31406; paralectotypes MCZ 298485. 

costulosa. Vertigo Plate 2, Figure 22 

1871d, PZS for 1871: 462 (Hawaii). Lectotype, here 
selected, MCZ 45244; paralectotypes MCZ 32294; 
not located in BMNH, teste Mordon (personal com- 
munication). 

coxi, Bulimus ("?" Borus) Plate 2, Figure 12 

1872b, AJC 7: 197 (Solomon Islands). Holotype MCZ 
86495, [New Hebrides Islands], teste Clench (1932: 
69). 

crassa 'Pease' Garrett, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1884, JANSP 
(2) 9: 49 (Faahuaite [Valley], Huaheine). Lectotype 
ANSP 59921a selected by Baker (1963: 204); para- 
lectotypes MCZ 25078. 

crassicostata, Borsonia 

1860b, PZS 28: 143 (Sandwich Islands). Lectotype 
BMNH 1962847 selected by Kay (1965: 17, pi. 2, 
figs. 5, 6). 

crassilabris, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 199, 201 ([Hapai Valley] Raiatea); 1867a, AJC 3: 
81, pi. 1, fig. 6, as crassilabrum. Holotype ANSP 
59477a, teste Pilsbry (1909: 226, pi. 21, fig. 10) se- 
lected as lectotype by Baker (1963: 205); paralec- 
totypes MCZ 25131, 25139, and 25292. 

crenulata, Dapyhnella Plate 7, Figure 3 

1868c, AJC 3: 221, pi. 15, fig. 20 ([Howland Island] 
Polynesia). Lectotype, here selected, ANSP 15694; 
paralectotype MCZ 221177. 

crenulata, Scalaria 

1868d, AJC 3: 290, pi. 24, fig. 13 (Tahiti). Holotype 
ANSP 19553, only specimen; refigured by DuShane 
(1990: 3, fig. 6). 

crispata, Scalaria Plate 9, Figure 13 

1868d, AJC 3: 289, pi. 24, fig. 12 (Paumotus). Lec- 
totype ANSP 19575 selected by DuShane (1988 (5): 
9, fig. [not numbered]) and refigured (1990: 8, fig. 
35); two paralectotypes ANSP 352472; paralecto- 
types MCZ 187528. 

crocata, Haminca 

1861b, PZS 28: 19, 432 (Sandwich Islands). Lecto- 
type BMNH 1961191 selected by Kay (1965: 7, pi. 
1, figs. 9, 10); paralectotypes MCZ 88127, 207387, 
and 297883. 

curta, Daphnella Plate 7, Figure 2 

1868a, AJC 3: 221, pi. 15, fig. 22 (Paumotus). Lec- 
totype, here selected, ANSP 16956; paralectotype 
MCZ 221178. 



MoLLUSKS Described by W. H. Pease • Johnson 11 



cylmdracca Tease' Nevill, Triiucatclla 

1878, HanJ List of Mollusca in the Indian Museum 1: 
253 [nonien nudum]. MCZ 161490. 

cyUndraceum, Cerithium 

' 1869d, AJC 5: 77 (Paumotus). Holotype ANSP 17703, 
teste, Houbrick (1992: 49, fig. 31j); paratypes MCZ 
297939 (Plate 6, Figure 6) and 298468. 

cylmdrata, Leptachatina 

1869c, JdeC 17: 168 (Kauai). Lectotype, here select- 
ed, ANSP 57806 is the syntype figured by Cooke 
(1 91 0: 18, pi. 8, figs. 63, 64); paralectotypes in MNHN, 
teste Fischer-Piette (1950: 72). 

cylindrica, Clathurella 

' 1860b, PZS 28: 143 (Sandwich Islands). Lectotype 
BMNH 1962765 selected by Kay (1965: 20, pi. 8, fig. 
8). 

cylindrica, Marginella 

' 1863c, PZS for 1862: 244 (Kingsmill Islands); 1868d, 
AJC 3: 280, pi. 23, fig. 19 [Tarawa Island] as Mar- 
ginella polita to replace M. cylindracea [sic] non Sow- 
erby 1846. Lectotype BMNH 1964300 selected by 
Kay (1965: 77, pi. 13, figs. 5, 6) is the syntype figured 
by Reeve (1865, Conchologia Iconica 15: Marginella, 
pi. 21, fig. 108) as Marginella peasei replacing polita 
Pease non Carpenter 1857; paralectotypes MCZ 
24965. 

cylindrica, Truncatella 
' 1865a, PZS for 1864: 676 [nomen nudum]. MCZ 181051. 

cylindricus, Triphorus 

' 1871a, PZS for 1870: 776 (Apaiang Island). Not lo- 
cated in BMNH, teste Way (personal communica- 
tion). 

cylindricus, Triton Plate 10, Figure 17 

' 1868g, AJC 4: 94, pi. 11, fig. 9 (Tahiti). Lectotype, 
here selected, MCZ 239749; paralectotypes MCZ 
288003; not located in ANSP. 

cytharoides 'Pease' Pace, Columbella 

1 902, Proceedings of the Malacological Society of London 
5: 74 [nomen nudum]. 

daedalea, Cithara [Cythara] Plate 6, Figure 1 1 

1868a, AJC 3: 218, pi. 15, fig. 13 (Paumotus); 1868h, 
AJC 4: 105 as Cythara debilis Pease to replace C. dae- 
dalea 1868 non Reeve 1846. Lectotype, here selected, 
ANSP 15663; paralectotype ANSP 391052; paralec- 
totypes MCZ 231920 and 231921. 

debilis, Atys 

1860a, PZS 28: 20 (Sandwich Islands). Lectotype 
BMNH 1961197 selected by Kay (1965: 11, pi. 1, 
figs. 5, 6); probable paralectotypes MCZ 31713 
though labeled as from Tahiti. See: Martens and 
Langkavel (1871, Donum Bismarckianum, p. 53), who 
also received specimens from Pease thus labeled. 

debilis, Cythara 

1868h, AJC 4: 105. New name for Cithara daedalea 
Pease 1861 non Reeve 1846. See under: daedalea, 
Cithara [Cythara]. 

debilis, Marginella 

1871e, AJC 7: 22. New name for Marginella oryza 
Pease 1860 non Lamarck 1822. 

debilis, Odostomia Plate 8, Figure 15 

1868d, AJC 3: 292, pi. 24, fig. 21 (Howland [Island]). 



Lectotype, here selected, MCZ 297933; not located 
in ANSP. 

decussata, Cithara [Cythara] Plate 6, Figure 7 

1868a, AJC 3: 217", pi. 15, fig. 10 ([Anaa Island] Pau- 
motus). Lectotype, here selected, ANSP 15651; para- 
lectotype ANSP 391051; paralectotypes MCZ 231922 
and 231923. 

decussata, Scalaria Plate 9, Figure 14 

1868d, AJC 3: 289, pi. 24, fig. 10 (Hawaii [Island]). 
Lectotype ANSP 19585 selected by DuShane (1988 
(7): 4, fig. [not numbered]) and refigured (1990: 7, 
fig. 30); paralectotype ANSP 391064; paralectotypes 
MCZ 187529 and 187530. 

decussata, Turbonilla 

1861b, PZS 28: 438 (Sandwich Islands). Lectotype 
BMNH 1962842 selected by Kay (1965: 68, pi. 5, 
figs. 15, 16). 

decussatula. Helix 

1866b, AJC 2: 291 (Sandwich Islands); 1871d, PZS 
for 1871: 474 as Pitys (Molokai). Specimens iden- 
tified by Pease [idiotypes] from Waimea or Wahimi, 
Kauai MCZ 17273 and 17274; not located in ANSP 
by Baker (1963: 232) or elsewhere by Solem (1976: 
377). 

deformis. Sty lifer Plate 9, Figure 15 

1868d, AJC 3: 293, pi. 24, fig. 23 (Paumotus). Lec- 
totype, here selected, ANSP 19836 is the figured 
type; three paralectotypes ANSP 391040; paralec- 
totypes MCZ 248839 and 248840. 

delicata, Narica Plate 8, Figure 3 

1868d, AJC 3: 282, pi. 23, fig. 25 (Paumotus). Lec- 
totype, here selected, ANSP 40201; paralectotypes 
MCZ 231415. 

delicatus, Pleurobranchus 

1861d, PZS for 1861: 245 (no locality); 1868e, AJC 
4: 79, pi. 9, fig. 1 (Huaheine). Possible syntype MCZ 
297873 though labeled as from Tahiti; not located 
in BMNH by Kay (1965: 86). 

dentata, Tornatellina Plate 4, Figure 9 

1871d, PZS for 1871: 460 (Hawaii [Island]). Lecto- 
type [so labeled], here selected, MCZ 28918; para- 
lectotypes MCZ 175730. 

dentifera, Vertigo 

1871d, PZS for 1871: 462 (Roratonga [error for Ai- 
tutaki. Cook Islands, teste Garrett (1881: 401), who 
collected the specimens]). Lectotype, here selected, 
MCZ 258352 figured by Pilsbry and Cooke (1920: 
329, pi. 30, fig. 14); paralectotype MCZ 48314. 

depressa, Siphonaria Plate 1, Figure 5 

1863d, PZS for 1862: 279 (Pacific Islands); 1868g, 
AJC 4: 98, pi. 1 1, fig. 23 (Apaiang Island). Lectotype 
ANSP 22199 selected by Baker (1964: 159); not lo- 
cated in BMNH by Kay (1965: 86). 

depressiformis. Helix Plate 3, Figure 1 

1865b, PZS for 1864: 670 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 475 (Tahiti). Lecto- 
type, here selected, MCZ 17342; paralectotypes MCZ 
297962. "Is Trochomorpha swainsoni (Pfeiffer), a Raia- 
tean (Society Island) species," teste Pilsbry and Cooke 
(1922: 17). 

depressiformis, Vitrina 

1865b, PZS for 1864: 675 [nomen nudum]. 



12 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



deshayesii, Neritina 

1868h, AJC 4: 130. New name for Neritma sand- 
wichensis Deshayes 1838 non Philippi 1843. 

discoidea, Helicina Plate 5, Figure 6 

1865a, PZS for 1864: 676 [nomen nudum]; 1868b, AJC 
3: 226 (Tahaa). Holotype ANSP 14398a (now 14398) 
is the measured type, teste Baker (1964: 161); prob- 
able paratypes MCZ 297927 though labeled as from 
Raiatea. 

discoidea, Torinia Plate 9, Figure 24 

1868g, AJC 4: 102, pi. 12, fig. 18 (Paumotus). Lec- 
totype, here selected, ANSP 38804; paralectotype 
ANSP 391039; paralectotype MCZ 297941. 

dispar, Neritina Plate 8, Figure 8 

1868d, AJC 3: 285, pi. 24, fig. 3 (Roratonga). Lec- 
totype ANSP 37714 selected by Baker (1964: 160); 
nine paralectotypes ANSP 358497; paralectotypes 
MCZ 73472. 

distans. Helix 

1866b, AJC 2: 290 (Sandwich Islands). Not located 
in ANSP. 

distorta, Leiostraca 

1861b, PZS 28: 438 (Sandwich Islands). Holotype 
BMNH 1962841 figured by Kay (1965: 67, pi. 11, 
figs. 12, 13); paratype MCZ 31706 figured by Pilsbry 
(1917: 229, figs. 13c, d). 

dubia, Partula affinis 

1865b, PZS for 1864: 675 [nomen nudum]. 

dubia Tease' Garrett, Partula otaheitana 

1865b, PZS for 1864: 675 [jwmen nudum]; 1884, J ANSP 
(2) 9: 49 (valley several miles from Papinoo [Tahiti]). 
Syntypes ANSP from Pease figured by Pilsbry (1909: 
188, pi. 25, figs. 10, 11) [not in Baker, 1963]; syntypes 
MCZ 25315. 



elegans, Clathurella 

1860b, PZS 28: 144 (Sandwich Islands). Lectotype 
BMNH 1961166 selected by Kay (1965: 21, pi. 2, 
figs. 21, 22; pi. 8, fig. 6); paralectotype MCZ 50009. 

elongata, Hydrocaena 

1865b, PZS for 1864: 676 [nomen nudum]. 

elongata, Partula 

1865b, PZS for 1864: 676 [nometi nudum]; 1866a, AJC 
2: 196 ([Vianae Ravine] Moorea); 1867a, AJC 3: 81, 
pi. 1, fig. 2. Lectotype ANSP 59477a selected by 
Baker (1963: 205) is the figured type; paralectotypes 
MCZ 24899, 24921, 25127, and 25294. 

elongata, Realia 

1868b, AJC 3: 225 (Raiatea); 1869a, JdeC 17: 152, pi. 
7, fig. 4 as Omphalotropis. Holotype in MNHN, teste 
Fischer-Piette (1950: 72); Baker (1964: 178), appar- 
ently unaware of this, selected as lectotype ANSP 
13350a; paratypes MCZ 187861 and 187862. 

elongata, Succtnea Plate 4, Figure 14 

1870a, JdeC 18: 96 (Kauai). Lectotype, here selected, 
MCZ 161665, probable measured type. 

elongata, Syphonota 

1860a, PZS 28: 24 (Sandwich Islands). Syntypes MCZ 
31442 and 298486; not located in BMNH by Kay 
(1965: 84). 



elongata, Turbonilla Plate 10, Figure 22 

1868d, AJC 3: 293, pi. 24, fig. 22 (Paumotus). Lec- 
totype, here selected, MCZ 10537; paralectotypes 
MCZ 10539; not located in ANSP. 

elongata, Volutella 

1868d, AJC 3: 281, pi. 23, fig. 23 (Fanning Island). 
Not located in ANSP. 

erecta, Laminella 

1869c, JdeC 17: 174 (Maui). Holotype in MNHN 
figured by Fischer-Piette (1950: 73, fig. 53); para- 
types MCZ 23338. 

ericea, Mitra 

1860b, PZS 28: 146 (Sandwich Islands); 1869f, AJC 
5: 85 is Mttra turgida Reeve 1845. Lectotype BMNH 
1961161 selected by Kay (1965: 28, pi. 3, figs. 3, 4); 
paralectotypes MCZ 260605. 

exaratus, Rhizochilus 

1861a, PZS 28: 399 (Sandwich Islands); 1868h, AJC 
4: 115 is Coralliophda deformis Lamarck 1816. Lec- 
totype BMNH 1961177 selected by Kay (1965: 41, 
pi. 4, figs. 10, 11); paralectotypes MCZ 297936. 

exilis, Clathurella 

1860b, PZS 28: 144 (Sandwich Islands). Lectotype 
BMNH 1961166 selected by Kay (1965: 21, pi. 2, 
figs. 21, 22; pi. 8, fig. 6); paralectotype MCZ 50009. 

exilis, DrUlia 

1868a, AJC 3: 220, pi. 15, fig. 19 (Tahiti). Lectotype, 
here selected, ANSP 15690 is the figured type; para- 
lectotype ANSP 316068. 

exilis, Eulima Plate 7, Figure 11 

1863b, PZS for 1862: 242 (Pacific Islands); 1868d, 
AJC 3: 294, pi. 24, fig. 25 (Paumotus). Lectotype, 
here selected, MCZ 187831; paralectotype MCZ 
187832; not located in BMNH by Kay (1965: 86). 

exilis, Trochus Plate 10, Figure 26 

1868d, AJC 3: 286, pi. 24, fig. 7 (Paumotus). Lecto- 
type, here selected, MCZ 104617; paralectotype MCZ 
303193; not located in ANSP. 

expansa, Auriculella 

1868f, JdeC 16: 343, pi. 14, fig. 8 (Hawaiian Islands). 
Holotype in MNHN, teste Fischer-Piette (1950: 71); 
paratypes MCZ 45155. 

expansa, Partula 

1871f, AJC 7: 26, pi. 9, fig. 3 (Tutuila). Holotype 
ANSP 59453, teste Baker (1963: 205). 

extensa, Leptachatina 

1870a, JdeC 18: 92 (Kauai). Not mentioned by Fi- 
scher-Piette (1950: 74). 

faba, Helicina 

1865a, PZS for 1864: 676 [nomen nudum]; 1868b, AJC 
3: 226 [nomen nudum] (neighboring island near Ta- 
haa). 

faba 'Pease' Garrett, Helicina 

1884, JANSP (2) 9: 105, pi. 3, figs. 61, 61a, b (Raiatea 
and Moorea). Holotype ANSP 14546a, teste Baker 
(1964: 161). 

fabrefacta, Helix Plate 3, Figure 2 

1865a, PZS for 1864: 669 ([Raiatea]). Lectotype, here 
selected, MCZ 17238; paralectotypes MCZ 176568. 
The neotype USNM 42427 selected by Solem (1976: 



MoLLUSKS Described by W. H. Pease • Johnson 13 



363) but not figured is considered here to be invalid 
since authentic types are available. 

fasciata, Partula 

1865a, PZS for 1864: 675 [nomcu nudum]; 1866a, AJC 
2: 202 (Marquesas Islands) is Partula ganymedes Pfeif- 
fer, teste Pease (1866, AJC 2: 293). Syntypes MCZ 
25113 and 25324. 

fascuita, Planaxis 

1868g, AJC 4: 102, pi. 12, fig. 17 (Paumotus). Ho- 
lotype ANSP 18286, only specimen; specimens sub- 
sequently identified by Pease [idiotypes] MCZ 
187838. 

ficta, Helix 

1865a, PZS for 1864: 669 (no locality, Tahaa [teste 
Garrett 1884: 38]). Lectotype USNM 24213 selected 
by Solem (1976: 362 [not figured]); paralectotypes 
MCZ 17240. 

filocostata, Pitys 

1871d, PZS for 1871: 454 (Kauai). Not located in 
BMNH, teste Mordan personal communication). 

fimbriatus, Euchelus 

1861c, PZS 28: 435 (Sandwich Islands). Holotype 
BMNH 1962823 figured by Kay (1965: 58, pi. 8, figs. 
14, 15); paratype MCZ 89896. 

fisheri Tease' Paetel, Dolahrifera 

1888, Catalog der Conchylien-Sammlung (Berlin) 1: 635 
[nomen nudum]. 

flammea, Rissoa Plate 9, Figure 7 

1868d, AJC 3: 297, pi. 24 [not 14], fig. 33 (Caroline 
[Islands]). Lectotype MCZ [not ANSP] 178868 [nor 
MCZ 178863 as on plate caption] selected by Ponder 
and de Keyzer (1992; 1058, fig. 3C) labeled as ho- 
lotype; six paralectotypes MCZ 178867 [not 178861] 
borrov/ed from the MCZ in 1990 but, "not exam- 
ined"; not located in ANSP. 

flammulata, Mitra 

1868a, AJC 3: 212 (Sandwich [Islands] and Pau- 
motus). Not located in ANSP. 

flammulata, Tnphoris 

1861b, PZS 28: 434 ([Haena, Kauai] Sandwich Is- 
lands). Lectotype BMNH 1961175 selected by Kay 
(1965: 52, pi. 6, figs. 15, 16); paralectotypes MCZ 
50065 and 50066. 

flai'escens, Helicwa 

1868b, AJC 3: 228, pi. 15, fig. 25 (Mangaia [Cook 
Islands]); 1871d, PZS for 1871: 476; here Pease ad- 
mits that Helicina flavescens is a redescription of his 
H. paafica. Holotype ANSP 14401a, teste Baker (1964: 
161); paratypes MCZ 297928. 

formosa, Cylmdra Plate 7, Figure 1 

1868d, AJC 3: 271, pi. 23, fig. 1 (Ascension [Island]). 
Holotype MCZ 260605 figured by Cernohorsky 
(1991: 132, pi. 130, fig. 2); not located in ANSP. 

formosa 'Pease' Garrett, Partula 

1884, JANSP (2) 9: 60, pi. 3, fig. 49 (Fatimu on the 
southwest part of Raiatea). Holotype ANSP 59453 
refigured by Pilsbry (1909: 218, pi. 20, fig. 5); para- 
types MCZ 25193. 

fortiplicata, Tumcula (Costellana) Plate 10, Figure 21 

1868a, AJC 3: 213, pi. 15, fig. 3 (Paumotus). Lecto- 



type, here selected, ANSP 28844; paralectotype 
ANSP 391043; paralectotypes MCZ 260609. 
foveolatus, Murex 

1869e, AJC 5: 83, pi. 8, fig. 3 (Gulf of [Golfo de] 
California, La Paz [Baja California Sur, Mexico]). 
Tryon, 1880, MofC (1)2: 129 as Murex peasei to re- 
place M. foveolatus non Hinds 1844. Lectotype ANSP 
36144 selected by Myers and D'Attilio (1989: 155, 
figs. 1, 2); two paralectotypes MCZ 304068. 

fragilts, Hydrocena 

1861c, PZS 28: 439 (Ebon Island); 1869a, JdeC 17: 
145, pi. 7, fig. 6 as Omphalotropis. Holotype in MNHN, 
teste Fischer-Piette (1950: 72); paratypes MCZ 297957. 

fragilis, Volvatella 

1860a, PZS 28: 20 (Sandwich Islands); 1868e, AJC 
4: 73, pi. 7, fig. 4. Holotype BMNH 1962753, teste 
Kay (1965: 12 [not figured]). 

fratercula, Helix 

1867b, AJC 3: 104. New name for Helix sculptilis 
Pease 1865 non Bland 1858. See under: sculptilis, 
Helix. 

fratercula, Pitys 

1871d, PZS for 1871: 452 [nomen nudum] (Hervey 
Islands). MCZ 176558 and 297860. 

frwola. Helix 

1866b, AJC 2: 290, pi. 21, fig. 3 (Polynesia); 1871d, 
PZS for 1871: 475 as Helicopsis (Oualan [Island]). 
Holotype ANSP 49296a, teste Baker (1963: 334); 
paratypes MCZ 135675. 

fucata, Scalaria 

1861a, PZS 28: 400 (Sandwich Islands). Lectotype 
BMNH 1961168 selected by Kay (1965: 43, pi. 6, 
figs. 11, 12); paralectotypes MCZ 187526. 

fucata, Triphoris 

1861b, PZS 28: 433 ([Kauai] Sandwich Islands). Lec- 
totype BMNH 1961171 selected by Kay (1965: 51, 
pl. 6, figs. 13, 14); paralectotypes MCZ 50067 and 
73736. 

fucatum, Cerithium 

1861b, PZS 28: 432 (Sandwich Islands). Lectotype 
BMNH 1962800 selected by Kay (1965: 46, pi. 5, 
figs. 7, 8). 

fusca, Dolabifera 

1868e, AJC 4: 76, pl. 8, fig. 4 ([Tahiti] Polynesia). 
Syntype MCZ 297870. 

fusca, Partula 

1865b, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 193 (Raiatea). Syntypes MCZ 25363; not located 
in ANSP by Baker (1965: 205). 

fusca, Vitrina 

1868), AJC 4: 155, pl. 12, fig. 6 (Marquesas [Islands]). 
Holotype ANSP 49270a, teste Baker (1963: 234). 

fuscata. Helix 

1865a, PZS for 1864: 675 [nomen nudum]. 

fuscata 'Pease' Pilsbry, Trochomorpha 

1896, Nautilus 9: 120 (Ponape, Caroline Islands). Ho- 
lotype ANSP 1934, teste Baker (1963: 238). 

fuscescens, Strigatella 

1860b, PZS 28: 146 (Sandwich Islands). Lectotype 
BMNH 1961184 selected by Kay (1965: 29, pl. 3, 
figs. 7, 8). 



14 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



fuscolineata, Neptunea 

1860c, PZS 28: 189 (Corea [Korea] Sea). Holotype 
BMNH 1964277, teste Kay (1965: 35, pi. 12, figs. 14, 
15). 

fuscomaculata, Clathurella 

1860b, PZS 28: 144 (Sandwich Islands). Lectotype 
BMNH 1961153 selected by Kay (1965: 23, pi. 2, 
figs. 7, 8). 

fuscomaculata, Cypraea 

1865d, PZS for 1865: 515 (Islands of the Central 
Pacific); 1868g, AJC 4: 95, pi. 1 1, figs. 10, 1 1 (Apaian 
Island [Garrett, 1879: 113, stated that he collected 
the types from the outer reefs of the Kingsmill 
Islands]). Lectotype BMNH 1964316 selected by Kay 
(1965: 83, pi. 14, figs. 1, 2). 

fusconigra, Vexilla 

1860b, PZS 28: 141 (Sandwich Islands); 1868h, AJC 
4: 115. Lectotype BMNH 1961164 selected by Kay 
(1965: 13, pi. 4, figs. 3, 4). 

fuscus Thilippi' Pease, Melampus 

1865a, PZS for 1864: 676 [rwmen nudum]. 

fusiformis, Columbella 

1868h, AJC 4: 122. New name for Columbella pusilla 
Pease 1862 non Sowerby 1844. See under: pusilla, 
Columbella. 

fusiformis, Conus 

1861a, PZS 28: 398 (Sandwich Islands); 1868h, AJC 
4: 126 non Fischer 1807. Changed to: Conus parvus. 
Lectotype BMNH 1962788 selected by Kay (1965: 
38, pi. 10, fig. 12); paralectotypes MCZ 197346 and 
197347. 

fusiformis, Engina 

1865d, PZS for 1865: 513 (Islands of the Central 
Pacific); 1868d, AJC 3: 273, pi. 23, fig. 5 (Howland 
Island). Lectotype BMNH 1964309 selected by Kay 
(1965: 81, pi. 13, figs. 15, 16); paralectotypes MCZ 
297954. 

fusiformis, Mitropsis 

1868a, AJC 3: 212, pi. 15, fig. 2 (Paumotus). Lecto- 
type, here selected, ANSP 28756 is the figured type. 

galba, Haminea 

1861b, PZS 28: 432 (Sandwich Islands). Lectotype 

BMNH 1961 194 selected by Kay (1965: 8, pi. 1, figs. 

11, 12). 
garrettii, Cythara 

1860b, PZS 28: 147 (Sandwich Islands); 1868h, AJC 

4: 105. Lectotype BMNH 1962780 selected by Kay 

(1965: 32, pi. 10, fig. 11); paralectotypes MCZ 49999 

and 298242. 
garrettii, Fossar 

1868h, AJC 4: 128. New name for Adeorbis costata 

Garrett 1857 non Philippi 1844. 
garrettii, Murex 

1868h, AJC 4: 103. New name for Murex exiguus 

Garrett 1857 non Broderip 1832. 
garrettii, Partula 

1865a, PZS for 1864: 672 (Islands of the Central 

Pacific [Vaioara on the west coast of Raiatea, teste 



Garrett, 1884: 56]). Syntype ANSP figured by Pils- 
bry (1909: 228, pi. 21, fig. 15), but not located by 
Baker (1963); syntypes MCZ 24894 and 25306. 
gibbus, Latirus Plate 7, Figure 16 

1865c, PZS for 1865: 54 (Pacific Islands); 1868d, AJC 
3: 279, pi. 23, fig. 17 (Howland Island). Lectotype, 
here selected, MCZ 261182; paralectotype MCZ 
303192; not located in BMNH by Kay (1965: 86) or 
in ANSP. 
glabra, Mitra 

1868d, AJC 3: 272, pi. 23, fig. 2 (Ascension [Island]); 
1869f, AJC 5: 85 is Mitra lubrica Pease 1869, new 
name for M. glabra Pease 1868 non Swainson 1821 
non Risso 1826. Lectotype, here selected, ANSP 
28854 is the figured type. 
globosa 'Pease' Pilsbry, Partula 

1865a, PZS for 1864: 675 [rwmen nudum]; 1909, MofC 
(2)20: 224. Manuscript name listed under the syn- 
onymy of Partula heba (Pfeiffer) (south end of Ra- 
iatea, Garrett ms); MCZ 25275. 
gracile, Cerithium 

1861b, PZS 28: 432 (Sandwich Islands). Lectotype 
BMNH 1961173 selected by Kay (1965: 45, pi. 5, 
figs. 5, 6). 
gracilior 'Pease' Hartman, Partula 

1881, BMCZ 9: 183 ([Moorea]). Syntype MCZ 25059. 
gracilis, Blauneria 

1860b, PZS 28: 145 (Sandwich Islands). Lectotype 
BMNH 1962770 selected by Kay (1965: 26, pi. 9, fig. 
3); paralectotypes MCZ 297790. 
gracilis, Citharopsis Plate 6, Figure 13 

1868g, AJC 4: 97, pi. 11, fig. 20 (Paumotus). Lecto- 
type, here selected, ANSP 16921. 
gracilis, Mucronalia Plate 7, Figure 24 

1868d, AJC 3: 295, pi. 24, fig. 27 (Tahiti). Lectotype 
MCZ 288506 selected by Waren (1980: 294, fig. 100); 
paralectotypes MCZ 248841 and ANSP 391065. 
gracilis, Nassa Plate 8, Figure 5 

1868d, AJC 3: 273, pi. 23, fig. 4 (Ascension [Island]). 
Lectotype, here selected, MCZ 228822; "Appears to 
be lost," Cernohorsky (1984: 66). 
gracilis, Odostomia 

1868d, AJC 3: 292, pi. 24, fig. 20 (Hawaii [Island]). 
Lectotype ANSP 19967 selected by Pilsbry (1917: 
321, fig. 18); paralectotypes MCZ 10489. 
gracilis, Partula 

1865b, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 197 (Tahitian Archipelago [upper portions of all 
the central valleys on both the east and west sides 
of Raiatea. It is more abundant in Tolo and Hapai 
Valleys than elsewhere. It occurs more rarely at 
Tahiti, teste Garrett, 1884: 70]); 1867a AJC 3: 81, pi. 
1, fig. 3. Not located in ANSP by Baker (1963: 205); 
syntypes MCZ 24905 and 25115 both from Raiatea. 
gracilis, Rissoa Plate 9, Figure 3 

1861b, PZS 28: 438 ([Kauai] Sandwich Islands); 1862b, 
JdeC 10: 382 as Rissoina. Lectotype, here selected, 
MCZ 178853; paralectotype MCZ 298460; not lo- 
cated in BMNH by Kay (1965: 86) or in MNHN by 
Fischer-Piette (1950: 20). 



MoLLUSKS Described by W. H. Pease • Johnson 15 



gracilis, Tornalcllina Plate 4, Figure 10 

1871d, PZS for 1871: 460 (Kauai). Lectotype, here 
selected, MCZ 302554. 

gracilis, Triphoris Plate 10, Figure 10 

1871a, PZS for 1870: 774 (Kauai). Lectotype, here 
selected, MCZ 50058; paralectotype MCZ 298493. 

gracilis, Triphoris 

1871a, PZS for 1870: 777 (Kauai). The description 
of this and the preceding Triphoris gracilis differ but 
slightly and though it appears that Pease described 
the same shells under the same name, Tryon (1872, 
AJC 7: 206) renamed the second gracilis, T. peasei. 

grandis, Syphonota 

1860a, PZS 28: 23 (Sandwich Islands). Syntype MCZ 
297866 [of questionable diagnostic value]; not lo- 
cated in BMNH by Kay (1965: 84). 

granifera, Fissurella 

1861d, PZS for 1861: 244 (Sandwich Islands). Lec- 
totype BMNH 1964278 selected by Kay (1965: 70, 
pi. 12, figs. 3, 4); paralectotypes MCZ 150701. 

granifera, Narica Plate 8, Figure 2 

1869d, AJC 5: 78, pi. 8, fig. 13 (Jarvis [Island]). Lec- 
totype, here selected, ANSP 37301; two paralecto- 
types ANSP 391041; paralectotypes MCZ 231414. 

grantferus, Vertagus 

1861b, PZS 28: 433 (Sandwich Islands). Holotype 
BMNH 1961208, teste Kay (1965: 49, pi. 5, figs. 9, 
10). 

granocostata, Scutellina Plate 5, Figure 1 

1868g, AJC 4: 100 (Hawaii [Island]). Lectotype, here 
selected, MCZ 83723 labeled as holotype. 

grariosus, Triphoris Plate 10, Figure 9 

1871a, PZS for 1870: 776 (Tahiti). Lectotype, here 
selected, MCZ 273207; paralectotypes MCZ 288954. 

granulata, Cypraea 

1863d, PZS for 1862: 278 (Pacific Islands). Lectotype 
BMNH 1964306 selected by Kay (1965: 79, pi. 14, 
figs. 17, 18); paralectotypes MCZ 297955 from An- 
naa Island, Tuamotu Archipelago. 

granulosa, Collonia Plate 6, Figure 20 

1868g, AJC 4: 92, pi. 11, fig. 4 (Ponape). Lectotype, 
here selected, MCZ 245264; paralectotypes MCZ 
297961; not located in ANSP. 

granulosa, Rissoina 

1862b, JdeC 10: 382, pi. 13, fig. 10 (Sandwich [Is- 
lands]). Holotype and paratype in MNHN, teste Fi- 
scher-Piette (1950: 20); paratype MCZ 178862. 

granulosus, Latirus Plate 7, Figure 17 

1868d, AJC 3: 279, pi. 23, fig. 10 [fig. 18 in error] 
(Paumotus). Lectotype MCZ 261181 selected by 
Cernohorsky (1987: 97, fig. 5) is the figured type; 
paralectotypes MCZ 297961. 

guppyi, Helicina 

1871d, PZS for 1871: 467. New name for Helicina 
humilis Guppy 1868 non Rousseau 1854. 

harpa, Clathurella 

1860b, PZS 28: 144 (Sandwich Islands). Lectotype 

BMNH 1961206 selected by Kay (1965: 22, pi. 2, 

figs. 23, 24). 
hawaiiensis, Perna 

1871f, AJC 7: 25. Invalid new name for Perna cali- 



f arnica Conrad 1834. Changed because calif arnica was 
a misnomer. 
hutchinsonii, Helicter Plate 4, Figure 2 

1862a, PZS for 1862: 7 (Maui). Lectotype, here se- 
lected, MCZ 45254; paralectotype MCZ 141500. 



ignominiosus 'Pease' Paetel, Achatinella 

1873, Catalog der Conchylien-Sammlung (Berlin), p. 
105 [nomem nudum]. 

imbricata, Vanikara 

1861b, PZS 28: 435 (Sandwich Islands). Holotype 
BMNH 1962820, teste Kay (1965: 57, pi. 8, fig. 11). 

imperforata 'Pease' Garrett, Partula 

1884d, JANSP (2) 9: 54, pi. 3, fig. 53 (Toloa and 
Hapai Valleys, west coast of Raiatea [Island], Society 
Islands). Holotype ANSP 59502a, teste Baker (1963: 
205); holotype and two paratypes figured by Pilsbry 
(1909: 221, pi. 20, figs. 13-15); paratypes MCZ 25262 
from Hapai Valley. 

imperforata, Pithys 

1870c, JdeC 18: 394 (Aitutake [Island, Cook Is- 
lands]); 1871, PZS for 1871: 453. Lectotype BPBM 
2322 selected by Solem (1976: 170, figs. 76e, f); para- 
lectotypes MCZ 17279. 

impressa 'Pease' Paetel, Achatinella 

1873, Catalog der Conchylien-Sammlung (Berlin), p. 
105 [nomen nudum]. 

incisa, Triphoris 

1861b, PZS 28: 434 ([Haena Point, Kauai] Sandwich 
Islands). Lectotype BMNH 1961151 selected by Kay 
(1965: 54, pi. 6, figs. 19, 20); paralectotypes MCZ 
50061 and 73738. 

"?" inconspicua, Helicina 

1865b, PZS for 1864: 676 [namen nudum]. 

inflexa, Eulima Plate 7, Figure 9 

1868d, AJC 3: 294, pi. 24, fig. 26 (Paumotus). Lec- 
totype, here selected, ANSP 59334 is the figured 
type; five paralectotypes ANSP 391056; paralecto- 
types MCZ 31703 and 31704. 

inflexa, Terebra szvainsonii 

1869d, AJC 5: 64. Based on specimen figured by 
Reeve, 1860, Conchologia Iconica 12, Terebra, pi. 22, 
fig. 118. Holotype BMNH 1979113 is the figured 
type. 

interlirata, Neritopsis Plate 8, Figure 9 

1868d, AJC 3: 282, pi. 23, fig. 26 (Annaa Island). 
Lectotype, here selected, MCZ 73479; not located 
in ANSP. 

intermedius, Tritan Plate 10, Figure 16 

1869d, AJC 5: 74 (Oahu). Lectotype MCZ 191331 
selected by Clench and Turner (1957: 217, pi. 122, 
fig. 2); paralectotypes MCZ 191330. 

interrupta, Daphnella 

1860b, PZS 28: 147 (Sandwich Islands). Lectotype 
BMNH 1962849 selected by Kay (1965: 34, pi. 5, fig. 
13); paralectotypes MCZ 50011. 



kauaiensis, Melania 

1870b, AJC 6: 7, pi. 3, fig. 6 (Kauai). Holotype ANSP 
26510, teste Baker (1964: 190); paratypes MCZ 74929. 



16 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



labiata Tease' Pilsbry, Partula 

1909, MofC (2) 20: 217, pi. 20, fig. 9 ([Vairahi Valley, 
Raiatea]). Holotype ANSP 59460, teste Baker (1963: 
205); paratype MCZ 24882. 

labiata Tease' Schmeltz, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1874, Mu- 
seum Godeffroy (Hamburg). Catalog 5: 207 [nomen 
nudum]. 

labiata, Succinea Plate 4, Figure 16 

1865a, PZS for 1864: 675 [nomen nudum]; 1868b, AJC 
3: 227 (Raiatea). Lectotype, here selected, MCZ 
298909; paralectotypes MCZ 31409; not located in 
ANSP by Baker (1963: 215). 

laevis, Gena Plate 6, Figure 2 

1868d, AJC 3: 283, pi. 23, figs. 7, 28, 29 (Tahiti). 
Lectotype, here selected, ANSP 40754; paralecto- 
types MCZ 297919. 

laevis, Lamellina Plate 2, Figure 16 

1865a, PZS for 1864: 672 (Islands of the Central 
Pacific); 1871, PZS for 1871: 473 (Tahiti). Lectotype, 
here selected, MCZ 154942 labeled "Hervey Is- 
lands"; syntype ANSP figured by Pilsbry (1915: 165, 
pi. 33, fig. 6), but not located in ANSP by Baker 
(1963). 

laevis, Leptachatina 

1870a, JdeC 18: 91 (Kauai); Crosse, 1876, JdeC 24: 
97, pi. 4, fig. 6. Holotype in MNHN, teste Fischer- 
Piette (1950: 149); paratypes MCZ 45173. 

laevis, Realia 

1865e, AJC 1: 289 (Polynesia); Tryon, 1866a, AJC 2: 
82, pi. 5, fig. 5 (Oualan Island); Pease, 1869a, JdeC 
17: 145 as Omphalotropis (Ponape [or] Ascension [Is- 
land]). Holotype ANSP 12225a, teste Baker (1964: 
178); paratypes MCZ 187921, 187922, and 187923. 

laminata. Helix Plate 3, Figure 4 

1866b, AJC 2: 292 (Sandwich Islands); 1871d, PZS 
for 1871: 474 as Endodonta (Kauai). Lectotype, here 
selected, MCZ 17233; paralectotypes MCZ 298477 
and BPBM, teste MCZ label; not located in ANSP 
by Baker (1963: 232) or elsewhere by Solem (1976: 
377). 

lateritia 'A. Adams' Pease, Assiminea 

1869b, JdeC 17: 164 [nomen nudum]. Error ioT Assimi- 
nea latericea H. and A. Adams 1864. 

lauta, Bullina 

1860a, PZS 28: 19 (Sandwich Islands). Holotype 
BMNH 1961201, teste Kay (1965: 6, pi. 1, figs. 3, 4). 

lauta, Drillia Plate 7, Figure 8 

1868a, AJC 3: 220, pi. 15, fig. 18 ([Anaa Island] Pau- 
motus). Lectotype, here selected, ANSP 15692 is the 
figured type; paralectotypes MCZ 49981. 

lauta, Terebra Plate 9, Figure 21 

1869d, AJC 5: 66 (Oahu). Holotype ANSP 33589 
figured by Tryon (1885: 33, pi. 10, fig. 91), teste 
Cernohorsky and Bratcher (1976: 139). 

lent a. Helix 

1865a, PZS for 1864: 675 [nomen nudum]. 

lenticulina, Helicina 

1865a, PZS for 1864: 676 [nomen nudum]. 

liberatus, Capulus 

1868d, AJC 3: 285, pi. 24, fig. 2 (Paumotus). Not 
located in ANSP. 



lignaria, Partula 
'^ 1865b, PZS for 1864: 671 (Islands of the Central 
Pacific [valley about two miles west of Fautaua [Ta- 
hiti], teste Garrett, 1884: 48]). Syntypes MCZ 25136, 
25142, and 25058. 

lineata, Alcyna Plate 6, Figure 4 

1869d, AJC 5: 69 ([Puuloa] Oahu). Holotype MCZ 
31724 figured by Pilsbry (1917: 212, pi. 15, fig. 4). 

Imeata, Columbella 

1861a, PZS 28: 399 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 85). 

lineolata, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1868b, AJC 
3: 224 (Tahiti). Holotype ANSP 59933a, teste Baker 
(1963: 205); paratypes MCZ 24933. 

lirata, Pleurotoma 

1869d, AJC 5: 68 (Oahu). Not located in ANSP. 

liratus, Latirus Plate 7, Figure 15 

1868i, AJC 4: 152, figured by Reeve, 1847, Conchol- 
ogia Iconica 4: Turbinella, pi. 12, figs. 61a, b (Mar- 
quesas [Islands]). Lectotype, here selected, MCZ 
302628; paralectotypes MCZ 297938; possible para- 
lectotypes BMNH 1979031; location of figured type 
unknown. 

lubrica, Mitra 

18691, AJC 5: 85. New name for Mitra glabra Pease 
1868 non Swainson 1821 non Risso 1826. See under: 
glabra, Mitra. 

lucida, Assiminea 

18691, JdeC 17: 166, pi. 7, fig. 10 (Annaa Island). 
Holotype in MNHN, teste Fischer-Piette (1950: 72); 
paratypes MCZ 74952. 

lucida, Leptachatina 

1869g, PZS for 1869: 650 [nomen nudum]; 1870a, JdeC 
18: 93 (Kauai). Not mentioned by Fischer-Piette 
(1950: 74). 

lucidus, Melampus Plate 4, Figure 22 

1 869d, AJC 5: 75 ([Honolulu] Oahu). Lectotype ANSP 
22284 selected by Baker (1964: 151); paralectotype 
ANSP 391058; paralectotypes MCZ 297789. 

lugubris, Partula "var.?" 

1865a, PZS for 1864: 672 (Islands of the Central 
Pacific [Hapai Valley, Raiatea, teste Garrett, 1884: 
77]); 1871d, PZS for 1871: 473. Syntypes MCZ 3641 
and 25288. 

lutea, Borsonia 

1860b, PZS 28: 143 (Sandwich Islands). Lectotype 
BMNH 1962756 selected by Kay (1965: 17, pi. 2, 
figs. 15, 16); paralectotypes MCZ 49978. 

luteostoma, Ranella 

1861a, PZS 28: 397 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 85). 

maculata, Engina lineata 

1869d, AJC 5: 76, pi. 8, fig. 12 (Apaian). Lectotype, 
here selected, ANSP 34538 is the figured type. 

maculata, Nerita Plate 8, Figure 7 

1868d, AJC 3: 286, pi. 24, fig. 6 (Tahiti). Lectotype, 
here selected, ANSP 37490; nine paralectotypes 
ANSP 391034. 



MoLLUSKs Described by W. H. Pease • Johnson 17 



niticulatuf. Tnphons Plate 10, Figure 13 

1871a, PZS for 1870: 777 (Kauai). Lectotype, here 
selected, MCZ 50069; paralectotype MCZ 298494. 

maculosa, Clathurella Plate 6, Figure 14 

1863b, PZS for 1862: 242 (Pacific Islands); 1868a, 
AJC 3: 219, pi. 15, fig. 16 (Paumotus). Lectotype, 
here selected, ANSP 48693; not located in BMNH 
by Kay (1965: 86). 

maculosa, Columbella 

1871e, AJC 7: 22. New name for Columbella dermes- 
toides Angas 1867 non Sowerby 1844. 

maculosa, Daphnella 

1860b, PZS 28: 148 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 85). 

maculosus Euchclus 

1863c, PZS for 1862: 243 (Pacific Islands); 1868g, 
AJC 4: 91, pi. 11, fig. 1 as Collotiia maculosa ([Anaa 
Island] Paumotus). Lectotype BMNH 1964298 se- 
lected by Kay (1965: 77, pi. 13, figs. 1, 2); figured 
paralectotype ANSP 38414; paralectotypes MCZ 
89828. 

mammillata, Succinea Plate 4, Figure 12 

1871b, PZS for 1871: 459 (Nukahiva). Lectotype, 
here selected, MCZ 155145; paralectotype MCZ 
298906; not located in BMNH, teste Mordan (per- 
sonal communication). 

marginatus, Pleurobranchus 

1860a, PZS 28: 25 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 84). 

marmorata. Purpura Plate 8, Figure 19 

1865d, PZS for 1865: 515 (Islands of the Central 
Pacific); 1868g, AJC 4: 92, pi. 11, fig. 5 (Apaian Is- 
land). Lectotype, here selected, MCZ 177824; para- 
lectotypes MCZ 302627; not located in BMNH by 
Kay (1965: 86). 

marmorata Tease' Martens and 

Langkavel, Triforis [sic] Plate 10, Figure 5 

1871, Donum Bismarckianum, p. 38, pi. 2, fig. 7 ([Kau- 
ai] Sandwich Islands). Lectotype, here selected, MCZ 
50055; paralectotypes MCZ 298492. 

marmorea. Bulla 

1861b, PZS 28: 431 (Sandwich Islands). Holotype 
BMNH 1961209 figured by Kay (1965: 44, pi. 1, figs. 
13, 14); paratypes MCZ 297878. 

marmorea 'Pease' Paetel, Dolabnfera 

1888, Catalog der Conchylietr-Sammlung (Berlin) 1: 635 
[nomen nudum]. 

marmorea, Margarita 

1861b, PZS 28: 435 (Sandwich Islands). Lectotype 
BMNH 1962824 selected by Kay (1965: 59, pi. 9, fig. 
5). 

marmoreus, Trochus Plate 9, Figure 25 

1868b, AJC 3: 287, pi. 24, fig. 9 (Paumotus). Lecto- 
type, here selected, ANSP 40614; paralectotype 
ANSP 391032; paralectotype MCZ 89894. 

marquesana, Helix Plate 3, Figure 5 

1868), AJC 4: 153, pi. 12, fig. 2 (Marquesas [Islands]). 
Lectotype, here selected, MCZ 302557; paralecto- 
types MCZ 297937; not located in ANSP by Baker 
(1963). 

megastoma 'Pease' Pilsbry, Partula 

1909, MofC (2) 20: 214, pi. 20, fig. 5 ([Raiatea: re- 



stricted to the higher portion of Haamoa Valley]). 
Holotype ANSP 5945a, teste Baker (1963: 205); para- 
type MCZ 25019. 

megastoma 'Pease' Schmeltz, Partula 

1865b, PZS for 1864: 675 [twmen nudum], 1874, Mu- 
seum Godeffroy (Hamburg). Catalog 5: 92 [nomen nu- 
dum]. 

micans 'Pease' Tryon, Columbella Plate 6, Figure 19 
1883, MofC (1) 5: 124, pi. 48, fig. 85 (Paumotus; Viti 
Isles). Lectotype, here selected, MCZ 304063 from 
the Paumotus. 

microstoma, Nassa 

1860b, PZS 28: 145 (Sandwich Islands). Holotype 
BMNH 1961458 figured by Kay (1965: 24, pi. 3, figs. 
5, 6); paratype MCZ 25341. 

microstoma 'Pease' Hartman, Partula 

1881, BMCZ 9: 184 ([Vairahi Valley] Raiatea). Syn- 
type ANSP figured by Pilsbry (1909: 233, pi. 27, fig. 
14); not located in ANSP by Baker (1963: 205); syn- 
types MCZ 25341. 

millecostata, Scalaria 

1861a, PZS 28: 400 (Sandwich Islands). Holotype 
BMNH 1961170, teste Kay (1965: 42, pi. 6, figs. 5, 
6). 

minimus, Tnphons Plate 10, Figure 7 

1871a, PZS for 1870: 774 (Howland Island; [Haena] 
Kauai). Lectotype, here selected, MCZ 50071; para- 
lectotypes MCZ 50070 and 298495 all from Kauai. 

modest a, Turricula Plate 10, Figure 20 

1868a, AJC 3: 212, pi. 15, fig. 6 ([Ponape] Polynesia). 
Lectotype, here selected, ANSP 28780; paralecto- 
types MCZ 260607. 

monilifera, Engina 

1860b, PZS 28: 142 (Sandwich Islands). Lectotype 
BMNH 1961460 selected by Kay (1965: 15, pi. 2, 
figs. 13, 14). 

monilifera, Pleurotoma 

1869d, AJC 5: 68 (Oahu). Not located in ANSP. 

monilifera. Turns 

1861a, PZS 28: 398 (Sandwich Islands). Holotype 
BMNH 1961196, teste Kay (1965: 39, pi. 5, figs. 17, 
18; pi. 8, fig. 7). 

moussoni, Omphalotropis 

1869a, JdeC 17: 147. New name for Omphalotropis 
ovata Mousson 1865 non Pease 1861. 

"?" multicolor, Helicina 

1865b, PZS for 1864: 676 [nomen nudum]. 

multicostatus, Fossar 

1861a, PZS 28: 398 (Sandwich Islands). Lectotype 
BMNH 1962790 selected by Kay (1965: 38, pi. 6, 
figs. 1, 2). 

multiplicata, Mitroidea 

1865d, PZS for 1865: 514 (Islands of the Central 
Pacific). Not located in BMNH by Kay (1965: 86) or 
elsewhere by Cernohorsky (1976: 471). 

multistriata 'Pease' Sowerby, Collonia 

1886, Thesaurus Conchyliorum 5: 212 [nomen nudum]. 

multistriatus 'Pease' Paetel Turbo 

1888, Catalog der Conchylien-Sammlung (Berlin) 1: 537 
[nomen nudum]. 



18 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



nebulosa, Borsonia 

1860b, PZS 28: 143 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 85). 

nebulosa, Omphalotropis Plate 4, Figure 4 

1872b, AJC 7: 197 ([Makela = San Cristobal Island] 
Solomon Islands). Lectotype, here selected, MCZ 
72347 labeled as holotype; paralectotype MCZ 72348. 

neglecta Nentina 

1861b, PZS 28: 435 (Sandwich Islands). Lectotype 
BMNH 1961186 selected by Kay (1965: 56, pi. 4, 
figs. 5, 6). 

neglectus, Cotius 

1861a, PZS 28: 398 (Sandwich Islands); 1871b, JdeC 
19: 99 is a variety of Conus flaindus Lamarck; not 
located in BMNH by Kay (1965: 85). 

neivcombii Tease' Brot, Melauia 

1872, Materiaux . . . des Melaniens III: 43 [nomen nu- 
dum]. Listed as a synonym of M. kauaiensis Pease, 
non neu'combii Lea 1866. 

newcombii, Mitra 

1869d, AJC 5: 69 (Oahu). Not located in ANSP. 

nigra, Philinopsis 
"l860a, PZS 28: 22 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 84). 

nigricans, Mitra 

1865d, PZS for 1865: 514 (Islands of the Central 
Pacific [Marquesas Islands]). Holotype BMNH 
1964312 figured by Kay (1965: 82, pi. 14, figs. 11, 
12); paratypes MCZ 260604. 

nigritella. Helix 

1865b, PZS for 1864: 675 [nomen nudum]. 

ntgropunctata, Hamwea 
'l868e, AJC 4: 71, pi. 7, fig. 1 (Raiatea). Probable 
syntypes MCZ 297876 though labeled from Tahiti. 

nitens. Vertigo Plate 2, Figure 21 

1861b, PZS 28: 439 (Ebon Island). Lectotype, here 
selected, MCZ 151650; paralectotypes MCZ 298456. 

nitida, Hydrocena 

1865b," PZS for 1864: 674 (Islands of the Central 
Pacific); 1869b, JdeC 17: 165, pi. 7, fig. 11 as/lss/m/wea 
(Huaheine). Holotype in MNHN, teste Fischer-Piette 
(1950: 72); paratypes MCZ 139120. 

nitida, Tornatellina Plate 4, Figure 7 

1861c, PZS 28: 439 (Ebon Island). Lectotype, here 
selected, MCZ 28921; paralectotypes MCZ 28922 
and 302556. 

nitidula, Mucronalia 

1861b, PZS 28: 437 (Sandwich Islands). Lectotype 
BMNH 1962835 selected by Kay (1965: 65, pi. 11, 
fig. 3); paralectotype MCZ 31711 figured by Pilsbry 
(1917: 226, fig. 12c); paralectotype MCZ 248842. 

nodicostata, Engina Plate 7, Figure 25 

1868d, AJC 3: 274, pi. 23, fig. 8 (Paumotus). Lecto- 
type, here selected, MCZ 260614; paralectotype MCZ 
260617; "type" ANSP 34543 with the note, "match- 
es the description but not the figure." 

nodifera, Drillia 

1860b, PZS 28: 145 ([Haena, Hawaii] Sandwich Is- 
lands). Lectotype BMNH 1961210 selected by Kay 
(1965: 24, pi. 5, figs. 1, 2); paralectotype MCZ 49982. 

nodulosa, Drillia 

1863d, PZS for 1862: 279 ([Sandwich Islands]). Ho- 



lotype BMNH 1964308 selected by Kay (1965: 80, 
pi. 14, figs. 9, 10); paratypes MCZ 49987. 

nodulosa, Engina 

1869d, AJC 5: 71, pi. 8, fig. 11 (Ebon Island). Lec- 
totype, here selected, ANSP 34513 is the figured 
type. 

nodulosa, Turricula (Pusia) Plate 10, Figure 24 

1868a, AJC 3: 214, pi. 15, fig. 5 (Paumotus). Lecto- 
type, here selected, ANSP 28713; four paralecto- 
types ANSP 391045; paralectotypes MCZ 260611. 

normalis. Helix Plate 3, Figure 9 

1865a, PZS for 1864: 669 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 475 (Moorea). Lecto- 
type, here selected, MCZ 11543; paralectotypes MCZ 
11544. 

nucea, Nassa 

1869d, AJC 7: 70, pi. 8, fig. 7 (Caroline Islands). Not 
located in ANSP. 

nucleola, Doris 

1860a, PZS 28: 29 (Sandwich Islands). Redescription 
and taxonomic reappraisal by Brodie and VVillan 
(1993: 124-133). 

nucleola 'Pease' Garrett, Partula 

1884, JANSP (2) 9: 72 ([Moorea]). Lectotype ANSP 
59531a selected by Baker (1963: 205); paralectotypes 
MCZ 24897, 24937, and 50854. 



oahuensis 'Pease' Brot, Melania 

1872, Materiaux . . . des Melaniens IIL- 43, pi. 3, fig. 
2 (Oahu). Type should be in Musee d'Histoire Na- 
turelle de Geneve [not confirmed]. 

obconica. Helix 

1865a, PZS for 1864: 669 (Islands of the Central 
Pacific [Raiatea, teste Garrett, 1884: 22, pi. 3, figs. 37, 
37a, b]); figured specimen was not located in ANSP 
by Baker (1964). Two syntypes ANSP 49337 from 
Pease; not located in BMNH, teste Mordan, personal 
communication. 

ohesa, Partula 

1868b, AJC 3: 223, pi. 15, fig. 12 (Polynesia). Ho- 
lotype, only specimen, not located in ANSP by Ba- 
ker (1963: 205). 

obliqua, Nassa Plate 8, Figure 6 

1865d, PZS for 1865: 513 (Islands of the Central 
Pacific). Lectotype, here selected, MCZ 228823 la- 
beled as probable measured holotype by Kay; not 
located in BMNH by Kay (1965: 86) or elsewhere 
by Cernohorsky (1984: 76). 

oblonga, Tornatellina Plate 4, Figure 6 

1865b, PZS for 1864: 673 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 473 (Tahiti). Lecto- 
type, here selected, MCZ 154941; paralectotypes 
MCZ 297947. 

oblonga, Tugalia 

1861b, PZS 28: 437 (Sandwich Islands). Lectotype 
BMNH 1962831 selected by Kay (1965: 64, pi. 11, 
figs. 4, 5); paralectotypes MCZ 298471. 

oceanica, Helicina Plate 5, Figure 9 

1868b, AJC 3: 226 (Kingsmill [Islands]). Lectotype, 
here selected, MCZ 176561; paralectotypes MCZ 
298458; not located in ANSP by Baker (1964: 162). 



MoLLUSKS Described by W. H. Pease • Johnson 19 



ochrostoma, Realia 

1865e, AJC 1: 287 (Polynesia); Tryon, 1866a, AJC 2: 
82, pi. 5, fig. 1 (Hervey Isles); Pease, 1869a, JdeC 
17: 147 as Omphalotropns. Holotype ANSP 13322a, 
teste Baker (1964: 178); paratypes MCZ 176573, 
187848, and 187849. 

olwacea, Carclia Plate 2, Figure 14 

1866b, AJC 2: 293 (Sandwich Islands); 1871d, PZS 
for 1871: 473 (Kauai). Holotype MCZ 57114, only 
specimen, teste Pease (1870c: 18: 402); two speci- 
mens subsequently identified by Pease [idiotypes] 
MCZ 23343 figured by Pilsbry and Cooke (1914: 16, 
pi. 9, figs. 11, 15). 

olivacea, Dolahrifera 

1860a, PZS 28: 22 (Sandwich Islands). Holotype 
BMNH 1964376, teste Kay (1965: 13 [not figured]). 

opprcssa, Trochomorpha mgritella Plate 2, Figure 6 

1870c, JdeC 18: 400 (Ponape); 1871d, PZS for 1871: 
457. Lectotype, here selected, MCZ 12187; paralec- 
totypes MCZ 12181 and 298478. 

ordmata 'Pease' Paetel, Mitra 

1887, Cat alogderConchiflien-Sammlung{BeThn) 1: 184 
[nomen nudum]. 

ornata, Citharopsis Plate 6, Figure 12 

1868g, AJC 4: 97, pi. 11, fig. 19 (Tahiti). Lectotype, 
here selected, ANSP 16919; paralectotype ANSP 
391046; paralectotypes MCZ 49994. 

oryza, Marginella 
"l860b, PZS 28: 147 (Sandwich Islands); 1871e, AJC 
7: 22 non Lamarck 1822. Changed to: Marginella 
debilis; not located in BMNH by Kay (1965: 85). 

oryza, Triphoris Plate 10, Figure 11 

"l871a, PZS for 1870: 776 (Kauai). Lectotype, here 
selected, MCZ 50072. 

oualanensis. Helix Plate 3, Figure 8 

1866b, AJC 2: 289, pi. 21, fig. 1 (Oualan [Island]). 
Lectotype, here selected, ANSP 47763; four para- 
lectotypes ANSP 391059. 

oualanensis 'Pease' Tryon, Melania Plate 4, Figure 20 
1866b, AJC 2: 299, pi. 20, fig. 4 (Oualan [Island]). 
Lectotype ANSP 26274 selected by Baker (1964: 191); 
paralectotypes MCZ 89614. 

ovalis, Hammea Plate 4, Figure 23 

1868e, AJC 4: 71, pi. 7, fig. 2 (Tahiti). Lectotype, 
here selected, MCZ 297877; paralectotypes MCZ 
303195. 

ovalis, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 194 (Raiatea). Holotype ANSP 59448a, teste Baker 
(1963: 205) figured by Pilsbry (1909: 172, pi. 19, fig. 
7); paratypes MCZ 25007. 

ovata, Engina Plate 7, Figure 4 

1865d, PZS for 1865: 513 (Islands of the Central 
Pacific); 1868d, AJC 3: 274, pi. 23, fig. 6 (Howland 
Island). Lectotype, here selected, ANSP 34536 is the 
figured type; paralectotypes MCZ 260598 and 
260599; not located in BMNH by Kay (1965: 86). 

ovata, Hydrocena 

1865a; PZS for 1864: 674 (Mangiers [error for Man- 
gaia. Cook Islands]); 1869a, JdeC 17: 148 as Omphalo- 
tropis. Not located in BMNH, teste Mordan (personal 



communication) or mentioned as in MNHN by Fi- 

scher-Piette (1950: 72). 
ovata, Mucronalia 

1861b, PZS 28: 437 (Sandwich Islands). Lectotype 

BMNH 1962810 selected by Kay (1965: 65, pi. 11, 

fig. 2). 
ovata, Succinea 

1865a, PZS for 1864: 675 [nomen nudum]; 1868b, AJC 

3: 227 "is Succinea papillata Pfeiffer." 



pachystoma, Labiella 

1869b, JdeC 17: 171 (Kauai). Measured holotype in 
MNHN, teste Fischer-Piette (1950: 73, fig. 52); para- 
types MCZ 45181. 

pacifica, Helicina Plate 2, Figure 3 

1865e, AJC 1: 291 (Polynesia); Tryon, 1866a, AJC 2: 
82, pi. 5, fig. 7 (Oualan Island). Lectotype ANSP 
14443 selected by Baker (1964: 162); paralectotype 
ANSP 358505. Redescribed by Pease. See under: 
flavescens, Helicina. 

pacifica, Marginella Plate 7, Figure 22 

1868d, AJC 3: 280, pi. 23, fig. 20 (Paumotus). Lec- 
totype, here selected, ANSP 29415; paralectotype 
ANSP 391023. 

pacifica, Pedicularia 

1865d, PZS for 1865: 516 (Islands of the Central 
Pacific); 1868g, AJC 4: 96, pi. 11, figs. 17, 18 (Apaian 
Island). Lectotype BMNH 1964312 selected by Kay 
(1965: 84, pi. 14, figs. 13, 14); paralectotypes MCZ 
297958. 

pacifica. Truncal ella Plate 10, Figure 19 

1868b, AJC 3: 230, pi. 15, fig. 27 (Oualan [Island]). 
Lectotype, here selected, MCZ 59799; two paralec- 
totypes ANSP 161791 ex MCZ. 

pallens, Trochomorpha trochiformis Plate 2, Figure 5 
1870c, JdeC 18: 399 (Tahiti; Moorea); 1871d, PZS 
for 1871: 457 (Tahaa). Lectotype, here selected, MCZ 
12188; paralectotypes MCZ 12188; both from Mo- 
orea. 

pallida 'Pease' Pace, Columbella 

1902, Proceedings of the Malacological Society of London 
5: 118 [nomen nudum]. 

pallida, Mitra 

1860b, PZS 28: 146 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 85); type lost, teste Cer- 
nohorsky (1976: 292). 

pallida, Taheitea [sic] Plate 9, Figure 17 

1868b, AJC 3: 229 (Tahiti; Huaheine). Lectotype, 
here selected, ANSP 12659 is the measured type 
specimen [as 13559a], teste Baker (1964: 175), no 
locality on label; four paralectotypes ANSP 391035; 
paralectotypes MCZ 178656 from Tahiti and 178660 
from Huaheine. 

pallidus, Triphoris Plate 10, Figure 8 

1871a, PZS for 1870: 774 (Kauai). Lectotype, here 
selected, MCZ 50073; paralectotypes MCZ 288955. 

parva, Engina 

1868d,'AJC 3: 276, pi. 23, fig. 11 (Paumotus). Lec- 
totype, here selected, ANSP 34542 is the syntype 



20 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



figured by Cernohorsky (1987: 100, figs. 17, 18); 
paralectotypes MCZ 49995. 

pari'a, Pterocyclos Plate 5, Figure 10 

1865e, AJC 1: 290 (Polynesia); Tryon, 1866a, AJC 2: 
82, pi. 5, fig. 8 (Hervey Isles); Pease, 1868j, AJC 4: 
158 (Aitutake, Hervey Isles). Lectotype ANSP 13406 
selected by Baker (1964: 178); two paralectotypes 
ANSP 372687; paralectotypes MCZ 141027, 141028, 
and 141029. 

parvidens, Helix 

1861d, PZS for 1861: 243 (Tahiti). Lectotype BPBM 
170888 selected by Solem (1976: 171, figs. 77a, b); 
paralectotypes MCZ 17267. 

parvula, Helicina 

1868), AJC 4: 156, pi. 12, fig. 10 (Atiu [Island, Cook 
Islands]). Holotype ANSP 14444a, teste Baker (1964: 
162); paratypes MCZ 297920. 

parvum, Cyclostoma Plate 2, Figure 15 

1865a, PZS for 1864: 674 (Islands of the Central 
Pacific). Lectotype, here selected, MCZ 74949; para- 
lectotypes MCZ 187914; both from Tahiti. 

parvus, Conus 

1868h, AJC 4: 126. New name for Conus fusiformis 
Pease 1861 non Fischer 1801 non Lamarck 1810. See 
under: fusiformis, Conus. 

paucicostata, Cithara [Ci/thara] Plate 6, Figure 8 

1868a, AJC 3: 217 (Tahiti). Lectotype, here selected, 
MCZ 231925. 

paucicostata, Clathurella 

1860b, PZS 28: 144 (Sandwich Islands). Holotype 
BMNH 1961158, teste Kay (1965: 22, pi. 2, figs. 11, 
12). 

paucicostata, Pithys Plate 2, Figure 7 

1870c, JdeC 18: 395 (Kauai). Lectotype, here select- 
ed, MCZ 17271; paralectotypes MCZ 298476; not 
mentioned by Fischer-Piette (1950: 76). 

paumotensis, Marginella Plate 7, Figure 23 

1868d, AJC 3: 281, pi. 23, fig. 22 (Paumotus). Lec- 
totype, here selected, ANSP 29497; paralectotype 
ANSP 391050; paralectotypes MCZ 297943. 

paumotensis, Scalaria Plate 9, Figure 16 

1868d, AJC 3: 289, pi. 24, fig. 11 (Paumotus). Lec- 
totype ANSP 19581 selected by DuShane (1988 (7): 
5, fig. [not numbered]) and refigured (1990: 9, fig. 
34); two paralectotypes ANSP 352473; paralecto- 
types MCZ 187394. 

paxillum, Cerithium 

1861b, PZS 28: 433 (Sandwich Islands). Lectotype 
BMNH 1964804 selected by Kay (1965: 47, pi. 10, 
fig. 6). 

peasei Martens and Langkavel, Columbella {Seminella) 
1871, Donum Bismarckianum, p. 23. New name for 
Cythara varia Pease 1860 non Columbella varia Sow- 
erby 1832. 

peasei Reeve, Marginella 

1865, Conchologia Iconica 15: Marginella, pi. 21, fig. 
108. New name for Marginella cylindrica non Sow- 
erby 1846 and M. polita Pease non Carpenter 1857. 
See under: cylindrica, Marginella. 

peasei Tryon, Triphoris 

1872d, AJC 7: 206. New name for T. gracilis Pease, 



1871: 777 non Pease, 1871: 774. See under: gracilis, 
Triphoris. 

pellucida, Columbella 

1861a, PZS 28: 399 (Sandwich Islands). Lectotype 
BMNH 1962794 selected by Kay (1965: 41, pi. 10, 
fig. 7). Is Columbella rorida Reeve, teste Pease (1868h: 
AJC 4: 122). 

pellucida, Partula 

1871d, PZS for 1871: 457 (Guadalcanal, Solomon 
Islands). Though not a type, MCZ 94837 from "Ysa- 
bel [Island] Solomon Is. (coll. Cox)" was figured by 
Hartman (1886: 35, pi. 2, fig. 17) as this species and 
by Pilsbry (1909: 297, pi. 36, fig. 6). The original 
lot, in the Hartman collection at the Carnegie Mu- 
seum 6246, consisted of one adult and two imma- 
ture examples. The former and one of the latter 
were transferred to the MCZ in 1935; not located 
in BMNH, teste Mordan (personal communication). 

pellucidus, Pleurobranchus 

1860b, PZS 28: 145 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 84). 

perfectus, Triphoris Plate 10, Figure 15 

1871a, PZS for 1870: 775 (Kauai). Lectotype, here 
selected, MCZ 302553; paralectotype MCZ 50075. 

perlonga. Vertigo 

1871d, PZS for 1871: 462 ([Nuuanu] Oahu). Holo- 
type MCZ 48063 figured by Pilsbry and Cooke (1920: 
258-259, pi. 23, figs. 1, 2). 

perplexa 'Pease' H. H. Smith, Partula 

1902, ACM 1: 463 (Huaheine). Syntypes MCZ 25358. 

perplexa, Scalaria Plate 9, Figure 1 1 

1868d, AJC 3: 288 (Hawaii [Island]). Lectotype [so 
labeled], here selected, MCZ 181978; paralectotypes 
MCZ 181979 and 181980. 

perversa 'Pease' H. H. Smith, Partula 

1902, ACM 1: 442 (Tahiti). Syntypes MCZ 25317. 

picea, Strigatella 

1860b, VZS 28: 146 (Sandwich Islands). Holotype 
BMNH 1962772 figured by Kay (1965: 28, pi. 9, fig. 
9). 

picta, Collonia Plate 6, Figure 21 

1868g, AJC 4: 91, pi. 11, figs. 2, 3 ([Anaa Island] 
Paumotus). Lectotype, here selected, MCZ 245268. 
Labeled, "Chosen [lectotype] by P. A. Maxwell, ms 
in prep. 1963" but apparently never published; 
paralectotypes MCZ 288001 and 288002; not located 
in ANSP. 

picta, Helicina 

1865b, PZS for 1864: 676 [nomen nudum]. MCZ 74428. 

plana, Libratula Plate 5, Figure 2 

1865d, PZS for 1865: 512 (Islands of the Central 
Pacific). Lectotype, here selected, MCZ 119162. 

planilabrum, Partula 

1865a, PZS for 1864: 671 (Islands of the Central 
Pacific [Haamene Valley, east coast of Tahaa, teste 
Garrett, 1884: 63]). Syntypes MCZ 25004 and 25298. 

plicatula, Turricula (Costellaria) Plate 10, Figure 27 
1868a, AJC 3: 213, pi. 15, fig. 4 (Paumotus). Lecto- 
type, here selected, ANSP 28845; four paralecto- 
types ANSP 391044; paralectotypes MCZ 260602. 



MoLLUSKS Described by W. H. Pease • Johnson 21 



plumbea, Plaiiaxis 

1861d, PZS for 1861: 244 (Sandwich Islands). Lec- 
totype BMNH 1964280 selected by Kay (1965: 71, 
pi. 13, figs. 11, 12); paralectotypes MCZ 39062, 
187835, and 207329. 

polita, Marginella 

1868d, AJC 3: 280, pi. 23, fig. 19 (Tarawa Island). 
New name for Margmella cylindracea [sic] Pease 1863 
non Sowerby 1846 non Carpenter 1857. See under: 
cylindrka, Marginella. 

polita, Odostowta Plate 8, Figure 10 

1868d, AJC 3: 291, pi. 24, fig. 17 (Tahiti). Lectotype, 
here selected, MCZ 10562; paralectotype MCZ 
298903; not located in ANSP. 

poryhyrostoma, A mastra 

1869c, JdeC 17: 172 (Oahu; Coll. [of] Pease). Two 
syntypes MCZ 45256, one of which was figured by 
Hyatt and Pilsbry (1911: 226, pi. 37, fig. 13). 

producta, Clathurella 

1860b, PZS 28: 143 (Sandwich Islands). Lectotype 
BMNH 1962761 selected by Kay (1965: 19, pi. 2, 
figs. 3, 4); paralectotypes MCZ 49974. 

producta, Omphalotropis 

1869a, JdeC 17: 151, pi. 7, fig. 8 (Tahaa); 1871d, PZS 
for 1871: 471 as Atropis (Raiatea; Tahaa). Holotype 
in MNHN, teste Fischer-Piette (1950: 72); paratypes 
MCZ 74932, 74933, and 187855 all from Tahaa. 

producta, Partula 

1865a, PZS for 1864: 671 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 473 (Tahiti). Syntypes 
MCZ 25016. 

producta, Ranella 

1861a, PZS 28: 397 (Sandwich Islands). Holotype 
BMNH 1961157, teste Kay (1965: 37, pi. 6, figs. 17, 
18). 

producta, Realia 

1865b, PZS for 1864: 673 (Islands of the Central 
Pacific); 1869a, JdeC 17: 151, pi. 7, fig. 8 as Om- 
phalotropis (Tahaa); 1871d, PZS for 1871: 476 (Ra- 
iatea) as Atropis. Holotype in MNHN, teste Fischer- 
Piette (1950: 72); paratypes MCZ 74943, 187855, and 
187856. 

propinqua 'Pease' Hartman, Partula 

1881, BMCZ 9: 185 (Tahaa). Syntypes MCZ 25124 
and 25308. 

propinqua, Terebra 

1869d, AJC 5: 66 (Hawaii [Island]). Not located in 
ANSP. 

prostrata. Helix 

1865a, PZS for 1864: 670 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 475 ("?" Lanai). Syn- 
types MCZ 17343 "are Planorbis opercularis Gould 
from the West Indies," teste Pilsbry and Cooke (1922: 
117). 

protea 'Pease' Schmeltz, Partula 

1865a, PZS for 1864: 675 [nomeyi nudum]; 1874, Mu- 
seum Godeffroy (Hamburg). Catalog 5: 92 (Raiatea) 
[nomen nudum]. MCZ 24996 and 25349. 

proximus, Helicter Plate 4, Figure 1 

1862a, PZS for 1862: 6 (Molokai). Lectotype, here 
selected, MCZ 25823. 



pudica, Mitra 

1860b, PZS 28: 146 (Sandwich Islands). Lectotype 
BMNH 1961190 selected by Kay (1965: 27, pi. 3, 
figs. 11, 12). 

pulchella, Clathurella 

1860b, PZS 28: 144 (Sandwich Islands). Lectotype 
BMNH 1962768 selected by Kay (1965: 22, pi. 2, 
figs. 19, 20); paralectotypes MCZ 50010. 

pulchra, Auriculella 

1868f, JdeC 16: 346, pi. 14, fig. 6 ([Oahu]). Holotype 
and four paratypes in MNHN, teste Fischer-Piette 
(1950: 71); paratypes MCZ 161609. 

pulchra, Partula varia 

1871d, PZS for 1871: 473 (Huaheine); Schmeltz, 1874, 
Museum Godeffroy (Hamburg). Catalog 5: 92; [both nom- 
ma nuda]. MCZ 24878 and 24920. 

punctata, Syphonota 

1868e, AJC 4: 77, pi. 9, fig. 2 (Huaheine). Probable 
syntype MCZ 297868 though labeled as from Tahiti. 

punctatus, Triphoris 

1871a, PZS for 1870: 775 (Annaa Island). Not located 
in BMNH, teste Way (personal communication). 

purus, Conus Plate 6, Figure 23 

1863d, PZS for 1862: 279 (Pacific Islands); 1871c, 
JdeC 19: 98 (Niihau Island). Holotype MCZ 72331, 
only specimen; not located in BMNH by Kay (1965: 
89). 

pusilla, Columbella 

1863b, PZS for 1862: 244 (Kingsmill Islands); 1868h, 
AJC 4: 1 22 as Columbella fusiformis to replace C. pusilla 
non Sowerby 1844. Lectotype BMNH 1964302 se- 
lected by Kay (1965: 78, pi. 13, figs. 9, 10); paralec- 
totypes MCZ 136617. 

pusilla, Cythara 

1860b,' PZS 28: 147 (Sandwich Islands). Lectotype 
BMNH 1962784 selected by Kay (1965: 33, pi. 10, 
fig. 14); paralectotypes MCZ 50003. 

pusilla, Distorsio 

1861a, PZS 28: 397. Holotype BMNH 1961155, teste 
Kay (1965: 37, pi. 3, figs. 15, 16). 

pusilla, Haminea 

1860a, PZS 28: 20 (Sandwich Islands). Lectotype 
BMNH 1962754 selected by Kay (1965: 9, pi. 9, fig. 
1); paralectotypes MCZ 297879. 

pusilla, Triton 

1861b, PZS 28: 434 (Sandwich Islands). Lectotype 
BMNH 1962818 selected by Kay (1965: 55, pi. 5, 
figs. 19, 20); paralectotypes MCZ 297942. 

pustulosus, Triphoris Plate 10, Figure 2 

1871a, PZS for 1870: 776 (Kauai). Lectotype, here 
selected, MCZ 50077. 

putillus, Turricula 

1865d, PZS for 1865: 514 (Central Pacific); 1868a, 
AJC 3: 214, pi. 15, fig. 24. Holotype BMNH 1964311, 
teste Kay (1965: 81, pi. 14, figs. 7, 8); paratypes MCZ 
260605. 

pyriformis, Marginella Plate 7, Figure 21 

' 1868d, AJC 3: 280, pi. 23, fig. 21 (Paumotus). Lec- 
totype, here selected, ANSP 29541; four paralec- 
totypes ANSP 391061; paralectotype MCZ 24968. 



22 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



pi/riformis, Voh^atella 

1868e, AJC 4: 73, pi. 7, fig. 5 (Huaheine). Syntypes 
MCZ 297944, mostly fragments. 

radiata, Aviciila Plate 1, Figure 4 

1863c, PZS for 1862: 244 (Kingsmill Islands). Lec- 
totype, here selected, MCZ 298466; paralectotypes 
MCZ 297882; not located in BMNH by Kay (1965: 
86). 

radiata Tease' Garrett, Partula 

1884, JANSP (2) 9: 74, pi. 3, fig. 45 (Hamoa Valley, 
east coast of Raiatea Island, Society Islands). Lec- 
totype ANSP 59409 selected by Baker (1963: 205) 
figured by Pilsbry (1909: 232, pi. 18, fig. 5); para- 
lectotypes MCZ 24957. 

radiata, Tectura 

1861b, PZS 28: 437 (Sandwich Islands). Lectotype 
BMNH 1962837 selected by Kay (1965: 66, pi. 11, 
figs. 6, 7). 

recta, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1868j, AJC 
4: 155, pi. 12, figs. 8, 9 ([Nukahiva] Marquesas [Is- 
lands]). Holotype ANSP 59789a fig. 8, teste Baker 
(1963: 205); paratypes MCZ 25338 and 25343. 

retkulatus, Pleurobranchus 

1860a, PZS 28: 25 (Sandwich Islands); 1864, PZS for 
1863: 510 is Pleurobranchus violaceus Pease 1864, new 
name for P. reticulatus Pease 1860 non Rang 1832 
non Kelaart 1859; not located in BMNH by Kay 
(1965: 84). 

retunsa. Helix 

1865b, PZS for 1864: 670 (Pacific Islands); 1871d, 
PZS for 1871: 475 (Tahiti). Lectotype BPBM 170913 
selected by Solem (1976: 412, figs. 178d-f); paralec- 
totypes MCZ 17224. 

robusta, Hi/drocaena 

1865b, PZS for 1864: 676 [twmen nudum]. 

robusta, Omphalotropis 

1869a, JdeC 17: 148, pi. 7, fig. 3 ([Raiatea]). Holotype 
in MNHN, teste Fischer-Piette (1950: 72); paratypes 
MCZ 74933 and 187853. 

robusta, 'Pease' H. H. Smith, Partula 

1865b, PZS for 1864: 675 [nomen nudum]; 1902, ACM 
1: 436 (Raiatea). Lectotype ANSP 59444a selected 
by Baker (1963: 205) figured by Pilsbry (1909: 248, 
pi. 18, fig. 14); paralectotypes MCZ 3651. 

robustus, Stylifer 

1861b, PZS 28: 437 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 85). 

robustus, Triphoris Plate 10, Figure 12 

1871a, PZS for 1870: 775 (Makaimo[Makemo Island, 
Taumotu Archipelago]). Lectotype, here selected, 
MCZ 73923; paralectotypes MCZ 298499. 

roratongensis, Pithys 

1870c, JdeC 18: 395 (Roratonga); 1871d, PZS for 
1871: 457. Lectotype BPBM 170885 selected by So- 
lem (1976: 174, figs. 77e, i) who changed it to rara- 
tongensis; paralectotypes MCZ 17345. 

rosacea, Gena Plate 6, Figure 1 

1868d, AJC 3: 284, pi. 24, fig. 1 (Paumotus). Lecto- 
type, here selected, ANSP 40756; two paralecto- 
types ANSP 391033; paralectotypes MCZ 297918. 



rosacea, Odostomia Plate 8, Figure 11 

1868d, AJC 3: 292, pi. 24, fig. 19 (Paumotus). Lec- 
totype, here selected, ANSP 19959; paralectotypes 
MCZ 391054. 

rosacea, Terebra 

1869d, AJC 5: 65 (Oahu). Not located in ANSP. 

rosea, Mucronalia 

1861b, PZS 28: 437 (Sandwich Islands). Holotype 
BMNH 1962809, teste Kay (1965: 64, pi. 11, fig. 1). 

rotella, Diadema 

1868), AJC 4: 158, pi. 12, fig. 13 (Atiu [Island, Cook 
Islands]). Holotype ANSP 13409a, teste Baker (1964: 
178). 

rotellina, Pithys 

1870c, JdeC 18: 393 (Aitutake [Island, Cook Is- 
lands]); 1871d, PZS for 1871: 453. Lectotype BPBM 
2312 selected by Solem (1976: 139, figs. 62e, f); para- 
lectotypes MCZ 176559. 

rubella, Succinea Plate 4, Figure 13 

1871d, PZS for 1871: 460 (Lanai). Lectotype, here 
selected, MCZ 161671; paralectotypes MCZ 298475. 

rubicunda, Helicina maugeriae Plate 5, Figure 12 

1865a, PZS for 1864: 676 [nomen nudum]; 1868d, AJC 
3: 227: 1871d, PZS for 1871: 466 (Raiatea). Lectotype 
ANSP 14512 selected by Baker (1964: 162) labeled 
as from Tahiti; paralectotypes MCZ 314014 from 
Raiatea. 

rubida, Catinella Plate 4, Figure 21 

1870a, JdeC 18: 97 (Kauai). Lectotype, here selected, 
MCZ 45252, probable measured type. 

rubida, Dentiora Plate 7, Figure 7 

1863b, PZS for 1862: 240 (Sandwich Islands). Lec- 
totype, here selected, MCZ 297951, probable mea- 
sured type; not located in BMNH by Kay (1965: 86). 

rubida, Neritina 

1865d, PZS for 1865: 514 (Islands of the Central 
Pacific); 1868d, AJC 3: 285, pi. 24, fig. 5 (Tahiti). 
Lectotype BMNH 1964313 selected by Kay (1965: 
82, pi. 14, figs. 15, 16); paralectotypes MCZ 89902; 
Baker (1964: 160) questions if ANSP 37675a is the 
figured type since the measurements are not close 
to those of Pease. 

rubra, Alcyna 

1861b, PZS 28: 436 (Sandwich Islands). Lectotype 
BMNH 1962828 selected by Kay (1965: 62, pi. 7, 
figs. 5, 6); paralectotypes MCZ 31720 and 205584. 

rubra, Odostomia Plate 8, Figure 12 

1868d, AJC 3: 291, pi. 24, fig. 18 ([Anaa Island] Pau- 
motus). Lectotype, here selected, ANSP 19955; para- 
lectotypes ANSP 391054; paralectotype MCZ 10488. 

rudis, Neritina 

1868d, AJC 3: 285, pi. 24, fig. 4 (Ponape). Holotype 
ANSP 37543a, teste Baker (1964: 160); paratypes MCZ 
73518. 

rufescens, Helicina 

1865a, PZS for 1864: 676 [nomen nudum]. MCZ 297926. 

rufus, Pleurobranchus 

1860a, PZS 28: 25 (Sandwich Islands). Syntype MCZ 
297872; not located in BMNH by Kay (1965: 84). 

rugata. Helix Plate 3, Figure 3 

'l866b, AJC 2: 291 (Sandwich Islands); 1871d, PZS 
for 1871: 474 as Endodonta (Maui). Lectotype, here 



MoLLUSKS Described by W. H. Pease • Johnson 



23 



selected, MCZ 17237; paralectotypes MCZ 298479; 
not located in ANSP by Baker (1963: 233) or else- 
where by Solem (1976: 377). 

rugulosa, Amastra 
\870a, JdeC 18: 95 (Kauai); Crosse, 1876, JdeC 24: 
99, pi. 1, figs. 4, 4a. Holotype and fragment of para- 
type in MNHN, teste Fischer-Piette (1950: 149 as 
second reference to A. A. rugulosa only); paratypes 
MCZ 45255. 

rugulosa, Hclicma Plate 5, Figure 7 

1868), AJC 4: 157, pi. 12, fig. 11 (Tahaa). Lectotype, 
here selected, MCZ 297922; paralectotypes MCZ 
298467; not located in ANSP by Baker (1964: 162). 

rugulosuni, Sistrum Plate 9, Figure 8 

'l868g, AJC 4: 93, pi. 11, fig. 7 (Howland [Island]). 
Lectotype, here selected, ANSP 36740; two para- 
lectotypes ANSP 391042; paralectotypes MCZ 
295581. 

rustica, Partula 

1865a, PZS for 1864: 675 [uomen nudum]; 1866a, AJC 
2: 199 (Tahitian Archipelago); 1867a, AJC 3: 81, pi. 
1, fig. 5; 1871d, PZS for 1871: 473 (Raiatea). Holo- 
type ANSP 59480 selected as lectotype by Baker 
(1963: 205); paralectotypes MCZ 25140 and 25302. 

rutella, Succinea 

1865b, PZS for 1864: 675[nomen nudum]. MCZ 155141. 



sagitta Tease' Pace, Columbella 

1902, Proceedings of the Malacological Society of London 
5: 132 [nomen nudum]. 

salt at a, Mitra Plate 7, Figure 18 

1865d, PZS for 1865: 512 (Islands of the Central 
Pacific). Lectotype MCZ 260613 [not 260605] se- 
lected by Cernohorsky (1976: 501, pi. 450, fig. 2); 
two paralectotypes MCZ 298462; not located in 
BMNH by Kay (1965: 86). 

sandivicensis, Dapbnella Plate 7, Figure 6 

1860b, PZS 28: 148 (Sandwich Islands). Lectotype, 
here selected, MCZ 49993; paralectotype MCZ 
298491; not located in BMNH by Kay (1965: 85). 

sandivicensis, Erato 

1860b, PZS 28: 146 (Sandwich Islands). Lectotype 
BMNH 1962776 selected by Kay (1965: 31, pi. 9, fig. 
10); paralectotypes MCZ 297959. 

sandwicensis, Marginella 

1860b, PZS 28: 146 (Sandwich Islands). Lectotype 
BMNH 1962778 selected by Kay (1965: 31, pi. 9, fig. 
11); paralectotype MCZ 24970. 

sandivicensis, Oliva 

1860b, PZS 28: 145 (Sandwich Islands). Lectotype 
BMNH 1961188 selected by Kay (1965: 25, pi. 3, 
figs. 9, 10). 

sandwicensis, Patella Plate 1, Figure 8 

1861b, PZS 28: 437 (Sandwich Islands). Lectotype, 
here selected, MCZ 304058; paralectotypes MCZ 
38803 and 150789; not located in BMNH by Kay 
(1965: 85). 

sandivicensis, Pedipes 

1860b, PZS 28: 146 (Sandwich Islands). Holotype 
BMNH 1962775 figured by Kay (1965: 30, pi. 9, fig. 
7); paratypes MCZ 74813. 



sandivicensis, Tornatina 

1860a, PZS 28: 19 (Sandwich Islands). Lectotype 
BMNH 1962751 selected by Kay (1965: 6, pi. 9, fig. 
4); paralectotypes MCZ 31712 with the note by Pils- 
bry "seems to be two species, but they are so worn 
I cannot be sure." 

sandwicensis, Turbo 

1861b, PZS 28: 436 (Sandwich Islands). Holotype 
BMNH 1961179, teste Kay (1965: 60, pi. 7, figs 7, 
8). 

sandwichensis, Columbella Plate 6, Figure 22 

1861d, PZS for 1861: 244 (Sandwich Islands); 1868h, 
AJC 4: 122 is Columbella turtunna Lamarck (1822). 
Lectotype, here selected, MCZ 297950; paralecto- 
types MCZ 298472; not located in BMNH by Kay 
(1965: 86). 

sandwichensis 'Pease' Nevill, Fossar 

1884, Hand List of Mollusca in the Indian Museum 2: 
165 [nomen nudum]. MCZ 242384. 

sandwichensis, Vitularia 

1861a, PZS 28: 397 (Sandwich Islands). Lectotype 
BMNH 1961182 selected by Kay (1965: 36, pi. 4, 
figs. 1, 2). 

scalariformis, "?" Cyclophorus 

1865e, AJC 1: 289 (Polynesia); Tryon, 1866a, AJC 2: 
82, pi. 5, fig. 6, as Pupoidea (Hervey Isles); Pease, 
187 Id, PZS for 1871: 465 as Palama (Caroline Is- 
lands). Holotype ANSP 13360, teste Baker (1964: 
165). 

scalariformis, Realia 

1865e, AJC 1: 288 (Polynesia); Tryon, 1866a, AJC 2: 
82, pi. 5, fig. 3 (Oualan Island); Pease, 1869a, JdeC 
17: 159 as Scallinella (Atiu). Holotype ANSP 13362a, 
teste Baker (1964: 178); paratypes MCZ 74935, 176572, 
and 187839 all from Atiu. 

sculpta, Nucula 

1860c, PZS 28: 189 (Corea [Korea] Sea). Not men- 
tioned by Kay (1965). 

sculptilis, Coralliobia 

1865d, PZS for 1865: 513 (Islands of the Central 
Pacific). Not located in BMNH by Kay (1965: 86). 

sculptilis. Helix 

1865a, PZS for 1864: 669 (Mangier [error for Man- 
gaia. Cook Islands]); 1867b, AJC 3: 104 as Helix fra- 
tercula to replace H. sculptilis 1865 non Bland 1868. 
Lectotype BMNH 1962705 selected by Solem (1976: 
423 [not figured]); paralectotypes MCZ 17206. 

sculptilis, Terebra Plate 8, Figure 18 

1869d, AJC 5: 64 ([Honolulu] Oahu). Lectotype MCZ 
248804, teste Bratcher and Cernohorsky (1987: 40) 
presumed here to have been selected by them; para- 
lectotype MCZ 248805. 

sculptum, Cerithium 

1869d, AJC 5: 77, pi. 8, fig. 8 (Paumotus). Holotype 
ANSP 17592, teste Houbrick (1992: 49, fig. 31h). 

scuta, Helix 

1865a, PZS for 1864: 675 [nomen nudum]. 

sectilis, Mitra 

1868d, AJC 3: 271 (Hawaii [Island]). Not located in 
ANSP. 



24 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



semicostata, Rissoa 

1868d, AJC 3: 296, pi. 24, fig. 32 (Paumotus). Type 
lost, label only in ANSP. 

semkostatus, Turbo 

1861b, PZS 28: 435 (Sandwich Islands). Lectotype 
BMNH 1961202 selected by Kay (1965: 60, pi. 7, 
figs. 3, 4); paralectotypes MCZ 297920. 

semiplicata, Melampus (Tralia) 

1860b, PZS 28: 146 (Sandwich Islands). Lectotype 
BMNH 1962773 selected by Kay (1965; 30, pi. 4, fig. 
7); paralectotypes MCZ 176566. 

semiplicata, Rissoina 

1863b, PZS for 1862: 242 (Pacific Islands); 1868d, 
AJC 3: 295, pi. 24, fig. 29 (Rowland Island). Lec- 
totype BMNH 1964294 selected by Kay (1965: 76, 
pi. 13, figs. 3, 4). 

semiplicata, Vanikoro 

1861b, PZS 28: 435 (Sandwich Islands). Holotype 
BMNH 1962819, teste Kay (1965: 57, pi. 6, figs. 7, 
8). 

semistriata, Amphiperas 

1863b, PZS for 1862: 241 (Pacific Islands); 1868g, 
AJC 4: 96, pi. 1 1, fig. 16 (Ponape). Lectotype 1964286 
selected by Kay (1965: 74, pi. 12, figs. 5, 6). Cate 
(1973: 112, fig. 138) apparently unaware that Kay 
had previously selected a lectotype for this species, 
or that Pease had later figured it and stated its lo- 
cality as Ponape Island, refigured ANSP 17042 as 
the holotype and restricted the type locality to: 73 
m, Kii Channel, Japan (24°00'N, 134°48'E). This re- 
striction is therefore invalid. 

semistriata, Atys 

1860a, PZS 28: 20 (Sandwich Islands). Holotype 
BMNH 1961459, teste Kay (1965: 10, pi. 1, figs. 7, 
8); paratypes MCZ 31716 largest figured by Pilsbry 
(1917: 217, fig. 5) and 31717. 

serrata, Lamellina Plate 2, Figure 17 

1861c, PZS 28: 439 (Ebon Island). Lectotype, here 
selected, MCZ 302555; paralectotypes MCZ 28926 
and 298484; paralectotype ANSP figured by Pilsbry 
and Cooke (1915: 165, pi. 33, figs. 1, 2); not located 
in ANSP by Baker (1963: 199). 

similaris, Amastra rugulosa Plate 2, Figure 11 

1870a, JdeC 18: 96 ([Waimea] Kauai). Lectotype, here 
selected, MCZ 45253; paralectotypes MCZ 58936 
and 298498. 

similis, Triphons Plate 10, Figure 14 

1871a, PZS for 1870: 774 ([Haena] Kauai). Lectotype, 
here selected, MCZ 50079. 

simillima, Haminea Plate 4, Figure 24 

1868e, AJC 4: 72, pi. 7, fig. 3 (Tahiti). Lectotype, 
here selected, MCZ 297875; paralectotypes MCZ 
303194. 

simillima, Helix 

1865a, PZS for 1864: 669 (Islands of the Central 
Pacific [Raiatea, teste Garrett, 1884: 19, pi. 2, figs. 32, 
32a, bj). Lectotype, here selected, ANSP 49288 is 
the specimen figured by Garrett; not located in 
BMNH, teste Mordan (personal communication). 

simplana. Vertigo 

187 Id, PZS for 1871: 461 (Marquesas Islands). Not 



located in BMNH, teste Mordan (personal com- 
munication). 

simplex, Leptachatina 

1869c, JdeC 17: 170 (Hawaii [Island]). Measured ho- 
lotype in MNHN, teste Fischer-Piette (1950: 73); 
paratype ANSP 57821 figured by Cooke (1910: 38, 
pi. 1, figs. 8, 9); paratypes MCZ 45176. 

simplex, Olivella (Callianax) Plate 8, Figure 16 

1868d, AJC 3: 281, pi. 23, fig. 24 (Paumotus). Lec- 
totype, here selected, ANSP 28969; paralectotype 
ANSP 391047. 

simplex, Tornatellma Plate 4, Figure 8 

1865a, PZS for 1864: 673 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 473 (Tahaa). Lecto- 
type, here selected, MCZ 175745; paralectotypes 
MCZ 298473. 

simulans, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 202 ([Moorea] Tahitian Archipelago); 1867a, AJC 
3: 81, pi. 1, fig. 11. Lectotype ANSP 59941 selected 
by Baker (1963: 205); paralectotypes MCZ 24881, 
24955, and 25116. 

stniatralis 'Pease' Pilsbry, Partula 

1909, MofC (2) 20: 185. Manuscript name under the 
synonymy of Partula otaheitana Bruguiere. 

sinistrosa 'Pease' Garrett, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; Garrett, 
1884, JANSP (2) 9: 49 (Papieri, Tahiti). Lectotype 
ANSP 59548a selected by Baker (1963: 205) figured 
by Pilsbry (1909: 191, pi. 26, fig. 1); paralectotypes 
MCZ 24997 and 297864. 

solid a, Amastra 

1869c, JdeC 17: 173 (Oahu; Coll. [of] Pease [and] 
Crosse). Three syntypes MCZ 23341 and 141338 fig- 
ured by Pilsbry and Cooke (1914: 28, 31, pi. 7, figs. 
1-3); syntype in MNHN, teste Fischer-Piette (1950: 
73). 

solida, Helictna Plate 5, Figure 3 

1865a, PZS for 1864: 673 (Islands of the Central 
Pacific [Tahiti]). Lectotype, here selected, MCZ 
297923; paralectotypes MCZ 302379. 

soliduscida 'Pease' Sowerby, Magilus 

1872 [in] Reeve, Conchologia Iconica 18: Magilus, pi. 
4, fig. 12 (Sandwich Islands). Holotype in BMNH, 
teste Sowerby. 

speciosa, Philinopsis 

1860a, PZS 28: 21 (Sandwich Islands). Syntype MCZ 
297874; not located in BMNH by Kay (1965: 84). 

sphaerica, Amastra 

1870a, JdeC 18: 94 ([Waimea] Kauai); Crosse, 1876, 
JdeC 24: 98, pi. 1, figs. 5, 5a. Holotype and paratypes 
in MNHN, teste Fischer-Piette (1950: 149 as first 
reference to y4. A. rugulosa); paratypes MCZ 45162. 

squamosum, Sistrum 

1868d, AJC 3: 277, pi. 23, fig. 14 (Kingsmill [Island]). 
Lectotype ANSP 29910 selected by Cernohorsky 
(1987: 97, figs. 10, 11) is the figured type; paralec- 
totypes MCZ 295583. 

squamosus, Latirus 

1863b, PZS for 1862: 240 (Pacific Islands); 1868d, 
AJC 3: 278, pi. 23, fig. 16 (Baker Island). Lectotype 



MoLLUSKS Described by W. H. Pease • Johnson 25 



BMNH 1964284 selected by Kay (1965: 72, pi. 12, 
figs. 12, 13); paralectotypes MCZ 2192 and 261183. 

squamulosa, Fastigiclla Plate 7, Figure 14 

1868d, AJC 3: 290, pi. 24, fig. 15 ([Anaa Island] Pau- 
motus). Lectotype, here selected, ANSP 36702; para- 
lectotype ANSP 391037; paralectotypes MCZ 297956. 

stolicia, Partula 

1866a, AJC 2: 198 (Tahitian); 1871d, PZS for 1871: 
473 (Raiatea). Syntypes MCZ 25311 and 104508 la- 
beled as from Tahiti; not located in ANSP by Baker 
(1963: 205). 

strammca, Helicina 

1865a, PZS for 1864: 676 [nomen nudum]. MCZ 176563. 

striata, Alcytia 

1869d, AJC 5: 70 (Hawaii [Island]). Not located in 
ANSP. 

striata, Chondrella Plate 2, Figure 19 

1871d, PZS for 1871: 477 (Roratonga [Raratonga Is- 
land, Cook Islands]). Lectotype, here selected, MCZ 
187917; paralectotypes MCZ 74942. 

striata, Etigina 

1868d, AJC 3: 275, pi. 23, fig. 10 (Paumotus). Not 
located in ANSP. 

striata, Odostomia Plate 8, Figure 14 

1868d, AJC 3: 291, pi. 24, fig. 16 (Paumotus). Lec- 
totype, here selected, MCZ 10491; paralectotypes 
MCZ 303452 and ANSP 58022. 

striatula, Rissoina Plate 9, Figure 4 

1868d, AJC 3: 296, pi. 24, fig. 31 (Paumotus). Lec- 
totype, here selected, MCZ 178844 labeled as ho- 
lotype; two paralectotypes ANSP 19254. 

striatula Vertigo 

1871d, PZS for 1871: 461 ([Kalapana, Puna] Hawaii 
[Island]). Lectotype, here selected, MCZ 45239 [not 
45234] figured by Pilsbry and Cooke (1926: 223, pi. 
28, figs. 1, 2); paralectotypes MCZ 151647 and BPBM 
ex MCZ. 

striatum, Sistrum Plate 9, Figure 9 

1868d, AJC 3: 276, pi. 23, fig. 12 (Kingsmill [Is- 
lands]). Lectotype, here selected, ANSP 36735; three 
paralectotypes ANSP 391063; paralectotypes MCZ 
178941. 

striatus, Melampms (Tralia) 

1861d, PZS for 1861: 244 (Tahiti); 1868g, AJC 4: 100, 
pi. 12, fig. 14. Lectotype ANSP 22356a selected by 
Baker (1964: 151); a lectotype BMNH 1964282 was 
later also selected by Kay (1965: 71, pi. 12, figs. 8, 
10). 

strigata, Cythara Plate 6, Figure 9 

1863b, PZS for 1862: 242 (Pacific Islands [Howland 
Island]). Lectotype, here selected, MCZ 49990; para- 
lectotypes MCZ 298490; not located in BMNH by 
Kay (1965: 86). 

strigata, Partula 

1868J, AJC 4: 1 55, pi. 12, fig. 7 (Marquesas [Islands]). 
Holotype ANSP 59510a, teste Baker (1963: 205). 

strigata, Pisania 

1863b, PZS for 1862: 241 (Pacific Islands); 1868g, 
AJC 4: 93, pi. 11, fig. 6 (Ponape). Lectotype BMNH 
1964288 selected by Kay (1965: 73, pi. 12, figs. 16, 
17). 



striolata. Helix Plate 2, Figure 2 

1861c, PZS 28: 439 (Ebon Island). Lectotype, here 
selected, MCZ 11563; paralectotypes MCZ 298474. 

striolata, Partula 

1865a, PZS for 1864: 675 [nomcti nudum]; 1866a, AJC 
2: 195 (Tahitian Archipelago [Moorea]); 1867a, AJC 
3: 81, pi. 1, fig. 4. Lectotype ANSP 59648a selected 
by Baker (1963: 205) is the figured type; paralec- 
totypes MCZ 24910, 25293, and 25303. 

subangulata, Alcyna 

1861b, PZS 28: 436 (Sandwich Islands). Holotype 
BMNH 1962830 figured by Kay (1965: 63, pi. 9, fig. 
6); paratype MCZ 31723 is now the figured holotype 
of Alcyna subangulata flammulata Pilsbry (1917: 213, 
pi. 15," figs. 5, 6). 

subangulata, Partula faba 

1870c, JdeC 18: 401 (Tahaa); 1871d, PZS for 1871: 
458. Syntypes MCZ 24948. 

subpellucida, Eulima Plate 7, Figure 12 

1868g, AJC 4: 94 (Tahiti). Lectotype, here selected, 
MCZ 31709; paralectotypes MCZ 303450. The type 
figured by Reeve (1865, Conchologia Iconica 15: Eu- 
lima, pi. 3, figs. 20a, b); was not located in BMNH 
by Kay (1965: 86). 

subrufa, Helicina 

1865a, PZS for 1864: 676 [nomen nudum]. 

subrufa 'Pease' Garrett, Helicina 

1884, JANSP (2) 9: 104, pi. 3, figs. 68, 68a, b (Raiatea 
and Borabora). Lectotype ANSP 14463a selected by 
Baker (1964: 162). 

subviridis, Vitrina 

1868), AJC 4: 154, pi. 12, fig. 5 (Marquesas [Islands]). 
Holotype ANSP 49269a, teste Baker (1963: 237). 

suffusa, Terebra Plate 9, Figure 18 

1869d, AJC 5: 67 ([Honolulu] Oahu). Lectotype MCZ 
248802 selected by Bratcher and Cernohorsky (1987: 
47, pi. 5, fig. 17b); paralectotype MCZ 248803. 

sulcata, Terebra Plate 8, Figure 1 

1869d, AJC 5: 67 ([Honolulu] Oahu). Lectotype, here 
selected, MCZ 49967; paralectotypes MCZ 248801. 

sulcifera, Torinia 

1869d, AJC 5: 79 (Kauai). Not located in ANSP. 

sulcosus, Triphoris Plate 10, Figure 3 

1871a, PZS for 1870: 774 (Kauai). Lectotype, here 
selected, MCZ 50080; paralectotype MCZ 298496. 

suturalis, Partula 

1865b, PZS for 1864: 675 [nomen nudum]. MCZ 24825. 

symmetrica, Scalaria 

' 1868d, AJC 3: 290, pi. 24, fig. 14 (Tahiti). Holotype 
ANSP 19586 refigured by DuShane (1988 (7): 5, fig. 
[not numbered]) and (1990: 9, fig. 44). 



tahitensis, Cyclostoma 

1861d, PZS for 1861: 243 (Huaheine); 1869b, JdeC 
17: 158, pi. 7, fig. 1, as Scalinella. Holotype in MNHN, 
teste Fischer-Piette (1950: 72); paratypes MCZ 74941, 
187858, 187859, and 187860. 

tahitensis, Dolabrifera 

1861d, PZS for 1861: 245 (Tahiti); 1868e, AJC 4: 77, 
pi. 8, fig. 5; not located in BMNH by Kay (1965: 86). 



26 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



tahitensis, Helicina 

1871d, PZS for 1871: 466. New name for Helicma 
pisum Rousseau 1854 non Philippi 1847. 

tahitensis 'Pease' Brot, Melama 

1877 [;>;] Martini and Chemnitz, Conchylien Cabinet 
(2) 1 pt. 24, Die Melaniaceen, p. 323 [nomen nudum]. 
Listed as a synonym of M. mamensis Lea. 

tahitensis, Tectura Plate 1, Figure 7 

1868g, AJC 4: 98, pL 11, fig. 21 (Tahiti). Lectotype, 
here selected, ANSP 38949; paralectotype MCZ 
150757. 

tenebrosa, Leptachatina 

1870a, JdeC 18: 92 ([Waimea] Kauai); Crosse, 1876, 
JdeC 24: 97, pL 3, fig. 5. Holotype in MNHN, teste 
Fischer-Piette (1950: 149); paratypes MCZ 45189 and 
50110. 

tenuicostata, Leptachatina 

1869c, JdeC 17: 170 (Hawaii Island). Holotype in 
MNHN figured by Fischer-Piette (1950: 72, pi. 3, 
fig. 51). 

tenuiscula, Helicina 

1865a, PZS for 1864: 676 [nomen nudum]; 1868b, AJC 
3: 226 [nomen nudum] (Tahaa). 

tenuistriata, Helicma 

1865a, PZS for 1864: 676 [nomen nudum]; 1868b, AJC 
3: 226 (Tahaa) [nomen nudum]. 

tenuistriata, Rissoina Plate 9, Figure 5 

1868d, AJC 3: 295, pi. 24, fig. 30 (Paumotus). Lec- 
totype, here selected, ANSP 19253; paralectotypes 
MCZ 178851. 

terreslris 'Pease' Garrett, Partula 

1884, J ANSP (2) 9: 75 (. . . Opoa Valley on the south- 
east coast [of Raiatea]). Lectotype ANSP 59450a se- 
lected by Baker (1963: 205) figured by Pilsbry (1909: 
243, pi. 17, fig. 11); paralectotypes MCZ 25296 and 
297859. 

tessellatus, Plcurobranchus 

1861d, PZS for 1861: 245 (Pacific Islands) [descrip- 
tion without name]; named, 1864, PZS for 1863: 510; 
1868e, AJC 4: 80, pi. 9, fig. 4; not mentioned by Kay 
(1965). 

triangulatum, Sistrum Plate 9, Figure 23 

1868d, AJC 3: 278, pi. 23, fig. 15 (Hawaii [Island]). 
Lectotype, here selected, MCZ 295580; paralecto- 
types MCZ 295589; not located in ANSP. 

tricarinatum, Bittium 

1861c, PZS 28: 433 (Sandwich Islands). Holotype 
BMNH 1962806, teste Kay (1965: 49, pi. 5, fig. 14). 

trigonalis. Pinna Plate 1, Figure 2 

1861a, PZS for 1861: 242 (Kingsmill Islands). Lec- 
totype, here selected, MCZ 225951; not located in 
BMNH by Kay (1965: 86). 

trilineata, Partula 

1865b, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 195 (Tahiti); 1867a, AJC 3: 81, pi. 1, fig. 1. Lec- 
totype ANSP 59563a selected by Baker (1963: 205) 
is the figured type; paralectotypes MCZ 24940 and 
25130. 

triplicata, Auriculella Plate 2, Figure 9 

1868f, JdeC 16: 346 ([Oahu]). Lectotype, here se- 



lected, MCZ 45150; paralectotypes MCZ 298487; not 
located in MNHN by Fischer-Piette (1950: 71). 

triticea, Rissoina 

1861b, PZS 28: 438 (Sandwich Islands). Holotype 
BMNH 1962844 figured by Kay (1965: 68, pi. 11, 
fig. 11); paratypes MCZ 178859. 

triticea, Tnphoris 

1861b, PZS 28: 433 (Sandwich Islands). Holotype 
BMNH 1962807, teste Kay (1965: 50, pi. 10, fig. 3). 

tuberculatus, Triphoris Plate 10, Figure 4 

1871a, PZS for 1870: 776 (Kauai). Lectotype, here 
selected, MCZ 50081; paralectotypes MCZ 298497. 

luberculiferum, Cerithium 

1869d, AJC 5: 76 (Paumotus). Figured by Sowerby 
[(>;] Reeve, 1865, Conchologia Iconica 15: Cerithium, pi. 
2, fig. 11 (Island of Annaa, Cuming) as Cerithium 
adansomi 'Bruguiere' Sowerby non Bruguiere 1792. 
Holotype BMNH 199021; paratypes MCZ 302380. 

tuberculosa, Engma 

1863b, PZS for 1862: 243 (Pacific Islands); 1867d, 
AJC 3: 274 (Baker Island). Lectotype BMNH 1964296 
selected by Kay (1965: 76, pi. 14, figs. 5, 6); para- 
lectotypes MCZ 260616. 

tumida, Clathurella Plate 6, Figure 18 

1868a, AJC 3: 218, pi. 15, fig. 14 (Paumotus). Lec- 
totype, here selected, ANSP 15813 is the figured 
type; two paralectotypes ANSP 391053; paralecto- 
types MCZ 231968 and 231969. 

tumida, Hyalopsis 

1871f, AJC 7: 27, pi. 9, fig. 6 (Solomon Islands). Not 
located in ANSP. 

turbinatus, Omphalius Plate 8, Figure 20 

1869e, AJC 5: 84, pi. 8, fig. 15 (Gulf of [Golfo de] 
California, La Paz [Baja California Sur, Mexico]). 
Lectotype ANSP 40820 selected by Pilsbry (1932: 
85, pi. 10, figs. 9, 9a, 9b); paralectotype ANSP 31053. 

turbinella, Helicma 

1865b, PZS for 1864: 676 [nomen nudum]. 

turgidula, Leptachatina 

i870a, JdeC 18: 89 ([Waimea] Kauai); Crosse, 1876, 
JdeC 24: 96, pi. 4, fig. 5. Holotype in MNHN, teste 
Fischer-Piette (1950: 149); paratypes MCZ 45182 and 
45183. 

turgidula, Limnaea Plate 4, Figure 1 1 

1870b, AJC 6: 5, pi. 3, fig. 3 (Oahu). Lectotype, here 
selected, MCZ 298901; paralectotype MCZ 298902; 
two paralectotypes ANSP 21932 though not located 
in ANSP by Baker (1964: 154). 

turgidus, Bulimus [Partula] 

1865a, PZS for 1864: 670 (Islands of the Central 
Pacific); 1871d, PZS for 1871: 473 (Tahiti). Probable 
syntypes MCZ 25367 and 26354 though both labeled 
as from Raiatea. 

turricula, Partula 

1865d, PZS for 1864: 675 [nomen nudum]; 1872b, AJC 
7: 196 ("?" New Hebrides). Holotype ANSP 59926a, 
/rsff Baker (1963: 205). 

turricula, Rissoina 

1861b, PZS 28: 438 (Sandwich Islands). Lectotype 
BMNH 1962845 selected by Kay (1965: 69, pi. 11, 
fig. 10); paralectotypes MCZ 178858. 



MoLLUSKS Described by W. H. Pease • Johnson 27 



ualanensis 'Pease' Martens and Langkavel, Melania 
1871, Donum Bismarcktatium, p. 38. Error for ouala- 
nensis 'Pease' Tryon, Melania. 

umbilicata, Cassis Plate 6, Figure 3 

1861b, PZS 28: 436 (Sandwich Islands). Lectotype, 
here selected, MCZ 298907; not located in BMNH 
by Kay (1965: 85). 

umbilicata, Partula 

1865a, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 200 ([Haamene Valley, Tahaa] Tahitian Archi- 
pelago); 1867a, AJC 3: 81, pi. 1, fig. 7. Lectotype 
ANSP 59452a selected by Baker (1963: 205) from 
one of the syntypes figured by Pilsbry (1909: 230, 
pi. 21, figs. 13, 14); paralectotypes MCZ 25300 and 
25310. 

umbilicata, Scalaria Plate 9, Figure 12 

1869d, AJC 5: 76 (Oahu). Holotype MCZ 187393 
figured by DuShane (1990: 4, fig. 9). 

undatolirata 'Pease' Dall, Patella 

1871, AJC 6: 279 (Sandwich Islands) [nomen nudum]; 
Pease, 1872c, AJC 7: 198 listed as a synonym of 
Helcionisus exaratus Nuttali. 

undulata 'Pease' Tryon, Natica 

1886, MofC (1) 8: 23 [nomen nudum]. 

unilineatum, Cerithium 

1861b, PZS 28: 432 (Sandwich Islands). Lectotype 
BMNH 1962798 selected by Kay (1965: 45, pi. 10, 
fig. 5); paralectotypes MCZ 297946. 

umplicata, Aunculella 

18681, JdeC 16: 344, pi. 14, figs. 7, 7a ([Lahaina] 
Maui). Two figured syntypes in MNHN, teste Fi- 
scher-Piette (1950: 71); syntypes MCZ 159563 and 
161636. 

I'aria, Cythara 

1860b, PZS 28: 147 (Sandwich Islands). Lectotype 
BMNH 1962782 selected by Kay (1965: 33, pi. 10, 
fig. 13); paralectotypes MCZ 50000, 50001, and 50002. 
See under: peasei, Martens and Langkavel, Colum- 
bella (Seminella). 

Z'ariabilis, Carelia 

1870c, JdeC 18: 402 (Kauai). Single specimen found 
by Pease; not mentioned by Fischer-Piette (1950: 
76). 

variabilis, Collonia 

1861b, PZS 28: 436 (Sandwich Islands). Lectotype 
BMNH 1963331 selected by Robertson [m] Kay (1965: 
61, pi. 7, figs. 1, 2); paralectotypes MCZ 119269. 

variabilis, Engina Plate 7, Figure 5 

1868d, AJC 3: 275, pi. 23, fig. 6 (Paumotus). Lecto- 
type MCZ 260618 selected by Cernohorsky (1987: 
99, figs. 11, 12); paralectotypes MCZ 260615. 

variabilis, Partula 

1865b, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 203 ([Vaioara Valley, on the west coast of Raiatea, 
teste Hartmann, 1884: 76] Tahitian Archipelago); 
1867a, AJC 3: pi. 1, figs. 12-14; 1871, PZS for 1871: 
473 (Raiatea). Lectotype ANSP 59452a selected by 
Baker (1963: 205) is the syntype represented on pi. 
1, fig. 12; paralectotypes MCZ 24994. 

variabilis, Realia 

1865e, AJC 1: 288 (Polynesia); Tryon, 1866a, AJC 2: 



82, pi. 5, fig. 2 (Hervey Isles); Pease, 1869a, JdeC 
17: 148 as Omphalotropis (Atiu); 1872b, AJC 7: 197 
(also Tongan Group). Holotype ANSP 13341a, teste 
Baker (1964: 179); paratypes MCZ 187844 and 187847 
from Atiu; paratypes MCZ 187846 from the Tongan 
Group. 

variatis, Pleurobranchus 

1860a, PZS 28: 25 (Sandwich Islands). Not located 
in BMNH by Kay (1965: 84). 

varicifera, Daphnella 

1868a, AJC 3: 221, pi. 15, fig. 21 (Paumotus). Ho- 
lotype ANSP 15726. 

variegata, Dolabella 

1860a, PZS 28: 22 (Sandwich Islands). Syntype MCZ 
297871; not located in BMNH by Kay (1965: 84). 

venosus. Helix 

1866b, AJC 2: 290, pi. 21, fig. 2 (Polynesia); 1871d, 
PZS for 1871: 475 as Helicopsis (Roratonga). Holo- 
type ANSP 49163a, teste Baker (1963: 237); paratypes 
MCZ 11529 and 297935. 

venusta 'Pease' Pace, Columbella 

1902, Proceedings of the Malacological Society of London 
5: 150 [nomen nudum]. 

venusta, Eulima Plate 7, Figure 10 

1868d, AJC 3: 294, pi. 24, fig. 24 (Tahiti). Lectotype, 
here selected, ANSP 19788; paralectotypes MCZ 
187748 and 187749. 

verecunda, Pithys 

1870c, JdeC 18: 397 (Marquesas [Islands]); 1871d, 
PZS for 1871: 454 as Pitys. Not mentioned by Fi- 
scher-Piette (1950: 76). 

verticillata, Helix Plate 3, Figure 7 

1865b, PZS for 1864: 675 [nomen nudum]; 1868b, AJC 
3: 228 as Nanina (Moorea). Holotype ANSP 49284, 
teste Baker (1963: 237). 

vescoi 'Dohrn' Pease, Helicina 

1865b, PZS for 1864: 676 [nomen nudum]. 

vexillum, Partula 

1865b, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 198 (Tahitian Archipelago); 1867a, AJC 3: 81, pi. 
1, fig. 8; 1871d, PZS for 1871: 474 (Moorea). Lec- 
totype ANSP 59646a selected by Baker (1963: 205); 
paralectotypes MCZ 24814, 24886, and 24936. 

violacea, Clathurella Plate 6, Figure 15 

1868a, AJC 3: 218, pi. 15, fig. 15 (Paumotus). Lec- 
totype, here selected, MCZ 231971; paralectotypes 
MCZ 303190; not located in ANSP. 

violaceus, Pleurobranchus 

1864, PZS for 1863: 510. New name for Pleurobran- 
chus reticulatus Pease 1860 non Rang 1832 non Ke- 
laart 1859. See under: reticulatus, Pleurobranchus. 

virescens, Pachypoma Plate 8, Figure 17 

1869d, AJC 5: 73, pi. 8, fig. 10 (Tarawa Island). Lec- 
totype, here selected, MCZ 302624; paralectotypes 
MCZ 302625; not located in ANSP. 

virginea, Partula 

1865a, PZS for 1864: 675 [nomen nudum]. 

virginea 'Pease' Garrett, Partula 

1884, JANSP (2) 9: 61, pi. 3, fig. 54 (Vaipiti Valley 
on the west coast of Tahaa). Lectotype ANSP 59474a 
selected by Baker (1963: 206) is the figured type; 
paralectotypes MCZ 25243. 



28 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



virgulata, Partula assimilis 

1870c, JdeC 18: 401 (Roratonga). Syntypes MCZ 
24939; not mentioned by Fischer-Piette (1950: 76). 

viridans, Cyclostoma 

1865b, PZS for 1864: 676 [nomen nudum]. 

viridescens, Cyclostoma 

1861d, PZS for 1861: 243 (Pacific Islands); 1869a, 
JdeC 17: 153, pi. 7, fig. 7, as Omphalotropis (Hua- 
heine). Holotype in MNHN, teste by Fischer-Piette 
(1950: 72). 

viridis, "7" Acanthochites 

1872a, AJC 7: 194 (Kauai). Not located in ANSP. 

viridis, Carelia variabilis 

1870c, JdeC 18: 402 (east side of Kauai). Pilsbry and 
Cooke (1914: 16, pi. 9, fig. 11) suggested that this 
figured specimen MCZ 23343 may be the type of 
viridis; not mentioned by Fischer-Piette (1950: 76). 

viridis, Lobiger Plate 4, Figure 18 

1864, PZS for 1863: 510 ([Tahiti] Pacific Islands); 
1861d, PZS for 1861: 246, description without a 
name. Lectotype, here selected, MCZ 297869; para- 
lectotypes MCZ 298463. 

viridis, Lophcercus 

1861d, PZS for 1861: 246 (Pacific Islands); 1868e, 
AJC 4: 74, pi. 8, figs. 1, 2. Lectotype BMNH 1964324 
selected by Kay (1965: 72, pi. 12, figs. 7, 8); para- 
lectotype MCZ 297932. 

vitrea, Bullina 

1860a, PZS 28: 19 (Sandwich Islands). Lectotype 
BMNH 1961456 selected by Kay (1965: 5, pi. 1, figs. 
1,2). 

vitrea 'Pease' Dohrn, Tornatellides 

1863, Malakozoologische Blatter 10: 162. Not recog- 
nizable, teste Pilsbry and Cooke (1915: 203). 

vittata, Partula 

1865b, PZS for 1864: 675 [nomen nudum]; 1866a, AJC 
2: 194 (Tahitian Archipelago [restricted to the high- 
er portions of Toloa Valley, on the west coast of 
Raiatea, teste Garrett, 1884: 75]). Syntypes MCZ 
24879. 

zigzac, Helicina Plate 5, Figure 8 

1868b, AJC 3: 229 (Oualan [Island]). Lectotype, here 
selected, ANSP 14485 is the measured type, teste 
Baker ( 1 964: 1 63); three paralectotypes ANSP 359504; 
paralectotypes MCZ 297924. 



The taxa for which no type material was 
located has been extracted from the pre- 
ceding alphabetical list and arranged here 
according to publications and dates. 

Proceedings of the Zoological 
Society of London 

nebulosa, Borsonia: 1860b, PZS 28: 143 (Sandwich 
Islands) 

pallida, Mitra: 1860b, PZS 28: 146 (Sandwich Is- 
lands) 



oryza, Marginella: 1860b, PZS 28: 147 (Sandwich 
Islands) 

maculosa, Daphnella. 1860b, PZS 28: 148 (Sandwich 
Islands) 

sculpta, Nucula: 1860b, PZS 28: 189 (Corea [Korea] 
Sea) 

luteostoma, Ranella. 1861a, PZS 28: 397 (Sandwich 
Islands) 

neglectus, Conus. 1861a, PZS 28: 398 (Sandwich Is- 
lands) 

cancellata, Coralliobia. 1861a, PZS 28: 399 (Sand- 
wich Islands); "Only a single dead specimen found." 

lineata, Cohimbella. 1861a, PZS 28: 399 (Sandwich 
Islands) 

costellifera, Anachis: 1863d, PZS for 1862: 279 (Pa- 
cific Islands) 

ovata, Hydrocena : 1865a, PZS for 1864: 674 [Mangaia 
Island] 

sculptilis, Coralliobia: 1865d, PZS for 1865: 513 (Is- 
lands of the Central Pacific) 

conica, Torinia: 1865d, PZS for 1865: 514 (Islands of 
the Central Pacific) 

multiplicata, Mitroidea. 1865d, PZS for 1865: 514 
(Islands of the Central Pacific) 

punctatus, Triphoris. 1871a, PZS for 1870: 775 (An- 
naa Island) 

cylindricus, Triphoris. 1871a, PZS for 1870: 776 
(Apaiang Island) 

filicostata, Pitys: 1871d, PZS for 1871 (Kauai) 

simplaria. Vertigo: 1871d, PZS for 1871: 461 (Mar- 
quesas Islands) 

bacca. Vertigo: 1871d, PZS for 1871; 461 (Kalapana 
[Puna District] Hawaii) 

costellifera, Truncatella: 1871d, PZS for 1871: 468 
(\'avau [Island]; "Collected at the above locality by 
Mr. John Brazier" 

Journal de Conchyliologie 

extensa, Leptachatina: 1870a, JdeC 18: 92 (Kauai) 
hicida, Leptachatina: 1870a, JdeC 18: 93 (Kauai) 
verectinda, Pithys: 1870c, JdeC 18: 397 (Marquesas 
Islands) 



American Journal of Conchology 

distans. Helix: 1866b, AJC 2: 290 (Sandwich Islands) 
flammulata. Mitra: 1868a, AJC 3: 212 (Sandwich and 

Paumotus Islands) 
.sectilis. Mitra: 1868d, AJC 3: 271 (Hawaii) 
aurantium. Opercidatum: 1868d, AJC 3: 287 (Ha- 
waii) 
rosacea. Terebra: 1869d, AJC 5: 65 (Oahu) 
propinqiia, Terebra: 1869d, AJC 5: 66 (Hawaii) 
lirata. Pleurotoma: 1869d, AJC 5: 68 (Oahu) 
monilifera. Pleurotoma: 1869d, AJC 5: 68 (Oahu) 
neivcombii. Mitra: 1869d, .^JC 5: 69 (Oahu) 
striata. .Mcijna: 1869d, AJC 5: 70 (Hawaii) 
balteata, Rissoina: 1869d, AJC 5: 72 (Hawaii) 
sulcifera, Torinia: 1869d, AJC 5: 79 (Kauai) 
viridis. "?" Acanthochites: 1872d, AJC 7: 194 (Kauai) 



MoLLUSKS Described by W. H. Pease • Johnson 



29 



obesa. Partula: 1868b, AJC 3: 223, pi. 15, fig, 12 

(Hab?); "We have but a single specimen" 
striata. Engina: 1868d, AJC 3: 275, pi. 23, fig. 10 

(Paumotus) 
elongata, Volutella: 1868d, AJC 3: 281, pi. 23, fig. 

23 (Fanning Island) 
semicostata. Rissoa: 1868d, AJC 3: 296, pi. 24, fig. 

32 (Caroline Islands) 
uttcea, Nassa: 1869d, AJC 5: 70, pi. 8, fig. 7 (Caroline 

Islands) 
tiimida, Hyalopsis: 1871, AJC 7: 27, pi. 9, fig. 6 

(Solomon Islands) 

ADDITIONAL OR CORRECTED 
LOCALITY DATA 

This list was based essentially on the Na- 
tional Geographic Atlas of the World (6th 
ed., 1990), but also useful were the Index 
to the Islands of the Pacific by Brigham 
(1900), the Hawaiian Gazetteer by Coul- 
ter (1935), and Pacific Island Names by 
Motteler (1986). 

Aitutake Island [Aitutaki Atoll, Hervey Group, Cook 
Islands] 

Annaa Island [Anaa, Tuamotu Archipelago] 

Apaian Island [Abaiang Island, Gilbert Islands, Ki- 
ribati] 

Apaiang Island [Abaiang Island, Gilbert Islands, Ki- 
ribati] 

Apiana Island [Abaiang Island, Gilbert Islands, Ki- 
ribati] 

Ascension Island [Pohnpei, Caroline Islands] 

Atiu Island [Hervey Group, Cook Islands] 

Baker Island [0°13'30"N, 176°29'30"W] 

Bolabola Island [Bora-Bora, Society Islands] 

Ebon Island [Atoll, Marshall Islands] 

Fanning Island [Tabuaeran, Kiribati; 3°5r25"N, 
159°22"W] 

Guam [Island, Mariana Islands] 

Hauheine Island [Society Islands] 

Hawaii [Island, Hawaiian Islands] 

Howland Island [0°49'N, 176°40'W] 

Huaheine Island [Society Islands] 

Jarvis Island [Line Islands; 0°22'33"S, 159°54'1I"W] 

Kauai [Island, Hawaiian Islands] 

Kingsmill Islands [Kingsmill Group, Gilbert Islands, 
Kiribati] 

Lanai [Island, Hawaiian Islands] 

Makaimo Island [Makemo Island, Tuamotu Archi- 
pelago] 

Mangaia Island [Hervey Group, Cook Islands] 

Mangia Island [Mangaia Island, Hervey Group, Cook 
Islands] 

Maui [Island, Hawaiian Islands] 

Molokai [Island, Hawaiian Islands] 

Niihau [Island, Hawaiian Islands] 

Niiie Island [south of the Samoa Islands; 19°S, 170°W] 

Nukahiva [Nuku Hiva Island, Marquesas Islands] 



Oahu [Island, Hawaiian Islands] 

Oualan [Kusaie or Kosrae Island, Caroline Islands] 

Paumotus [Tuamotu Archipelago] 

Philip Island [south of Norfolk Island] 

Ponape [Pohnpei Island, Caroline Islands] 

Raiatea [Island, Society Islands] 

Roratonga [Rarotonga Island, Hervey Group, Cook 

Islands] 
Sandwich [Hawaiian Islands] 
Tahaa [Island, Society Islands] 
Tahiti [Island, Society Islands] 
Tarawa [Island, Gilbert Islands, Kiribati] 
Tutuila [Island, Samoa Islands] 
Vavau [Island, Vava'u Group, Tonga Islands] 
Viti [Fiji] Islands] 

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. 1964. Type land snails in the Academy of 

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Bratcher, T., and W. Cernohorsky. 1987. Liv- 
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Brot, a. 1872. Notice sur quelques especes de Me- 
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. 1877. [in] Martini and Chemnitz (Kuester's 

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Gate, C. N. 1973. A systematic revision of the 
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Cernohorsky, W. O. 1976. The Mitridae of the 
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30 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



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. 1984. Systematics of the family Nassariidae 

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. 1987. Type specimens of Pacific Mollusca 

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Cernohorsky, W. O., and T. Bratcher. 1976. 
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Clench, VV. J. 1932. Diplomorpha coxi (Pease). 
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. 1975. Catalogue of species and bio-bibli- 
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1979. A biography of Andrew Garrett, ear- 



ly naturalist of Polynesia: Part 2. Catalogue of 
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Clench, W. J., and R. D. Turner. 1948. A cat- 
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. 1957. The family Cymatiidae in the West- 
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Cooke, C. M., and Y. Kondo. 1960. Revision of 
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Coulter, J. W. 1935. A gazetteer of the territory 
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. 1932. Studies on the variation, distribution, 

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. 1881. The terrestrial Mollusca inhabiting 

the Cook's or Hervey Islands. Journal of the 
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. 1884. The terrestrial Mollusca inhabiting 

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. 1887. On the terrestrial mollusks of the V'iti 



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some facts — some theories. Ibid. 8(8): 1, 3, 4, 8 
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last few years and death at 47. Ibid. 8(10): 5, 8 
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Hartman, W. D. 1881. Observations on the species 
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(November). 

. 1886. New species of Partula from the New 

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HouBRiCK, R. S. 1992. Monograph of the genus 
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MOLLUSKS Described by W. H. Pease • Johnson 31 



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Martens, E. von, and B. Langkavel. 1871. Do- 
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Nevill, G. 1878. Hand List of Mollusca in the 
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1884. Hand List of Mollusca in the Indian 

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. 1861b. Descriptions of forty-seven new 

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. 1861d. Descriptions of new species of Mol- 
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Proceedings of the Zoological Society of London 



32 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



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. 1865c. Description of a new species of La^i- 

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. 1869e. Remarks on marine Gastropodae 

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. 1869f. Corrections and additions to "Syn- 

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. 1869g. On the classification of the Helic- 



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. 1870a. Observations sur les especes de co- 

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. 1870b. Remarks on the species of Melania 

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. 1870c. Remarques sur certaines especes de 

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. 1871a. Remarks on the genus Triphoris 

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. 1871b. Synonymic de quelques genres et 

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. 1871c. Remarques sur le genre Dibaphus 

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. 187 Id. Catalogue of the land-shells inhab- 
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distribution, and variation, and descriptions of 
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. 187 le. Notes on the synonymy and distri- 



MoLLUSKS Described by W. H. Pease • Johnson 



33 



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. 1872a. Polynesian Chitonidae. American 



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. 1872b. Descriptions of four species of land 

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. 1872c. Svnonvmv of Patella exarata, Rve. 



American Journal of Conchology, 7(3): 198-200 
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. 1896. Trochomorpha fuscata Pease. Nau- 
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. 1917. Marine mollusks of Hawaii, I-III. 



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. 1920. Lyropiipa and Nesopupa. Manual of 

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. 1922. The identity of Helix depressiformis 

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34 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



INDEX 

This index lists all of the names introduced by Pease included in the catalog by Clench (1975) as well as 
those mistakenly attributed to him by other authors. Here the species are placed under the original genera 
in which they were included. References to all of the numbered notes and notes additionalles made on the 
opisthobranch species studied by Pruvot-Fol (1947: 107-114) are included in this index, but only those 
mentioned by Kay (1965) or those that had shells, and were subsequently located, are mentioned in the 
preceding list of taxa. 

All taxa for which type material is extant is indicated here by **. A single * indicates that either no type 
material was located or that the taxon was included for some other reason such as a replacement for a 
preoccupied one. No * indicates that the taxa are opisthobranchs that occur only in the Clench list; several 
taxa followed by C were not in the latter's list. 

The following names mentioned here are not listed by Clench: Amastra rugulosa similaris. Helix fraterciila, 
Melania neivcombii, Melania oualaensis, Melania valanensis, Partula approximata, and Partula microstoma. 



Acanthochites "?" armattis** 

viridis 
Aclesia areola 16 

producta 66 
Aeolis parvula 47 

semidecora 46 
Alcyna lineata** 

rubra ** 

striata* 

stibangulata 
Amastra porphyrostoma** 

rugulosa** 

rugulosa similaris; not in C 

solida** 

sphaerica** 
Amathina bicarinata** 
Amphiperas semistriata** 
Anachis costellifera* 
Assiminea lateritia* 

lucida ** 
Atropis 
Atys costulosa 80** 

debilis 7** 

semistriata 6** 
Auriculella ambusta** 

expansa** 

ptdchra** 

triplicata** 

uniplicata** 
Avicula bru tinea** 

radiata** 
Bittium tricarinatum** 
Blauneria gracilis** 
Bornella arborescens 83 
Borsonia bifasciata** 

corrugata* 

crassicostata** 

lutea** 

nebulosa* 
Bulimus annectens** 

argutus** 

("?" Borus) coxi** 

turgidus** 
Bulla conspersa 79** 

marmorea** 



Bullina lauta 2** 

vitrea 1** 
Capidus liberatus* 
Carelia adusta angulata** 

olivacea** 

variabilis* 

variabilis viridis** 
Cassis umbilicata** 
Catinella rubida** 
Cerithium asperum** 

boeticum** 

cylindraceum** 

fucatum** 

gracile** 

paxillum** 

sculptum** 

tuberculiferum** 

unilineatum** 
Chondrella striata** 
Chromodoria godeffroijana 
Chromodoris inornata 102 

lentiginosa 103 

maculosa 99 

rufomaculata 100 

simplex 101 

varians 104 

variegata 98 
Cirsotrema attenuatum** 
CAthara. See also Cythara 

brevis** 

[Cythara] daedalea** 

[Cythara] decussata** 

paucicostata** 
Citharopsis gracilis** 

ornata** 
Clathurella bait eat a** 

bicarinata** 

brunnea** 

buccinoides** 

canaliculata** 

cylindrica** 

elegans** 

exilis** 

fuscomaculata** 

harpa** 



MoLLUSKS Described by W. H. Pease • Johnson 35 



mactilosa** 

patu'icostata** 

producta** 

ptilchella** 

tiimida** 

violacea** 
CoUonia "?" Candida** 

granulosa** 

multistriata* 

picfa** 

variabilis** 
ColumheUa cytharoides* 

fiisiforjins* 

lineata* 

maculosa* 

micans** 

pallida* 

peasei* 

pelliicida** 

pusilla** 

sagitta* 

sandwichensis** 

venusta* 
CoTius fusiformis** 

neglectus* 

parvus* 

purus** 
Coralliobia cancellata* 

sculptilis* 
Crypt ophthalmus cylindricus 56, 56 bis 
Cyclophorus "?" scalariformis** 
Cyclostonm biangidatum** 

parvum** 

tahitensis** 

viridans* 

viridescens** 
Cylindra formosa** 
Cypraea Candida** 

compta** 

fuscomaculata** 

granulate** 
Cythara. See also Cithara 

angiostoma* 

debilis* 

garrettii** 

pusilla ** 

strigata** 

varia** 
Daphnella bella** 

crenulata** 

curt a** 

interrupta** 

maculosa* 

sandwicensis** 

varicifera** 
Dentiora rubida** 
Diadema rotella** 
Distorsio pusilla** 
Dolabella oariegata 11** 
Dolabrifera fischeri* 

fusca 73** 



marniorea* 

olivacea 12** 

tahitensis 57, 57 bis* 
Doriopsis 100 

granulosa 41 

scabra 41, 81 

viridis 41, 53, 53 bis 
Doris albopustulosa 35 

cinerosa 94 

compta Note 5 

debilis 91, Note 4 

decora 32 

echinata 25 

excavata 23 

foetida 38 

fuscescens 97 

grandiflora 36 

marginata 33 

nubilosa 95 

nucleola 31* 

oreosoma 23 

papillata 

papillosa 34 

picta 30 

pilosa 27 

prismatica imperialis 35, 39 

prismatica lineata 40 

propinquata 29, 35 

pulchra 60 

reticulata 24 

rubrilineata 93 

rugosa 37 

scabriuscula 26 

setosa 22 

sordida 96 

tmcfa 

vibrata 28, 35 

villosa 90 
Drillia exilis** 

lauta** 

nodifera** 

nodulosa** 
Eburnella 
Elysia ocellata 48 
Emarginula clathrata** 
Endodonta celsa* 
Engina albocincta** 

costata** 

fusiformis** 

lineata maculata** 

monilifera** 

nodicostata** 

nodulosa** 

ovata** 

parva** 

striata* 

tuberculosa** 

variabilis** 
Erato sandtvicensis** 
Euchelus angulatus** 

corrugatus** 



36 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



fimbriatus** 

maculosus** 
Eulima aciculata** 

exilis** 

inflexa ** 

subpelhtcida** 

venusta** 
Fastigiella squamulosa** 
Fissurella granifera** 
Fossa r garret tii* 

nuilticostatus** 

sandwichensis* 
Gena laevis** 

rosacea** 
Goniobranchus p. 106 

albomacitlatus 63, 63 bis 

reticulatus 64 
Haminea aperta 70** 

crocata 4** 

galba** 

nigropunctata 67** 

ovalis 68** 

pusilla 5** 

simillima 69** 
Helicina Bella* 

hrazieri** 

calliostoma** 

cincta* 

color ata** 

corrugata** 

discoidea ** 

faha** 

flavesceiis** 

guppyi* 

inconspicua* 

lenticulina* 

maugeriae albinea** 

maugeriae rubicunda** 

"?" multicolor* 

oceanica** 

pacifica** 

parvula** 

picta* 

rufescens* 

rugulosa** 

solida** 

straminea* 

stilmija** 

tahitensis* 

tenuiscula* 

tenuistriata* 

turbinella* 

vescoi* 

zigzac** 
Helicter hutchinsonii** 

proximus** 
Helix acetabulum** 

aha** 

capillata** 

congrua** 

consimilis** 



conula** 

decussatula* 

depressiformis** 

distans* 

fabrejacta** 

ficta** 

fratercula; not in C* 

frivola** 

ftiscata* 

laminata** 

lent a* 

marquesana** 

nigritella* 

normalis** 

obconica** 

oualanensis** 

parvidens** 

prostrata** 

retunsa** 

rugata** 

sculptilis** 

scuta* 

simillima** 

striolata* 

venosus** 

verticillata** 
Hexabranchus nebidosus 43 

pulchellus 42 
Hindsia angicostata** 
Histiophorus p. 99 

maculatiis 51 
Hyalopsis tumida* 
Hijdrocaena costata* 

elongata* 

robusta* 
Hijdrocena fragilis** 

nitida** 

ovata* 
Labiella compact a** 

pachystoma** 
Laimodonta conica** 
Lamellina laevis* 

serrata** 
Laminella erect a** 
Lampania baetica* 
Latirus gibbus** 

granulosus** 

liratus** 

squamosus** 
Leiostraca distorta** 
Leptachatina antiqua** 

balteata** 

brevicula** 

costulosa** 

cylindrata** 

extensa* 

laevis** 

lucida* 

simplex** 

tenebrosa** 

tenuicostata** 



MoLLUSKs Described by W. H. Pease • Johnson 37 



turgidula** 
Leptothijra costata** 
Lihraiula plana** 
Lirunaea amhigiia** 

compacta** 

turgidula** 
Littorina cinerea** 
Lobifera p. 101 

nigricans 61 

papillosa 62 
Lobiger picta 58, 58 bis 

viridis 58** 
Lophocerctis viridis 59, 59 bis* 
Magilus soliduscula** 
Margarita marmorea** 
Marginella cijlindracea** 

cijlindrica* 

debilis* 

orijza* 

pacifica** 

paumotensis* 

peasei* 

polita* 

pyriformis** 

sandwicensis** 
Melampus cinctus* 

fusciis* 

liicidiis** 

(Tralia) semiplicata** 

{Tralia) striatus** 
Melania contigua** 

kauaiensis** 

newcombii; not in C* 

oualaensis., not in C** 

tahitensis* 

valanensis; not in C* 
Melibe pilosa 
Mitra assimilis** 

ericea** 

flammulata* 

glabra** 

lubrica* 

netvcombii* 

nigricans** 

ordinata* 

pallida* 

pudica** 

saltata** 

sectilis* 
Mitroidea multiplicata* 
Mitropsis ftisiformis** 
Mucronalia gracilis** 

nitidula** 

ovata** 

rosea** 
Murex foveolatus** 

garettii* 
Narica delicata** 

granifera** 
Nassa approximata** 

balteata** 



gracilis** 

microstoma** 

nticea* 

obliqua** 
Natica undtdata* 
Neptunea fuscolineata** 
Nerita maculata** 
Neritina deshaijesii* 

dispar* 

neglecta** 

rubida** 

rudis** 
Neritopsis interlirata** 
Nucula sculpta* 
Odostomia debilis** 

gracilis** 

polita** 

rosacea** 

rubra** 

striata** 
Oliva sandwicensis** 
Olivella (Callianax) simplex* 
Omphalius turbinatus** 
Omphalotropis moussoni* 

nebulosa** 

producta** 

robust a** 
Operculatum aurantium* 
Pachypoma virescens** 
Partula abbreviata* 

ajfinis** 

affinis dubia* 

alternata** 

approximata; not in C** 

assimilis** 

attenuata** 

bella* 

bicolor** 

bilineata** 

brazieri** 

brevicula** 

castanea** 

citrina** 

clara** 

cognata** 

compacta** 

concinna* 

crassa** 

crassilabris** 

crassilabrun\* 

elongata** 

expansa** 

faba subangulata** 

fasciata** 

formosa** 

fusca ** 

garrettii** 

globosa* 

gracilior** 

gracilis** 

imperforata** 



38 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



labiata** 

lignaria** 

lineolaia** 

Itigubris** 

megastoma** 

microstoma; not in C** 

nucleola** 

obesa* 

otaheitana dubia** 

ovalis** 

peUucida * 

perplexa** 

perversa** 

planilabrum** 

producta** 

propinqua** 

protea* 

pulchra* 

pulchra varia* 

radio t a** 

recta** 

robust a** 

rustica** 

simulans** 

sinistralis* 

sinistrorsa** 

stolida** 

strigata** 

striolata (2)** 

subangidata** 

suturalis* 

terrestris** 

trilineata** 

turricula** 

umbilicata** 

variabilis** 

vexillum** 

virginea** 

virgulata** 

vittata** 
Parttilina compta** 
Patella sandwicensis** 

undatolirata* 
Pedicularia pacifica** 
Pedipes sandwicensis** 
Perdicella 
Perna hatvaiiensis* 
Philinopsis p. 99 

nigra 10* 

speciosa 9** 
Phyllidia nigra 78 
Pinna trigonalis** 
Pisania strigata** 
Pithy s analogica** 

atien-sis** 

celsa ** 

imperforata** 

paucicostata** 

rarotongensis* 

roratongensis** 

rotellina** 



verecunda* 
Pitys filocostata* 

fratercula* 
Placobranchus gracilis 85 

variegatus 86 
Planaxis abbreviata** 

atra** 

fasciata** 

plumbea** 
Pleurobranchus delicatus 54, 53 bis* 

grandis 76 

marginatus 18* 

ovalis 11 

pelhicidus 17 

reticulatus 21* 

rufus 19** 

tessellatus* 

varians 20* 

violaceiis 21* 
Pleurotoma lirata* 

monilifera* 
Polybranchia p. 100 

pelhicida 52* 
Pterocyclos parva** 
Pterogasteron p. 104 

belliim 50 

marginatus 87 

nigropunctatus 89 

ornatum 49 

rujescens 88 
Pupoidea 

Purpura marmorata** 
Ranella luteostoma* 

producta** 
Realia abbreviata** 

affinis** 

costata** 

elongata** 

laevis** 

ochrostoma** 

producta** 

scalariformis** 

variabilis** 
Registoma complanatum** 
Rhizochilus exaratus** 
Rissoa flammea** 

gracilis** 

semicostata* 
Rissoina angasii* 

balteata* 

cerithiopsis* 

costulata** 

granulosa** 

semiplicata** 

striatula** 

tenuistriata** 

triticea** 

turricula** 
Scalaria crenulata** 

crispata** 

decussata** 



MoLLUSKS Described by W. H. Pease • Johnson 



39 



fucata** 
millecostata** 
patimotensis** 
perplexa** 

symmetrica** 
iimhilicata** 
Scutellina aculeata** 

cancellata** 

compressa** 

granocostata** 
Seminella 

Siphonaria depressa** 
Sistrum affine** 

rugulosum** 

squamosum** 

striatum** 

triangulatum* 
Stenodoris p. 99 

rubra 65 
Strigatella brunnea** 

fiiscescens** 

picea** 
Strombus cancellatus* 
Stylifer deformis** 

robustus* 
Succinea costulosa* 

elongata** 

labiata** 

mammillata** 

ovata* 

rubella** 

rutella* 
Syphonota p. 104 

bipes 13** 

elongata 15** 

grandis 14** 

punctata 75** 

viridescens 74 
Taheitea pallida** 
Tectura conoidalis** 

radiata** 

tahitensis** 
Terebra assimilis* 

contigua* 

costellifera** 

lauta** 

propinqua* 

rosacea * 

sculptilis** 

suffusa** 

sulcata** 

swainsonii inflexa** 
Thala alba** 

angiostoma** 
Torinia conica** 

discoidea ** 

sulci f era* 
Tornatellides vitrea* 
Tornatellina aperta** 

dentata** 

gracilis** 



nit Ida** 

oblonga ** 

simplex** 
Tornatina sandwicensis 3** 
Trevelyana picta 84 
Triforis marmorata** 
Triopa "?" gracilis 82 
Triphoris affinis** 

alternata** 

bicolor* 

brunneus** 

cingulifera** 

clavata** 

costatus** 

cylindricus* 

flammulata** 

fucata** 

gracilis** 

granosus** 

incisa** 

maculatus** 

minimus** 

oryza** 

pallidus** 

peasei* 

prefectus** 

punctatus* 

pustulosus** 

robustus** 

similis** 

sulcosus** 

triticea** 

tuberculatus** 
Triton cylindricus** 

intermedius** 

pusilla** 
Tritonia hawaiiensis 44 
Tritonidea australis* 
Trivia corrugata** 
Trochomorpha contigua* 

fuscata** 

nigritella oppressa** 

trochiformis pollens** 
Trochus conoidalis* 

exilis** 

marmoreus** 
Truella 
Truncatella concinna** 

costellifera* 

cylindracea* 

cylindrica * 

pacifica** 
Tugalia oblonga** 
Turbo multistriatus* 

sandwicensis** 

semicostatus** 
Turbonilla decussata** 

elongate** 
Turcica coreensis** 
Turricula approximata** 

bella** 



40 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



fortipHcata** ""^"^ „ 

modesta** perlonga** 

nodulosa** simplaria* 

phcatula** stnatula** 

putillus** Vexilla fusconigra** 

Turns monilifera** ^'f" depressiformis* 
Vanikoro imbricata** jusca 

semiplicata** subviridis** 

Vertagus graniferus** Vitularia sandwicensis** 

Vertigo arnmta** J^« "^^/ « ''"""fT.o.. 

Ijacca* Volvatella Candida 72** 
costata** f'^'Si'ts 8, 8 bis** 

costulosa** pyriformis 71** 

dentijera** 



42 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



Plate 1 

Figure 1. "?" Acanthochltes armatus Pease. [Pauloa] Oahu [Hawaiian Islands]. Holotype MCZ 73452. Length 10 mm, width 
6 mm. 

Figure 2. Pinna trigonalis Pease. Kingsmill Islands [Kiribati]. Lectotype MCZ 225951 . Length 217 mm, width 94 mm. 

Figure 3. Avicula brunnea Pease. [Moiokai] Sandwich [Hawaiian] Islands. Lectotype MCZ 298465. Length 141 mm, width 
73 mm. 

Figure 4. Avicula radiata Pease. Kingsmill Islands [Kiribati]. Lectotype MCZ 298466. Length 84.5 mm, width 42 mm. 

Figure 5. Siphonaria depressa Pease. Apaiang [Abaiang] Island [Kiribati]. Lectotype ANSP 22199. Length 18 mm, width 14 
mm, height 3 mm. 

Figure 6. Tectura conoidalis Pease. Roratonga [Rarotonga Island, Cook Islands] labeled as from the Paumotus. Lectotype 
MCZ 302558. Length 18 mm, width 14 mm, height 7 mm. 

Figure 7. Tectura tahitensis Pease. Tahiti [Society Islands]. Lectotype ANSP 38949. Length 38 mm, width 12 mm, height 
3.5 mm. 

Figure 8. Patella sandwicensis Pease. Sandwich [Hawaiian] Islands. Lectotype MCZ 304058. Length 37 mm, width 30 mm, 
height 18.5 mm. 

Figure 9. Scutellina aculeata Pease. Hawaii [Hawaiian Islands]. Lectotype MCZ 83720. Length 4.5 mm, width 4 mm, height 
3.4 mm. 

Figure 10. Scutellina compressa Pease. Tahiti [Society Islands]. Lectotype MCZ 302552. Length 4 mm, width 1.5 mm, height 
1.4 mm. 



MoLLUSKS Described by W. H. Pease • Johnson 43 




Plate 1 



44 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 

Plate 2 

Figure 1. Helix capillata Pease. Kauai, Sandwich [Hawaiian] Islands. Lectotype ANSP 1975a. Width 4 mm, height 2 mm. 

Figure 2. Helix striolata Pease. Ebon Island [Atoll], Marshall [Islands]. Lectotype MCZ 11563. Width 3 mm, height 2 mm. 

Figure 3. Helicina pacifica Pease. Oualan [Kusaie or Kosrae Island, Caroline Islands]. Lectotype ANSP 14443. Width 6 mm, 
height 4 mm. 

Figure 4. Helix congrua Pease. Ponape [Pohnpei Island, Caroline Islands]. Lectotype MCZ 12161. Width 11.5 mm, height 

8.5mm. 

Figure 5. Trochomorpha trochiformis pallens Pease. Moorea [Society Islands]. Lectotype MCZ 12188. Width 18 mm, height 
12 mm. 

Figure 6. Trochomorpha nigritella oppressa Pease. Ponape [Pohnpei Island, Caroline Islands]. Lectotype MCZ 12187. Width 
15 mm, height 8.5 mm. 

Figure 7. Pithys paucicostata Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 17271. Width 3.5 mm, height 2 mm. 

Figure 8. Pithys atiensis Pease. Atiu [Island, Cook Islands]. Lectotype MCZ 17335. Width 3 mm, height 1.5 mm. 

Figure 9. Auhculella triplicata Pease. Oahu [Hawaiian Islands]. Lectotype MCZ 45150. Width 5 mm, height 10 mm. 

Figure 10. Auhculella ambusta Pease. [Waianae Mountains, Oahu, Hawaiian Islands]. Lectotype MCZ 45152. Width 5 mm, 
height 8.5 mm. 

Figure 11. Amastra rugulosa similahs Pease. [Waimea] Kauai [Hawaiian Islands]. Lectotype MCZ 45253. Width 8 mm, height 
13,5 mm. 

Figure 12. Bulimus ("?" Borus) coxi Pease. [New Hebrides Islands]. Holotype MCZ 86495. Width 15 mm, length 25 mm. 

Figure 13. Cyclostoma biangulatum Pease. Aitutaki [Atoll], Cook Islands. Lectotype MCZ 141031. Width 2 mm, height 3 mm. 

Figure 14. Carelia olivacea Pease. Kauai [Hawaiian Islands]. Holotype MCZ 571 14. Width 18 mm, height 41 mm. It is assumed 
that of the original measurements given by Pease — width 19 mm, height 69 mm — the latter is a misprint. 

Figure 15. Cyclostoma parvum Pease. [Tahiti, Society Islands]. Lectotype MCZ 74949. Width 2 mm, height 2.5 mm. 

Figure 16. Lamellina laevis Pease. Hervey Islands [Cook Islands]. Lectotype MCZ 154942. Width 1.5 mm, height 3.15 mm. 

Figure 17. Lamellina serrata Pease. Ebon Island [Atoll], Marshall [Islands]. Lectotype MCZ 302555. Width 1.5 mm, height 
3 mm. 

Figure 18. Tornatellina aperta Pease. Tahiti [Society Islands]. Lectotype MCZ 175722. Width 2.5 mm, height 3 mm. 

Figure 19. Chondrella striata Pease. Roratonga [Rarotonga Island, Cook Islands]. Lectotype MCZ 187917. Width 1.5 mm, 
height 2 mm. 

Figure 20. Vertigo costata Pease. [Kona] Hawaii [Hawaiian Islands]. Holotype MCZ 45238. Width 1 mm, height 2 mm. 

Figure 21. Vertigo nitens Pease. Ebon Island [Atoll], Marshall [Islands]. Lectotype MCZ 151650. Width 1 mm, height 2 mm. 

Figure 22. Vertigo costulosa Pease. Hawaii [Hawaiian Islands]. Lectotype MCZ 45244. Width 1 mm, height 1.5 mm. 



MoLLUSKS Described by W. H. Pease • Johnson 45 




Plate 2 



46 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 

Plate 3 

Figure 1. Helix depressiformis Pease. Tahiti [Society Islands]. Lectotype MCZ 17342. Width 7 mm, height 2.5 mm. 

Figure 2. Helix fabrefacta Pease. [Raiatea, Society Islands]. Lectotype MCZ 17238. Width 8 mm, height 4 mm. 

Figure 3. Helix rugata Pease. Maui [Hawaiian Islands]. Lectotype MCZ 17237. Width 5 mm, height 3 mm. 

Figure 4. Helix laminate Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 17233. Width 7 mm, height 2.5 mm. 

Figure 5. Helix marquesana Pease. Marquesas [Islands]. Lectotype MCZ 302557. Width 10 mm, height 7.5 mm. 

Figure 6. Helix conula Pease. Tahiti [Society Islands]. Lectotype MCZ 297945. Width 6.5 mm, height 5 mm. 

Figure 7. Helix verticillata Pease. Moorea [Society Islands]. Holotype ANSP 49284. Width 5 mm, height 3 mm. 

Figure 8. Helix oualanensis Pease. Oualan [Kusaie or Kosrae Island, Caroline Islands]. Lectotype ANSP 47763. Width 5.5 mm, 
height 2 mm. 

Figure 9. Helix normalis Pease. Moorea [Society Islands]. Lectotype MCZ 11543. Width 4.5 mm, height 3 mm. 



MoLLUSKS Described by W. H. Pease • Johnson 47 




Plate 3 



48 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 

Plate 4 

Figure 1. Helicter proximus Pease. Molokai [Hawaiian Islands]. Lectotype MCZ 25823. Width 15 mm, height 29 mm. 

Figure 2. Helicter hutchinsonii Pease. Maui [Hawaiian Islands]. Lectotype MCZ 45254. Width 7 mm, height 16 mm. 

Figure 3. Registoma complanatum Pease. Ebon Island [Atoll], Marshall [Islands]. Lectotype MCZ 141018. Width 3 mm, height 
6 mm. 

Figure 4. Omphalotropis nebulosa Pease. [Makela = San Cristobal Island] Solomon Islands. Lectotype MCZ 72347. Width 5.5 
mm, height 9 mm. 

Figure 5. Labiella compacta Pease. [Palauea] Maui [Hawaiian Islands]. Lectotype MCZ 45196. Width 4.5 mm, height 9 mm. 

Figure 6. Tornatellina oblonga Pease. Tahiti [Society Islands]. Lectotype MCZ 154941. Width 1.5 mm, height 3 mm. 

Figure 7. Tornatellina nitida Pease. Ebon Island [Atoll], Marshall [Islands]. Lectotype MCZ 28921. Width 2 mm, height 4 mm. 

Figure 8. Tornatellina simplex Pease. Tahaa [Island, Society Islands]. Lectotype MCZ 175745. Width 2 mm, height 3 mm. 

Figure 9. Tornatellina dentata Pease. Hawaii [Hawaiian Islands]. Lectotype MCZ 28918. Width 1 mm, height 2.5 mm. 

Figure 10. Tornatellina gracilis Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 302554. Width 1.5 mm, height 4 mm. 

Figure 11. Limnaea turgidula Pease. Oahu [Hawaiian Islands]. Lectotype MCZ 298901. Width 8 mm, height 14.5 mm. 

Figure 12. Succinea mammlllata Pease. Nukahiva [Nuku Hiva, Marquesas Islands]. Lectotype MCZ 155145. Width 7.5 mm, 
height 12.5 mm. 

Figure 13. Succinea rubella Pease. Lanai [Hawaiian Islands]. Lectotype MCZ 161671. Width 7.5 mm, height 11.5 mm. 

Figure 14. Succinea elongata Pease. Kauai [Hawaiian Islands]. Holotype MCZ 161665. Width 9 mm, height 15.5 mm. 

Figure 15. Succinea costulosa Pease. Tahiti [Society Islands]. Lectotype MCZ 31406. Width 5 mm, height 7 mm. 

Figure 16. Succinea labiata Pease. Raiatea [Society Islands]. Lectotype MCZ 298909. Width 12 mm, height 19.5 mm. 

Figure 17. Haminea aperta Pease. Tahiti [Society Islands]. Lectotype ANSP 57575. Width 11 mm, height 15 mm. 

Figure 18. Lobiger viridis Pease. Tahiti [Society Islands]. Lectotype MCZ 297869. Width 8 mm, height 12 mm. 

Figure 19. Melania contigua Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 74887. Width 10 mm, height 20 mm. 

Figure 20. Melania oualanensis 'Pease' Tryon. Oualan [Kusaie or Kosrae Island, Caroline Islands]. Lectotype ANSP 26274. 
Width 12 mm, height 29 mm. 

Figure 21 . Catinella rubida Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 45252. Width 9 mm, height 1 1 mm. 

Figure 22. Melampus lucidus Pease. [Honolulu] Oahu [Hawaiian Islands]. Lectotype ANSP 22284. Width 3.5 mm, height 5 mm. 

Figure 23. Haminea ovalis Pease. Tahiti [Society Islands]. Lectotype MCZ 297877. Width 8 mm, height 10 mm. 

Figure 24. Haminea simillima Pease. Tahiti [Society Islands]. Lectotype MCZ 297875. Width 8 mm, height 10 mm. 



MoLLUSKS Described by W. H. Pease • Johnson 49 




Plate 4 



50 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



Plate 5 

Figure 1 . Scutellina granocostata Pease. Hawaii [Hawaiian Islands]. Lectotype MCZ 83723. Width 6 mm, length 8 mm, height 
3 mm. 

Figure 2. Libratula plana Pease. Islands of the Central Pacific. Lectotype MCZ 1 1 91 62. Width 3 mm, length 7 mm, height 1 mm. 

Figure 3. Heliclna solida Pease. [Tahiti, Society Islands]. Lectotype MCZ 297923. Width 8 mm, height 5 mm. 

Figure 4. Helicina corrugata Pease. Raiatea [Society Islands]. Lectotype MCZ 297925. Width 5 mm, height 3 mm. 

Figure 5. Helicina calliostoma Pease. Marquesas [Islands]. Lectotype MCZ 74950. Width 9 mm, height 6 mm. 

Figure 6. Helicina discoidea Pease. Tahaa [Island, Society Islands]. Holotype ANSP 14398. Width 6 mm, height 3 mm. 

Figure 7. Helicina rugulosa Pease. Tahaa [Island, Society Islands]. Lectotype MCZ 297922. Width 3 mm, height 2 mm. 

Figure 8. Helicina zigzac Pease. Oualan [Kusaie or Kosrae Island, Caroline Islands]. Lectotype ANSP 14485. Width 6 mm, 
height 4.5 mm. 

Figure 9. Helicina oceanica Pease. Kingsmill [Islands, Kiribati]. Lectotype MCZ 176561. Width 4 mm, height 3 mm. 

Figure 10. Petrocyclos parva Pease. Aitutake [Aitutaki Atoll], Hervey Islands [Cook Islands]. Lectotype ANSP 13406. Width 
3 mm, height 3 mm. 

Figure 1 1 . Helicina brazieri Pease. Niue [Island; 19°S, 170°W]. Lectotype MCZ 74947. Width 6 mm, height 5 mm. 

Figure 12. Helicina maugeriae rubicunda Pease. [Tahiti, Society Islands]. Lectotype ANSP 14512. Width 13 mm, height 8 mm. 



MoLLUSKS Described by W, H. Pease • Johnson 51 




^gf JP '^^JI^SJpT-v-K . 





. »'>5^ V 



.^iltaai9>^ 




>' 




#11 r: i 



A 



X 



8 



# 



^iL 



^K 



.\ 



^- "^"^ 



.V 



11 




12 



Plate 5 



52 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 

Plate 6 

Figure 1. Gena rosacea Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 40756. Length 8.5 mm, width 5 mm. 

Figure 2. Gena laevls Pease. Tahiti [Society islands]. Lectotype ANSP 40754. Length 9 mm, width 5.5 mm. 

Figure 3. Cassis umbllicata Pease. Sandwich [Hawaiian] Islands. Lectotype MCZ 298907. Width 43 mm, height 64.5 mm. 

Figure 4. Alcyna lineata Pease. [Puuioa] Oahu [Hawaiian Islands]. Holotype MCZ 31724. Width 1.5 mm, height 2 mm. 

Figure 5. Atys costulosa Pease. [Waimalu] Oahu [Hawaiian Islands]. Holotype MCZ 31714. Width 2 mm, height 3 mm. 

Figure 6. Cerlthium cylindraceum Pease. Paumotus [Tuamotu Archipelago]. Paratype MCZ 297939. Width 10 mm, height 
20.5mm. 

Figure 7. Cithara [Cythara] decussata Pease. [Anaa Island] Paumotus [Tuamotu Archipelago]. Lectotype ANSP 1 5651 . Width 
4 mm, height 9 mm. 

Figure 8. Cithara [Cytt^ara] paucicostata Pease. Tahiti [Society Islands]. Lectotype MCZ 231925. Width 3.5 mm, height 8 mm. 

Figure 9. Cythara strigata Pease. [Howland Island; 0°49N, 176°40'W]. Lectotype MCZ 49990. Width 3 mm, height 7 mm. 

Figure 10. Cithara [Cythara] brevis Pease. [Anaa Island] Paumotus [Tuamotu Archipelago]. Lectotype ANSP 15652. Width 
3.5 mm, height 7 mm. 

Figure 1 1 . Cithara [Cythara] daedalea Pease. Paumotus [Tuamotu Archipelago). Lectotype ANSP 15663. Width 2 mm, height 
4.5 mm. 

Figure 12. Citharopsis ornata Pease. Tahiti [Society Islands]. Lectotype ANSP 16919. Width 1.5 mm, height 3.5 mm. 

Figure 13. Citharopsis gracilis Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 16921 . Width 1 .5 mm, height 4 mm. 

Figure 14. Clathurella maculosa Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 48693. Width 2 mm, height 5 mm. 

Figure 1 5. Clathurella violacea Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 231 971 . Width 2 mm, height 4.5 mm. 

Figure 16. Clathurella canaliculata Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 231978. Width 5 mm, height 
10mm. 

Figure 17. Clathurella bicarinata Pease. Kingsmill Islands [Kiribati]. Lectotype ANSP 15806. Width 5 mm, height 10 mm. 

Figure 1 8. Clathurella tumida Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 1 581 3. Width 2.5 mm, height 6.5 mm. 

Figure 19. Columbella micans Pease' Tryon. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 304063. Width 5.5 mm, height 
10 mm. 

Figure 20. Collonia granulosa Pease. Ponape [Pohnpei Island, Caroline Islands]. Lectotype MCZ 245264. Width 3 mm, height 
2.5 mm. Photographed by James H. McLean. 

Figure 21 Collonia picta Pease. [Anaa Island] Paumotus [Tuamotu Archipelago]. Lectotype MCZ 245268. Width 4.5 mm, 
height 4 mm. Photographed by James H. McLean. 

Figure 22. Columbella sandwichensis Pease. Sandwich [Hawaiian] Islands. Lectotype MCZ 297950. Width 8 mm, height 1 2 mm. 

Figure 23. Conus purus Pease. Niihau Island [Hawaiian Islands]. Holotype MCZ 72331. Width 22.5, height 41 mm. 



MoLLUSKS Described by W. H. Pease • Johnson 53 







I 



% 




12 ''* 14 ^ 16 




17 




19 




Plate 6 



54 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



Plate 7 

Figure 1. Cylindra formosa Pease. Ascension [Pohnpei Island, Caroline Islands]. Holotype MCZ 260605. Width 6 mm, height 
14 mm. 

Figure 2. Daphnella curta Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 16956. Width 2 mm, height 4 mm. 

Figure 3. Daphnella crenulata Pease. [Howland Island; 0°49'N, 176°40'W]. Lectotype ANSP 1 5694. Width 2 mm, height 7 mm. 

Figure 4. Engina ovata Pease. Howland Island [0°49'N, 176°40'W]. Lectotype ANSP 34536. Width 8 mm, height 12 mm. 

Figure 5. Engina variabilis Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 260618. Width 4 mm, height 7 mm. 

Figure 6. Daphnella sandwicensis Pease. Sandwich [Hawaiian] Islands. Lectotype MCZ 49993. Width 3.5 mm, height 8 mm. 

Figure 7. Dentlora rubida Pease. Sandwich [Hawaiian] Islands. Lectotype MCZ 297951. Width 2 mm, height 3 mm (2 views). 

Figure 8. Drillia lauta Pease. [Anaa Island] Paumotus [Tuamotu Archipelago]. Lectotype ANSP 15692. Width 4 mm, height 
8.5mm. 

Figure 9. Eulima inflexa Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 59334. Width 4 mm, height 10 mm. 

Figure 10. Eulima venusta Pease. Tahiti [Society Islands]. Lectotype ANSP 19788. Width 2 mm, height 6 mm. 

Figure 11. Eulima exilis Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 187831. Width 4 mm, height 9 mm. 

Figure 12. Eulima subpellucida Pease. Tahiti [Society Islands]. Lectotype MCZ 31709. Width 7 mm, height 16 mm. 

Figure 13. Euchelus angulatus Pease. Annaa [Anaa] Island [Tuamotu Archipelago]. Lectotype ANSP 40671. Width 5 mm, 
height 5 mm. 

Figure 14. Fastigiella squamulosa Pease. [Anaa Island] Paumotus [Tuamotu Archipelago]. Lectotype ANSP 36702. Width 1 1 
mm, height 27 mm. 

Figure 15. Latirus liratus Pease. Marquesas [Islands]. Lectotype MCZ 302628. Width 15 mm, height 28.5 mm. 

Figure 16. Latirus gibbus Pease. Howland Island [0°49'N, 176°40'W]. Lectotype MCZ 261182. Width 8 mm, height 13 mm. 

Figure 1 7. Latirus granulosa Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 261 1 81 . Width 1 1 mm, height 20.5 mm. 

Figure 18. Mitra saltata Pease. Islands of the Central Pacific. Lectotype MCZ 260613. Width 3 mm, height 7.5 mm. 

Figure 19. Leptothyra costata Pease. Maui [Hawaiian Islands]. Lectotype MCZ 245261 . Width 4 mm, height 4 mm. 

Figure 20. Mitra assimilis Pease. [Huahine [sic] Island, Society Islands]. Lectotype ANSP 28718. Width 7 mm, height 17 mm. 

Figure 21 . Marginella pyriformis Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 29541 . Width 2 mm, height 4 mm. 

Figure 22. Marginella pacifica Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 2941 5. Width 2.5 mm, height 4 mm. 

Figure 23. Marginella paumotensis Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 29497. Width 2.5 mm, height 
5 mm. 

Figure 24. Mucronalia gracilis Pease. Tahiti [Society Islands]. Lectotype MCZ 288506. Width 2 mm, height 4 mm. 

Figure 25. Engina nodicostata Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 260614. Width 3.5 mm, height 6 mm. 
Photographed by Kenneth J. Boss. 



MoLiAJSKS Described by W. H. Pease • Johnson 55 





^m 1 




/ 



I 




18 



Plate 7 



56 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 

Plate 8 

Figure 1 . Terebra sulcata Pease. [Honolulu] Oahu [Hawaiian Islands]. Lectotype MCZ 49967. Width 5 mm, height 22.5 mm. 

Figure 2. Narica granifera Pease. Jarvis [Island, Line Islands. 0°22'33"S, 159°54'11"W]. Lectotype ANSP 37301. Width 8.5 
mm, height 9 mm. 

Figure 3. Narica delicate Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 40201. Width 6.5 mm, height 6 mm. 

Figure 4. Nassa approximata Pease. [Ascension (Pohnpei) Island, Caroline Islands]. Lectotype MCZ 151800. Width 9.8 mm, 
height 24.5 mm. 

Figure 5. Nassa gracilis Pease. Ascension [Pohnpei Island, Caroline Islands]. Lectotype MCZ 228822. Width 11 mm, height 
19.5 mm. 

Figure 6. Nassa obliqua Pease. Islands of the Central Pacific. Lectotype MCZ 228823. Width 12 mm, height 15.5 mm. 

Figure 7. Nehta maculata Pease. Tahiti [Society Islands]. Lectotype ANSP 37490. Width 15.6 mm, height 16.5 mm. 

Figure 8. Neritina dispar Pease. Roratonga [Rarotonga, Cook Islands]. Lectotype ANSP 37714. Width 8 mm, height 9 mm. 

Figure 9. Neritopsis interlirata Pease. Annaa [Anaa] Island, [Tuamotu Archipelago]. Lectotype MCZ 73479. Width 10 mm, 
height 10 mm. 

Figure 10. Odostomia pollta Pease. Tahiti [Society Islands]. Lectotype MCZ 10562. Width 7 mm, height 7 mm. 

Figure 1 1 . Odostomia rosacea Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 1 9959. Width 2 mm, height 6 mm. 

Figure 12. Odostomia rubra Pease. [Anaa Island] Paumotus [Tuamotu Archipelago]. Lectotype ANSP 19955. Width 2 mm, 
height 8 mm. 

Figure 13. Planaxis abbreviate Pease. Tahiti [Society Islands]. Paralectotype MCZ 187833. Width 6 mm, height 9.5 mm. 

Figure 14. Odostomia striata Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 10491 . Width 1 .5 mm, height 4.5 mm. 

Figure 15. Odostomia debilis Pease. Howland [Island; 0°49N, 176°40W]. Lectotype MCZ 297933. Width 2 mm, height 9 mm. 

Figure 16. Olivella {Callianax) simplex Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 28969. Width 2 mm, height 
4.5 mm. 

Figure 17. Pachypoma virescens Pease. Tarawa Island [Kiribati]. Lectotype MCZ 302624. Width 26 mm, height 28 mm. 

Figure 1 8. Terebra sculptilis Pease. [Honolulu] Oahu [Hawaiian Islands]. Lectotype MCZ 248804. Width 4.5 mm, height 24 mm. 

Figure 1 9. Purpura marmorata Pease. Apaian [Abaiang] Island [Kiribati]. Lectotype MCZ 1 77824. Width 27 mm, height 45 mm. 

Figure 20. Omphialius turbinatus Pease. Gulf of [Golfo de] California, La Paz [Baja California Sur, Mexico]. Lectotype ANSP 
40820. Width 16.5 mm, height 14 mm. 



MoLLUSKS Described by W. H. Pease • Johnson 57 




Plate 8 



58 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 

Plate 9 

Figure 1. Planaxis atra Pease. Marquesas [Islands]. Lectotype ANSP 18282. Width 5.5 mm, height 9 mm. 

Figure 2. Planaxis abbreviata Pease. Tahiti [Society Islands]. Lectotype ANSP 18261 . Width 6 mm, height 8 mm. 

Figure 3. Rissoa gracilis Pease. [Kauai] Sandwich [Hawaiian] Islands. Lectotype MCZ 178853. Width 1 mm, height 2.5 mm. 

Figure 4. Rissolna striatula Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 178844. Width 4 mm, height 8 mm. 

Figure 5. Rissolna tenulsthata Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 19253. Width 4 mm, height 9 mm. 

Figure 6. Rissolna costulata Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 19241. Width 2 mm, height 4 mm. 

Figure 7. Rissoa flammea Pease. Caroline [Islands]. Lectotype MCZ 178868. Width 1 mm, height 2.68 mm. From Ponder and 
de Keyzer. 

Figure 8. Slstrum rugulosum Pease. Howland [Island; 0°49'N, 176°40'W]. Lectotype ANSP 36740. Width 5.5 mm, height 8 mm. 

Figure 9. Slstrum striatum Pease. Kingsmill [Islands, Kiribati]. Lectotype ANSP 36735. Width 8.5 mm, height 16.5 mm. 

Figure 10. Strlgatella brunnea Pease. Polynesia [(Paumotus) Tuamotu Archipelago]. Lectotype ANSP 29722. Width 10.5 mm, 
height 21.5 mm. 

Figure 11. Scalaria perplexa Pease. Hawaii [Hawaiian Islands]. Lectotype MCZ 181978. Width 12.5 mm, height 27 mm. 

Figure 12. Scalaria umblllcata Pease. Oahu [Hawaiian Islands]. Holotype MCZ 187393. Width 4 mm, height 9.5 mm. 

Figure 13. Scalaria chspata Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 19575. Width 7 mm, height 16 mm. 

Figure 14. Scalaria decussata Pease. Hawaii [Hawaiian Islands]. Lectotype ANSP 19585. Width 4 mm, height 10 mm. 

Figure 1 5. Styllfera deformis Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 19836. Width 4 mm, height 1 1 .5 mm. 

Figure 1 6. Scalaria paumotensis Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 1 9581 . Width 5 mm, height 9 mm. 

Figure 17. Taheltea [sic] pallida Pease. Tahiti; Huaheine [both Society Islands; localities not differentiated on label]. Lectotype 
ANSP 12659. Width 2.5 mm, height 6.5 mm. 

Figure 18. Terebra suffusa Pease. [Honolulu] Oahu [Hawaiian Islands]. Lectotype MCZ 248802. Width 10 mm, height 30 mm. 

Figure 19. Terebra costelllfera Pease. [Honolulu] Oahu [Hawaiian Islands]. Lectotype MCZ 248800. Width 6 mm, height 20 mm. 

Figure 20. Thala alba Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 28755. Width 2.5 mm, height 8 mm. 

Figure 21 . Terebra lauta Pease. Oahu [Hawaiian Islands]. Holotype ANSP 33589. Width 5.5 mm, height 20 mm. 

Figure 22. Thala angiostoma Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 28754. Width 4 mm, height 12 mm. 

Figure 23. Slstrum triangulatum Pease. Hawaii [Island]. Lectotype MCZ 295580. Width 14 mm, height 24 mm. Photographed 
by Kenneth J. Boss. 

Figure 24. Torinia discoldea Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 38804. Width 5.5 mm, height 3 mm. 

Figure 25. Trochus marmoreus Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 4061 4. Width 4.5 mm, height 6 mm. 

Figure 26. Trochus conoidalis Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 18868. Width 4 mm, height 4 mm. 



MoLLUSKS Described by W. H. Pease • Johnson 59 




60 Bulletin Museum of Comparative Zoology, Vol. 154, No. 1 



Plate 10 

Figure 1 . Triphoris costatus Pease. Annaa [Anaa Island, Tuamotu Archipelago]. Lectotype MCZ 273206. Width 8 mm, height 
25 mm. 

Figure 2. Triphoris pustulosus Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 50077. Width 5.3 mm, height 8 mm. 

Figure 3. Triphoris sulcosus Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 50080. Width 2 mm, height 8 mm. 

Figure 4. Triphoris tuberculatus Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 50081. Width 2 mm, height 8 mm. 

Figure 5. Triforis marmorata 'Pease' Martens and Langkavel. [Kauai] Sandwich [Hawaiian] Islands. Lectotype MCZ 50055. 
Width 3 mm, height 10.5 mm. 

Figure 6. Triphoris brunneus Pease. Apaiang [Abaiang] Island [Kiribati]. Lectotype MCZ 73922. Width 2 mm, height 7 mm. 

Figure 7. Triphoris minimus Pease. [Haena] Kauai [Hawaiian Islands]. Lectotype MCZ 50071. Width 1 mm, height 3 mm. 

Figure 8. Triphoris pallidus Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 50073. Width 1 .5 mm, height 6.5 mm. 

Figure 9. Triphoris granosus Pease. Tahiti [Society Islands]. Lectotype MCZ 273207. Width 2 mm, height 7 mm. 

Figure 10. Triphohs gracilis Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 50058. Width 1.5 mm, height 7.5 mm. 

Figure 11. Triphoris oryza Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 50072. Width 2 mm, height 7.5 mm. 

Figure 12. Triphoris robustus Pease. Makaimo [Makemo Island, Tuamotu Archipelago]. Lectotype MCZ 73923. Width 2 mm, 
height 5.5 mm. 

Figure 13. Triphoris maculatus Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 50069. Width 2 mm, height 4 mm. 

Figure 14. Triphoris similis Pease. [Haena] Kauai [Hawaiian Islands]. Lectotype MCZ 50079. Width 1.5 mm, height 4.5 mm. 

Figure 15. Triphoris perfectus Pease. Kauai [Hawaiian Islands]. Lectotype MCZ 302553. Width 1 mm, height 4 mm. 

Figure 16. Triton intermedius Pease. Oahu [Hawaiian Islands]. Lectotype MCZ 191331 . Width 22.5 mm, height 42 mm. 

Figure 17. Triton cylindricus Pease. Tahiti [Society Islands]. Lectotype MCZ 239749. Width 4.5 mm, height 1 1 mm. 

Figure 18. Truncatella concinna Pease. Apaiang [Abaiang Island, Kingsmill Islands, Kiribati]. Lectotype MCZ 178650. Width 3 
mm, height 7 mm. 

Figure 19. Truncatella pacifica Pease. Oualan [Kusaie or Kosrae Island, Caroline Islands]. Lectotype MCZ 59799. Width 3 mm, 
height 8 mm. 

Figure 20. Turricula modesta Pease. [Ponape Island, Caroline Islands] Polynesia. Lectotype ANSP 28780. Width 1 2 mm, height 
38 mm. 

Figure 21. Turricula (Costellaria) fortiplicata Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 28844. Width 3 mm, 
height 7 mm. 

Figure 22. Turbonilla elongata Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 10537. Width 5 mm, height 18 mm. 

Figure 23. Turcica coreensis Pease. Corea [Korea] Sea. Lectotype MCZ 104609. Width 21.5 mm, height 25 mm. 

Figure 24. Turricula (Pusia) nodulosa Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 28713. Width 4 mm, height 
9 mm. 

Figure 25. Trivia corrugata Pease. Paumotus [Tuamotu Archipelago]. Lectotype MCZ 303451 . Width 4.5 mm, height 6 mm. 

Figure 26. Trochus excilis Pease. Pautomus [Tuamotu Archipelago]. Lectotype MCZ 104617. Width 4 mm, height 6 mm. 

Figure 27. Turricula (Costellaria) plicatula Pease. Paumotus [Tuamotu Archipelago]. Lectotype ANSP 28845. Width 3 mm, 
height 7 mm. 



MoLLusKs Described by W. H. Pease • Johnson 61 




(US ISSN 0027-4100) 



Bulletin OF THE 

Museum of 

Comparative 

Zoology 



MCZ 
LIBRARY 



The Neotropical Orb-Weaver Genus 
Metazygia (Araneae: Araneidae) 



HERBERT W.LEVI 



HARVARD UNIVERSITY 

CAMBRIDGE, MASSACHUSETTS, U.S.A. 



VOLUME 154, NUMBER 2 
8 MAY 1995 



(US ISSN 0027-4100) 



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Occasional Papers on Mollusks, 1945- 

SPECIAL PUBLICATIONS. 

1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963. Phylogeny and 
Evolution of Crustacea. 192 pp. 

2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Tere- 
dinidae (Mollusca: Bivalvia). 265 pp. 

3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- 
derms. 284 pp. 

4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the 
Present Day. 236 pp. 

5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology 
and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 
725 pp. 

6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. 

Other Publications. 

Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. 
Reprinted 1964. 

Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of 
Insects. (Bulletin of the M.C.Z., Vol. 108.) Reprinted 1971. 

Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. 

Lyman, C. P., and A. R. Davve (eds.), 1960. Proceedings of the First Inter- 
national Symposium on Natural Mammalian Hibernation. (Bulletin of 
the M.C.Z., Vol 124.) 

Ornithological Gazetteers of the Neotropics (1975-). 

Peters' Check-list of Birds of the World, vols. 1-16. 

Proceedings of the New England Zoological Club 1899-1947. (Complete 
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© The President and Fellows of Harvard College 1995. 



THE NEOTROPICAL ORB-WEAVER GENUS METAZYGIA 
(ARANEAE: ARANEIDAE) 



HERBERT W. LEVM 

Abstract. Metazygia are Neotropical nocturnal orb 
weavers. Many species are small, less than 5 mm total 
length. They are found from the southeastern United 
States to Argentina, but most species occur in the 
Amazon area and southeastern South America. Al- 
though lacking a paramedian apophysis in the male 
palpus, they exhibit other characters that suggest that 
they be grouped (together with Eustala) close to Al- 
paida. 

There are 86 Neotropical species: 68 new (79%) 
and only 18 previously known (21%). Also, there are 
two Nearctic species, M. carolinalis and M. calix, 
making a total of 88 species of Metazygia. Of the 68 
new species, 18 are known from both sexes, 16 from 
the male only, and 34 from the female only. Six names 
are synonymized for the first time. 

The females of all species are believed to rest in a 
retreat at the side of the web during daytime, and 
many build the orb with an open sector adjacent to 
the retreat. 

INTRODUCTION 

This is one of a series of revisions of 
Neotropical orb weavers (complete list in 
Levi, 1993b). These revisions should make 
it possible for researchers to identify Neo- 
tropical orb weavers, not possible earlier 
as some previously described species had 
never been illustrated and males had not 
been matched to females. Examining and 
illustrating the holotype specimens of old 
names is one of the most important tasks 
of the revisor. After all the genera of the 
family have been revised, the relationship 
of the genera to each other can be studied. 

METHODS AND MATERIALS 

The methods have been described in de- 
tail in Levi (1993b). As in previous papers, 



' Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge, Massachusetts 02138. 



eye sizes are expressed as ratios, comparing 
the diameter of the measured eye (with 
cornea in profile) to that of the anterior 
median eyes (Levi, 1993b, figs. 27, 28). 
Distances between eyes of the anterior row 
are expressed as diameters of the anterior 
median eyes (in profile); distances between 
eyes of the posterior row are given as di- 
ameters of the posterior median eyes. The 
height of the clypeus (the distance be- 
tween the lower edge of the anterior me- 
dian eyes and the edge of the carapace) is 
given in diameters of an anterior median 
eye (Levi, 1993b, fig. 28f). These mea- 
surements are approximate, as araneid eyes 
are variable and difficult to measure ac- 
curately. 

Secondary Homonyms. The superb spi- 
der catalogs by Petrunkevitch, Roewer, and 
Bonnet, which so greatly facilitate the work 
of systematists, lumped genera. As a result 
of lumping genera, secondary homonyms 
are created: specific names that are unique 
in their own genera turn out to be hom- 
onyms when placed in the large genera 
Aranea or Araneus, having been used with 
Aranea or Araneus previously. 

Petrunkevitch (1911) and also Roewer 
(1942) made new names for the secondary 
homonyms. I have dismissed these new 
names when returning species to their pre- 
vious genera. In this I have followed other 
authors. For example, Petrunkevitch (1911) 
lumped 18 genera, replacing Singa moesta 
with metuens, Singa maculata with tusus, 
and Singa variabilis with varians, among 
others. These Petrunkevitch replacement 
names have not been used in Comstock 
(1912), Gertsch (Comstock, rev. edit. 



Bull. Mus. Comp. Zool., 154(2): 63-151, May, 1995 



63 



64 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Gertsch 1940), or Kaston (1948). All have 
tried to avoid name changes. 

According to Article 59 of the Interna- 
tional Code of Zoological Nomenclature 
(ICZN, 1985), a junior secondary hom- 
onym replaced before 1961 is permanently 
invalid. Article 59b says that if the re- 
placement name for such a junior second- 
ary homonym is the cause of confusion, 
then the case is to be referred to the Com- 
mission. Article 59d says "a species group 
name rejected after 1960 on grounds of 
junior secondary homonymy is to be re- 
instated by anyone who believes the two 
species group taxa in question are not con- 
generic. . . ." The newer method (59d) has 
been used by American arachnologists all 
along and has avoided confusion. 

Roewer (1942, 1955) replaced some sec- 
ondary homonyms cited in this revision 
(e.g., Aranea palloides Roewer for Me- 
tazygia pallidula, Aranea errans for Me- 
tazygia erratica). But Roewer had the good 
judgment not to use Petrunkevitch's re- 
placement names (e.g., for Singa). Also, 
Bonnet (1955-59) did not use the replace- 
ment names of Petrunkevitch when re- 
turning species to their original genera 
(e.g., Petrunkevitch changed Zilla guttata 
to gemellus when he placed it in Araneus, 
but the name is back to Zilla guttata in 
Bonnet, 1959). Perhaps the International 
Commissioners should make the rules more 
flexible. 

It is unfortunate that many younger ar- 
achnologists still give their new species 
overworked names such as pallida, ma- 
culata, and variabilis, which are likely to 
have been used before in the same family 
or in related families and may be the cause 
for later discovery of secondary homony- 
my. 

Lectotypes. As in previous papers, lec- 
totypes have been designated when syn- 
types belonged to different species. They 
were not indicated routinely as an aspect 
of the revision; there is no requirement to 
do so (ICZN, 1985: Art. 74). A decision has 
to be made on whether to designate a male 
or a female as the lectotype. This choice 



may become critical later, if it is found 
that the presumed species actually consists 
of sibling species recognizable only in one 
sex, not in the other. 

Collections. Specimens from the follow- 
ing collections were used. I thank the cu- 
rators for making the material available 
for this study: 



AD A. Dean, Texas A and M Uni- 

versity, College Station, Texas, 
United States 

AMNH American Museum of Natural 
History, New York, United 
States; N. Platnick, L. Sorkin 

ANSP Academy of Natural Science, 

Philadelphia, Pennsylvania, 
United States 

BMNH Natural History Museum, 
London, England; P. Hillyard, 
F. Wanless 

CAS California Academy of Sci- 

ences, San Francisco, United 
States; W. J. Pulawski, D. 
Ubick 

CD C. Deeleman, Ossendrecht, 

Netherlands 

cue Cornell University Collection, 

kept in the AMNH; N. Plat- 
nick 

CV Carlos Valderrama A.; Bogota, 

Colombia 

DU D. Ubick, San Francisco, Cal- 

ifornia, United States 

FSCA Florida State Collection of Ar- 

thropods, Gainesville, Florida, 
United States; G. B. Edwards 

HECO Hope Entomology Collec- 
tions, Oxford University, Ox- 
ford, England; J. Lansbury 

IBNP Inventario Biologico Nacional, 

San Lorenzo, Paraguay; J. A. 
Kochalka 

lELP Instituto de Ecologia, Casilla 

20127, La Paz, Bolivia; E. For- 
na, J. Coddington 

lESC Instituto Ecologia y Systema- 

tica, Cuban Academy of Sci- 
ence, Havana, Cuba, L. Armas 



METAZYGIA'Levi 



65 



INPA Institute Nacional de Pesquis- 

as da Amazonia, Manaus, Est. 
Amazonas, Brazil; C. Magal- 
haes 

IRSNB Institut Royal des Sciences Na- 

turelles de Belgique, Brussels, 
Belgium; L. Baert 

JEC J. Carico, Lynchburg, Virgin- 

ia, United States 

JMM J. Maes, Leon, Nicaragua 

MACN Museo Argentine de Ciencias 
Naturales, Buenos Aires, Ar- 
gentina; E. A. Maury 

MCN Museu de Ciencias Naturais, 

Funda9ao Zoobotanica do Rio 
Grande do Sul, Porto Alegre, 
Rio Grande do Sul, Brazil; E. 
H. Buckup 

MCP Pontificia Universidade Cato- 

lica do Rio Grande do Sul, Por- 
to Alegre, RS, Brazil; A. A. Lise 

MCZ Museum of Comparative Zo- 

ology, Cambridge, Massachu- 
setts, United States 

MECN Museo Ecuatoriano de Cien- 
cias Naturales, Quito, Ecua- 
dor; L. Aviles 

MEG M. E. Galiano; Buenos Aires, 

Argentina 

MIUP Museo de Invertebrados, Uni- 

versidad de Panama, Panama 
City, Panama; D. Quintero A. 

MLP Museo de Universidad Na- 

cional, La Plata, Argentina; R. 
F. Arrozpide 

MNHN Museum National d'Histoire 
Naturelle, Paris, France; J. 
Heurtault, C. Rollard 

MNHNC Museum Nacional de Historia 
Natural, Havana, Cuba; G. 
Alayon 

MNRJ Museu Nacional, Rio de Ja- 

neiro, Brazil; A. Timotheo da 
Costa 

MNSD Museo Nacional de Historia 

Natural, Santo Domingo, Re- 
publica Dominicana; Felix Del 
Monte 

MUSM Museo de Historia Natural, 
Universidad Nacional Mayor 



de San Marcos, Lima, Peru; D. 
Silva D. 

MZSP Museu de Zoologia, Universi- 

dade de Sao Paulo, Sao Paulo, 
SP, Brazil; P. Vanzolini, J. L. 
Neme 

MZUF Museo Zoologico de "La Spe- 

cola" Universita di Firenze, 
Florence, Italy; S. Mascherini 

NMW Naturhistorisches Museum, 

Vienna, Austria; J. Gruber 

NRMS Naturhistoriska Riksmuseet, 

Stockholm, Sweden; T. Kro- 
nestedt 

PAN Polska Akademia Nauk, War- 

szawa, Poland; A. Riedel, J. 
Proszynski, A. Slojewska, E. 
Kierych 

PMY Peabody Museum, Yale Uni- 

versity; C. Remington, D. G. 
Furth 

REL R. E. Leech; Edmonton, Al- 

berta, Canada 

SMF Forschungsinstitut Sencken- 

berg, Frankfurt am Main, 
Germany; M. Grasshoff 

SR Susan Riechert; Knoxville, 

Tennessee, United States 

USNM National Museum of Natural 

History, Smithsonian Institu- 
tion, Washington, D.C., Unit- 
ed States; J. Coddington 

ZMB Zoologisches Museum der 

Humbolt Universitat, Berlin, 
Germany; M. Moritz 

ZMK Zoologisk Museum, Copenha- 

gen, Denmark; H. Enghoff 

ZSM Zoologisches Staatssammlung, 

Munich, Germany 

I would also like to thank R. Buskirk 
and W. Eberhard for natural history in- 
formation and G. B. Edwards and P. Van- 
zolini for information on collecting sites. 
C. D. Dondale gave advice on a nomen- 
clatural problem. Laura Leibensperger 
helped thoughout and read the manu- 
script. L. R. Levi improved the wording. 
E. H. Buckup and two anonymous readers 
carefully read the manuscript and made 



66 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



numerous corrections and improvements. 
D. Sherry helped with the final word pro- 
cessing. The research was started with the 
help of NSF grant BMS 75-05719. Publi- 
cation costs of this study were covered in 
part by the Wetmore Colles Fund. 

Metazygia F. P.-Cambridge 

Metazygia F. P.-Cambridge, 1903: 501. Type species 
by original designation M. wittfeldae (McCook). 
The gender of the name is feminine. 

Diagnosis. Metazygia differs from oth- 
er araneid genera by having a glabrous 
carapace with posterior median eyes that 
face up and almost touching each other, 
and an oval abdomen, that is widest at the 
middle (Figs. 5, 12). 

In many species, but not all, the female 
has a round, laterally compressed scape on 
the ventral face of the epigynum (Figs. 1- 
4). 

The palpus of the male has only one 
patellar seta and lacks a paramedian 
apophysis (Figs. 45, 112). 

Contiguous posterior median eyes are 
found also in Larinia and Cyclosa and 
among some species of Alpaida, Araneus, 
Aculepeira, and Dubiepeira. Larinia dif- 
fers from Metazygia by having an elon- 
gate abdomen. These genera (except Al- 
paida) differ from Metazygia by having 
the carapace setose and by the annulate 
scape. All except Alpaida and Cyclosa have 
two macrosetae on the palpal patella, and 
the male palpus has a distal hematodocha. 
Cyclosa differs by having the carapace 
narrow in the cephalic region. Both Al- 
paida and Cyclosa have a paramedian 
apophysis in the male palpus. Metazygia 
females can easily be confused with those 
of Singa, Nuctenea (Larinioides) , and 
Zygiella because of similar abdomen col- 
oration and shape. These three genera are 
not found in the Neotropical region, al- 
though Zygiella x-notata (Clerck) and 
Larinioides sclopetaria (Clerck) are com- 
mon in Chile where they have been intro- 
duced. No Metazygia species are known 
from Chile. Zygiella males differ from Me- 



tazygia by having a cone-shaped palpal 
tibia, as is common in tetragnathids; Nuc- 
tenea and Larinioides males have two pal- 
pal patellar macrosetae. Some Metazygia 
females have been confused with Chry- 
someta, but this has tetragnathid charac- 
ters (Levi, 1986). The species of these sim- 
ilar genera all make a tube-shaped, silken 
retreat. 

It is possible to have doubts and misplace 
Metazygia species if only a female is avail- 
able. 

Relationship. Metazygia is closest to the 
genus Eustala. Shared apomorphies in- 
clude the anteriorly projecting scape in 
some Metazygia females (Figs. 263, 270), 
the lateral placement of the median 
apophysis in the male palpus (M in Figs. 

44, 45), and, in the palpus of larger species, 
a semitransparent blister below the distal 
prong of the terminal apophysis (B in Figs. 

45, 46). In spite of these unusual shared 
characters, Eustala is distinguished by the 
position of the posterior median eyes, on 
a slight swelling and facing laterally, and 
by the shape of the abdomen, subtrian- 
gular to elongate, widest anteriorly and 
often with a median white streak on the 
underside. Also, the carapace of Eustala is 
setose, while that of Metazygia is glabrous 
(Table 1). 

Both Metazygia and Eustala, although 
lacking a paramedian apophysis in the male 
palpus (Figs. 45, 112), have to be grouped 
near Alpaida and other genera having a 
paramedian apophysis. They mostly have 
only a single palpal patellar seta and the 
position of the conductor of the male pal- 
pus is usually on the inside face of the 
tegulum (and not on the outside rim of the 
tegulum as in Araneus-related genera). 
There is a relative absence of the distal 
hematodocha in the male palpus, a struc- 
ture also prominent in Araneus-relaied 
genera. There is abundant pigment around 
the eyes in Metazygia species, as there is 
in Alpaida species. Many of the genera 
related to Alpaida (but not Alpaida or 
Metazygia) have abdomens with two or 
three posterior, median humps on the ab- 



METAZYGIA'Levi 



67 



Table 1. Differential characters of Parawixia (PARW), Eriophora (ERIO), Acanthepeira (ACAN), 
Wagneriana (WAGN), Eustala (EUST), Acacesia (ACAC), Alpaida (ALPA), Ocrepeira (OCRE), 

Cyclosa (CYCL), and Metazygia (METZ). 



















CYCL 




EUST 




PARW 


ERlO 


acan 


WAGN 


ACAC 


ALPA 


OCRE 


(prov.) 


METZ 


(prov) 


Pattern 






















Carap. glabrous 


- 


- 


- 


- 


— 


[ + *] 


— 


_ 


[ + ] 


_ 


Paired spots on carap. 


+ * 


- 


+ 


- 


- 




— 


- 




_ 


Marks between ME and LE 


+ * 


_* 


+ 


- 


- 


- 


_* 


- 


— 


— 


Black eye rings 


- 


- 


- 


- 


- 


[ + *] 


- 


- 


+ * 


— 


Sides of thoracic reg. black 


- 


- 


- 


[ + *] 


- 




_* 


— 


— 


— 


Pattern on sternum 


[ + *] 


- 


- 




- 


— 


— 


— 


_ 


_ 


Abd. V. black with white streak 




- 


- 


- 


- 


- 


- 


- 


- 


[ + ] 


Female morphology 






















Eye reg. narrow 


- 


- 


- 


- 


[ + ] 


- 


— 


+ 


- 


— 


LE on sides of tuber. 


+ 


- 


+ 


- 




- 


- 


- 


- 


— 


PME on slight swelling 


_* 


- 


- 


- 


+ 


- 


+ * 


- 


— 


+ 


PME touch 


- 


- 


- 


- 


- 


- 


— 


+ 


[ + ] 


— 


Carap. swollen behind eyes 


+ 


+ 


+ 


+ 


- 


+ 


+ * 


- 




- 


Abd. with tubers 


4-15 


0-3 


12 + 


9-15 


- 


_* 


[2*] 


0-2 


- 


_* 


Ant. median abd. tuber 


_* 


- 


+ 


- 


- 


- 




— 


— 


- 


Abd. subspherical 


+ * 


+ 


+ 


- 


- 


- 


— 


— 


+ 


+ * 


Abd. oval 


_* 


- 


- 


+ 


+ 


+ 


_* 


+ 


[ + ] 


+ * 


Abd. with tail 


- 


- 


- 


+ * 


- 


- 


— 


+ * 




+ * 


Three median post, tubers 


+ 


_* 


+ 


+ 


— 


- 


— 


— 


— 


— 


Abd. glabrous 


- 


- 


- 


- 


- 


[ + ] 


- 


- 


[ + ] 


- 


Epigynum 






















Scape 


+ 


+ 


+ 


- 


+ 


— 


+* 


+ 


+ * 


+ 


Lat. flattened scape 


- 


- 


- 


- 


- 


— 


- 


- 


[ + *] 


- 


Lobe 


- 


- 


- 


+ 


- 


+ 


_* 


— 


+ * 


— 


Knob at tip 


_* 


- 


— 


+ 


- 


_* 


- 


— 


— 


— 


Notch on face 


- 


- 


- 


- 


- 


[ + *] 


- 


- 


— 


— 


Scape ant. projection 


- 


- 


- 


- 


- 




- 


- 


_* 


[ + ] 


Male morphology 






















Ceph. reg. wide 


- 


- 


[ + ] 


- 


- 


- 


- 


- 


- 


- 


Hook on coxa I 


+ 


+ 


- 


+ 


+ 


+ * 


+ 


+* 


+ 


+ 


Macrosetae on coxa III, IV 


+ * 


+ * 


+ * 


— 


+ * 


+ * 


+* 


+ * 


+ * 


— 


Trochanter IV macrosetae 


+ * 


+ * 


— 


— 


- 


_* 


- 


- 


- 


- 


Tibia II modified 


- 


- 


— 


— 


+ 


_* 


- 


- 


- 


- 


Carapace with lobes 


- 


- 


- 


- 


- 


- 


- 


- 


[ + *] 


- 


Fangs clasping 


- 


- 


- 


- 


- 


- 


- 


- 


[ + *] 


- 


Palpus 






















Much wider than long 


- 


- 


- 


- 


- 


- 


- 


[ + ] 


- 


- 


Patella macrosetae 


1** 


[2**] 


1 


1 


1 


1* 


1 


1 


1 


1 


Y narrow 


- 


[ + ] 


- 


- 


- 


- 


- 


- 


- 


- 


PM present 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


- 


+ 


M lateral 


- 


- 


- 


- 


- 


- 


+ 


- 


+ 


- 


SA blister-shaped 


— 


- 


- 


- 


- 


- 


- 


- 


+ 


- 


M soft 


- 


- 


- 


- 


- 


- 


- 


- 


_* 


- 



* There are exceptions. 

** P. bistriata and E. nephiloides (Levi, 1971) have a large macroseta and a smaller one on the male* 
palpal patella. 

Bracketed characters are autapomorphies for the genus. 

Abbreviations: abd., abdomen; ant., anterior; carap., carapace; ceph., cephalic; post., posterior(ly); prov., 
provisionally; reg., region; tuber(s)., tubercle(s); v., venter. LE, lateral eyes; M, median apophysis; ME, median 
eyes; PM, paramedian apophysis; PME, posterior median eyes; Y, cymbium. 

Data from Levi, 1971, 1976, 1977, 1988, 1991b, 1992, 1993b. 



68 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



domen (Eriophora, Parawixia, Wagner- 
iana, Acanthepeira, and also Eustala), a 
feature uncommon in any other spider. 
Perhaps the lack of paramedian apophysis 
and the sometimes intermediate position 
of the conductor (Fig. 112) indicate an 
intermediate relationship (Table 1). 

Description. The cephalothorax is or- 
ange to orange-brown in alcohol, the legs 
rarely with dark rings. The abdomen has 
a characteristic folium pattern consisting 
of pairs of brackets (Figs. 5, 124) or some- 
times a Zygiella-\ike pattern (Figs. 58, 110). 
The smallest species have a white (green 
or silvery when alive) abdomen with a 
black band around the anterior margin 
(Figs. 325, 330). The green color, which 
washes out in alcohol, is known for M. 
octama, M. serian, and M. lopez (Eber- 
hard, personal communication). The black 
band around the anterior is not found in 
other Neotropical araneids. The carapace 
has few setae, and the median eye quad- 
rangle is always narrower posteriorly (ex- 
cept in M. vaupes, where it is square). The 
anterior median eyes are slightly larger 
than the posterior medians, and the laterals 
are always the smallest. The height of the 
clypeus is less than the diameter of an an- 
terior median eye. The eye region is usu- 
ally all black but in some species lacks pig- 
ment (Fig. 336). The cephalic region of 
the carapace is more than half the width 
of the thoracic region; in M. uma (Fig. 
224), it is almost equal in width. 

The abdomen is oval, widest in the mid- 
dle. In some small species, it is almost 
spherical (Figs. 306, 307); in some large 
species, almost cylindrical (Fig. 228). 
Sometimes the abdomen is anteriorly flat- 
tened (Fig. 317) or anteriorly projecting 
(Fig. 242) or has an anterior notch (M. 
vaupes. Fig. 301). 

The males of three species have modi- 
fied fangs (see later). 

Genitalia. In the larger species, the epi- 
gynum has a laterally flattened scape, 
which is round in lateral view (Figs. 1-4, 
69-71). The greatest diversity of epigyna 
occurs in the small species, where some 



even have a long scape (Figs. 322, 328). 
In many species, the scape appears to be 
torn off by the male after mating, and it 
may be unusual to find a female with the 
scape intact (Figs. 277-280). (Removal of 
the scape by the male may protect its sperm 
by preventing additional female matings.) 
Often it is not known whether there is a 
scar from a torn scape or the middle area 
is sculptured (Figs. 285-287, 365, 366). Part 
of the base of the epigynum is torn off in 
M. mundulella (Fig. 234) and may be 
missing also in M. saturnino (Figs. 197, 
198) and M. amalla (Figs. 247, 248). While 
it is common for an araneid male to re- 
move the scape, destruction of the base of 
the epigynum is not otherwise known in 
araneid spiders. In M. erratica, the open- 
ing of the epigynum is sealed with an 
amorphous black secretion that is difficult 
to remove (Figs. 370-372). A similar brown 
exudate may be present on the epigynum 
of M. lopez. In still other species, part of 
the male embolus breaks and plugs the 
opening of the epigynum (Fig. 40). 

Internal female genitalia were exam- 
ined in two pairs of species: M.wittfeldae 
and M. dubia, and M. zilloides and M. 
keyserlingi. No differences were found in 
the similar species that might be useful for 
determinations. 

Male. Males of some small species have 
the carapace margin lobed above the first 
coxae (Figs. 384, 390), a modification not 
seen in males of other genera. All males 
have one macroseta on the palpal patella. 
Males of all except three species (M. gre- 
galis, M. benella, and M . yobena) have a 
tooth on the endite that faces a tooth or 
tubercle at the proximal end of the palpal 
femur. These three species also have mod- 
ified fangs: the fangs have a lobe (Figs. 
261, 262, 269, 276), presumably to hold 
the female during mating. Metazygia gre- 
galis also has the distal end of the chelic- 
erae modified as a protecting flange (Fig. 
261). 

All species have a small hook on the 
distal margin of the first coxa, but if very 
small, the hook may have moved, to face 



Metazygia 'Levi 



69 



the second coxa. The second tibiae are usu- 
ally thicker than the first and are armed 
with macrosetae. Males may have macro- 
setae on the first tibia, also. Some males 
have a short macroseta on the fourth coxa 
and sometimes on the third, as in related 
genera (those with one patellar macroseta). 

Some species have a large terminal 
apophysis (A) in the palpus, with a ter- 
minal prong above a transparent blister (B 
in Figs. 45, 112); others have lost the ter- 
minal apophysis and have only one sclerite 
beyond the embolus (E) or none (Fig. 260). 
The part remaining with the embolus (Figs. 
221, 230, 237) might be a reduced terminal 
apophysis or the embolus lamella (L of 
Figs. 45, 54). There is no way at present 
to determine the homology. Here it is called 
the "lamella" (in keys and descriptions). 
None of the Metazygia species has a para- 
median apophysis. As in Eustala, the me- 
dian apophysis has moved ventrally to the 
side of the palpus (M in Figs. 44, 45, 112, 
113) and has lost its sclerotization, often 
becoming soft and white. The conductor 
(C) has moved in the same direction, and 
there is no projection from the conductor 
toward the cymbium, as in Alpaida, to 
form a paramedian apophysis. 

The position of the conductor in ara- 
neids may be on the rim of the tegulum, 
visible in ventral view as in Neoscona (Levi, 
1993a, C in figs. 6, 7, 16), in Dubiepeira 
(Levi, 1991a, center of fig. 525, C in fig. 
526), and in many species of Araneus (Levi, 
1991a, figs. 3, 14, the light, round sclerite 
in center of fig. 22), or it may be on the 
inside face of the tegulum, closer to the 
cymbium, as in most species with a para- 
median apophysis. In the latter case, it is 
also surrounded on the outside by the wall 
of the tegulum, as in Micrathena (Levi, 
1985, C in figs. 6, 9), Alpaida (Levi, 1988, 
C in fig. 10), and Wagneriana (Levi, 1991b, 
fig. 19). In Metazygia, which lack a para- 
median apophysis, the conductor may be 
on the rim, as in M. crewi (C in Fig. 113) 
and M. isabelae (at 8 hr in Fig. 92), or 
inside, below the rim of the tegulum, as 
in M. castaneoscutata (center of Fig. 308), 



or below the rim but hanging over it, as 
in M. zilloides (C in Figs. 44, 45) and M. 
gregalis (at 10 hr in Fig. 259, C in Fig. 
260). 

In addition to the blister-shaped subter- 
minal apophysis and the lateral position of 
the median apophysis, there are additional 
peculiarities in the palpus of the larger 
species. The radix is pushed "down," out 
of the way, and is in a much "lower" po- 
sition (R in Figs. 44, 45, 112) than in spe- 
cies of other genera. Also, there is a stipes 
(I) in the form of a band that overlaps the 
dorsal surface of the palpal bulb (bottom 
third of Fig. 46), to which the embolus and 
its lamella are attached (bottom left of Fig. 
46, and also Fig. 112). Finally, the median 
apophysis may be in a more common po- 
sition (at 5 hr in Fig. 243; M in Fig. 245), 
the radix farther "up" in the palpus (below 
the embolus in Fig. 308). 

There are many small species in Me- 
tazygia. As is common in spiders, the 
smallest species display the greatest diver- 
sity in genitalia. Great diversity in geni- 
talia is also known for Micrathena species, 
mostly medium-sized. Matching males 
with females of the same species is difficult 
because so many species are similar in ap- 
pearance, differing only in genitalic struc- 
tures. 

It has not been possible to clearly group 
Metazygia species into subgenera because 
the diversity of characters does not fall into 
corrolative patterns. The larger species 
have a pattern of brackets on the abdomen 
(Fig. 5), have a terminal apophysis in the 
palpus (A in Fig. 45), and have the median 
apophysis (M in Figs. 44, 45) soft and to- 
ward the side. Small species have a black 
band around the anterior of the abdomen 
(Figs. 381, 404), a diversity of female epi- 
gyna, have the male palpus without ter- 
minal apophysis, and the median apoph- 
ysis in the more common araneid position 
at 4-5 hr in the left palpus (Figs. 383, 389). 
However, the Metazygia curari female 
(Figs. 144-146) has the characteristic flat- 
tened, round scape, as does M. wittfeldae 
(Figs. 1-4), and the male lacks a terminal 



70 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



apophysis (Fig. 148). Metazygia mundu- 
lella also has this kind of scape (Figs. 231- 
234), but the male has a median apophysis 
with sclerotized points at 5 hr in the left 
palpus (Fig. 237). Metazygia serian (Fig. 
175) and M. adisi (Fig. 141), with a flat- 
tened round "Metazygia" scape, have the 
black band of small species around the an- 
terior of the abdomen. 

Natural History. All species build a ver- 
tical orb and have a retreat that is usually 
above the web and attached to a branch, 
wall, ceiling, or curled leaf. The spider 
rests in the retreat during the day and in 
the center of the web at night. Some webs 
have a vacant sector in the part of the orb 
adjacent to the retreat (Plate 1): the vacant 
sector orb is known for M. wittfeldae, zil- 
loides, keyserlingi, chicanna, and incerta. 
The webs of M. keyserlingi and M. lati- 
ceps are kept up during the day and have 
a signal line (Eberhard, personal com- 
munication). "All other species built at 
night, many (all?) quite early in the eve- 
ning and not generally have either an open 
sector or a recognizable retreat" (Eber- 
hard, personal communication). Metazy- 
gia incerta rebuilds orbs every two to four 
days (Buskirk, personal communication). 
Some species take down their webs during 
the day (Lubin, 1978). 

While Metazygia wittfeldae is usually 
solitary, the web size and structure are the 
same when they aggregate and the angle 
of the orb continues to vary from just hor- 
izonal to vertical at Monteverde, Costa Rica 
(Buskirk 1986). 

Eberhard (personal correspondence) 
writes, 

I have watched both gregalis and octama 
build in great hurry (rapidly, with little 
exploratory behavior) just as the light is 
failing, and have web photos of chenevo 
... at 6 pm; serian at 5 pm, lopez at 7 pm; 



wittfeldae here [Costa Rica] also builds ear- 
ly in the evening. Thus I suspect these spe- 
cies are working on the flush of insects which 
fly just at dusk. One keyserlingi also had a 
web up at night and since I saw another 
web of this species which was rebuilt around 
noon after rain, I suspect it is like gregalis 
in having not one but a series of webs dur- 
ing each 24 hour period. . . . 

Species occur often in great abundance, 
females and males together. But because 
they are difficult to collect by sweeping or 
beating, many species are present in col- 
lections only as single individuals. 

The following observations are excerpt- 
ed from Eberhard (personal communica- 
tion): 

Relatively open habitat (rel. early sec- 
ondary growth, grass): lopez, gregalis, oc- 
tama, pallidula, wittfeldae, yobena, be- 
nella, lazepa, serian, chenevo. I suspect 
some species at least of preferring to be near 
water (esp. pallidula), and of liking twigs, 
barbed wire or other relatively rigid sup- 
ports for their webs, but have seen yobena 
and chenevo on webs in tall grass. On build- 
ings (especially near lights): wittfeldae, 
gregalis, dubia. Silk retreats, more or less 
cylindrical (open at both ends — spider will 
leave on rear if bothered from front side) 
during the day: octama, gregalis, (in this 
case, often in cracks or other protected 
sides); the retreat of this species generally 
has no connection whatsoever with the web, 
which is often left intact during the day 
when the spider is in its retreat, and it is 
thus generally impossible to associate a giv- 
en spider with the vestiges of a given web 
during the day. In contrast, octama re- 
moves the web completely during the day, 
I think usually without a single line being 
left up, and its retreat is at least sometimes 
on a green leaf in relatively exposed posi- 



Plate 1 . A, Metazygia chicanna n. sp. B, M. dubia (Keyserling). C, photograph of web of M. crew/ (Banks); horizontal diameter 
6 cm. D, photograph of web of M. keyserlingi Banks, horizontal diameter in middle about 12 cm. 



M £ TAZYCIA • Levi 7 1 




72 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 




Map 1 . The number of species of Metazygia in various areas. 



tion. I don't know much about the retreats 
of the other nocturnal species (and since 
retreats may be disassociated from webs, I 
was unhkely to be aware of the spider's 
retreat when I found the spider on a web 
at night). 

Dynamics of webs: I can only give you 
details for gregalis and octama. The oc- 
tama web seems relatively fragile, and the 
spiders had often torn down the web and 
were feeding on a ball of prey not more 
than a couple of hours after dusk. I never 
saw one of these (they lived in our yard in 
Cali) put up another web, but might have 
missed it (espcially if a second web was put 
up just before dawn and then soon torn 
down). I had some students do all night 
projects with gregalis, and they found that 
the same individual built two to three webs 
per night. Generally the first web was just 
at dusk, and the others substantially later 
at night. 

M. gregalis is a generalist when it comes 
to feeding. I have the impression that in- 
traspecific variation in the form in the Me- 
tazygia orb is relatively high in M. gregalis. 

Sizes of prey items are reported in Cas- 
tillo and Eberhard (1983). 



Distribution. Metazygia is known only 
from the Americas. There are several pairs 
of very similar allopatric species, one north, 
the other south, without overlap: wittfel- 
dae and dubia, and zilloides and keyser- 
lingi. 

Metazygia species of the southeastern 
United States, not otherwise cited in this 
paper, are as follows: M. carolinalis (Arch- 
er) (the male is unknown); and M. calix 
(Walckenaer), NEW COMBINATION. 
Metazygia calix (Levi, 1976, figs. 137-144) 
was placed in Alpaida but has genitalia 
similar to those of M. laticeps (Figs. 226, 
227, 230) and M. sendero (Figs. 216, 217, 
221). 

Misplaced Species. Metazygia livida 
Mello-Leitao, 1941: 151, 12. Female from 
Argentina is a Dictyna (Dictynidae). 

Metazygia unquiformis: — Valle and 
Valle, 1972: 33 is Alpaida veniliae (Key- 
serling) (Levi, 1988: 402). 

Keys. Keys are difficult to construct for 
species of which only one or a few indi- 
viduals are known. With few specimens, 
one does not know whether or not the col- 
oration is characteristic, whether or not the 
epigynum has been torn apart by a male 
when mating, whether the male has a vir- 
gin embolus with a cap or if he has mated, 
and whether all males of the species have 
a macroseta on the fourth coxa or only the 
one sampled. 

For using the key, the female's epigyn- 
um has to be slightly pulled out with a 
curved needle to see the posterior face of 
the structure. 

Key to Female Metazygia 

1. Epigvnum with an anterior projection 

(Figs. 254, 263, 270) _ 2 

Epigynum otherwise 4 

2(1). Epigynum in ventral view wider than 
long (Fig. 254); Central America, West 
Indies to northern Argentina (Map 3E) 

gregalis 

Epigynum in ventral view longer than 
wide (Figs. 263, 270) 3 

3(2). Epigynum with a deep notch on the pos- 
terior border (Fig. 270); anterior pro- 
jection without pair of wings (Fig. 270); 

Amazon area; Sao Paulo (Map 3C) 

yobena 



METAZYGIA'Levi 



73 




Map 2. Distribution of Metazygia species. 



- Epigynum without notch on posterior 5(4). 
border (Fig. 263); anterior projection 
with a pair of wings (Fig. 263); Pan- 
ama, Colombia (Map 3C) benella 

4(1). Epigynum in posterior view with a me- - 

dian plate forming a septum in an hour 
glass-shaped depression as in Figure 

108; Greater Antilles (Map 21) crewi 6(5). 

Epigynum with median posterior plate 
otherwise 5 



Epigynum in ventral view with a scape 
that extends beyond the posterior mar- 
gin of the base (Figs. 193, 277, 298, 
303, 314, 322, 328) 6 

Epigynum with scape not extending be- 
yond posterior margin (Figs. 2, 55, 231) 
12 

In ventral view scape extending from 
epigynum's posterior margin (Figs. 
298, 314) 7 



74 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



- Scape extending from middle or anterior 

of basal plate (Figs. 193, 277, 303, 322, 

328) 8 

7(6). Epigynum with notch on each side (Fig. 
298), anterior of abdomen indented 
(Fig. 301); western Amazon area (Map 
4 A) vaupes 

- Epigynum without notch (Fig. 314), ab- 

domen oval to subspherical (Fig. 317); 
Rio de Janeiro State, Brazil (Map 4B) 

floresta 

8(6). Scape attached on anterior of base (Figs. 

193, 277, 328) 9 

- Epigynum with scape attached in mid- 

dle of plate (Figs. 303, 322) 11 

9(8). Scape thick and with a deep groove (Fig. 

193); Bahia State, Brazil (Map 3A) 

uratron 

- Scape otherwise 10 

10(9). Scape thin and transparent (Fig. 328); 

Amazon area to Misiones Prov., Ar- 
gentina (Map 4C) lagiana 

- Scape thick, opaque (Fig. 277); Amazon 

area to northern Argentina (Map 3D) 

voluptifica 

11(8). Epigynum with notch on each side in 
posterior view (Fig. 323); Mexico, 
Central America (Map 4G) .... nigrocincta 

- Epigynum with posterior view otherwise 

(Fig. 305); Amazon area (Map 4B) 

castaneoscutata 

12(5). Abdomen with paired, dark patches (Fig. 

90); Goias State, Brazil (Map 2C) 

redfordi 

Abdomen marked otherwise (Figs. 58, 

175) 

13(12). Epigynum with scape round, laterally 

flattened (Figs. 1-4, 62-64, 158-160, 

1 64- 1 66 ) 

- Epigynum otherwise (Figs. 114, 133, 189, 

370) 

14(13). Epigynum with bordered depression or 

opening, visible in ventral view (Figs. 

40, 55, 74, 87, 158, 168, 172) 

Epigynum otherwise (Figs. 2, 35, 129, 

1 44, 23 1 ) 22 

15(14). Openings oval, very large and on each 

side of epigynum (Fig. 74); Mexico, to 

Guianas and Peru (Map 2E) palliduta 

Openings otherwise (Figs. 40, 48, 55, 121, 

1 58, 1 59) 1 6 

16(15). Openings along anterior of plate (Figs. 

40, 49, 158, 168) 17 

- Openings in middle or posterior 20 

17(16). In posterior view epigynum longer than 

wide; with narrow median plate (Fig. 
169); northern Amazon region, Guian- 
as (Map 2K) uraricoera 

In posterior view epigynum wider than 
long, median plate wide (Figs. 41, 50, 
159) _.._ 18 



18(17). Openings anterior, lateral (Figs. 39, 48) 



19 



13 



14 



42 



19(18). 



20(16). 



Openings near median, indistinct (Fig. 
158); western Amazon area (Map 2K) 
yucumo 

United States to Honduras, West Indies 
(Map 2G) zilloides 

Costa Rica, Trinidad to Colombia (Map 
2G) keyserlingi 

Abdomen with black band around an- 
terior (Fig. 175); Costa Rica (Map 2) 
serian 

Abdomen otherwise 21 

21(20). Openings round in center of each side 
(Fig. 55), scape small, light, indistinct 
(Fig. 55); southern Mexico to Hon- 
duras, Jamaica (Map 2B) chicanna 

- Openings tiny notches toward posterior 

of base, scape large, distinct (Fig. 121); 

Mexico (Map 21) taman 

22(14). In posterior view of epigynum, width of 
median plate equal to or less than that 
of laterals (Figs. 63, 95) or median plate 
T-shaped with vertical piece narrow 
(Figs. 139, 165) 23 

- In posterior view median plate wider than 

laterals (Figs. 30, 36, 70, 126) or oth- 
erwise 

23(22). A scale on each side of epigynum as in 
Figures 94 and 95; southeastern Brazil 

(Map 2C) rogenhoferi 

Epigynum without scale (Figs. 62, 139, 
1 65 ) 24 

24(23). Median plate T-shaped (Figs. 139, 165) 

25 



26 



Median plate otherwise (Fig. 63); west- 
ern Amazon area (Map 2B) tapa 

25(24). Arms of T-shaped median plate con- 
stricted at base and pointed (Fig. 139); 
Amazon area (Map 2H) adisi 

Arms not constricted (Fig. 165); southern 
Brazil, northern Argentina (Map 2K) 

ipanga 

15 26(22). Base of epigynum on each side with shal- 
low lateral notch as in Figures 231 and 

234; southeastern Brazil (Map 3F) 

mundulella 

Base without such notches 27 

27(26). Margin of base in ventral view entire, 
without notches on sides (Fig. 69); 
Central America (Map 2J) incerta 

- Margin of base otherwise (Figs. 22, 100, 
121, 181) 28 

28(27). Folds posterior to scape in ventral view 
(Fig. 100); Sao Paulo State, Brazil (Map 
2F) harueri 

- Epigynum otherwise 29 

29(28). A transverse bar posterior to scape in 

ventral view as in Figure 181; Colom- 
bian Amazon area (Map 3A) lazepa 

- Epigynum otherwise 30 



Metazygia 'Levi 



75 




Map 3. Distribution of Metazygia species. 



76 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



30(29). A dark area on each side of scape in 
ventral view as in Figure 125; Amazon 
area (Map 2H) paquisha 

- Epigynum otherwise 31 

31(30). In lateral view scape about twice as long 

as wide (Figs. 83, 146) 32 

- In lateral view scape about as long as 

wide (Figs, 18, 37, 123, 131) 33 

32(31). Abdomen with a pair of dorsal, longi- 
tudinal white lines (Fig. 147); Amazon 
area (Map 2H) curari 

- Abdomen with a pair of dorsal, longi- 

tudinal dusky bands (Fig. 84); Neblina 

area of Amazon (Map 2E) enabla 

33(31). Abdomen oval, about three quarters as 

wide as long 34 

- Abdomen elongate, about two thirds as 

wide as long (Fig. 153), epigynum as 
in Figures 150-152; Bolivia (Map 2F) 

bolivia 

34(33). Posterior median plate with a pair of 
notches on each side as in Figures 30 
and 1 78 35 

- Posterior median plate otherwise 36 

35(34). In ventral view length of scape about two 

thirds length of base (Fig. 177); Guy- 
ana (Map 3A) chenevo 

- In ventral view length of scape less than 

half length of base (Fig. 29); Peruvian 
Amazon to Bahia State, Brazil (Map 
2D) patiama 

36(34). Median plate in posterior view heart- 
shaped (Fig. 130); Ecuador (Map 2H) 

nobas 

Median plate otherwise (Figs. 23, 36, 122) 
37 

37(36). Posterior median plate hexagonal (Fig. 

122); Mexico (Map 21) taman 

Posterior median plate otherwise (Figs. 
23, 36) 38 

38(37). Epigynum in ventral view with a slit 
depression on each side as in Figure 
35; Pernambuco, Brazil (Map 2D) .. ipago 

- Epigynum otherwise (Figs. 1, 2, 9, 16, 

22) 39 

39(38). Abdomen with paired spots (Fig. 25); 
posterior median plate about as long 
as wide (Fig. 23); Venezuela, Peru 

(Map 2D) pimentel 

Abdomen with foHum (Figs. 5, 12, 19); 
posterior median plate slightly wider 

than long (Figs. 3, 10, 17) 40 

40(39). Bahama Islands (Map 2A) bahama 

- United States to northern South Ameri- 

ca, West Indies 41 

41(40). Base of epigynum almost twice as wide 
as long in ventral view (Fig. 2); United 
States to Costa Rica (Map 2A) 

wittfeldae 

Base of epigynum narrower, about three 
eighths as long as wide (Fig. 16); Costa 



Rica, West Indies, Galapagos to north- 
ern Brazil and Peru (Map 2A) dubia 

42(13). Epigynum with set off scape (Figs. 133, 

185, 208, 309, 378) 43 

- Epigynum without scape or with only 

scars of torn scape (Figs. 280, 285, 292, 

360, 370, 391) 49 

43(42). Scape with transverse wrinkles (Fig. 208); 

Cuba (Map 2J) matanzas 

Scape smooth (Figs. 133, 185, 189) 44 

44(43). Scape with dent on each side as in Figure 

185; Peruvian Amazon (Map 3 A) 

atalaya 

- Scape otherwise 45 

45(44). Scape ventrally flattened (Figs. 189, 378) 

46 

Scape knob-like (Figs. 114, 133, 309) 47 

46(45). Scape subtriangular (Fig. 378); Peruvian 

Amazon region (Map 4E) genaro 

Scape oval (Fig. 189); Colombian Am- 
azon region (Map 3B) corima 

47(45). Posterior median plate with concave sides 
(Fig. 310); Panama, Colombia (Map 

4B) octama 

Posterior median plate convex on each 

side (Figs. 115, 134) 48 

48(47). Posterior median plate wider than long 

(Fig. 115); Guatemala (Map 21) 

vaurieorum 

- Posterior median plate almost as wide as 

long (Fig. 134); Lower Amazon area 

(Map 2K) goeldii 

49(42). Epigynum in posterior view longer than 

wide (Figs. 240, 338) 50 

- Epigynum in posterior view wider than 

long (Figs. 361, 386) 51 

50(49). Posterior median plate narrower dorsal- 
ly than ventrally (Fig. 338); Bolivian 
Amazon area (Map 4C) loque 

- Posterior median plate almost rectan- 

gular (Fig. 240); coast of southeastern 

Brazil (Map 3F) genialis 

51(49). Epigynum with scars, scape usually torn 

off' (Figs. 197, 198, 248, 280, 292) 52 

Epigynum without distinct scars (Figs. 

216, 226, 365, 391) 58 

52(51). Scars in midline only (Figs. 203, 280, 

285, 289) 54 

- Whole venter of epigynum seemingly 

torn off (Figs. 197, 247) 53 

53(52). Posterior median plate wider than long 
(Fig. 248); southern Brazil (Map 3D) 

amalla 

Posterior median plate square (Fig. 198); 

southern Brazil (Map 3A) saturnino 

54(52). Posterior median plate in a depression 
(Fig. 286); southern Brazil (Map 3A) 
viriosa 

- Posterior median plate otherwise (Figs. 

204, 281, 290, 293) 55 

55(54). Posterior median plate much wider than 



METAZYGiA'Levi 77 




Map 4. Distribution of Metazygia species. 



78 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



long (Fig. 290); Mato Grosso, Brazil 

(Map 3D) ituari 



Posterior median plate otherwise (Figs. 
204, 281, 293) 



56 



56(55). Posterior median plate Y-shaped as in 
Figure 293; Peruvian Amazon (Map 

4A) limonal 

Posterior median plate otherwise 57 

57(56). Epigynum in ventral view with a round, 
shallow depression on each side as in 
Figure 280; Amazon area to northern 
Argentina (Map 3D) voluptifica 

- Epigynum in ventral view with a raised 

area on each side as in Figure 203; 

southeastern Brazil (Map 3B) 

crabroniphila 

58(51 ). Cephalic area of carapace almost as wide 
as thoracic area; abdomen with lon- 
gitudinal stripes as in Figure 224; Am- 
azon area (Map 3B) uma 

Carapace otherwise 59 

59(58). Epigynum in ventral view pentagonal as 
in Figure 333; Colombian Amazon 
(Map 4C) carimagua 

- Epigynum otherwise 60 

60(59). Epigynum subtriangular in ventral view 

with shallow median groove and with- 
out distinct lip as in Figures 216 and 
226; cephalic region of carapace rel- 
atively wide (Figs. 219, 228) 61 

Epigynum otherwise (Figs. 346, 360, 391) 

62 

61(60). Posterior median plate longer than wide 
(Fig. 226); Panama to Rio de Janeiro 
State, Brazil (Map 3F) laticeps 

Posterior median plate wider than long 
(Fig. 217); Ecuador, Peru (Map 3B) . 

sendero 

62(60). Epigynum in ventral view with posterior 
margin lobed and median area swollen 
as in Figures 342 and 351 63 

Epigynum otherwise (Figs. 346, 360, 365, 

370) 64 

63(62). Epigynum with depression on ventral 
face (Figs. 351, 354), posterior median 
plate dumb-bell-shaped (Fig. 352); 
Venezuela to Peru (Map 4D) lopez 

- Epigynum swollen on ventral face (Fig. 

342), posterior median plate triangu- 
lar (Fig. 343); Hispaniola (Map 4D) ... 

_ _ cienaga 

64(62). In ventral view posterior margin of epi- 
gynum with a swollen lip as in Figures 
360, 365, 370 and 405 65 

- Posterior margin of epigynum without 

swollen lip (Figs. 346, 385, 409) 68 

65(64). Lip a horizonatal bar as in Figure 405; 
posterior median plate T-shaped (Fig. 
406); Rio Grande do Sul, Brazil (Map 

4F) ._ „ _ valentim 

Lip V-, U-, or T-shaped (Figs. 360, 365, 
370) „ _ 66 



66(65). Posterior median plate pentagonal (Fig. 
371); black amorphous material on 
each side posteriorly (Figs. 370, 372); 
Amazon area (Map 4E) erratica 

- Posterior median plate not pentagonal 

(Figs. 361, 366); without black amor- 
phous material 67 

67(66). Posterior median plate triangular (Fig. 

361); Peruvian Amazon (Map 4D) 

samiria 

- Posterior median plate square, anterior 

to it a textured area (Fig. 366); Am- 
azon region, Bolivia (Map 4D) ducke 

68(64). Posterior margin with a notch in middle 
and a lobe extending each side as in 
Figure 346; Amazon area (Map 4D) .. 
souza 

- Posterior margin otherwise, often with a 

pair of lobes (Figs. 385, 391, 401, 409) 

69 

69(68). In ventral view a pair of lobes as in Fig- 
ure 385; posterior median plate dumb- 
bell-shaped (Fig. 386); Mato Grosso, 

Brazil (Map 4E) voxanta 

Ventral view otherwise (Figs. 391, 401, 

409) 70 

70(69). A median ventral notch in posterior view 

of epigynum (at 12 hr in Figs. 402, 

410) 7 1 

No notch visible in posterior view (Figs. 

392, 395); Colombia, Ecuador to mouth 

of Amazon (Map 4F) peckorum 

71(70). Posterior median plate constricted ven- 
trally (at 12 hr in Fig. 402); Ecuador- 
an, Peruvian Amazon (Map 4F) . moldira 
Posterior median plate as in Figure 410; 
Bahia to Sao Paulo States, Brazil (Map 
4F) bahia 

Key to Male Metazygia 

1. Cheliceral bases or fangs modified with 

transparent lobes (Figs. 261, 262, 269, 

276) 2 

Cheliceral bases or fangs not modified .. 

4 

2(1). Median apophysis (M in Fig. 260), in 
mesal view short (Figs. 258, 260); Cen- 
tral America, West Indies, South 

America (Map 3E) _ gregalis 

Median apophysis, in mesal view, longer 
(Figs. 267, 274) 3 

3(2). Median apophysis with a black wall (at 
4 hr in Fig. 274, at 6 hr in Fig. 275); 
Amazon area, Sao Paulo State, Brazil 
(Map 3C) yobena 

- Median apophysis without black wall 
(Figs. 267, 268); Panama, Colombia 
(Map 3C) benella 

4(1). Fourth coxae with a macroseta or point- 
ed tubercle 5 

Fourth coxae without macroseta or tu- 
bercle 15 



METAZYGIA'Levi 



79 



5(4). 

6(5). 

7(6). 
8(7). 

9(8). 

10(8). 



Fourth coxae with a tubercle; embohis 
of palpus thorn-like (Fig. 357, E in Fig. 
359); Venezuela, Amazon area (Map 
4D) lopez 

Fourth coxae with macroseta 6 

Carapace with a lobe above first coxae 
(Fig. 390) 7 

Carapace without lobe above first coxae 

13 



Abdomen posteriorly black (Fig. 388); 
Mato Grosso, Brazil (Map 4E) voxanta 

Abdomen posteriorly light 

In mesal view, median apophysis pro- 
jecting beyond tegulum (at 3 hr in Fig. 
318, at 3 hr in Fig. 423) 9 

Median apophysis smaller and not pro- 
jecting beyond tegulum (Figs. 313, 383, 
4 1 8, 425 ) 1 

Median apophysis very large, facing 
cymbium at 3 hr in Figure 423; Bahia, 
Brazil (Map 4H) atama 

Median apophysis distally rectangular in 
mesal view at 3 hr in Figure 318; Am- 
azon area (Map 4C) mariahelenae 

Embolus S-shaped as seen through trans- 
parent lamella (between 11 hr and at 

center in Fig. 313) octama 

- Embolus otherwise, usually hidden (Figs. 

383, 418, 425) 11 

11(10). Median apophysis with bulge on side (at 

3 hr in Fig. 425, right of center in Fig. 
426); coastal Ecuador (Map 41) oro 

Median apophysis without bulge (Figs. 

383, 4 1 8 ) .' 1 2 

12(11). Median apophysis subtriangular (at 3 hr 
in Fig. 418); Rio Grande do Sul, Brazil 
(Map 4H) cunha 

Median apophysis distally enlarged (at 4 
hr in Fig. 383); Peruvian Amazon (Map 

4E) genaro 

13(6). Embolus a thread with a transverse loop 
as in Figures 112 and 113; Greater An- 
tilles (Map 21) crewi 

Embolus otherwise (Figs. 327, 332) 14 

14(13). Median apophysis distally tapering to a 
point (at 3 hr in Fig. 327); Mexico, 
Central America (Map 4G) .... nigrocincta 

Median apophysis distally bulging as in 

4 hr in Figure 332; southern Amazon 
region to Misiones Prov., Argentina 
(Map 4C) lagiana 



420); eastern Para State, Brazil (Map 

41) aldela 

18(15). Abdomen with median, transverse light 

band (Fig. 417); palpus as in Figure 

416; Amazon area (Map 41) cazeaca 

Abdomen and palpus otherwise 19 

Palpus with terminal apophysis (A in Figs. 

45, 112; top of Figs. 162, 201, 207) 20 

Palpus without terminal apophysis, only 

a lamella (Figs. 142, 148, 213, 221, 

225, 230, 237, 243-246) 38 

8 20(19). Terminal apophysis with distal straight 

or slightly curved prong (Figs. 6, 13) 



15(4). 

16(15). 

17(16). 



Carapace with small lobe above first coxa 

(Figs. 376, 415, 421) 



Carapace without lobe 18 

Abdomen with ventral black band as in 

Figure 377; Peru (Map 4E) manu 

Abdomen marked otherwise 17 

Sickle-shaped embolus (Fig. 414); Co- 
lombian Amazon (Map 4F) rothi 

Embolus barely visible, hidden by large 
sclerotized lamella (at 11 hr in Fig. 



19(18). 



21 



- Terminal apophysis with short distal 

prong (between center and 2 hr in Fig. 

91); Mato Grosso, Brazil (Map 2C) 

_ isabelae 

21(20). In mesal view terminal apophysis with 
two prongs (between center and 2 hr 
in Fig. 6); southeastern United States 
to Costa Rica (Map 2A) wittfeldae 

- Terminal apophysis otherwise (Figs. 13 

27) 

22(21). A comb projecting beyond prong in mes 

al view (at 1 hr in Figs. 20, 27, 66) ... 

No comb-like projection (Figs. 13, 201 



22 



23 



25 



23(22). Comb-like projection longer than wide 
in mesal view (at 1 hr in Fig. 20); te- 
gulum with pointed spine (at 3 hr in 
Fig. 21); Costa Rica to Guianas and 
northern Peru, Galapagos Islands and 
West Indies (Map 2A) dubia 

- Comb-like projection in mesal view wid- 
er than long (at 1 hr in Figs. 27, 66); 
tegulum without pointed spine (Figs. 
28, 67 ) 24 

24(23). Comb small (at 1 hr in Fig. 66), most of 
embolus hidden behind conductor 
(Figs. 66, 67); northern Peruvian Am- 
azon (Map 2B) pastaza 

Comb large (at 1 hr in Fig. 27), most of 
embolus exposed, only tip of conduc- 
tor hidden (at 3 hr in Fig. 27); Ven- 
ezuela to Peruvian coast (Map 2D) ._. 
pimentel 

25(22). A pointed tooth projecting beyond prong 
of terminal apophysis in mesal view 
(at 1 hr in Fig. 13, center in Fig. 14); 

Bahama Islands (Map 2A) bahama 

_ 26 



No such tooth 



16 26(25). Median apophysis projecting beyond 
other sclerites toward 4 hr, conductor 
toward 3 hr with triangular space be- 
tween these sclerites in mesal view 

(Figs. 201, 207); southern Brazil 27 

Median apophysis and conductor oth- 
erwise (Figs. 60, 99, 104, 213) 28 

27(26). Conductor with a knob at tip (at 3 hr in 
Fig. 201); median apophysis straight 



80 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



(at 4 hr in Fig. 201); Rio Grande do 

Sul, Brazil (Map 3A) saturnino 39(38). 

Conductor tapering to tip (at 3 hr in Fig. 
207); median apophysis with elbow (at 
4 hr in Fig. 207); southeastern Brazil 
(Map 3B) crabroniphila 

28(26). Tip of embolus with horseshoe-shaped 
structure (Fig. 104, center of Fig. 105); 
Guianas, Amazon area (Map 2F) . jamari 40(38). 
Tip of embolus otherwise, or hidden by 

conductor (Figs. 60, 99, 162) 29 - 

29(28). Embolus long, saber-shaped, and curved 41(40). 

up (Fig. 99); southeastern Brazil (Map 

2C) rogenhoferi 

Embolus otherwise (Figs. 73, 119, 162) 
30 

30(29). Embolus U-shaped (with dark cap as in 

Fig. 119); Guatemala (Map 21) carrizal 42(40). 

Embolus otherwise _ 31 

31(30). Embolus a small hook as in center of 

Figure 162; western Amazon area (Map 43(42). 

2K ) yiicumo 

Embolus otherwise 32 

32(31). Embolus almost straight structure, tip 
hidden by terminal apophysis prong 
as in Figure 85; Neblina area of Am- 
azon (Map 2E) enabla 

- Embolus otherwise 33 - 

33(32). Embolus thorn-shaped as in Figure 33; 

Peruvian Amazon to Bahia, Brazil 

(Map 2D) patiama 

Embolus otherwise; Mexico, Central 44(42). 

America 34 

34(33). Embolus lamella (L in Figs. 45, 54) cov- 
ering most of embolus with one large - 
point and a forked tooth as in Figure 

73; Central America (Map 2J) incerta 45(44). 

Embolus lamella otherwise (Figs. 44, 54, 

60, 79) 35 - 

35(34). Embolus lamella triangular tip covered 

by blister-like part of terminal apoph- 46(45). 
ysis (center in Fig. 60); southern Mex- 
ico to Honduras, Jamaica (Map 2B) ... 
chicanna - 

- Embolus lamella otherwise 36 

36(35). Embolus lamella rounded as in Figure 47(45). 

44; Florida, Texas to Honduras, Cuba, 

Jamaica (Map 2G) zilloides 

Embolus lamella otherwise 37 

37(36). Embolus lamella with axis of tip at right 

angle to axis of cymbium (Fig. 54); 48(47). 

embolus without lobes (Fig. 54); Costa 
Rica to Trinidad and Colombia (Map 

2G) „ keyserlingi 

Embolus lamella with tip pointing to- 
ward 1 hr in Figure 79; embolus with 
two lobes below (Figs. 79, 80); Mexico 49(48). 

to Guianas and Peru (Map 2E) . pallidula 

38(19). Median apophysis long, thumb-shaped - 

and projecting beyond tegulum (at 3 
hr in Figs. 302, 308) 39 



Median apophysis otherwise 40 

Embolus lamella projecting beyond 
cymbium edge as at 11 hr in Figure 

308; Amazon area (Map 4B) 

castaneoscutata 

Cymbium edge extending beyond la- 
mella as at 11 hr in Figure 302; Am- 
azon area (Map 4A) vaupes 

Tegulum with a large fold having a comb 
(at 12 hr in Figs. 283, 295) 41 

Tegulum otherwise 42 

Embolus axis at right angle to axis of 
cymbium (Fig. 283); Amazon area to 
southeastern Brazil (Map 3D) voliiptifica 

Embolus axis forming an acute angle with 
that of cymbium (Fig. 295); Guyana 
(Map 4A) tanica 

Tegulum with a large distal lobe (at 12 
hr in Figs, 237, 243) 43 

Tegulum otherwise or only small lobe 44 

Lamella with subparallel sides distally 
concave (at 3 hr in Fig. 243, at 10 hr 
in Fig. 244, L in Fig. 245); median 
apophysis with one point (at 5 hr in 
Fig. 243, M in Fig. 245); Bahia State 
to Rio Grande do Sul, Brazil (Map 3F) 
genialis 

Lamella distally narrowing (at 3 hr in 
Fig. 237); median apophysis with two 
points (at 5 hr in Fig. 237); southeast- 
ern Brazil (Map 3F) mundiilella 

Median apophysis "hanging down"; up- 
side-down T-shaped (at 6 hr in Fig. 
250); Guianas (Map 3D) ikuruwa 

Median apophvsis otherwise (Figs. 142, 
149, 213, 22 i, 230) 45 

Lower edge of median apophysis semi- 
circular (at 5 hr in Figs. 213, 230) 46 

Median apophysis otherwise (Figs. 142, 
148, 221, 225) 47 

Palpus as in Figure 230; Panama to Rio 

de Janeiro State, Brazil (Map 3F) 

laticeps 

Palpus as in Figure 213; Bolivia to Mato 
Grosso do Sul (Map 3B) corumba 

Embolus thread-like curving "above" te- 
gulum (at 12 hr in Fig. 142); Neblina 
area of Amazon (Map 2H) arnoi 

Embolus otherwise (Figs. 148, 221, 225) 
48 

Median apophysis having a "bottom" 
bulge and tip hidden by conductor in 
mesal view (Figs. 148, 149); Amazon 
area (Map 2H) curari 

Median apophvsis otherwise (Figs. 221, 
225) ' 49 

Median apophysis as in Figure 225; Am- 
azon area (Map 3B) uma 

Median apophysis as in Figure 221; Ec- 
uador to Amazonian Peru (Map 3B) . 
sendero 



Metazygia 'Levi 



81 



Metazygia wittfeldae (McCook) 
Figures 1-7; Map 2A 

Epeira wittfeldae McCook, 1894: 168, pi. 7, figs. 6, 
7, 2, 6. Three female, two male, and one imm. male 
syntypes from Florida in ANSP, examined. 

Metazygia wittfeldae:— F. P.-Cambridge, 1904: 501, 
pi. 47, figs. 22, 23, 2, 6. Roewer, 1942: 368. Bonnet, 
1957: 2820. Levi, 1977: 92, pi. 6, figs. 90-103, 2, S. 

Description. Female from Tabasco, 
Mexico. Carapace orange, cephalic region 
dark orange. Chelicerae dark orange. La- 
bium, endites, sternum orange. Coxae, legs 
orange. Dorsum of abdomen with gray 
pattern on white pigment spots (Fig. 5); 
venter light orange-gray, without marks. 
Posterior median eyes 0.5 diameter of an- 
terior medians, laterals 0.7 diameter. An- 
terior median eyes 0.7 diameter apart, 1.3 
diameters from laterals. Posterior median 
eyes 0.2 diameter apart. Height of clypeus 
equals 0.5 diameter of anterior median eye. 
Total length 8.0 mm. Carapace 4.2 mm 
long, 3.1 wide, 1.9 behind lateral eyes. First 
femur 3.2 mm, patella and tibia 4.1, meta- 
tarsus 3.0, tarsus 1.2. Second patella and 
tibia 3.8 mm, third 2.3, fourth 3.1. 

Male from Tabasco, Mexico. Color as in 
female but cephalic region lighter. Pos- 
terior median eyes 0.7 diameter of anterior 
medians, laterals 0.7 diameter. Anterior 
median eyes 0.7 diameter apart, 1 diam- 
eter from laterals. Posterior median eyes 
0.3 diameter apart, 2.1 diameters from lat- 
erals. Height of clypeus equals 0.5 diam- 
eter of anterior median eye. Second tibia 
thicker than first, both first and second 
with macrosetae. Total length 5.2 mm. 
Carapace 2.9 mm long, 2.1 wide, 1.1 be- 
hind lateral eyes. First femur 2.7 mm, pa- 
tella and tibia 3.5, metatarsus 3.0, tarsus 
1.1. Second patella and tibia 3.1 mm, third 
1.7, fourth 2.1. 

Note. Males and females are commonly 
collected together. 

Variation. Total length of females 7.2 
to 11.1 mm, males 4.2 to 7.2. Illustrations 
were made from specimens from Tabasco 
State, Mexico. 

Diagnosis. In ventral view the epigyn- 



um is twice as wide as long and has pos- 
terior swellings on each side (Fig. 2); in 
related species M. dubia (Fig. 16) and M. 
bahama (Fig. 9), it is narrower and lacks 
these swellings. Males are separated from 
M. dubia and M. bahama by the soft prong 
parallel to the sclerotized prong of the ter- 
minal apophysis (at 1 hr in Fig. 6), which 
is absent in the other two. The distribution 
of M. wittfeldae is allopatric with respect 
to related species (Map 2A). 

Natural History. Specimens were col- 
lected under eaves of buildings and in brush 
and are commonly found in mud-dauber 
wasp nests. 

Distribution. From southeastern United 
States, Virginia to Costa Rica. Its distri- 
bution does not overlap that of M. bahama 
and M. dubia (Map 2A). United States and 
some Mexican records on Map 2 come from 
Levi (1977). 

Specimens Examined. MEXICO Tamaulipas: 
Ciudad Mante (AMNH); Tampico (AMNH). San Luis 
Potosi: Tamazuchale (AMNH, CAS); Valles (AMNH). 
Zacatecas: Tabasco (MCZ). Nayarit: Tepic (AMNH); 
27 km S Acaponeta (CAS). Colima: Santiago, NW 
Manzanillo (AMNH). Veracruz: Acayucan (CAS); 
Catemaco (AMNH, CAS); 7.5 km W Catemaco; 17 
km W Cerro Azul; Cordoba; Fortin de las Flores; 
Jalapa; La Palma; Lago Catemaca (all AMNH); Mo- 
cambo (CAS); Papantla; Tecolutla (all AMNH); Ve- 
racruz (AMNH, MCZ); Orizaba (MCZ). Guerrero: 13 
km W Acapulco (AMNH). Oaxaca: Tehuantepec; 3.2 
km NE Tehuantepec (all AMNH). Tabasco: Villa 
Hermosa (AMNH). Yucatan: Chetumal (MCZ). Chia- 
pas: Palenque Ruins (MCZ); Prusia (AMNH). BE- 
LIZE Stann Creek: Dangriga (MCZ); 80 km S Stann 
Creek (MCZ); Twin Cays, W of Swamp Dock (USNM). 
GUATEMALA Antigua; Moca; Suchitepequez; San 
Julian; Tiquizate; Variedades; Zacapa (all AMNH). 
HONDURAS Lancetilla, nr. E Tela (MCZ). NICA- 
RAGUA Lago Jiloi (SR). COSTA RICA Cartago: 
Turrialba (MCZ). Guanacaste: 4 km NW Caiias 
(MCZ); nr. Cafias (MCZ). San fose: San Jose, common 
in city (AMNH, MCZ). 

Metazygia baliama new species 
Figures 8-14; Map 2A 

Holotype. Male holotype, one male, two female para- 
types from South Bimini, Bahama Islands, June 
1951 (M. A. Cazier), in AMNH. The specific name 
is a noun in apposition after the type locality. 

Description. Female paratype. Cara- 
pace orange-brown with sides of thoracic 



82 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



region orange. Chelicerae red-brown. La- 
bium, endites dark orange. Sternum, cox- 
ae, legs orange. Dorsum of abdomen with 
pairs of black brackets (Fig. 12); venter 
light gray without marks. Posterior me- 
dian eyes 0.7 diameter of anterior medi- 
ans, laterals 0.7 diameter. Anterior median 
eyes 0.8 diameter apart, 2 diameters from 
laterals. Posterior median eyes 0.3 diam- 
eter apart, 4 diameters from laterals. 
Height of clypeus equals 0.8 diameter of 
anterior median eye. Total length 8.0 mm. 
Carapace 3.9 mm long, 2.7 wide, 1.7 be- 
hind lateral eyes. First femur 2.9 mm, pa- 
tella and tibia 3.7, metatarsus 2.5, tarsus 
1.1. Second patella and tibia 3.4 mm, third 
2.1, fourth 2.7. 

Male holotype. Color as in female. Pos- 
terior median eyes 0.6 diameter of anterior 
medians, laterals 0.6 diameter. Anterior 
median eyes 0.8 diameter apart, 1 diam- 
eter from laterals. Posterior median eyes 
0.3 diameter apart, 2.2 diameters from lat- 
erals. Height of clypeus equals 0.5 diam- 
eter of anterior median eye. Second tibia 
slightly thicker than first, with stronger 
macrosetae. Abdomen widest in middle. 
Total length 5.1 mm. Carapace 2.8 mm 
long, 2.1 wide, 1.1 behind lateral eyes. First 
femur 2.8 mm, patella and tibia 3.7, meta- 
tarsus 2.8, tarsus 1.1. Second patella and 
tibia 3.1 mm, third 1.7, fourth 2.2. 

Note. Males and females were collected 
together. 

Variation. Total length of females 6.7 
to 10.7 mm, males 4.8 to 5.7. Illustrations 
were made from the male holotype and a 
female paratype collected with it. 

Diagnosis. Epigynum of the female (Fig. 
9) is narrower than that of M. wittfeldae 
in ventral view (Fig. 2), and the posterior 
median plate is wider dorsally (at 6 hr in 
Fig. 10) than that of M. dubia (at 6 hr in 



Fig. 17) in posterior view. The male differs 
from both of these species by having a 
spine on the subterminal apophysis (at 1 
hr in Fig. 13 and center of Fig. 14). 

Natural History. This species probably 
has habits similar to M. wittfeldae and M. 
dubia. 

Distribution. Bahama Islands. The dis- 
tribution does not overlap that of M. dubia 
and M. wittfeldae (Map 2A). 

Paratypes. From type locality: June 
1950, 62, 13, imm. (M. A. Cazier, F. Rindge, 
AMNH); May 1951, 59, 43, imm. (M. Ca- 
zier, W. J. Gertsch, AMNH); June 1951, 
302, 13, imm. (M. A. Cazier; C, P. Vaurie, 
AMNH); July 1951, 13 (C, P. Vaurie, 
AMNH). 

Specimen Examined. BAHAMA ISLANDS Dog 
Key, N Andros Island, 13 May 1904, 16 (AMNH). 

Metazygia dubia (Keyserling) 
Plate 1 ; Figures 15-21 ; Map 2A 

Epeira dubia Keyserling, 1864: 123, pi. 4, figs. 12, 
13, 9. Two female syntypes from Sta. Fe de Bogota, 
N. Granada [Bogota, Colombia], in BMNH, ex- 
amined. Keyserling, 1892: 187, pi. 9, fig. 138, 9. 

Epeira moraballii Kingston, 1932: 363, figs. 53, 54, 
web. Specimens from the Essquibo River, Guyana, 
lost [not in BMNH]. NEW SYNONYMY. 

Aranea dubia: — Roewer, 1942: 841. 

A. moraballii: — Roewer, 1942: 847. 

Araneus moraballicus Bonnet, 1955: 546. 

Metazygia dubia: — Levi, 1991a: 179. 

Synonymy. Hingston's E. moraballii is 
synonymized with M. dubia because 
Hingston described the proximity of the 
posterior median eyes, the oval abdomen, 
and the Zygiella x-notata-\ike web. Also, 
only a few Guianan Metazygia are 11 mm 
total length. (The other large Metazygia, 
M. laticeps, has spinnerets anterior of the 
posterior tip, a fact noticed by Hingston 
for his Epeira folisecens, but not here.) 



Figures 1-7. Metazygia wittfeldae (McCook). 1-5, female. 1-4, epigynum. 1, anterior. 2, ventral. 3, posterior. 4, lateral. 5, 
dorsal. 6, 7, left male palpus. 6, mesal. 7, apical. 



Figures 8-14. M. bahama n. sp. 8-1 2, female. 8-1 1 , epigynum. 8, anterior. 9, ventral. 1 0, posterior. 1 1 , lateral. 1 2, dorsal. 1 3, 
14, male palpus. 13, mesal. 14, apical. 



METAZYGiA'Levi 83 




Figures 15-21, M. dubia (Keyserling). 15-19, female. 15-18, epigynum. 15, anterior. 16, ventral. 17, posterior. 18, lateral. 19, 
dorsal. 20, 21, male palpus. 20, mesal. 21, apical. 

Figures 22-28. M. pimentel n. sp. 22-26, female. 22-24, epigynum. 22, ventral. 23, posterior. 24, lateral. 25, dorsal. 26, 
abdomen, ventral. 27, 28, male palpus. 27, mesal. 28, apical. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



84 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Bonnet (1955: 466) lists the name Epeira 
dubia erroneously as a synonym of Ara- 
neus cornutus [= Larinioides cornutus 
(Clerck 1758)]. 

Description. Female from Gamboa, 
Panama. Carapace brown, sides of thorac- 
ic region orange. Chelicerae brown. La- 
bium, endites light brown. Sternum or- 
ange. Coxae light orange, legs orange. Dor- 
sum of abdomen whitish with gray marks 
(Fig. 19); venter light gray without marks. 
Posterior median eyes 0.7 diameter of an- 
terior medians, laterals 0.7 diameter. An- 
terior median eyes their diameter apart, 
two diameters from laterals. Posterior me- 
dian eyes 0.3 diameter apart. Height of 
clypeus equals 0.5 diameter of anterior 
median eye. Total length 9.5 mm. Cara- 
pace 4.1 mm long, 2.9 wide, 2.2 behind 
lateral eyes. First femur 3.4 mm, patella 
and tibia 4.0, metatarsus 3.0, tarsus 1.2. 
Second patella and tibia 3.8 mm, third 2.1, 
fourth 3.3. 

Male from Gamboa. Color as in female, 
but carapace all orange. Posterior median 
eyes 0.7 diameter of anterior medians, lat- 
erals 0.7 diameter. Anterior median eyes 
their diameter apart, 1.3 diameters from 
laterals. Posterior median eyes 0.2 diam- 
eter apart. Height of clypeus equals 0.4 
diameter of anterior median eye. Total 
length 6.7 mm. Carapace 3.5 mm long, 2.5 
wide, 1.5 behind lateral eyes. First femur 
3.5 mm, patella and tibia 4.4, metatarsus 
3.7, tarsus 1.4. Second patella and tibia 3.8 
mm, third 2.1, fourth 2.7. 

Note. Males and females were collected 
together. 

Variation. Total length of females 7.2 
to 1 1.7 mm, males 4.8 to 6.2. The abdomen 
may have dark marks or sometimes none 
at all. The scape is torn off the epigynum 
in some females. The epigynum of a fe- 
male from Depto. Huila, Colombia, is 
heavily sclerotized. A male from Vene- 
zuela had the tegulum spine short. Illus- 
trations were made from a female and male 
from Gamboa, Panama; the specimen on 
Plate 1 is from Negril, Jamaica. 

Diagnosis. The Metazygia dubia epi- 



gynum (Fig. 16) is narrower in ventral view 
than that of M. wittfeldae (Fig. 2) and 
lacks the swelling on each side. It has a 
narrower scape (Figs. 16, 17) than that of 
M. bahama (Figs. 9, 10). Males can be 
separated from the more northern M. witt- 
feldae by the distal comb-like projection 
of the male terminal apophysis (at 1 hr in 
Fig. 20, center of Fig. 21) and by the long 
black spine on the tegulum (at 3 hr in Fig. 
21). 

Natural History. This species was col- 
lected from disturbed areas outside and 
sometimes inside buildings. It was found 
under bark in Cuba and was collected from 
coral just above the high tide mark in Ja- 
maica; from pasture and from dense veg- 
etation in Jamaica; on a wire fence at Coa- 
mo, Puerto Rico; and under a roof over- 
hang and on a boat dock on Barro Colorado 
Island, Panama. Specimens from Galapa- 
gos were found in seashore vegetation. The 
spiders are nocturnal and sit in a silk re- 
treat during the day. There is no signal 
line to the retreat. The vertical web is re- 
built every evening and has a partly closed 
hub (Lubin, personal communication). 
Some specimens come from mud-dauber 
wasp nests. 

Distribution. Costa Rica, West Indies to 
Brazil and Peru and Galapagos Islands. It 
does not overlap M. wittfeldae (Map 2A). 

Specimens Examined. COSTA RICA Limon: Li- 
mon (DU). PANAMA Herrera: Sarigua (MIUP). Co- 
de: Rio Hato (MIUP). Colon: Santa Rosa (AMNH). 
Panama: very common (AMNH, MIUP, MCZ). CUBA 
Archip. Canarreos: Cayo Cantiles, Bajo Corteza 
(lESC); Cayo Rosario, Bajo Corteza (lESC); Cayo 
Avalos (MNHNC). JAMAICA very common (AMNH, 
MCZ). HISPANIOLA Dominican Republic: Samana: 
Las Terrenas (MNSD). Independencia: Bano de Zorsa 
(MNHNC); betw. Neiba and Duverge (MNSD). Dis- 
trito Nacional: Acuario Nacional, Santo Domingo 
(MNSD). PUERTO RICO Laguna Cartagena, 10 km 
SW Lajas (MCZ); Banos de Coamo (MCZ). CURA- 
gAO Hato (AMNH). VENEZUELA Sucre: Cumani 
(MCN). Monagas: Caripito (AMNH). Bolivar: Ca- 
naima [PCanaime] (AD). COLOMBIA Magdalena: 
Cienaga (IBNP); Pozo Colorado, 11 km W Santa Mar- 
ta (AMNH); San Pablo (IBNP). Atldntico: Barran- 
quilla (AMNH, IBNP). Antioquia: Mutata (MCZ). 
Huila: 10 km E Santa Leticia, Finca Meremberg, 
2,300 m (MCZ). ECUADOR Sucumbios: Cuyabeno 



Metazygia 'Levi 



85 



Reserv. (MCZ). Pastaza: Rio Pastaza, Rio Verde, Mara 
Trail, 1,200 m (AMNH). Giiayas: 3 km NE La Lib- 
ertad (CAS). Galapagos IsL: W coast Albemarle Isl. 
(AMNH): Bahia Borrero, Santa Cruz (MCZ). PERU 
Libertad: Pacasma>o (PAN); Guadalupe (PAN). 
BRAZIL Amazonas: Rio Autas, Sta. Amelia (NRMS). 
Ceard: Pacajus Guarani, 3 July 1972, 29 (Exped. Acad. 
Bras. Cienc, MZSP 12408). ' 

Metazygia pimentel new species 
Figures 22-28; Map 2D 

Holotype. Male holotype and four female paratypes 
from Pimentel, Depto. Lambayeque, Peru, 21 Sept. 
1988 (D. Silva D.), in MUSM; one paratype in MCZ. 
The specific name is a noun in apposition after the 
type locality. 

Description. Female paratype. Cara- 
pace orange, darkest in eye region. Che- 
licerae dark orange-brown Labium, en- 
dites, sternum light orange. Coxae light 
orange, legs orange. Dorsum of abdomen 
whitish with five pairs of black spots (Fig. 
25); venter with indistinct white patch sur- 
rounded by dusky area (Fig. 26). Posterior 
median eyes 0.8 diameter of anterior me- 
dians, laterals 0.8 diameter. Anterior me- 
dian eyes 0.8 diameter apart, 2.2 diameters 
from laterals. Posterior median eyes 0.2 
diameter apart. Height of clypeus equals 
0.7 diameter of anterior median eye. Total 
length 8.2 mm. Carapace 4.5 mm long, 3.4 
wide, 2.1 behind lateral eyes. First femur 
3.4 mm, patella and tibia 4.1, metatarsus 
2.7, tarsus 1.2. Second patella and tibia 3.8 
mm, third 2.5, fourth 3.4. 

Male holotype. Color as in female but 
white patch in dusky area on venter of 
abdomen is more distinct. Posterior me- 
dian eyes 0.7 diameter of anterior medi- 
ans, anterior laterals 0.7 diameter, poste- 
rior laterals 0.6. Anterior median eyes 0.4 
diameter apart, 1.2 diameters from later- 
als. Posterior median eyes 0.2 diameter 
apart. Height of clypeus equals 0.3 di- 
ameter of anterior median eye. Total length 
5.7 mm. Carapace 3.1 mm long, 2.4 wide, 
1.4 behind lateral eyes. First femur 2.9 
mm, patella and tibia 3.7, metatarsus 2.7, 
tarsus 1.2. Second patella and tibia 3.0 mm, 
third 1.9, fourth 2.3. 

Note. Males and females were collected 
together. 



Variation. Total length of males 5.7 to 
6.3. Illustrations were made from the type 
specimens. 

Diagnosis. The median, less sclerotized 
triangular area of the epigynum is more 
pointed (Fig. 22) than that of M. dubia 
(Fig. 16). The male has a comb-like pro- 
jection of the subterminal apophysis (at 1 
hr in Fig. 27) but lacks the spine on the 
tegulum that is present in M. dubia (at 3 
hr in Fig. 21). The venter of the abdomen 
has a white patch (Fig. 26) absent in sim- 
ilar species. 

Natural History. Specimens were abun- 
dant in branches of locust "algarrobos," 
Prosopis, SL leguminous tree growing in 
sand dunes in Peru, and dry to very dry 
tropical forest in Venezuela. 

Distribution. Venezuela, Peru in arid 
areas (Map 2D). 

Specimen Examined. VENEZL^ELA Falcon: Par- 
aguana Peninsula, 6 km W Nuevo Pueblo, 26 Nov.- 
4 Dec. 1960, \i (A. L. Markezich, MCZ). 

Metazygia patiama new species 
Figures 29-34; Map 2D 

Holotype. Female holotype from Fazenda Matiapa, 
Camacan, Bahia, Brazil, 16 Oct. 1978 (J. S. Santos), 
in MCN no. 11116; male paratype, same locality 
and collector, 14 Oct. 1978, in MCN no. 10182. 
The specific name is an arbitrary combination of 
letters. 

Description. Female holotype. Cara- 
pace orange. Chelicerae orange-brown. 
Labium, endites dark orange. Sternum or- 
ange. Coxae light orange; legs dark orange, 
distal parts of articles darker. Dorsum of 
abdomen setose, with anterior pair of dark 
patches on dusky white and posterior quar- 
ter dark gray (Fig. 32). Venter black, fad- 
ing toward sides. Posterior median eyes 0.7 
diameter of anterior medians, laterals 0.7 
diameter. Anterior median eyes 0.5 di- 
ameter apart, 1.2 diameters from laterals. 
Posterior median eyes 0.2 diameter apart, 
2.4 diameters from laterals. Height of 
clypeus equals 0.6 diameter of anterior 
median eye. Total length 5.6 mm. Cara- 
pace 2.8 mm long, 2.2 wide, 1.4 behind 
lateral eyes. First femur 2.5 mm, patella 



86 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



and tibia 2.7, metatarsus 1.9, tarsus 0.9. 
Second patella and tibia 2.6 mm, third 1.6, 
fourth 2.3. 

Male paratype. Color light orange, ex- 
cept for abdomen, which has tiny white 
pigment spots dorsally and lacks all gray 
or black marks. Posterior median eyes 0.8 
diameter of anterior medians, laterals 0.6 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 0.8 diameters from laterals. 
Posterior median eyes 0.2 diameter apart, 
1.3 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Total length 4.4 mm. Cara- 
pace 2.3 mm long, 1.6 wide, 0.8 behind 
lateral eyes. First femur 2.1 mm, patella 
and tibia 2.5, metatarsus 1.7, tarsus 0.7. 
Second patella and tibia 2.3 mm, third 1.3, 
fourth 1.6. 

Note. Males and females were collected 
at the same locality. 

Variation. Total length of females 5.6 
to 5.7 mm. Illustrations were made from 
the type specimens. 

Diagnosis. The epigynum (Fig. 29) is 
wider than that of M. pimentel in ventral 
view (Fig. 22), and in posterior view (Fig. 
30) the lateral plates appear wider than 
those of M. pimentel (Fig. 23) and M. 
dubia (Fig. 17). The male palpus (Figs. 33, 
34) lacks the comb-like projection of the 
terminal or subterminal apophysis found 
in M. dubia (at 1 hr in Fig. 20) and M. 
pimentel (at 1 hr in Fig. 28) and also lacks 
the spine on the tegulum. The pattern on 
the abdomen (Fig. 32) is not a complete 
folium as in related species. 

Natural History. Specimens were col- 
lected at night in Peru. 

Distribution. Amazon region of Peru to 
Bahia State, Brazil (Map 2D). 



Specimen Examined. PERU Madre de Dios: Zona 
Reservada de Manu, Puesto de Vigilancia Pakitza, 6 
Oct. 1987, 12 (D. Silva D., J. Coddington, USNM). 

Metazygia ipago new species 
Figures 35-38; Map 2D 

Holotype. Female holotype from Igarapeagu, igapo 
capim flutuante (periodically flooded forest), Est. 
Pernambuco, Brazil, 12 July 1980, in MNRJ. The 
specific name is an arbitrary combination of letters. 

Description. Female holotype. Cara- 
pace orange, cephalic region dark orange. 
Chelicerae dark orange. Labium, endites 
dark orange. Sternum orange. Coxae light 
orange, legs orange. Dorsum of abdomen 
with a faint gray folium on white (Fig. 
38); venter with some white pigment spots 
on light gray behind epigynum. Posterior 
median eyes 0.7 diameter of anterior me- 
dians, laterals 0.7 diameter. Anterior me- 
dian eyes 0.6 diameter apart, 1.2 diameters 
from laterals. Posterior median eyes 0.3 
diameter apart. Height of clypeus equals 
0.4 diameter of anterior median eye. Total 
length 6.8 mm. Carapace 3.4 mm long, 2.7 
wide, 1.5 behind lateral eyes. First femur 
3.1 mm, patella and tibia 4.0, metatarsus 
2.7, tarsus 1.1. Second patella and tibia 3.5 
mm, third 1.9, fourth 2.7. 

Diagnosis. In ventral view there is a 
groove to each side of the median area of 
the epigynum (Fig. 35). The groove is ab- 
sent in the epigynum of M. patiama (Fig. 
29) and similar species. 

Metazygia zilloides (Banks) 
Figures 39-47; Map 2G 

Epeira zilloides Banks, 1898: 255, pi. 15, fig. 2, 2, 6. 
Three female, one male, and one juvenile syntype 
from Tepic, Nayarit, Mexico, in MCZ, examined. 

Aranea dilatata F. P.-Cambridge, 1904: 513, pi. 49, 



Figures 29-34. Metazygia patiama n. sp. 29-32, female. 29-31, epigynum. 29, ventral. 30, posterior. 31, lateral. 32, dorsal. 
33, 34, left male palpus. 33, mesal. 34, apical. 

Figures 35-38. M. ipago n. sp.. female. 35-37, epigynum. 35, ventral. 36, posterior. 37, lateral. 38, dorsal. 



Figures 39^7. M. zilloides (Banks). 39-43, female. 39^2, epigynum. 39, anterior. 40, ventral. 41, posterior. 42, lateral. 43, 
dorsal. 44-46, male palpus. 44, mesal. 45-47, pulled apart. 45, mesal. 46, dorsal, cymbium removed. 47, emtxjius and lamella. 



METAZYCiA'Levi 87 




Figures 48-54. M. keyserlingi Banks. 48-53, female. 48-51 , epigynum. 48, anterior. 49, ventral. 50, posterior. 51 , lateral. 52, 
dorsal. 53, atxiomen, ventral. 54, male palpus, mesal. 

Abbreviations. A, tenninal apophysis; B, blister-like subterminal apophysis; C, conductor; E, embolus; H, hematodocha; I, stipes; 
L, embolus lamella; M, median apophysis; P, paracymbium; R, radix; Y, cymbium. 



Scale lines. 1.0 mm, genitalia 0.1 mm. 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



fig. 9, $. Male lectotype designated Levi, 1977: 92 
from Guatemala, in BMNH, examined. Roewer, 
1942: 841. Synonymized by Levi, 1977. 

Metazygia a/foonigrc— Bryant, 1940: 339, figs. 107- 
109, 111,2, (5. Erroneous determination, noi Larinia 
albonigra Franganillo. 

Aranea zilloides: — Roewer, 1942: 857. 

Araneus pallidulus:—KTaus, 1955: 24, fig. 66, S. Er- 
roneous determination. 

Araneus dilatatus: — Bonnet, 1955: 497. 

Araneus zilloides: — Bonnet, 1955: 632. 

Metazygia zilloides: — Levi, 1977: 92. 



Description. Nontype female from Te- 
pic, Nayarit, Mexico. Carapace orange, 
dusky in midline. Chelicerae, labium or- 
ange-brown. Endites, sternum orange, sides 
darker. Coxae, legs orange, distal ends of 
femora and tibiae darker. Dorsum of ab- 
domen with white pigment spots and an- 
terior black marks (Fig. 43); venter with 
white pigment spots. Eyes subequal. An- 
terior median eyes 0.8 diameter apart, 1.2 
diameters from laterals. Posterior median 
eyes 0.3 diameter apart, 2 diameters from 
laterals. Height of clypeus equals 0.4 di- 
ameter of anterior median eye. Total length 
7.4 mm. Carapace 3.0 mm long, 2.6 wide, 
1.3 behind lateral eyes. First femur 3.6 
mm, patella and tibia 4.6, metatarsus 3.1, 
tarsus 1.1. Second patella and tibia 3.7 mm, 
third 2.1, fourth 3.0. 

Male from Tepic, Mexico. Color as in 
female, but abdomen darker with poste- 
rior transverse bars, and venter with a 
transverse white patch behind genital 
groove, which is surrounded by black. Pos- 
terior median eyes same diameter as an- 
terior medians, laterals 0.9 diameter. An- 
terior median eyes 0.9 diameter apart, 1 
diameter from laterals. Posterior median 
eyes 0.2 diameter apart, 1.8 diameters from 
laterals. Height of clypeus equals 0.4 di- 
ameter of anterior median eye. Total length 
4.8 mm. Carapace 2.6 mm long, 2.1 wide, 
1.0 behind lateral eyes. First femur 3.2 
mm, patella and tibia 4.5, metatarsus 3.6, 
tarsus 1.1. Second patella and tibia 3.4 mm, 
third 1.7, fourth 2.4. 

Note. Males and females were collected 
together. 

Variation. Total length of females 3.8 



to 6.7 mm, males 3.0 to 5.0. Illustrations 
were made from nontype specimens from 
Tepic, Mexico. 

Diagnosis. Females are difficult to sep- 
arate from M. keyserlingi. The embolus 
part stuck in the opening is smaller (Fig. 
40), and there is a median ventral groove 
in posterior view of the epigynum (arrow 
in Fig. 41). The male differs from that of 
M. keyserlingi (Fig. 54) by the round shape 
of the embolus lamella (L in Fig. 44). 

Natural History. Specimens were col- 
lected in second-growth forest edge in 
Mexico and from beach grape, in a hotel, 
and in a citrus orchard and pasture in Ja- 
maica. Others came from a Sceliphron 
wasp nest in Jamaica. 

Distribution. Florida, central Texas to 
Honduras, Bahamas, Cuba, Cayman Is- 
lands, and Jamaica (Map 2G). United States 
and some Mexican records of Map 2 come 
from Levi (1977). 

Specimens Examined. MEXICO Tamaulipas: Li- 
mon (AMNH); Mante (AMNH). Nuevo Leon: Linares 
(AMNH): Los Cristales (AMNH). San Luis Potosi: 
Tamazunchale (AMNH); Valles (AMNH). Nayarit: 3 
km N Compostela (AMNH); Tepic (AMNH). Jalisco: 
Chapala (CAS); Puerto Vallarta (AMNH); Tizapan 
(AMNH). Veracruz: Catemaco (AMNH); Lago Ca- 
temaco (AMNH); Rio Blanco (MCZ); 4 km N Son- 
tecomapan (REL); Veracruz (USNM). Hidalgo: Ixmi- 
quilpan, Rio Tula (AMNH). Distrito Federal: 
(AMNH). Michoacan: Jiquilpan (AMNH); Lago Cha- 
pala (AMNH); Ciudad Michoacan (AMNH). Morelos: 
Cuernavaca (AMNH, MCZ); Tehui.xtla (AMNH). Oa- 
xaca: Temascal (MCZ); Tolosa (AMNH). Tabasco: 3 
km NE Comalcalco (AMNH); Villa Hermosa 
(AMNH). Campeche: Ciudad del Carmen (AMNH). 
Yucatan: Chetumal (MCZ); Chicxulub (CAS). Chia- 
pas: N Arriaga Mtns. (AMNH); Cacahuatan (AMNH); 
24 km SW Cintalapa (AD); 45 km SE Comitan 
(AMNH); Las Cruces (AMNH); Mapastepec (AMNH); 
Prusia (AMNH); Tonala (AMNH). GUATEMALA 
Guatemala: Amatitlan (AMNH). Quiche: Chichicas- 
tenango (AMNH). Sacatepequez: Antigua (AMNH); 
Capetillo, 1,500 m (AMNH). Suchitepequez: Moca 
(AMNH); Nebaj (AMNH); San Julian (AMNH); Var- 
iedades, 300 m (AMNH). Chimaltenango: Yepocapa 
(AMNH); San Pedro (AMNH). EL SALVADOR Can- 
delaria (AMNH), HONDURAS Copan (AMNH); 27 
km S Tegucigalpa (MCZ). BAHAMA ISLANDS An- 
dros Island: Coakley Town (AMNH), CUBA Pinar 
del Rio: Cabafias (AMNH); S Pinar del Rio (AMNH); 
San Vicente (AMNH), La Habana: Habana (MCZ, 
USNM), Matanzas: Cienaga de Zapata (MCZ); Ma- 
tanzas (AMNH). Villa Clar-a: Vega Alta (MCZ). Cien- 



Metazygia 'Levi 



89 



fuegos: Soledad (MCZ); Trinidad Mtns., Mina Carlota 
(MCZ); Topes de Collantes (AMNH). Camagiiey: 
Agramonte (AMNH); San Bias (MCZ). Holguin: Banes 
(AMNH): Valle de Maybe (MNHNC). Santiago: Si- 
bonev (AMNH); Santiago (AMNH); coast below Pico 
Turquino (MCZ). CAYMAN ISLANDS Grand Cay- 
man (MCZ). JAMAICA Christiana (AMNH); Clare- 
mont (MCZ); Evanton (MCZ); Fort Henderson 
(AMNH); Hope Gardens (AMNH); Kingston (MCZ); 
Long Mtn. (MCZ); Lucea (AMNH); Mandreville 
(AMNH); Mona (MCZ); Negril (MCZ); Old Harbour 
(MCZ); Port Henderson (MCZ); 1 km E Reading 
(MCZ); Spanish Town (MCZ). 

Metazygia keyserlingi Banks 
Plate 1 ; Figures 48-54; Map 2G 

Metazygia keyserlingi Banks, 1929: 94, pi. 4, fig. 63, 
9. Five female syntypes from Barro Colorado Is- 
land, Canal Zone [Lago Gatun, Panama Prov., Pan- 
ama], in MCZ, examined. Roewer, 1942: 868. Bon- 
net, 1957: 2820. 

Synonymy. Banks designated two vials 
with females as types, two females col- 
lected on 20-24 June and three females on 
13 July (both without year). This species 
had been erroneously synonymized with 
M. zilloides (Levi, 1977). When describing 
M. keyserlingi. Banks compared it to M. 
pallidula but not to his own M. zilloides, 
which is more similar to M. keyserlingi 
than is M. pallidula. 

Description. Female from Barro Colo- 
rado Island. Carapace orange. Chelicerae, 
labium, endites orange. Sternum orange. 
Legs orange. Dorsum of abdomen whitish 
with two indistinct black longitudinal 
bands (Fig. 52). Venter with some white 
pigment behind epigynum, dark dusky on 
each side between epigynum and spin- 
nerets, and with a faint white line on each 
side (Fig. 53). Posterior median eyes same 
diameter as anterior medians, laterals 0.8 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 1 diameter from laterals. 
Posterior median eyes 0.3 diameter apart, 
1.3 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Total length 4.5 mm. Cara- 
pace 1.9 mm long, 1.5 wide, 0.9 behind 
lateral eyes. First femur 2.1 mm, patella 
and tibia 2.5, metatarsus 1.7, tarsus 0.7. 
Second patella and tibia 1.9 mm, third 1.1, 
fourth 1.7. 



Male from Barro Colorado Island. Color 
as in female. Eyes subequal. Anterior me- 
dian eyes 1.2 diameters apart, 1 diameter 
from laterals. Posterior median eyes 0.5 
diameter apart. Height of clypeus equals 
0.6 diameter of anterior median eye. Total 
length 3.6 mm. Carapace 1.8 mm long, 1.3 
wide, 0.7 behind lateral eyes. First femur 
2.0 mm, patella and tibia 2.5, metatarsus 
2.0, tarsus 0.8. Second patella and tibia 2.0 
mm, third 1.0, fourth 1.4. 

Note. Males and females were collected 
together. 

Variation. Total length of females 3.5 
to 5.7 mm, males 2.3 to 3.6. Illustrations 
were made from specimens collected from 
the type locality: Barro Colorado Island, 
Gatun Lake, Panama. The web photo- 
graph (Pi. 1) is also from Barro Colorado 
Island, Panama. 

Diagnosis. The white patch on the ven- 
ter of the abdomen (Fig. 53) is more dis- 
crete and the groove present in M. zilloides 
(arrow in Fig. 41) is smaller or absent (Fig. 
50). The male can be told from M. zilloides 
(Fig. 44) by the embolus lamella (L in Fig. 
54), which is pointed "above" the embolus. 
The females are difficult to separate from 
M. zilloides, but all the ones collected with 
males had the central swollen part of the 
epigynum 0.32 mm wide, whereas those 
of M. zilloides had the swollen part of the 
epigynum 0.40 mm wide. The epigynum 
of M. keyserlingi has a slightly smaller 
scape than that of M. zilloides. 

Natural History. This species was col- 
lected in moist tropical forest in Costa Rica, 
in leaf litter in Panama, and in a garden 
in Call, Colombia. Males are uncommon 
in collections. 

Distribution. Costa Rica to southern Co- 
lombia, Trinidad (Map 2G). 

Specimens Examined. COSTA RICA Limon: Ca- 
huita, 31 Mar. 1979, 25 (J. Coddington, MCZ). Pun- 
tarenas: Osa, Parque Nacional Corcovado, 15 Aug. 
1978, 12 (J, Coddington, MCZ); Manuel Antonio Na- 
tional Park, 24-26 Mar. 1983, 29, 1<5 (D. Ubick, DU); 
Osa Peninsula, Llorona Station, 6 Aug. 1980, 19 (J. 
Coddington, USNM). PANAMA Colon: Fort Sher- 
man, Aug. 1939, 19 (A. M. Chickering, MCZ); Fort 
Gulick, 23 Feb. 1980, IS (Harlan, AMNH). Panama: 



90 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Barro Colorado Island, Lago Gatun, very common, 
19, 13 (AMNH, MCZ); Pipeline Road, Soberania Natl. 
Park, June 1978, 59 (F. Vollrath, MCZ), 7 Aug. 1983, 
19 (H., L. Levi, MCZ); Toboga Island, 23 Aug. 1946, 
29 (N. L. H. Krauss, AMNH). TRINIDAD St. George 
Co.: Simla, 6.4 km N Arima, 2 May 1967, 19 (C. T. 
Collins, AMNH); Arima Valley, 10-22 Feb. 1964, IS 
(P. Wygodzinsky, AMNH). COLOMBIA Valle: Cali, 
19 Oct. 1969, 19, 2 Oct. 1969, 19; Rio Jamundi, 1,000 
m, 18 km S Cali, 9 July 1969, 19, 14 Jan. 1970, 19, 
17 June 1970 (all W. Eberhard, MCZ); 4 Mar. 1973, 
39 (W. Eberhard, H. Levi, MCZ); 10 km N Piendamo, 
1,700 m, Feb. 1974, 19, 16 (W. Eberhard, MCZ). 
Narino: nr. Barbacoas, 20 m, 20 Mar. 1974, 19 (W. 
Eberhard 727, MCZ). 

Metazygia chicanna new species 
Plate 1 ; Figures 55-61 ; Map 2B 

Holotype. Female holotype and female and two male 
paratypes from Chicanna Ruins, 8 km W Xpujii, 
ca. 18°32'N, 89''31'W, Campeche, Mexico, 12-14 
July 1983 (W. Maddison), in MCZ. The specific 
name is a noun in apposition after the type locality. 

Description. Female holotype. Cara- 
pace orange, with median dusky band. 
Chelicerae, labium, endites, sternum or- 
ange. Coxae, legs orange. Dorsum of ab- 
domen with two pairs of black bands, sides 
black (Fig. 58); venter with a central white 
patch (Fig. 59). Posterior median eyes 0.8 
diameter of anterior medians, laterals 0.8 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 1 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
1.6 diameters from laterals. Height of 
clypeus equals 0.6 diameter of the anterior 
median eyes. Total length 5.5 mm. Cara- 
pace 2.2 mm long, 1.5 wide, 0.8 behind 
lateral eyes. First femur 2.1 mm, patella 
and tibia 2.7, metatarsus 1.8, tarsus 0.7. 
Second patella and tibia 2.3 mm, third 1.3, 
fourth 1.9. 



Male paratype. Color as in female, but 
legs indistinctly ringed darker, and ab- 
domen with two pairs of longitudinal 
bands. Posterior median eyes 0.9 diameter 
of anterior medians, laterals 0.8 diameter. 
Anterior median eyes 0.9 diameter apart, 
0.8 diameter from laterals. Posterior me- 
dian eyes 0.2 diameter apart, 1 .8 diameters 
from laterals. Height of clypeus equals 0.6 
diameter of anterior median eye. Total 
length 3.5 mm. Carapace 1.9 mm long, 1.4 
wide, 0.7 behind lateral eyes. First femur 
2.1 mm, patella and tibia 2.6, metatarsus 
2.1, tarsus 0.9. Second patella and tibia 2.1 
mm, third 1.1, fourth 1.6. 

Note. Males and females were collected 
together. 

Variation. Total length of females 2.5 
to 5.8 mm, males 2.5 to 4.2. Illustrations 
were made from paratypes; the photo- 
graph (Pi. 1) was made in Negril, Jamaica. 

Diagnosis. All specimens have a white 
spot on the venter of the abdomen (Fig. 
59). Females can be distinguished by the 
ventral view of the epigynum, which has 
a cone-shaped scape with openings on each 
side (Fig. 55). The scape also has a kink in 
lateral view (Fig. 57). Unlike other species, 
the male has a cone-shaped embolus la- 
mella (at center of Fig. 60). 

Natural History. The holotype came 
from short tropical rain forest. Other spec- 
imens came from moist forest border in 
Quintana Roo, Mexico, border of forest 
road; from roadside, on top of woody shrubs 
without leaves; and from beach grape in 
Jamaica. 

Distribution. From Yucatan Peninsula, 
Mexico, to Honduras, Jamaica (Map 2B). 



Figures 55-61 . Metazygia ctiicanna n. sp. 55-59, female. 55-57, epigynum. 55, ventral. 56, posterior. 57, lateral. 58, dorsal. 
59, atxjomen, ventral. 60, 61 , left male palpus. 60, mesal. 61 , apical. 

Figures 62-65. W. tapa n. sp., female. 62-64, epigynum. 62, ventral. 63, posterior. 64, lateral. 65, dorsal. 

Figures 66-68. M. pastaza n. sp., male. 66, 67, male palpus. 66, mesal. 67, apical. 68, dorsal. 



Figures 69-73. M. incerta (O. P.-Cambridge). 69-72, female. 69-71 , epigynum. 69, ventral. 70, posterior. 71 , lateral. 72, dorsal. 
73, male palpus. 



METAZYCiA'Levi 91 




Figures 74-80. M. pallidula (Keyserling). 74-78, female. 74-76, epigynum. 74, ventral. 75, posterior. 76, lateral. 77, dorsal. 78, 
abdomen, ventral. 79, male palpus. 80, embolus, with and without cap. 

Abbreviations. C, conductor; L, embolus lamella; M, median apophysis. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



92 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Specimens Examined. MEXICO Campeche: Be- 
can, 18°33'N, 89°30'W, 31 July 1991, 16 (W. Piel, G. 
S. Bodner, MCZ); Ceiba Playa [?], 2 Aug. 1949, 12 
(C. J. Goodnight, AMNH). Yucatan: Chichen Itza, 
Nov. 1945, 19 (H. Wagner, AMNH). Quintana Roo: 
Chetumal, 28 June 1975, 15 (W. C. Sedgwick, MCZ); 
Chancanah Cozumel, 8 Aug. 1949, 19 (C. J. Good- 
night, AMNH); Reserva de Sian Ka'an, km 5, 4 June 
1991, many 9, 1<? (G. Alayon, L. F. Armas (lESC); 31 
km NE Fehpe Carillo Puerto, Highway 307, 17 July 
1983, 19 (W. Maddison, R. S. Anderson, MCZ); X-Can, 
6-7 June 1959, 19 (C, P. Vaurie, AMNH). BELIZE 
Stann Distr.: W Possum Point Biol. Sta., 16°49'N, 
88°20'\V, 2 July 1991, 19 (W. H. Piel, G. S. Bodner, 
MCZ). HONDURAS Lancetilla nr. Tela, July 1929, 
19; Tela beach 26 July 1929, U (A. M. Chickering, 
MCZ). JAMAICA 4.8 km E May Pen, 22 Nov. 1957, 
19 (A. M. Chickering, MCZ); Kingston, 6 Dec. 1954, 
15 (A. M. Nadler, AMNH); Kinloss, 23 Mar. 1955, 19 
(A. M. Nadler, AMNH); 22 km E Kingston, 11 July 
1960, 19 (C, P. Vaurie, AMNH); Long Mtn., 22 Oct. 
1957, 16, 26 Oct., 15 (A. M. Chickering, MCZ), 11 
July 1960, 19 (C, P. Vaurie, AMNH); Negril, 23-31 
Mar, 1981, 109, 45 (H., L. Levi, MCZ). 

Metazygia tapa new species 
Figures 62-65; Map 2B 

Holotype. Female holotype and two female para- 
types from Zona Reservada Tambopata, 290 m, 
12''50'S, 69°17"W, hotel at night, Depto. Madre de 
Dios, Peru, 8 June 1988 (D. Silva D.), in MUSM, 
one paratype in MCZ. The specific name is an 
arbitrary combination of letters. 

Description. Female holotype. Cara- 
pace orange. Chelicerae orange, distally 
brown. Labium, endites, sternum, legs or- 
ange. Abdomen whitish with white pig- 
ment spots underneath exoskeleton (Fig. 
65). Posterior median eyes 0.7 diameter of 
anterior medians, laterals 0.6 diameter. 
Anterior median eyes 0.5 diameter apart, 
0.7 diameter from laterals. Posterior me- 
dian eyes 0.3 diameter apart, 1 .8 diameters 
from laterals. Height of clypeus equals 0.5 
diameter of anterior median eye. Total 
length 5.4 mm. Carapace 2.3 mm long, 1.9 
wide, 1.0 behind lateral eyes. First femur 
2.3 mm, patella and tibia 2.5, metatarsus 
1.6, tarsus 0.8. Second patella and tibia 2.1 
mm, third 1.5, fourth 2.0. 

Variation. Total length of females 4.9 
to 6.0 mm. Illustrations were made from 
the holotype. 

Diagnosis. The abdomen is oval and 



slightly pointed anteriorly, widest in mid- 
dle (Fig. 65). Metazygia tapa is separated 
from M. ipago and M. patiama by the 
dark patches in ventral view of the epi- 
gynum (Fig. 62) and the narrow posterior 
median plate (Fig. 63). 

Natural History. Specimens were col- 
lected by fogging at night in Depto. Lo- 
reto, Peru. 

Distribution. Amazon drainage, Peru 
(Map 2B). 

Paratype. PERU Madre de Dios: Zona 
Reservada Tambopata, 290 m, 12°50'S, 
69°17'W, 3 June 1988, 19 (J. Coddington, 
USNM). 

Specimens Examined. PERU Loreto: Rio Samiria, 
12-28 May 1990, 29 (T. Erwin, D. Silva D,, MUSM). 
Madre de Dios: 15 km E Puerto Maldonado, 200 m, 
12°33'S, 69°03'W, 21 Feb.-8 Mar. 1989, 59 (D. Silva 
D., MUSM). 

Metazygia pastaza new species 
Figures 66-68; Map 2B 

Holotype. Male holotype from Pastaza, Depto. Lo- 
reto, Prov. Alto Amazonas, Peru, swamp plants, 
Aug. 1973 (J. C. Olin), in MCZ. The specific name 
is a noun in apposition after the type locality. 

Description. Male holotype. Carapace 
orange, cephalic region darker orange. 
Chelicerae orange-brown. Labium, en- 
dites dark orange. Sternum orange. Coxae, 
legs orange. Dorsum of abdomen with fo- 
lium as in related species (Fig. 68); venter 
with scattered white pigment spots. Pos- 
terior median eyes 0.7 diameter of anterior 
medians, laterals 0.7 diameter. Anterior 
median eyes 0.6 diameter apart, 1.8 di- 
ameters from laterals. Posterior median 
eyes 0.2 diameter apart. Height of clypeus 
equals 0.4 diameter of anterior median eye. 
Second tibia as thick as first. Total length 
6.7 mm. Carapace 3.5 mm long, 2.7 wide, 
1.6 behind lateral eyes. First femur 4.6 
mm, patella and tibia 5.2, metatarsus 4.2, 
tarsus 1.4. Second patella and tibia 4.5 mm, 
third 2.1, fourth 3.1. 

Note. This might be the male of M. 
tapa. 

Diagnosis. The palpus of this species 
(Fig. 67) lacks the spine on the tegulum of 



METAZYGIA'Levi 



93 



M. dubia (at 3 hr in Fig. 21) and has a 
smaller comb (at 1 hr in Fig. 66) than M. 
pimentel (Fig. 27). Most palpal sclerites 
differ slightly from those two similar spe- 
cies. 

Metazygia incerta (O. P.-Cambridge) 
Figures 69-73; Map 2J 

Epeira incerta O. P.-Cambridge, 1889: 23, pi. 4, fig. 
15, 9. Female syntypes from Costa Rica, in BMNH, 
examined. Keyserling, 1892: 163, pi. 8, fig. 120, 9. 

Epeira fecunda O. P.-Cambridge, 1889: 26, pi. 6, 
figs. 9, 10, 9, (5. Numerous syntypes from Guatemala 
in BMNH, examined. Keyserling, 1892: 164, pi. 8, 
fig. 121, 9, S. First synonymized by F. P.-Cam- 
bridge, 1904. 

Epeira maculata: — Keyserling, 1892: 242. Six female 
and one male paralectotype from Baltimore. The 
female lectotype is a Mangora (see Levi, 1975). 

Aranea incerta: — F. P.-Cambridge, 1904: 512, pi. 49, 
figs. 7, 8, 9, S. Roewer, 1942; 845. 

Araneus incertus: — Bonnet, 1955: 521. Kraus, 1955: 
22, figs. 62-65, 9, <?. 

Metazygia incerta: — Levi, 1991a: 179. 

Synonymy . There are male specimens 
in the vial with the female syntypes of 
Epeira incerta, which may have been add- 
ed later. Keyserling reports Epeira incerta 
as coming from Guatemala, but both Key- 
serling and O. P.-Cambridge examined the 
same females. Paralectotypes of Epeira 
maculata (Levi, 1975) are this species; they 
are marked as coming from Baltimore, but 
this is an erroneous G. Marx locality. 

Description. Female from San Jose, Cos- 
ta Rica. Carapace orange-brown. Chelic- 
erae, labium, endites orange. Sternum, 
coxae orange, legs brown. Dorsum of ab- 
domen black and white (Fig. 72); venter 
dusky, some white pigment spots in center. 
Eyes subequal. Anterior median eyes 0.8 
diameter apart, 1 diameter from laterals. 
Posterior median eyes 0.3 diameter apart. 
Height of clypeus equals 0.4 diameter of 
anterior median eye. Total length 6.5 mm. 
Carapace 2.7 mm long, 2.0 wide, 1.1 be- 
hind lateral eyes. First femur 2.9 mm, pa- 
tella and tibia 3.6, metatarsus 2.6, tarsus 
0.9. Second patella and tibia 2.9 mm, third 
1.7, fourth 2.4. 

Male from Monteverde, Costa Rica. 
Color as in female, but abdomen much 



darker. Posterior median eyes same di- 
ameter as anterior medians, laterals 0.9 di- 
ameter. Anterior median eyes their di- 
ameter apart, their diameter from laterals. 
Posterior median eyes 0.3 diameter apart, 
2 diameters from laterals. Height of clyp- 
eus equals 0.3 diameter of anterior median 
eye. Endite with small, sharp tooth, palpal 
femur with facing tubercle. First coxa with 
small hook. Second tibia thicker than first, 
without macrosetae. Total length 4.6 mm. 
Carapace 2.8 mm long, 2.2 wide, 1.2 be- 
hind lateral eyes. First femur 4.4 mm, pa- 
tella and tibia 5.7, metatarsus 4.5, tarsus 
1.2. Second patella and tibia 4.5 mm, third 
2.2, fourth 4.0. 

Note. Males and females were collected 
together. 

Variation. Total length of females 4.5 
to 9.0 mm, males 3.7 to 6.3. Illustrations 
were made from a female from San Jose, 
Costa Rica, and male from Puntarenas 
Prov., Costa Rica. 

Diagnosis. The female differs from oth- 
er Metazygia species by having the epi- 
gynum in ventral view an entire, rectan- 
gular plate, not showing openings (Fig. 69): 
the openings are anterior of the plate and 
not visible in ventral view. The male dif- 
fers by having two teeth on the embolus 
lamella (L in Fig. 73), below the lamella's 
sclerotized tip. 

Natural History. Specimens were found 
in Trypargilum nitidum and T. tenoctitla 
wasp nests at Cafias, Costa Rica. Most ob- 
servations come from R. Buskirk (personal 
communication, 28 Feb. 1972). The ob- 
servations were made in Monteverde, 1,380 
m, Costa Rica. The orb is made in tree 
branches about 1 m above ground. Spiders 
are active in the hub at night from dusk 
to about 30 minutes to one hour after sun- 
rise; during the day they are in folded-leaf 
retreats near or attached to the orb. The 
orbs are rebuilt every 2 to 4 days, de- 
pending on the damage to the web. In the 
Monteverde area, M. incerta is most abun- 
dant in moist habitats such as a river valley, 
and in places they are only a few deci- 
meters above the water. Individual orbs 



94 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



may be quite close to one another, nearly 
adjacent, but not continuous. Other col- 
lecting localities were rain forest, humid 
forest, cloud forest, a grapefruit orchard 
near a river, a porch at night, and mixed 
vegetation. 

Distribution. Common from Belize to 
Panama (Map 2J). 

Specimens Examined. BELIZE San Ignacio 
[IT'lO'N, 89°04'W] (MCZ). GUATEMALA Peten: E 
end Lago Peten Itza (AMNH). Alta Vera Paz: Coban 
(AMNH), HONDURAS Copan (AMNH); Lancetilla 
nr. Tela (MCZ); Siguatepeque, 1,100-1,200 m 
(AMNH). EL SALVADOR (Desague, Laguna Guija, 
460 m, Kraus, 1955). NICARAGUA Granada (MCZ); 
Managua (JMM); Musawas, Rio Waspuc (AMNH); 5 
km N Matagalpa (JMM); Islas de Solentiname (JMM); 
5 km E Jinotepe (JMM); 122 km S Managua, W shore 
Lago Nicaragua, 32 m (USNM). COSTA RICA He- 
redia: Heredia (AMNH); San Jose de la Montana (DU). 
Alajuela: Grecia (AMNH); Palmares (AMNH); road 
to Volcan Poas, 1,500 m (MCZ). Guanacaste: 4 km 
NW Canas, La Pacifica (MCZ); Penas Blancas (S. 
Riechert, AMNH); La Pacifica, 160 km SW, Finca 
Palo Verde (MCZ); Tilaran (MNRJ). Limon: Pens- 
hurst 10 km N Cahuita (DU). Cartago: Cartago, 1,400 
m (AMNH); Moravia (CAS); Turrialba (CAS, CUC); 
Paraiso (AMNH). San Jose: Braulio Carillo Natl. Park 
(DU); Escazii (MCZ); San Jose, 1,200 m (AMNH, DU, 
MCZ); Ciudad Universitaria (USNM); La Verbena 
(MCZ); San Pedro de Montes (USNM); Zapote (FSCA). 
Puntarenas: Monteverde, 1,380 m (AMNH, DU, 
MCZ). PANAMA Bocas del Toro: Changuinola 
(AMNH, CUC). 

Metazygia pallidula (Keyserling) 
Figures 74-80; Map 2E 

Epeira pallidula Keyserling, 1864: 124, pi. 4, figs. 14, 
15, 9. Female holotype from St. Fe de Bogota, New 
Granada [Bogota, Colombia], in BMNH, examined. 
Keyserling, 1892: 158, pi. 8, fig. 116, 2. 

Epeira simplicissima Keyserling, 1883: 203, pi. 15, 
fig. 8, 2. Female holotype from Tumbez [Tumbes, 
Depto. Tumbes], Peru, in PAN, examined. Key- 
serling, 1892: 169, pi. 8, fig. 125, 2. NEW SYN- 
ONYMY 

Singa mollybyrnae McCook, 1894: 229, pi. 19, fig. 1, 
2. One female from District of Columbia, United 
States, in ANSP, examined. Roewer, 1942: 878. Syn- 
onymized by Levi, 1972: 231. 

Aranea pallidula: — F. P.-Cambridge, 1904: 514, pi. 
49, fig. 13, 2. Roewer, 1942: 849. 

Aranea simplicissima: — Roewer, 1942: 852. 

Araneus mollybyrnae: — Bonnet, 1955: 546. 

Araneus pallidulus: — Bonnet, 1955: 562. 

Araneus simplicissimus: — Bonnet, 1955: 600. 

Araneus palloides Roewer, 1955: 1715. New name 



for Epeira pallidula Keyserling, thought to be pre- 
occupied by Araneus pallidula Clerck, 1758 {=Clu- 
biona pallidula). 
Metazygia pallidula: — Levi, 1991: 179. 

Synonymy. The holotypes of Epeira 
pallidula and Singa mollybyrnae have the 
same large lateral openings of the epigyn- 
um. The holotype of S. mollybyrnae comes 
from Washington, D.C., and another spec- 
imen from Biscay ne Bay, Florida. Each is 
an erroneous G. Marx locality. d 

Description. Female from Chiriqui ~ 
Prov., Panama. Carapace orange, cephalic 
area dusky. Chelicerae, labium, endites, 
sternum orange. Coxae, legs orange, dis- 
tally darker. Dorsum of abdomen whitish 
with anterior pair of black marks and in- 
distinct dusky folium (Fig. 77); venter a 
white transverse patch posterior to the epi- 
gynum (Fig. 78). Posterior median eyes 0.8 
diameter of anterior medians, laterals 0.8 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 1 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
2.8 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Total length 5.6 mm. Cara- 
pace 2.3 mm long, 1.9 wide, 1.1 behind 
lateral eyes. First femur 2.7 mm, patella 
and tibia 3.1, metatarsus 2.3, tarsus 0.9. 
Second patella and tibia 2.7 mm, third 1.5, 
fourth 2.2. 

Male from Chiriqui Prov., Panama. Col- 
or as in female, but legs with indistinct 
dark rings and no white on venter. Pos- 
terior median eyes 0.8 diameter of anterior 
medians, laterals 0.8 diameter. Anterior 
median eyes 0.8 diameter apart, 0.8 di- 
ameter from laterals. Posterior median eyes 
0.2 diameter apart, 1.6 diameters from lat- 
erals. Height of clypeus equals 0.4 diam- 
eter of anterior median eye. Total length 
4.5 mm. Carapace 2.5 mm long, 1.9 wide, 
0.9 behind lateral. First femur 3.5 mm, 
patella and tibia 4.5, metatarsus 3.7, tarsus 
1.2. Second patella and tibia 3.6 mm, third 
1.8, fourth 2.5. 

Note. Males and females are commonly 
collected together. 

Variation. Total length of females 3.8 



METAZYGIA'Levi 



95 



to 6.5 mm, males 2.5 to 4.2. Illustrations 
were made from specimens from Chiriqui 
Prov., Panama. 

Diagnosis. The large pair of ear-like de- 
pressions on the ventral view of the epi- 
gynum (Fig. 74) separates females from 
similar species. Males can be separated by 
the shape of the embolus with two rounded 
lobes "below" (Figs. 79, 80) and by the 
two pointed lobes of the embolus lamella 
(Fig. 79). 

Natural History. Specimens were col- 
lected from a wet area near Acapulco, 
Mexico; from wet forest in Limon Prov., 
Costa Rica; on a bridge, banks of a river 
bed, and a rock wall in a garden in Chi- 
riqui Prov., Panama; in a banana grove in 
Sullana, Peru; in light under the eaves of 
a house in northern Colombia; and in low 
shrubs and large herbs at night in Tur- 
rialba, Costa Rica. Many collections were 
made at night. 

Distribution. From central Mexico to 
French Guiana and Ecuadorian and Pe- 
ruvian coast (Map 2E). 

Specimens Examined. MEXICO Veracruz: Coat- 

zacoalcos (AMNH); Tlacotalpan (AMNH). Guerrero: 
13 km W Acapulco (AMNH, MCZ); 2.4 km W Ac- 
apulco (AMNH); Acapulco (MCZ). Tabasco: Coma- 
calco (AMNH). Chiapas: La Zacualpa (AMNH). EL 
SALVADOR San Salvador (AMNH). HONDURAS 
Comavagua (FSCA). NICARAGUA Managua 
(AMNH, cue). COSTA RICA Heredia: W Alajuela 
(Riechert Coll.); La Selva (AMNH, MCZ). Alajuela: 
Fortuna [Rio Fortuna], nr. Esparta (MCZ). Limon: 
nr. Cahuita (MCZ); 5 km E Guapiles (DU); Tortu- 
guero Natl. Park (DU), Cartago: Turrialba (CAS, 
MCZ). San Jose: San Jose (MCZ); San Isidro (MCZ). 
Puntarenas: Corcovado Natl. Park, Sirena (MCZ); 
Jaco (MCZ); Parrita (MCZ); San Isidro del General 
(MCZ); Tarcoles (MCZ); 10 km N Mai Pais (MCZ), 
PANAMA Boga del Tow: Changuinola (MCZ), Chi- 
riqui: Bambito nr. Cerro Punta, 1,400 m (CAS, MCZ); 
Cerro Punta (AMNH); Boquete (AMNH, MCZ); 9 
km N David (USNM); David (MCZ, MIUP); La For- 
tuna (MCZ, MIUP); Volcan (AMNH, MCZ); Volcan 
Baru (FSCA), Colon: Gamboa (MCZ); France Field 
(MCZ); Frijoles (MCZ). Code: El Cope (MIUP); El 
Valle (AMNH, MCZ). Panama: Balboa (MCZ); Barro 
Colorado Island (AMNH, MCZ); road to Chiva (MCZ); 
Cerro Azul (MIUP); Experimental Gardens (MCZ); 
Forest Reserve (MCZ); Madden Dam (MCZ); Pedro 
Miguel (MCZ); Pipeline Road (MCZ); Playa Corona 
nr. San Carlos (MCZ); Summit (MCZ). VENEZUELA 
Delta Amacuro: Rio Orinoco delta (MCZ). GUYANA 



Bartica: Kartabo (CUC). FRENCH GUIANA St, 
Laurent du Maroni (PAN), COLOMBIA La Guajira: 
Rio Guatapuri, 1,100 m, Magdalena: above Minca 
Valley, 880 m (IBNP); East Cerro Dunarua, 1,300 m 
(IBNP); Rio Domachui, 1,700 m (IBNP); Rio Cordua, 
800 m (IBNP); Rio Frio, 530 m (IBNP); stream betw, 
Cerros Chivolo and Chumchuruba, 1,100 m (IBNP); 
Rio Gaira (SMF); San Pablo (IBNP); San Sebastian de 
Rebango, 3,300 m. Sierra Nevada de Santa Marta 
(AMNH); Serra Nueva Granada, 1,310 m (IBNP), 
Cesar: Escorpa Mission, Sierra de Parija, 1,300-1,400 
m (AMNH). Cundinamarca: 22 km SE Caqueza (CAS) 
Cordoba: Ayapel nr. Cienaga (MCZ). Meta: Villa- 
vicencio (AMNH); 5 km W Villavicencio (CAS). An- 
tioquia: Mutata (MCZ). Valle: Cali (MCZ); Lago Co- 
lima nr. Darien (MCZ). Narino: Barbacoas (MCZ). 
ECUADOR Pichincha: Santo Domingo de las Colo- 
rades (FSCA). Manabi: El Carmen (AMNH); road 
betw. Crucita and Charapoto (MCZ). Guayas: 4.8 km 
N Manglar Alto (CAS); Guayaquil (CAS); Milagro 
(CAS); Puna Island (CAS). Azuaij: Jubones (CAS), El 
Oro: Buena Vista, 20 km Machala (CAS), PERU Lo- 
reto: Barranca (CAS), Piura: Negritos (CAS); Mal- 
lares, Rio Chira (CAS); nr. Miramar (CAS); Porta 
Chuala [? Portachuelo] (CAS); 6 km W Sullana (CAS); 
Sullana (CAS); Las Lomas (CAS), Lambayeque: 10 
km S Chiclayo (CAS), Libertad: Pacasmayo (PAN). 
Lima: Lima (CAS), Ayacucho: Monterrico (PAN). 

Metazygia enabia new species 
Figures 81-86; Map 2E 

Holotype. Female holotype from Cerro de la Neblina, 
base camp, 140 m, 0°5'0'N, 66°10'W, Territ. Federal 
Amazonas, Venezuela, 21-28 Feb. 1985, male para- 
type, 9 Feb. 1985, both from low foliage (W. E. 
Steiner), in USNM. The specific name is an arbi- 
trary combination of letters. 

Description. Female holotype. Cara- 
pace light orange, eye region black. Che- 
licerae distally much darker orange. La- 
bium, endites dusky orange. Sternum, legs 
orange. Dorsum of abdomen with dense 
white pigment spots and two dark gray 
longitudinal bands; bands are most distinct 
posteriorly and become indistinct anteri- 
orly (Fig. 84). Venter light dusky. Posterior 
median eyes 0.8 diameter of anterior me- 
dians, laterals 0.7 diameter. Anterior me- 
dian eyes 0.3 diameter apart, 0.3 diameter 
from laterals. Posterior median eyes 0.2 
diameter apart. Height of clypeus equals 
0.2 diameter of anterior median eye. Total 
length 3.7 mm. Carapace 1.8 mm long, 1.4 
wide, 0.8 behind lateral eyes. First femur 
1.6 mm, patella and tibia 1.8, metatarsus 



96 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



1.1, tarsus 0.6. Second patella and tibia 1.6 
mm, third 0.9, fourth 1.4. 

Male paratype. Color as in female, but 
orange parts shghtly dusky and dark bands 
of dorsum of abdomen more distinct. Pos- 
terior median eyes 0.7 diameter of anterior 
medians, laterals 0.7 diameter. Anterior 
median eyes 0.3 diameter apart, 0.2 di- 
ameter from laterals. Posterior median eyes 
0.2 diameter apart, 1 diameter from lat- 
erals. Height of clypeus equals 0.3 diam- 
eter of anterior median eye. Total length 
2.8 mm. Carapace 1.53 mm long, 1.17 
wide, 0.57 behind lateral eyes. First femur 
1.43 mm, patella and tibia 1.67, metatarsus 
1.14, tarsus 0.59. Second patella and tibia 
1.33 mm, third 0.88, fourth 1.14. 

Note. Males and females were matched 
because both were collected at the same 
locality, in the same habitat, and have sim- 
ilar markings: longitudinal dark bands, 
darkest posteriorly (Fig. 84). 

Diagnosis. The female epigynum dif- 
fers from that of others by the ventrally 
projecting median plate (at 6 hr in Fig. 
81, at center of Fig. 82) and the sclerotized 
edge of the lateral plates, as seen on each 
side in ventral view (Fig. 81). The male 
differs by the long, exposed embolus, which 
is not covered by the embolus lamella (Fig. 
85). 

Metazygia redfordi new species 
Figures 87-90; Map 2C 

Holotype. Female holotype from Parque Nacional 
das Emas, nr. Mineiros, Est. Goias, Brazil, associ- 
ated with a termite mound, Sept. -Oct. 1981 (K. H. 
Bedford) in MZSP ex MCZ. The species is named 
after the collector. 

Description. Female holotype. Cara- 
pace orange. Chelicerae, labium, endites 
orange. Sternum orange. Coxae orange. 



legs black. Dorsum of abdomen whitish 
with five pairs of black patches (Fig. 90); 
venter black except for book-lungs and area 
in-between, sides black. Posterior median 
eyes 0.6 diameter of anterior medians, an- 
terior laterals 0.6 diameter, posterior lat- 
erals 0.5. Anterior median eyes 0.4 di- 
ameter apart, 1.5 diameters from laterals. 
Posterior median eyes 0.3 diameter apart. 
Height of clypeus equals 0.6 diameter of 
anterior median eye. Total length 7.5 mm. 
Carapace 3.7 mm long, 2.7 wide, 1.5 be- 
hind lateral eyes. First femur 2.8 mm, pa- 
tella and tibia 3.5, metatarsus 2.2, tarsus 
1.1. Second patella and tibia 3.1 mm, third 
2.0, fourth 2.9. 

Diagnosis. The contrasting markings on 
the abdomen (Fig. 90) would suggest that 
this species belongs in Alpaida; the epi- 
gynum (Figs. 87-89), however, is that of 
a Metazygia. Unlike that of other species, 
the scape is wide anteriorly (Fig. 87). The 
shape of the median plate in ventral and 
posterior view (Figs. 87, 88) is unlike that 
of other species. 

Metazygia isabetae new species 
Figures 91-93; Map 2C 

Holotype. Male holotype from Santa Isabel do Morro, 
Ilha do Bananal, Est. Tocantins, Brazil, June 1961 
(M. Alvarenga), in AMNH. The species is named 
after the local saint. 

Description. Male holotype. Cephalo- 
thorax orange, the third and fourth coxae 
and legs lightest. Dorsum of abdomen light, 
with three faint, dusky, longitudinal bands 
(Fig. 93); venter light. Posterior median 
eyes 0.6 diameter of anterior medians, lat- 
erals 0.6 diameter. Anterior median eyes 
0.5 diameter apart, 1.3 diameters from lat- 
erals. Posterior median eyes 0.2 diameter 
apart, 2.8 diameters from laterals. Height 



Figures 81-86. Metazygia enabia n. sp. 81-84, female. 81-83, epigynum. 81 , ventral. 82, posterior. 83, lateral. 84, dorsal. 85- 
86, male left palpus. 85, mesal. 86, apical. 

Figures 87-90. M. redfordi n. sp., female. 87-89, epigynum. 87, ventral. 88, posterior. 89, lateral. 90, dorsal. 

Figures 91-93. M. isat)elae n. sp., male. 91 , 92, palpus. 91 , mesal. 92, ventral. 93, dorsal. 



METAZYGiA'Levi 97 




Figures 94-99. M. rogenrtofer/ (Keyserling). 94-98, female. 94-96, epigynum. 94, ventral. 95, posterior. 96, lateral. 97, dorsal. 
98, abdomen, ventral. 99, male palpus. 

Figures 100-103. M. baruerin. sp., female. 100-102, epigynum. 100, ventral. 101, posterior. 102, lateral. 103, dorsal. 

Figures 104-106. M. jamah u. sp., male. 104, 105, palpus. 104, ventral. 105, apical. 106, dorsal. 

Abbreviations. A, terminal apopfiysis; C, conductor; M, median apopfiysis. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



98 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



of clypeus equals 0.6 diameter of anterior 
median eye. Total length 4.8 mm. Cara- 
pace 2.8 mm long, 2.1 wide, 1.1 wide be- 
hind lateral eyes. First femur 3.1 mm, pa- 
tella and tibia 3.9, metatarsus 3.4, tarsus 
1.1. Second patella and tibia 3.2 mm, third 
1.8, fourth 2.4. 

Diagnosis. This male has an enlarged 
bubble-like subterminal apophysis (Figs. 
91, 92) unlike that of any other species. 

Metazygia rogenhoferi (Keyserling) 
Figures 94-99; Map 2C 

Zilla rogenhoferi Keyserling, 1878: 578, pi. 14, fig. 
6, 9. Keyserling, 1892: 296, pi. 15, fig. 219. Female 
holotype from Brazil in NMW, examined. 

Zygiella rogenhoferi: — Roewer, 1942: 887. Bonnet, 
1959: 5005. 

Metazygia rogenhoferi: — Levi, 1974: 271. 

Description. Female from Guaiba, Rio 
Grande do Sul. Carapace light orange. 
Chelicerae, labium, endites light orange. 
Sternum light orange. Legs light orange. 
Dorsum of abdomen white, with indistinct 
anterior dusky patches (Fig. 97). Venter 
with white pigment patches behind epi- 
gynum, on the sides and behind spinnerets 
(Fig. 98). Posterior median eyes 0.9 di- 
ameter of anterior medians, laterals 0.9 
diameter. Anterior median eyes 1 diam- 
eter apart, 0.8 diameter from laterals. Pos- 
terior median eyes 0.3 diameter apart, 1.2 
diameters from laterals. Height of clypeus 
equals 0.7 diameter of anterior median eye. 
Total length 3.3 mm. Carapace 1.8 mm 
long, 1.4 wide, 0.7 behind lateral eyes. First 
femur 1.8 mm, patella and tibia 2.1, meta- 
tarsus 1.6, tarsus 0.7. Second patella and 
tibia 1.8 mm, third 1.1, fourth 1.5. 

Male from Guaiba, Rio Grande do Sul. 
Color as in female, but with pairs of dusky 
brackets on dorsum of abdomen and ab- 
domen with less white pigment. Posterior 
median eyes 0.9 diameter of anterior me- 
dians, laterals 0.9 diameter. Anterior me- 
dian eyes their diameter apart, 0.5 diam- 
eter from laterals. Posterior median eyes 
0.3 diameter apart, 1.4 diameters from lat- 
erals. Height of clypeus equals 0.6 diam- 
eter of anterior median eye. Second tibia 



thinner than first, with few macrosetae. 
Total length 4.1 mm. Carapace 1.9 mm 
long, 1.6 wide, 0.7 behind lateral eyes. First 
femur 2.2 mm, patella and tibia 3.1, meta- 
tarsus 2.4, tarsus 0.9. Second patella and 
tibia 2.1 mm, third 1.2, fourth 1.6. 

Note. Males and females were collected 
together. 

Variation. Total length of females 3.4 
to 5.6 mm, males 3.4 to 4.2. The lateral 
bracts framing the epigynum on each side 
(Fig. 94) are transparent and variable in 
shape. Illustrations were made from spec- 
imens from Guaiba, Rio Grande do Sul, 
Brazil. 

Diagnosis. Females differ from other fe- 
male Metazygia by having transparent 
bracts on each side of the epigynum, hav- 
ing a short scape (Fig. 94), and by the flask- 
shaped median plate as seen in posterior 
view (Fig. 95). Males are separated from 
others by the long sword-shaped embolus 
(Fig. 99). 

Natural History. Specimens were col- 
lected with bromeliads in Sao Paulo state. 

Distribution. From Bahia State to Rio 
Grande do Sul, Brazil (Map 2C). 

Specimens Examined. BRAZIL Bahia: Uruq'uca, 
Fazenda Almada (MCN); Fazenda Jacaranda, Ita- 
maraju (MCN). Rio de Janeiro: Rio de Janeiro, Pin- 
heiro (MNRJ). Sao Paulo: Alto da Serra (MZSP); Bar- 
ueri (MZSP); Itanhaem, Baixo Rio Branco (MZSP); 
Campo Limpo, EFSD (AMNH); Juquia, Fazenda P090 
Grande (MZSP); Jurubatuba (MZSP); Osasco (MZSP); 
Ribeirao Pires (AMNH); Ilha de S5o Sebastiao (MZSP); 
Sao Bernando (MZSP). Rio Grande do Sul: Canela 
(MCN); Rio Grande, Esta9ao Ecologica do Taim 
(MCN); Santa Vitoria do Palmar, Esta^ao. Ecologica 
do Taim (MCN); Guaiba, Granja Carola (MCN); Praia 
do Curumim (MCN); Torres (MCN); Triunfo (MCN); 
Viamao, Aguas Belas (MCN); Viamao, Lagoa do Cas- 
amento (MCN). 

Metazygia barueri new species 
Figures 100-103; Map 2F 

Holotype. Female hoIot> pe and male paratype from 
Barueri, Est. Sao Paulo, Brazil, 8 Sept. 1965 (K. 
Lenko), in MZSP no. 4026. The specific name is a 
noun in apposition after the t\pe locality. 

Description. Female holotype. Cara- 
pace light orange, cephalic region orange. 
Chelicerae brown. Labium, endites or- 



Metazygia 'Levi 



99 



ange. Sternum light orange. Coxae light 
orange, legs dusky orange. Dorsum of ab- 
domen with folium outlined by dusky 
brackets and with a median longitudinal 
dusky line (Fig. 103). Venter light, without 
marks. Posterior median eyes 0.6 diameter 
of anterior medians, laterals 0.7 diameter. 
Anterior median eyes 0.8 diameter apart, 
1 diameter from laterals. Posterior median 
eyes 0.4 diameter apart. Height of clypeus 
equals 0.5 diameter of anterior median eye. 
Total length 5.5 mm. Carapace 2.4 mm 
long, 1.9 wide, 1.1 behind lateral eyes. First 
femur 2.1 mm, patella and tibia 2.3, meta- 
tarsus 1.7, tarsus 0.9. Second patella and 
tibia 2.1 mm, third 1.3, fourth 1.9. 

Diagnosis. Unlike the epigynum of oth- 
er species, the area behind and on the side 
of the scape is wrinkled (Figs. 100-102). 

Metazygia jamari new species 
Figures 104-106; Map 2F 

Holotype. Male holotype from Jamari, Rondonia, 
Brazil, 23 Jan. 1989 (Equipe Operacjao Jamari), in 
MCN no. 18550. The specific name is a noun in 
apposition after the type locality. 

Description. Male holotype. Carapace 
orange. Chelicerae, labium, endites or- 
ange. Sternum orange. Legs orange. Dor- 
sum of abdomen white with a pair of lon- 
gitudinal dark bands (Fig. 106); venter 
gray. Carapace rebordered above first coxa. 
Posterior median eyes 0.8 diameter of an- 
terior medians, laterals 0.7 diameter. An- 
terior median eyes 0.8 diameter apart, 0.6 
diameter from laterals. Posterior median 
eyes 0.2 diameter apart, 1.4 diameters from 
laterals. Height of clypeus equals 0.6 di- 
ameter of anterior median eye. Second tib- 
ia thinner than first. Total length 2.5 mm. 
Carapace 1.5 mm long, 1.1 wide, 0.5 be- 
hind lateral eyes. First femur 1.7 mm, pa- 
tella and tibia 2.1, metatarsus 1.6, tarsus 
0.7. Second patella and tibia 1.6 mm, third 
0.8, fourth 1.2. 

Note. I was not successful matching this 
male to a female. 

Variation. Total length of males 2.4 to 
2.8. The Porto Velho male differed slightly 
in the shape of the embolus. It is not known 



if the horse shoe-shaped structure above 
the embolus (Figs. 104, 105) breaks off 
when mating. Illustrations were made from 
the holotype. 

Diagnosis. Metazygia jamari differs 
from other species by the U-shaped tip of 
the embolus and by the straight terminal 
apophysis (at 12 hr to 2 hr in Fig. 104, left 
in Fig. 105). 

Distribution. Amazon region (Map 2F). 

Specimens Examined. SURINAM Marowijne: 
Lawa River, Anapaike Village, 8-29 Nov. 1963, 16 
(B. Malkin, AMNH). BRAZIL Roraima: Ilha de Mar- 
aca, Rio Uraricoera, 29 Mar. 1987, IS (A. A. Lise, 
MCN 20060). Rondoina: Porto Velho, Rio Tapirape, 
Feb., Mar. 1963, 16 (P. Pinheiros, AMNH). 

Metazygia crew! (Banks) 
Plate 1; Figures 107-113; Map 21 

Singa crewi Banks, 1903: 342, pi. 15, fig. 8, 9. Female 
holotype from Haiti, lost (not in AMNH, ANSP, 
cue, MCZ, USNM). Roewer, 1942: 877. 

Larinia coamensis Petrunkevitch, 1930: 335, figs. 221- 
224, 9. Female holotype from Coamo Springs, Puer- 
to Rico, in PMY, examined. Roewer, 1942: 771. 
Bonnet, 1957: 2348. First synonymized by Bryant 
1945. 

Aranea crewi: — Bryant, 1945: 364, figs. 1-3, S. 

Araneus crewi: — Bonnet, 1955: 471. 

Metazygia crewi: — Harrod, Levi, and Leibensper- 
ger, 1991: 245. 

Description. Female from Puerto Plata, 
Dominican Republic. Carapace dusky or- 
ange, black between eyes, margin of tho- 
racic region darker. Chelicerae dusky or- 
ange. Labium, endites orange. Sternum 
light yellow-brown, black on each side, 
black fusing posteriorly to form a V shape. 
Legs light orange, distally with some dark 
rings. Dorsum of abdomen with dusky me- 
dian band, a white cardiac mark, a white 
band on each side and black on sides (Fig. 
110). Venter black with a square white 
patch posterior to genital groove, black 
surrounding spinnerets, sides light (Fig. 
111). Carapace narrow in front without 
thoracic depression. Posterior median eyes 
0.6 diameter of anterior medians, laterals 
0.7 diameter. Anterior median eyes 0.6 di- 
ameter apart, 0.7 from laterals. Posterior 
median eyes 0.4 diameter apart, 2 diam- 



100 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



eters from laterals. Height of clypeus equals 
0.4 diameter of anterior median eye. Total 
length 5.0 mm. Carapace 2.1 mm long, 1.6 
wide, 0.7 behind lateral eyes. First femur 
2.3 mm, patella and tibia 2.5, metatarsus 
1.9, tarsus 0.8. Second patella and tibia 1.9 
mm, third 1.2, fourth 1.8. 

Male specimen from Puerto Plata, Do- 
minican Republic. Color as in female. 
Thoracic depression a longitudinal line. 
Posterior median eyes 0.7 diameter of an- 
terior medians, anterior laterals 0.7, pos- 
terior 0.5. Anterior median eyes 1 diam- 
eter apart, slightly less from laterals. Pos- 
terior median eyes 0.4 diameter apart, 2 
diameters from laterals. Height of clypeus 
equals 0.8 diameter of anterior median eye. 
First coxa with hook, fourth with a small 
macroseta. Total length 3.7 mm. Carapace 
1.9 mm long, 1.5 wide, 0.6 behind lateral 
eyes. First femur 2.3 mm, patella and tibia 
2.9, metatarsus 2.2, tarsus 0.9. Second pa- 
tella and tibia 2.0 mm, third 1.1, fourth 
1.7. 

Note. Males and females were collected 
together. 

Variation. Total length of females 4.0 
to 5.5 mm, males 3.3 to 3.7. Some females 
have three teeth on the anterior margin of 
the chelicerae, two on the posterior; others 
have four and three. The epigynum's 
transparent scape can be short (Fig. 107) 
or long and overhanging the anterior bor- 
der of the epigynum; often it appears torn 
off with only a dark round scar. Illustra- 
tions were made from specimens collected 
in Puerto Plata, Dominican Republic; the 



web (Pi. 1) was photographed near Mar- 
icao, Puerto Rico. 

Diagnosis. The female is separated from 
other species by having an epigynum that, 
when viewed posteriorly, has two long slits, 
separated by a narrow median plate (Fig. 
108). The male is distinguished by having 
a palp with filamentous embolus and over- 
hanging terminal apophysis (A) (Figs. 112, 
113). 

Natural History. Specimens were col- 
lected sweeping in forest on St. Johns, Vir- 
gin Islands, and in a coffee plantation at 
Jayuya, Puerto Rico. Many specimens came 
from an abandoned, dry, sunny road on a 
south-facing slope near Maricao, Puerto 
Rico, at 800 m elevation. The spiders had 
retreats in the heads of grass above a small 
transparent web (Pi. 1). These spiders could 
not be dislodged with a sweep net but had 
to be collected individually. 

Distribution. Greater Antilles, Virgin 
Islands (Map 21). 

Specimens Examined. CUBA [no local], 19 (R. V. 
Chamberlin, AMNH). HAITI Cap Haitien, Mar. 1934, 
29 (Utowana Exped., MCZ); Dame Marie, 1941, 29, 
IS (A. Audant, MCZ); Port-au-Prince, July 1941, 139, 
2(5 (A. Audant, MCZ); 18-21 July 1955, 49, 2S, 8 imm. 
(A. F. Archer, AMNH); 21, 22 Mar. 1969, 29; 30 Aug- 
7 Sept. 1969, 29, U (L. Reynolds, MCZ), DOMINI- 
CAN REPUBLIC Balneario Saladilla, S Barahona, 8 
Aug. 1958, 19 (A. F. Archer, AMNH); 5 km NW Las 
Matas de Farfan, 25 Aug, 1970, 19 (B, Patterson, 
MCZ); Loma de Los Pinos, Colonia Ramfis, T. Valdez, 
700-900 m, 7 Aug, 1958, 29 (A, F, Archer, E, Boynie 
Moya, AMNH); Puerto Plata, July-Aug, 1941, 239, 
2(5 (D. Hurst, MCZ); La Gran Chorra, Altagracia, 11 
Apr, 1992, 39 (F, Del Monte, MNSD), PUERTO RICO 
Jayuya, 1,000 m, 20-26 Mar, 1986, 29, imm, (H., L. 



Figures 107-113. Metazygia crew/ (Banks), 107-111, female. 107-109, epigynum, 107, ventral, 108, posterior, 109. lateral, 
110, dorsal. Ill, atxjomen, ventral, 112, 113, left male palpus. 111, mesal, 112, ventral. 

Figures 114-118, M. vaurieorum n, sp,, female, 114-116, epigynum, 114, ventral, 115, posterior, 116, lateral, 117, dorsal, 118, 
abdomen, ventral. 

Figures 119, 120. M. carrizaln. sp., male. 119, palpus. 120, dorsal. 

Figures 121-124, M. taman n. sp., female. 121-123, epigynum. 121, ventral. 122, posterior. 123, lateral. 124, dorsal. 

Figures 125-128, M. paquisha female. 125-127, epigynum. 125, ventral. 126, posterior. 127, lateral, 128, dorsal. 

Figures 129-132. M. nobasn. sp., female. 129-131, epigynum, 129, ventral, 130, posterior, 131, lateral, 132, dorsal. 



METAZYCIA'Levi 101 




Figures 133-137. M. goeldiin. sp., female. 133-136, epigynum. 133, ventral. 134, posterior. 135, lateral. 136, paratype, ventral. 
137, dorsal. 

Abbreviations. A, terminal apophysis; C, conductor; M, median apophysis; R, radix. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



102 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Levi, MCZ); Reserva Forestal, Maricao, Monte de 
Estado, 800 m, 4-6 Apr. 1989, 132, 16 imm. (H., L. 
Levi, MCZ). U.S. VIRGIN ISLANDS St. Johns, forest 
above Cinnamon Bav, 17 Mar. 1970, 12, 13 (H., L., 
F. Levi, MCZ), 

Metazygia vaurieorum new species 
Figures 114-118; Map 21 

Holotype. Female holotype and one paratype from 
\ariedades, H-Ol'N, 90°30'W, Suchitepequez, 120- 
170 m, Guatemala 27-31 Aug. 1947, and two fe- 
male paratypes, 1-4 July 1947 (C. and P. Vaurie), 
in AMNH. The species is named after the collectors. 

Note. The location of the type locaUty 
is pubhshed in Vaurie and Vaurie (1949). 

Description. Female holotype. Cara- 
pace orange, cephalic region darker. Che- 
licerae dark orange-brown. Labium, en- 
dites, sternum orange. Coxae legs orange. 
Dorsum of abdomen with gray folium pat- 
tern on white (Fig. 117). Venter with trans- 
verse white patch behind epigynum and 
semicircular white line in front of spin- 
nerets (Fig. 118). Posterior median eyes 
0.7 diameter of anterior medians, laterals 
0.7 diameter. Anterior median eyes 0.7 di- 
ameter apart, 1.3 diameters from laterals. 
Posterior median eyes 0.2 diameter apart. 
Height of clypeus equals 0.5 diameter of 
anterior median eye. Total length 6.0 mm. 
Carapace 3.5 mm long, 2.7 wide, 1.5 be- 
hind lateral eyes. First femur 3.4 mm, pa- 
tella and tibia 4.1, metatarsus 3.2, tarsus 
IT. Second patella and tibia 3.6 mm, third 
2.0, fourth 2.7. 

Diagnosis. In ventral view the epigyn- 
um (Fig. 114) resembles that of M. crewi 
(Fig. 107), but the scape is a large knob 
rather than a flat plate (perhaps a piece 
has broken off). It differs in posterior view 
by having a wide median plate (Fig. 115). 

Metazygia carrizat new species 
Figures 119, 120; Map 21 

Holotype. Male holotype from Mataquescuintla, El 
Carrizal, Depto. Jalapa, Guatemala, beating foliage 
at river, 25 Apr. 1982 (S. Fend), in CAS. The spe- 
cific name is a noun in apposition after the type 
locality. 

Description. Male holotype. Carapace 
dusky orange to black. Chelicerae dusky 



orange. Labium black, endites orange. 
Sternum black. Coxae light orange, legs 
orange with dark rings. Dorsum of abdo- 
men with median longitudinal light band 
and gray band to the side, whitish color to 
the sides of gray (Fig. 120). Venter dark 
with indistinct white pigment spots in cen- 
ter. Posterior median eyes 0.7 diameter of 
anterior medians, laterals 0.7 diameter. 
Anterior median eyes 0.8 diameter apart, 
0.8 diameter from laterals. Posterior me- 
dian eyes 0.3 diameter apart, 2 diameters 
from laterals. Height of clypeus equals 0.3 
diameter of anterior median eye. Second 
tibia thicker than first, with long macro- 
setae on both first and second tibiae. Total 
length 4.2 mm. Carapace 2.1 mm long, 1.5 
wide, 0.8 behind lateral eyes. First femur 
2.3 mm, patella and tibia 3.1, metatarsus 
2.3, tarsus 0.9. Second patella and tibia 2.3 
mm, third 1.2, fourth 1.8. 

Note. This was collected near a M. vau- 
rieorum female. However, it is considered 
to represent another species, as it has a 
different coloration: black sternum in male, 
light in female; white above cardiac area 
in male (Fig. 120), dark in female (Fig. 
117). 

Diagnosis. This species differs from all 
others by the terminal apophysis hanging 
"down" with its axis at an acute angle to 
the axis of the cymbium and by the shape 
of the sclerotized embolus lamella (at cen- 
ter of Fig. 119) and by the U-shaped em- 
bolus with a pointed cap (Fig. 119). 

Metazygia taman new species 
Figures 121-124; Map 21 

Holotype. Female holotype from near Taman, ca. 16 
km SW of Tamazunchale on Highway 85, 21°irN, 
98°53'W, ca, 300 m, San Luis Potosi State, Mexico, 
11 June 1983 (W, Maddison, R. S. Anderson), in 
MCZ. The specific name is a noun in apposition 
after the t\pe locality , 

Description. Female holotype. Cara- 
pace orange, cephalic area dusky. Chelic- 
erae, labium, endites orange-brown. Ster- 
num dusky orange. Coxae, legs orange, dis- 
tal articles darker. Dorsum of abdomen 
with paired dark gray marks on orange- 



Metazycia 'Levi 



103 



gray (Fig. 124); venter uniformly dark 
gray. Posterior median eyes 0.8 diameter 
of anterior medians, laterals 0.7 diameter. 
Anterior median eyes 0.7 diameter apart, 
1.4 diameters from laterals. Posterior me- 
dian eyes 0.3 diameter apart. Posterior me- 
dian eyes oval, the long diameter from 
median posterior, to lateral and anterior. 
Height of clypeus equals 0.6 diameter of 
anterior median eye. Total length 7.7 mm. 
Carapace 3.9 mm long, 3.0 wide, 1.6 be- 
hind lateral eyes. First femur 3.3 mm, pa- 
tella and tibia 3.9, metatarsus 2.5, tarsus 
1.1. Second patella and tibia 3.4 mm, third 
2.1, fourth 3.1. 

Diagnosis. This species has a thinner 
scape (Figs. 121-123) than does M. pa- 
quisha (Figs. 125-127), but the scape is 
rounded (Fig. 123) and the posterior me- 
dian plate is wider (Fig. 122) than that of 
M. paquisha (Fig. 126). 

Metazygia paquisha new species 
Figures 125-128; Map 2H 

Holotype. Female holotype from Alto Rio Comaina, 
Puesto de Vigilancia 22, "Falso Paquisha, " Cor- 
dillera del Condor, border with Ecuador, 05°02'S, 
78°5rW, Depto. Amazonas, Peru, night collecting, 
23 Oct. 1987 (D. Silva D.), in MUSM. The specific 
name is a noun in apposition after the type locality. 

Description. Female holotype. Cara- 
pace orange, darkest anterior. Chelicerae 
orange. Labium, endites, sternum dusky 
orange. Legs dusky orange, darkest dis- 
tally. Abdomen gray without marks (Fig. 
128). Carapace with a median line of setae. 
Posterior median eyes 0.8 diameter of an- 
terior medians, laterals 0.7 diameter. An- 
terior median eyes 0.6 diameter apart, 1.1 
diameters from laterals. Posterior median 
eyes 0.3 diameter apart. Height of clypeus 
equals 0.6 diameter of anterior median eye. 
Total length 7.2 mm. Carapace 3.1 mm 
long, 2.3 wide, 1.3 behind lateral eyes. First 
femur 2.5 mm, patella and tibia 3.3, meta- 
tarsus 2.3, tarsus 0.9. Second patella and 
tibia 3.1 mm, third 2.1, fourth 2.5. 

Illustration. Figures 125-128 were made 
from the holotype. 

Diagnosis. The female differs from oth- 



ers by the wide massive scape (Figs. 125- 
127) and from M. taman (Figs. 121-124) 
by the more pointed scape and narrower 
posterior plate (Figs. 126, 127). 

Distribution. Amazon Region (Map 2H). 

Specimen Examined. VENEZUELA Amazonas: 
Cerro de la Neblina, base camp, 140 m, 0°50'N, 
66°10'W, low foliage, 21-28 Feb. 1985, 12 (W. E. 
Steiner, USNM). 



Metazygia nobas new species 
Figures 129-132; Map 2H 

Holotype. Female holotype from Bafios, 1,600 m, 
Tungurahua Prov., Ecuador, July 1938, and para- 
type from Banos, 2,000 m, July-Aug. 1938 (W. C. 
Macintyre), in MCZ. The specific name is an ar- 
bitrary combination of letters. 

Description. Female holotype. Cara- 
pace orange. Chelicerae, labium, endites 
orange-brown. Sternum dark orange. Legs 
orange. Dorsum of abdomen light with a 
faint folium and spots (Fig. 132); venter 
gray without marks. Posterior median eyes 
0.8 diameter of anterior medians, laterals 
0.8 diameter. Anterior median eyes 0.8 di- 
ameter apart, 1.7 diameters from laterals. 
Posterior median eyes 0.2 diameter apart, 
2.5 diameters from laterals. Height of 
clypeus equals 0.6 diameter of anterior 
median eye. Total length 8.6 mm. Cara- 
pace 4.0 mm long, 2.9 wide, 1.9 behind 
lateral eyes. First femur 3.0 mm, patella 
and tibia 3.6, metatarsus 2.3, tarsus 1.1. 
Second patella and tibia 3.2 mm, third 2.2, 
fourth 2.9. 

Diagnosis. The epigynum (Figs. 129- 
131) differs from that of the Mexican M. 
taman (Figs. 121-123) and the Colombian 
M. chenevo (Figs. 177-179) by having the 
median anterior edge of the base sclero- 
tized (at 12 hr in Fig. 129) and the pos- 
terior median plate heart-shaped (Fig. 
130). Metazygia nobas differs from M. pa- 
quisha (Fig. 126) in the wider posterior 
median plate (Fig. 130). The posterior 
margin of the epigynum base in ventral 
view has a notch on each side (at 7 hr and 
5 hr in Fig. 129) resembling that of M. 
voluptifica (Fig. 280). 



104 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Metazygia goeldii new species 
Figures 133-137; Map 2K 

Holotype. Female holotype from Goeldi Museum, 
Belem, Est. Para, Brazi'l, 10 Feb. 1959 (A. M. Nad- 
ler), in AMNH. The species is named after the 
BraziUan naturaUst E. Goldi. 

Description. Female holotype. Cara- 
pace orange. Chelicerae, labium, endites 
dark orange. Sternum orange. Coxae light- 
est orange, legs dusky orange. Dorsum of 
abdomen light with gray folium (Fig. 137); 
venter light dusky. Posterior median eyes 
0.7 diameter of anterior medians, laterals 
0.7 diameter. Anterior median eyes 0.6 di- 
ameter apart, 1.2 diameters from laterals. 
Posterior median eyes 0.2 diameter apart, 
2 diameters from laterals. Height of clyp- 
eus equals 0.3 diameter of anterior median 
eye. Total length 6.0 mm. Carapace 2.8 
mm long, 2.3 wide, 1.2 behind lateral eyes. 
First femur 2.4 mm, patella and tibia 2.9, 
metatarsus 2.0, tarsus 0.9. Second patella 
and tibia 2.5 mm, third 1.7, fourth 2.3. 

Variation. Total length of females 4.7 
to 6.0 mm. The shape of the scape in the 
two specimens differs (Figs. 133, 136). Il- 
lustrations (Figs. 133-135, 137) were made 
from the female holotype. 

Diagnosis. The abdomen is oval, widest 
in anterior half (Fig. 137). Metazygia goel- 
dii differs from others by the knob-shaped 
scape (Figs. 133-135), which appears 
curved in lateral view (Fig. 135). 

Specimen Examined. BRAZIL Pard: Belem, Inst. 
Agron., 11 Feb. 1959, 19 (A. Nadler, AMNH). 

Metazygia adisi new species 
Figures 138-141; Map 2H 

Holotype. Female holotype from Lago do Jose, Ma- 
naus, Amazonas State, Brazil, 9 Aug. 1987 (J. Adis 
at al), in MCN no. 20058. The species is named 
after the collector. 

Description. The female holotype has 
lost all white pigment including the silver 
tapetum; it apparently was collected in a 
preservative other than alcohol (Levi, 
1989). The specimen is all orange, except 
for a black band around anterior of ab- 
domen which is in the middle (Fig. 141). 
The eye region is black. Posterior median 



eyes same diameter as anterior medians, 
laterals 0.7 diameter. Anterior median eyes 
0.7 diameter apart, 0.5 diameter from lat- 
erals. Posterior median eyes 0.2 diameter 
apart, 1 diameter from laterals. Height of 
clypeus equals 0.3 diameter of anterior 
median eye. Abdomen slightly flattened 
anteriorly (Fig. 141). Total length 3.7 mm. 
Carapace 1.8 mm long, 1.3 wide, 0.7 be- 
hind lateral eyes. First femur 1.9 mm, pa- 
tella and tibia 2.5, metatarsus 1.8, tarsus 
0.6. Second patella and tibia 2.1 mm, third 
1.2, fourth 1.8. 

Diagnosis. Unlike other species, Meta- 
zygia adisi has framed bulges on each side 
of the scape of the epigynum (Figs. 138, 
139). 

Metazygia arnoi new species 
Figures 142, 143; Map 2H 

Holotype. Male holotype from Morro dos Seis Lagos, 
Parque Nacional do Pico da Neblina, Est. Ama- 
zonas, Brazil, 3 Oct. 1990 (A. A, Lise), in MCP. 
The species is named after the collector. 

Description. Male holotype. Carapace 
orange. Chelicerae orange. Labium, en- 
dites, orange. Sternum orange, slightly 
dusky. Coxae; legs orange. Dorsum of ab- 
domen light gray with faint dusky marks, 
pairs of spots and outline of folium (Fig. 
143); venter light gray. Carapace with me- 
dian longitudinal line. Posterior median 
eyes 0.8 diameter of anterior medians, lat- 
erals 0.7 diameter. Anterior median eyes 
0.6 diameter apart, 0.4 diameter from lat- 
erals. Posterior median eyes 0.3 diameter 
apart, 1.5 diameters from laterals. Height 
of clypeus equals 0.5 diameter of anterior 
median eye. Total length 5.1 mm. Cara- 
pace 2.6 mm long, 2.1 wide, behind lateral 
eyes 0.8 wide. First femur 2.4 mm, patella 
and tibia 2.7, metatarsus 2.0, tarsus 0.8. 
Second patella and tibia 2.3 mm, third 1.4, 
fourth 1.9. 

Note. This might be the male of M. 
paquisha. 

Diagnosis. The palpus (Fig. 142) lacks 
a terminal apophysis and differs from all 
others by the loop of the thread-shaped 
embolus (at 12 hr in Fig. 142). 



Met AZYGiA* Levi 



105 



149 




Figures 138-141. Metazygia adisin. sp., female. 138-140, epigynum. 138, ventral. 139, posterior. 140, lateral. 141, dorsal. 

Figures 142, 143. M. arnoin. sp., male. 142, left male palpus. 143, dorsal. 

Figures 144-149. M. curarin. sp. 144-147, female. 144-146, epigynum. 144, ventral. 145, posterior. 146, lateral. 147, dorsal. 
148, 149, male palpus. 148, mesal. 149, ventral. 

Figures 150-153. M. bolivia n. sp., female. 150-152, epigynum. 150, ventral. 152, posterior. 152, lateral. 153, dorsal. 

Figures 154-163. M. yucumon. sp., 154-161, female. 154-156, 158-160, epigynum. 154, 158, ventral. 155, 159, posterior. 
156, 160, lateral. 157, 161, dorsal. 154-157, (Colombia). 158-161, (Bolivia). 162, 163, male. 162, palpus. 163, left femur, 
subventral. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



106 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Metazygia curari new species 
Figures 144-149; Map 2H 

Holotype. Female holotype from Ilha de Curari, Ma- 
naus, Est. Amazonas, Brazil, 3 Aug. 1987 (J. Adis 
et al), in MCN no. 20055. The specific name is a 
noun in apposition after the type locality. 

Description. Female holotype. Cara- 
pace yellowish white, little black between 
eyes. Chelicerae, labium, endites yellowish 
white. Sternum, legs yellowish white. Dor- 
sum of abdomen with two longitudinal 
white lines on dusky white (Fig. 147); ven- 
ter yellowish white. Posterior median eyes 
0.8 diameter of anterior medians, laterals 
0.7 diameter. Anterior median eyes 1.2 di- 
ameters apart, 0.4 diameter from laterals. 
Posterior median eyes 0.3 diameter apart, 
1.3 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Total length 2.7 mm. Cara- 
pace 1 .25 mm long, 0.97 wide, 0.49 behind 
lateral eyes. First femur 1.38 mm, patella 
and tibia 1.61, metatarsus 1.11, tarsus 0.49. 
Second patella and tibia 1.32 mm, third 
0.82, fourth 1.28. 

Male. Color as in female but distal ar- 
ticles of legs dusky. Posterior median eyes 
0.9 diameter of anterior medians, laterals 
0.8 diameter. Anterior median eyes 0.4 di- 
ameter apart, 0.2 diameter from laterals. 
Posterior median eyes 0.1 diameter apart, 
1.2 diameters from laterals. Height of 
clypeus equals 0.6 diameter of anterior 
median eye. Endite with pointed tooth. 
Second tibia thicker than first, proximally 
swollen with macrosetae. Total length 2.5 
mm. Carapace 1.32 mm long, 1.05 wide, 
0.66 behind lateral eyes. First femur 1.30 
mm, patella and tibia 1.61, metatarsus 1.13, 
tarsus 0.59. Second patella and tibia 1.32 
mm, third 0.84, fourth 1.09. 

Note. Males and females were matched 
because both have similar median white 
lines on the abdomen and little black pig- 
ment between eyes. Their collecting lo- 
calities are within 10 km of each other. A 
recently collected female had a green ab- 
domen. 

Diagnosis. The pair of longitudinal 
white lines on the abdomen (Fig. 147) is 
a distinctive character. In ventral view of 



the epigynum, the median plate overlaps 
the lateral plates along their posterior mar- 
gins (at 8 hr, and at 4 hr in Fig. 144). The 
palpus lacks a terminal apophysis and the 
embolus has a distinctive shape (Fig. 148). 

Specimens Examined. BRAZIL Amazonas: Ma- 
naus. Canal Januari, mixed water forest, 16, 17 June 
1987, IS (H. Hofer, IXPA); Ilha de Marchantaria, Rio 
Solimoes, 3°15'S, 59°58'W. 2 Sept. 1992, 19 (J. Adis 
et al., INPA). 

Metazygia botivia new species 
Figures 150-153; Map 2F 

Holotype. Female holotype from Est. Biologico Beni, 
Zone 3, ca. 14°47'S, 66°15'W, ca. 225 m, Depto. 
Beni, Bolivia, 8-14 Nov. 1989 (J. Coddington, S. 
Larcher, C. Griswold, D. Silva D., E. Panaranda), 
in lELP. The specific name is a noun in apposition 
after the type locality. 

Description. Female holotype. Cara- 
pace orange, sides of thoracic region light- 
est. Chelicerae orange-brown. Labium, 
endites dark orange. Sternum orange. Cox- 
ae light orange, legs dusky orange. Dorsum 
of abdomen with white pigment spots and 
pairs of gray brackets (Fig. 153); venter 
light gray. Posterior median eyes 0.7 di- 
ameter of anterior medians, laterals 0.7 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 1.3 diameters from laterals. 
Posterior median eyes 0.4 diameter apart. 
Height of clypeus equals 0.3 diameter of 
anterior median eye. Total length 6.7 mm. 
Carapace 3.0 mm long, 2.0 wide, 1.5 be- 
hind lateral eyes. First femur 2.2 mm, pa- 
tella and tibia 2.5, metatarsus 1.8, tarsus 
0.8. Second patella and tibia 2.3 mm, third 
1.5, fourth 2.0. 

Diagnosis. The elongate abdomen (Fig. 
153) suggests that this species may belong 
to another genus, as yet unnamed, whose 
species have a cylindrical abdomen. How- 
ever, the epigynum is of the characteristic 
shape found in Metazygia, although the 
flat scape is larger than in other species 
(Figs. 150-152). 

Metazygia yucumo new species 
Figures 154-163; Map 2K 

Holotype. Female holotype, female and male para- 
types from 26.9 km SW Yucumo, 500 m, ca. 15°23'S, 
66°59'W, Depto. Beni, BoHvia, 15-19 Nov. 1989 (J. 



METAZYGIA'Levi 



107 



Coddington, C. Griswold, D. Silva D., S. Larcher, 
E. Panaranda), in USNM. The specific name is a 
noun in apposition after the type locafity. 

Description. Female holotype. Cara- 
pace light orange-yellow. Chelicerae, la- 
bium, eridites orange-yellow. Sternum, 
coxae orange-yellow. Legs orange-yellow, 
except distal half of first femora with brown 
ring, underside black, and first two tibiae 
and distal articles brown, underside darker. 
Dorsum of abdomen white without pat- 
tern (Figs. 157, 161); vente^ light gray. 
Posterior median eyes 0.8 diameter of an- 
terior medians, laterals 0.8 diameter. An- 
terior median eyes their diameter apart, 
0.7 diameter from laterals. Posterior me- 
dian eyes 0.3 diameter apart, 2 diameters 
from laterals. Height of clypeus equals 0.4 
diameter of anterior median eye. Total 
length 4.2 mm. Carapace 2.0 mm long, 1.4 
wide, 0.8 behind lateral eyes. First femur 
2.3 mm, patella and tibia 2.7, metatarsus 
1.8, tarsus 0.7. Second patella and tibia 2.2 
mm, third 1.1, fourth 1.8. 

Male. Color as in female. Posterior me- 
dian eyes 0.8 diameter of anterior medi- 
ans, laterals 0.8 diameter. Anterior median 
eyes their diameter apart, 0.5 diameter 
from laterals. Posterior median eyes 0.3 
diameter apart, 1 diameter from laterals. 
Height of clypeus equals 0.4 diameter of 
anterior median eye. First femur with 
many macrosetae (Fig. 163). Total length 
4.2 mm. Carapace 1.9 mm long, 1.3 wide, 
0.6 behind lateral eyes. First femur 2.3 
mm, patella and tibia 2.9, metatarsus 2.3, 
tarsus 0.9. Second patella and tibia 2.3 mm, 
third 1.1, fourth 1.7. 

Note. Males and females were collected 
together. 

Variation. Total length of females 3.7 
to 4.5 mm. The openings of the epigynum 
may be indistinct and hard to see (Fig. 
154); some have a lip laterally (Fig. 158). 
Illustrations were made from the female 
holotype (Figs. 158-161) and a female from 
Mitu, Colombia (Figs. 154-157), and the 
male paratype. 

Diagnosis. The female is separated from 
others by the epigynum, which has an in- 
distinct pair of anterior median openings 



on each side of a thin scape (at 11 hr and 
at 1 hr in Figs. 154, 158). The male has a 
brush of ventral setae on the underside of 
the femur (Fig. 163), and the tip of the 
embolus is hook-shaped (Fig. 162). 

Distribution. Upper Amazon region 
(Map 2K). 

Specimens Examined. COLOMBIA Vaupes: Mitii, 
200 m, Feb. 1975, 19 (P. A. Schneble, MCZ). PERU 
Hudnuco: Divisoria, 1,700 m, 23 Sept. to 3 Oct. 1946, 
19 (F. Woytkowski, AMNH). Madre de Dios: El Li- 
monal, Alto Rio Madre de Dios, 21 July 1988, night 
collecting, 19 (P. Lozada, MUSM). 

Metazygia ipanga new species 
Figures 164-167; Map 2K 

Holotype. Female holotype from Museum Park, Ipi- 
ranga, Sao Paulo, Est. Sao Paulo, Brazil, 6 Dec. 
1960 (J. Luiz), in MZSP no. 7642. The specific name 
is an arbitrary combination of letters. 

Note. Ipiranga is the district of the city 
of Sao Paulo in which the MZSP is located. 

Description. Female holotype. Cepha- 
lothorax very light orange, only eyes with 
black pigment. Dorsum of abdomen whit- 
ish with faint black band around anterior 
(Fig. 167). Venter whitish without pig- 
ment. Posterior median eyes same diam- 
eter as anterior medians, laterals 0.8 di- 
ameter. Anterior median eyes 0.7 diame- 
ter apart, 0.5 diameter from laterals. Pos- 
terior median eyes 0.2 diameter apart, 1 
diameter from laterals. Height of clypeus 
equals 0.3 diameter of anterior median eye. 
Total length 4.5 mm. Carapace 1.9 mm 
long, 1.4 wide, 0.8 behind lateral eyes. First 
femur 2.2 mm, patella and tibia 2.5, meta- 
tarsus 2.0, tarsus 0.6. Second patella and 
tibia 2.0 mm, third 1.2, fourth 1.8. 

Variation. The holotype lacks white 
pigment; it appears washed out by the pre- 
serving fluid (Levi, 1989). The specimen 
from the Chaco Prov., Argentina, had both 
sides of the abdomen white and white pig- 
ment spots anterior to the black band. The 
Bolivian specimen had black pigment be- 
tween the eyes and a square white pigment 
area on the venter of the abdomen. The 
holotype was illustrated. 

Diagnosis. The abdomen is spherical 
(Fig. 167). This species differs from M. 



108 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



uraricoera by the shape of the posterior 
median plate of the epigynum (Fig. 165). 
Distribution. Sao Paulo State, northern 
Argentina to Bolivia (Map 2K). 

Specimens Examined. BOLI\'IA Beni: Est. Biol. 
Beni, HMT'S, 66°15'W, ca. 225 m, 8-14 Nov. 1989, 
19 (J. Coddington et al., USNM). ARGENTINA Cha- 
co: Selvas del Rio Oro, 27 Jan. 1965, 19 (M. E. Galiano, 
MEG). 

Metazygia uraricoera new species 
Figures 168-171; Map 2K 

Holotype. Female holotype from llha de Maraca, Rio 
Uraricoera, Est. Roraima, Brazil, 20 Mar. 1987 (A. 
A. Lise), in MCN no. 20064. The specific name is 
a noun in apposition after the type locality. 

Description. Female holotype. Cara- 
pace orange, eye region black. Chelicerae 
dark orange. Labium, endites dark orange. 
Sternum orange. Legs orange. First femur 
with a distal black spot on underside, distal 
end of first tibia dark. Dorsum of abdomen 
white with an anterior pair of black spots 
(Fig. 171). Venter with a white longitu- 
dinal band, starting at the side of book- 
lung covers. Posterior median eyes 0.8 di- 
ameter of anterior medians, laterals 0.7 
diameter. Anterior median eyes 0.3 di- 
ameter apart, 0.2 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
1.5 diameters from laterals. Height of 
clypeus equals 0.3 diameter of anterior 
median eye. All patellae with a posterior, 
blunt tubercle. Total length 3.1 mm. Car- 
apace 1.4 mm long, 1.1 wide, 0.5 behind 
lateral eyes. First femur 1.4 mm, patella 
and tibia 1.6, metatarsus 1.2, tarsus 0.5. 
Second patella and tibia 1.3 mm, third 0.8, 
fourth 1.1. 

Variation. Total length of females 3.1 
to 4.4 mm. The specimen from Surinam 
had the epigynum slightly shorter in pos- 



terior view and the abdomen more elon- 
gate. Illustrations were made from the ho- 
lotype. 

Diagnosis. The epigynum of this spe- 
cies, unlike that of others, has a sclerotized 
notch anteriorly on each side (at 10 hr and 
at 2 hr in Fig. 168) and a long narrow 
posterior median plate (Fig. 169). 

Natural History. The specimen collect- 
ed in Guyana came from forest savanna. 

Distribution. Guianas to northern Brazil 
(Map 2K). 

Specimens Examined. GUYANA Canje River, Iku- 
ruvva, 5°50'N, 57°50'W, Aug.-Sept. 1961, 19 (G. Bent- 
ley, AMNH). SURINAM Marowijne: Lawn River, 
Anapaike Village, Nov, 1963, 19 (B. Malkin, AMNH). 
BRAZIL Amazonas: Morro dos Seis Lagos, Parque 
Nacional do Pico da Neblina, 5 Oct. 1990, 19 (A. A. 
Lise, MCP). 

Metazygia serian new species 
Figures 172-176; Map 21 

Holotype. Female holotype from La Selva, 4 km SE 
of Puerto Viejo, Heredia Prov., Costa Rica, 7 Aug. 
1980, probably from wasp nest (R. Coville), in MCZ. 
The specific name is an arbitrary combination of 
letters. 

Description. Female holotype. Cara- 
pace orange, cephalic region brown, eye 
region black. Chelicerae dark brown to 
black. Labium, endites brown. Sternum 
orange. Coxae, legs orange. Dorsum of ab- 
domen with dense white pigment spots and 
transverse black band around anterior (Fig. 
175). Venter white with a large white 
square between epigynum and spinnerets 
(Fig. 176). Posterior median eyes 0.9 di- 
ameter of anterior medians, laterals 0.8 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 0.3 diameter from laterals. 
Posterior median eyes 0.3 diameter apart, 
1.2 diameters from laterals. Height of 



Figures 164-167. M. ipanga n. sp., female. 164-166, epigynum. 164, ventral. 165, posterior. 166, lateral. 167, dorsal. 
Figures 168-171. M. uraricoera n. sp.. female. 168-170, epigynum. 168, ventral. 169, posterior. 170, lateral. 171, dorsal. 



Figures 172-176. M. serian n. sp., female. 172-174, epigynum. 172, ventral. 173, posterior. 174, lateral. 175, dorsal. 176, 
abdomen, ventral. 



Metazygia • Levi 



109 




Figures 177-180. M. chenevon. sp., female. 177-179, epigynum. 177, ventral. 178, posterior. 179, lateral. 180, dorsal. 
Figures 181-184. M. lazepa n. sp., female. 181-183, epigynum. 181, ventral. 182, posterior. 183, lateral. 184, dorsal. 
Figures 185-188. M. atalaya n. sp., female. 185-187, epigynum. 185, ventral. 186, posterior. 187, lateral. 188, dorsal. 
Figures 189-192. M. corima n. sp., female. 189-191, epigynum. 189, ventral. 190, posterior. 191, lateral. 192, dorsal. 
Figures 193-196. M. uratron n. sp., female. 193-195, epigynum. 193, ventral. 194, posterior. 195, lateral. 196, dorsal. 
Scale lines. 1 .0 mm, genitalia 0.1 mm. 



110 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



clypeus equals 0.7 diameter of anterior 
median eye. Total length 5.0 mm. Cara- 
pace 1.8 mm long, 1.5 wide, 0.9 behind 
lateral eyes. First femur 2.2 mm, patella 
and tibia 2.6, metatarsus 1.8, tarsus 0.6. 
Second patella and tibia 2.2 mm, third 1.2, 
fourth 1.9. 

Variation. Total length of females 4.8 
to 5.0 mm. Living specimens are green 
(Eberhard, in letter). Illustrations were 
made from the holotype. 

Diagnosis. The abdomen is subspheri- 
cal, widest in anterior half (Fig. 175). This 
species resembles the tetragnathid genus 
Chrysometa in general appearance and 
epigynum (but not the coloration). In ven- 
tral view, the epigynum has openings close 
to the posterior margin (Fig. 172), unlike 
any other Metazygia species. 

Paratype. COSTA RICA Heredia: La 
Selva, Jan. 1978, 1$ (W. Eberhard no. 1279, 
MCZ). 

Metazygia chenevo new species 
Figures 177-180; Map 3A 

Holotype. Female holotype from Finca Chenevo, 175 
m elev., 20 km S El Porvenir, 20 km N Rio Muco, 
Depto. Meta, Colombia, 1978 (W. Eberhard, no. 
1386), in MCZ. The specific name is a noun in 
apposition after the type locality. 

Description. Female holotype. Cara- 
pace orange, cephalic region darkest. Che- 
licerae orange-brown. Labium, endites or- 
ange. Sternum orange. Coxae lighter or- 
ange, legs dusky orange. Dorsum of ab- 
domen with three longitudinal dusky bands 
on white (Fig. 180), sides and venter gray. 
Posterior median eyes 0.6 diameter of an- 
terior medians, laterals 0.6 diameter. An- 
terior median eyes 0.8 diameter apart, 1.5 
diameters from laterals. Posterior median 
eyes 0.2 diameter apart, 3 diameters from 
laterals. Height of clypeus equals 0.5 di- 
ameter of anterior median eye. Total length 
6.0 mm. Carapace 2.9 mm long, 2.2 wide, 
1.3 behind lateral eyes. First femur 2.7 
mm, patella and tibia 3.1, metatarsus 2.1, 
tarsus 0.9. Second patella and tibia 2.7 mm, 
third 1.7, fourth 2.2. 

Diagnosis. Unlike the Ecuadorian M. 



nobas (Fig. 130), the posterior median plate 
of the epigynum has a lateral constriction 
in posterior view (Fig. 178). 

Distribution. Amazon drainage, Colom- 
bia to Guyana (Map 3A). 

Specimen Examined. GUYANA Rupununi Savan- 
na, swamp, Sept. -Oct. 1989, 12 (S. Djojosudharmo, 
F. Mees, CD). 

Metazygia lazepa new species 
Figures 181-184; Map 3A 

Holotype. Female holotype from Hacienda Mozam- 
bique, 15 km SW Puerto Lopez, 200 m, Depto. 
Meta, Colombia, Aug. 1978 (W. Eberhard, no. 1812), 
in MCZ. The specific name is an arbitrary com- 
bination of letters. 

Description. Female holotype. Cara- 
pace orange, cephalic region orange- 
brown. Chelicerae brown. Labium, en- 
dites orange brown. Sternum, coxae or- 
ange; legs dark orange. Dorsum of abdo- 
men light gray with pairs of gray brackets 
(Fig. 184); venter gray without markings. 
Posterior median eyes 0.7 diameter of an- 
terior medians, laterals 0.7 diameter. An- 
terior median eyes 0.5 diameter apart, 1 
diameter from laterals. Posterior median 
eyes 0.2 diameter apart, 2.5 diameters from 
laterals. Height of clypeus equals 0.6 di- 
ameter of anterior median eye. Total length 
8.8 mm. Carapace 3.8 mm long, 3.1 wide, 
2.0 behind lateral eyes. First femur 3.4 
mm, patella and tibia 3.9, metatarsus 2.9, 
tarsus 1 .2. Second patella and tibia 3.4 mm, 
third 2.1, fourth 3.1. 

Diagnosis. Unlike other Metazygia, the 
posterior plate of the epigynum is wide 
and completely covers the lateral plates 
(Figs. 181, 182). The holotype was illus- 
trated. 

Distribution. Colombia, Venezuela 
(Map 3A). 

Specimen Examined. VENEZUELA Carabobo: San 
Esteban, 26 Jan. 1940, 19 (P. Andruze, AMNH). 

Metazygia atalaya new species 
Figures 185-188; Map 3A 

Holotype. Female holotype from Atalaya, Rio Car- 
bon, night collecting, Depto. Cuzco, Peru, 23 Sept. 



Metazygia 'Levi 



111 



1987 (D. Silva D.), in MUSM. The specific name 
is a noun in apposition after the type locality. 

Description. Female holotype. Cara- 
pace orange. Chelicerae, labium, endites 
dark orange. Sternum orange. Legs dusky 
orange. Abdomen gray without any marks 
(Fig. 188). Posterior median eyes 0.7 di- 
ameter of anterior medians, anterior lat- 
erals 0.7 diameter, posterior 0.6. Anterior 
median eyes 0.7 diameter apart, 1.1 di- 
ameters from laterals. Posterior median 
eyes 0.2 diameter apart. Posterior median 
eyes slightly oval. Height of clypeus equals 
0.3 diameter of anterior median eye. The 
abdomen is slightly pointed anteriorly (Fig. 
188). Total length 7.5 mm. Carapace 3.2 
mm long, 2.4 wide, 1.4 behind lateral eyes. 
First femur 2.9 mm, patella and tibia 3.4, 
metatarsus 2.5, tarsus 1.2. Second patella 
and tibia 3.1 mm, third 1.9, fourth 2.7. 

Diagnosis. The abdomen is slightly 
pointed anteriorly (Fig. 188). The epigyn- 
um of this species differs from that of oth- 
ers by the unusual shape of the scape (Figs. 
185-187). 

Metazygia corima new species 
Figures 189-192; Map 3B 

Holotype. Female holotype from Carimagua, 100 m, 
Depto. Meta, Colombia, grass, brush along fence, 
Oct. 1973 (W. Eberhard), in MCZ. The specific 
name is an arbitrary combination of letters. 

Description. Female holotype. Cara- 
pace orange, cephalic region brown. Che- 
licerae brown. Labium, endites orange. 
Sternum dark orange. Coxae, legs orange. 
Dorsum of abdomen with a median lon- 
gitudinal light band that is bordered by 
darker ones (Fig. 192). Sides, venter gray. 
Posterior median eyes 0.7 diameter of an- 
terior medians, anterior laterals 0.7 di- 
ameter, posterior 0.6. Anterior median eyes 
0.6 diameter apart, 1 diameter from lat- 
erals. Posterior median eyes 0.4 diameter 
apart, 2 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Total length 5.5 mm. Cara- 
pace 2.7 mm long, 1.7 wide, 1.1 wide be- 
hind lateral eyes. First femur 2.3 mm, pa- 



tella and tibia 2.7, metatarsus 1.9, tarsus 
0.8. Second patella and tibia 2.4 mm, third 
1.5, fourth 2.1. 

Diagnosis. The epigynum and scape re- 
sembles that of Araneus tiganus (Cham- 
berlin) (Levi, 1991a, figs. 9-11); the scape 
is oval in ventral view (Fig. 189). 

Metazygia uratron new species 
Figures 193-196; Map 3A 

Holotype. Female holotype from Fazenda Santo An- 
tonio, Uruguca, Bahia, Brazil, 24 Oct. 1978 (J. S. 
Santos), in MCN no. 1097. The specific name is an 
arbitrary combination of letters. 

Description. Female holotype. Cara- 
pace orange. Chelicerae dark orange. La- 
bium, endites orange. Sternum light or- 
ange, with irregular dusky spots. Coxae 
light orange, legs orange. Dorsum of ab- 
domen dusky with darker gray folium out- 
line (Fig. 196). Venter gray without marks. 
Posterior median eyes 0.8 diameter of an- 
terior medians, laterals 0.7 diameter. An- 
terior median eyes 0.7 diameter apart, 1.5 
diameters from laterals. Posterior median 
eyes 0.2 diameter apart. Height of clypeus 
equals 0.3 diameter of anterior median eye. 
Total length 7.0 mm. Carapace 3.4 mm 
long, 2.5 wide, 1.5 behind lateral eyes. First 
femur 3.1 mm, patella and tibia 3.3, meta- 
tarsus 2.3, tarsus 1.1. Second patella and 
tibia 2.7 mm, third 1.9, fourth 2.7. 

Diagnosis. This species differs from oth- 
er Metazygia by the shape of the large 
scape (Figs. 193-195). 

Metazygia saturnine new species 
Figures 197-202; Map 3A 

Holotype. Male holotype and immature male from 
Barragem Saturnine de Brito, Santa Maria, Rio 
Grande do Sul, Brazil, 7 July 1982 (M. Rosenau), 
in MCN no. 10596. The specific name is a noun in 
apposition after the type locality. 

Description. Female from Rio Grande, 
Rio Grande do Sul. Carapace with cephalic 
region brown, thoracic region light orange. 
Chelicerae brown. Labium, endites, brown. 
Sternum dusky orange. Coxae light or- 
ange; legs orange to brown, first two legs 



112 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



and distal articles darkest. Dorsum ot ab- 
domen with gray pattern on light gray 
(Fig. 200); venter light gray. Posterior me- 
dian eyes 0.6 diameter of anterior medi- 
ans, laterals 0.6 diameter. Anterior median 
eyes 0.7 diameter apart, 0.7 diameter from 
laterals. Posterior median eyes 0.3 diam- 
eter apart. Height of clypeus equals 0.6 
diameter of anterior median eye. Total 
length 5.3 mm. Carapace 2.5 mm long, 2.1 
wide, 1.2 behind lateral eyes. First femur 
2.1 mm, patella and tibia 2.5, metatarsus 
1.7, tarsus 0.8. Second patella and tibia 2.3 
mm, third 1.3, fourth 1.8. 

Male holotype. Color as in female but 
sternum and coxae are both orange. Pos- 
terior median eyes 0.8 diameter of anterior 
medians, laterals 0.8 diameter. Anterior 
median eyes 0.7 diameter apart, 0.7 di- 
ameter from laterals. Posterior median eyes 
0.2 diameter apart, 2 diameters from lat- 
erals. Height of clypeus equals 0.7 diam- 
eter of anterior median eye. First coxa with 
minute hook. Second tibia as thick as first. 
Total length 4.5 mm. Carapace 2.3 mm 
long, 1.9 wide, 0.8 wide behind lateral eyes. 
First femur 2.7 mm, patella and tibia 3.3, 
metatarsus 2.5, tarsus 1.1. Second patella 
and tibia 2.8 mm, third 1.4, fourth 1.9. 

Note. Males and females were matched 
because of similar size and dark cephalic 
region of carapace. 

Variation. The epigynum of the female 
is probably broken off (Figs. 197-199). 

Diagnosis. The base of the epigynum 
appears broken. The epigynum has a long, 
wide posterior median plate (Fig. 198). The 
male differs from that of M. crabroniphila 
(Fig. 207) by the shape of the conductor 
and the much larger radix (Fig. 201). 

Specimen Examined. BRAZIL Rio Grande do Siil: 
Rio Grande, Est. Ecologica do Taim, 15 Oct. 1985, 
19 (H. Buckup, MCN 13559). 

Metazygia crabroniphila Strand 
Figures 203-207; Map SB 

Aranea (Metazygia) crabroniphila Strand, 1915: 117. 
Female and male syntypes from Joinville, Est. Santa 
Catarina, Brazil, in SMF, examined. 

Metazygia crabroniphila: — Roewer, 1942: 867. Bon- 
net, 1957: 2819. 



Description. Female from Jurubatuba, 
Est. Sao Paulo. Carapace light orange, ce- 
phalic region orange. Chelicerae brown. 
Labium, endites dark orange. Sternum or- 
ange. Coxae light orange; legs orange, dis- 
tal articles darker. Dorsum of abdomen 
with a pair of anterior dusky patches and 
pairs of diagonal marks (Fig. 206); venter 
light, without marks. Posterior median eyes 
0.8 diameter of anterior medians, laterals 
0.8 diameter. Anterior median eyes 0.8 di- 
ameter apart, 1 diameter from laterals. 
Posterior median eyes 0.3 diameter apart. 
Height of clypeus equals 0.6 diameter of 
anterior median eye. Total length 5.5 mm. 
Carapace 2.4 mm long, 1.8 wide, 1.1 wide 
behind lateral eyes. First femur 2.1 mm, 
patella and tibia 2.6, metatarsus 1.6, tarsus 
0.5. Second patella and tibia 2.3 mm, third 
1.3, fourth 1.8. 

Male from Pinhal, Santa Catarina, Bra- 
zil. Color as in female. Posterior median 
eyes 0.8 diameter of anterior medians, an- 
terior laterals 0.8 diameter, posterior lat- 
erals 0.6. Anterior median eyes 0.8 di- 
ameter apart, 0.6 diameter from laterals. 
Posterior median eyes 0.3 diameter apart. 
Height of clypeus equals 0.6 diameter of 
anterior median eye. First coxa without 
hook. Total length 4.0 mm. Carapace 2.5 
mm long, 1.8 wide, 0.9 wide behind lateral 
eyes. First femur 2.4 mm, patella and tibia 
3.1, metatarsus 2.2, tarsus 0.9. Second pa- 
tella and tibia 2.7 mm, third 1.3, fourth 
1.8. 

Variation. Total length of females 4.5 
to 8.0 mm, males 4.0 to 4.3. It is uncertain 
whether the scape of the epigynum is torn 
off in all examined or is a minute bulge 
(Fig. 203). Illustrations were made from a 
female and male from Pinhal. The dorsal 
view is of a female from Jurubatuba. 

Diagnosis. The female can be separated 
from others by the two oval sclerotized 
plates of the epigynum (Fig. 203). The 
male can be distinguished from M. satur- 
nino by having a straight conductor (at 3 
hr in Fig. 207) and a longer median apoph- 
ysis (at 4 hr and at 5 hr in Fig. 207). 

Natural History. Most examined spec- 



Metazygia • Levi 



113 




Figures 197-202. Metazygia saturnino n. sp. 197-200, female. 197-199, epigynum. 197, ventral. 198, posterior. 199, lateral. 
200, dorsal. 201, 202, left male palpus. 201, mesal. 202, apical. 

Figures 203-207. M. crabroniphila Strand. 203-206, female. 203-205, epigynum. 203, ventral. 204, posterior. 205, lateral. 
206, dorsal. 207, male palpus. 

Figures 208-21 2. M. matanzas n. sp., female. 208-21 0, epigynum. 208, ventral. 209, posterior. 21 0, lateral. 21 1 , dorsal. 21 2, 
abdomen, ventral. 

Figures 213-215. M. corumba n. sp., male. 213, palpus. 214, dorsal. 215, abdomen, ventral. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



imens are in poor condition and may all 
have come from wasp nests. 

Specimens Examined. BRAZIL Sdo Paulo: Juru- 
batuba, 6 July 1941, 19 (F. Lane, MZSP 9630); Jardim 
Botanico, Agua Funda, 7 July 1962, 19, 13 (A. F. 
Archer, AMNH). Parana: Curitiba, 2 Feb. 1988, 3$ 
(D. H. Habeck, FSCA). Santa Catarina: Pinhal, Dec. 
1947, 79, 16; Jan. 1948, 19; Dec. 1948, 29, 16; Jan. 
1949, 39 (A. Mailer, AMNH). 



Metazygia matanzas new species 
Figures 208-212; Map 2 J 

Holotype. Female holotype from Pan de Palenque, 
Matanzas, Cuba, 11 Aug. 1955 (A. F. Archer), in 
AMNH. The specific name is a noun in apposition 
after the type locality. 

Description. Female holotype. Cara- 
pace dusky orange, darkest in eye region. 



114 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Chelicerae, labium, endites dusky orange. 
Sternum orange with sides black. Coxae 
orange, legs dusky orange. Dorsum of ab- 
domen with black bands on sides, bordered 
by white bands, and an indistinct dusky 
folium containing anterior black marks 
(Fig. 211). Venter with white patch on 
black (Fig. 212). Posterior median eyes 0.8 
diameter of anterior medians, laterals 0.8 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 0.6 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
1.7 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Total length 3.9 mm. Cara- 
pace 1.6 mm long, 1.2 wide, 0.7 wide be- 
hind lateral eyes. First femur 1.4 mm, pa- 
tella and tibia 1.8, metatarsus 1.1, tarsus 
0.7. Second patella and tibia 1.5 mm, third 
0.9, fourth 1.2. 

Diagnosis. This species may belong to 
Araneus because it has an annulate, flat, 
round scape (Fig. 208). It is placed here 
because of the shape of the abdomen and 
the position of the posterior median eyes 
(Fig. 211). 

Metazygia corumba new species 
Figures 213-215; Map 3B 

Holotype. Male holotype and male paratype (with 
one female of M. voluptifica) from Corumba, Mato 
Grosso do Sul, Brazil, 28-29 May 1960 (B. Malkin), 
in AMNH. The specific name is a noun in appo- 
sition after the type locality. 

Description. Male holotype. Cephalo- 
thorax orange. Dorsum of abdomen white 
with sides dusky. Venter with white patch 
behind genital groove (Fig. 215). Posterior 
median eyes 0.8 diameter of anterior me- 
dians, anterior laterals 0.8 diameter, pos- 
terior laterals 0.7. Anterior median eyes 
0.6 diameter apart, 0.6 diameter from lat- 
erals. Posterior median eyes 0.3 diameter 
apart, 1.7 diameters from laterals. Height 
of clypeus equals 0.5 diameter of anterior 
median eye. Endite without tooth. First 
coxa with small hook. Total length 5.2 mm. 
Carapace 2.5 mm long, 2.0 wide, 1.0 wide 
behind lateral eyes. First femur 2.6 mm, 
patella and tibia 3.2, metatarsus 2.4, tarsus 



0.9. Second patella and tibia 2.7 mm, third 
1.5, fourth 2.0. 

Note. The two males from Corumba 
were collected with a female of M. mun- 
dula. 

Variation. Total length of males 3.7 to 
5.2 mm. Illustrations were made from the 
holotype. 

Diagnosis. This species differs from M. 
crabroniphila by having the two edges of 
the embolus straight and parallel (Fig. 213). 

Specimen Examined. BOLIVIA Smita Cruz: Ma- 
taral, 14 Dec. 1984, 16 (L. Pena, AMNH). 

Metazygia sendero new species 
Figures 216-221 ; Map SB 

Holotype. Female holotype from Sendero Campa- 
mento, Laguna Grande, PUCE Field Station, Re- 
serva Faunistica Cuyabeno, 00°00'N, 76°10-irW, 
31 July to 5 Aug. 1988, Sucumbios Prov., Ecuador, 
31 July to 5 Aug. 1988 (W. Maddison), in MECN. 
The specific name is a noun in apposition after the 
type locality; sendero is the Spanish word for path. 

Description. Female holotype. Cara- 
pace orange and black. Chelicerae brown- 
ish black. Labium, endites brown. Sternum 
brown. Coxae dusky orange, legs dark or- 
ange-brown. Dorsum of abdomen with a 
series of pairs of black brackets (Fig. 219). 
Venter with a black square (Fig. 220). Pos- 
terior median eyes 0.7 diameter of anterior 
medians, laterals 0.7 diameter. Anterior 
median eyes 0.5 diameter apart, 2.2 di- 
ameters from laterals. Posterior median 
eyes 0.2 diameter apart. Height of clypeus 
equals 0.5 diameter of anterior median eye. 
Total length 11.0 mm. Carapace 4.6 mm 
long, 3.2 wide, 2.5 behind lateral eyes. First 
femur 3.5 mm, patella and tibia 4.5, meta- 
tarsus 3.1, tarsus 1.4. Second patella and 
tibia 3.9 mm, third 2.2, fourth 3.4. 

Male. Coloration as in female, but ven- 
ter of abdomen with scattered white pig- 
ment spots. Posterior median eyes 0.7 di- 
ameter of anterior medians, laterals 0.7 
diameter. Anterior median eyes 0.4 di- 
ameter apart, 1.6 diameters from laterals. 
Posterior median eyes 0.4 diameter apart, 
3.5 diameters from laterals. Height of 
clypeus equals 0.7 diameter of anterior 



Metazygia • Levi 115 




Figures 216-221. Metazygia sendero n. sp. 216-220, female. 216-218, epigynum. 216, ventral. 217, posterior. 218, lateral. 
219, dorsal. 220, abdomen, ventral. 221, left male palpus. 

Figures 222-225. M. uma n. sp. 222-224, female. 222, 223, epigynum. 222, ventral. 223, posterior. 224, right, dorsal; left, 
abdomen, ventral. 225, male palpus. 

Figures 226-230. M. laticeps (O. P.-Cambridge). 226-229, female. 226-228, epigynum. 226, ventral. 227, posterior. 228, dorsal. 
229, abdomen, ventral. 230, male palpus. 

Figures 231-238. M. mundulella Strand. 231-236, female. 231-235, epigynum. 231 , ventral. 232, posterior. 233, lateral. 234, 
broken ventral. 235, broken posterior. 236, dorsal. 237, 238, palpus. 237, mesal. 238, ventral. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



116 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



median eye. Total length 6.5 mm. Cara- 
pace 3.2 mm long, 2.3 wide, 1.5 behind 
lateral eyes. First femur 2.5 mm, patella 
and tibia 3.1, metatarsus 2.3, tarsus 1.2. 
Second patella and tibia 2.7 mm, third 1.5, 
fourth 2.1. 

Note. Males and females were matched 
because of similar coloration, a similar wide 
cephalic region and swelling behind the 
eyes, and similarly shaped abdomen. The 
only other species with these characters is 
M. laticeps. Male and female come from 
the opposite slopes of the Andes. 

Variation. Total length of females 8.0 
to 11.0 mm. Illustrations were made from 
the female holotype and a male from Tin- 
alandia, Ecuador. 

Diagnosis. The abdomen is elongate, 
oval, slightly overhanging spinnerets (Fig. 
220). This species differs from M. laticeps 
by having the epigynum with a narrower 
triangle in ventral view (Fig. 216); in pos- 
terior view, the median plate is as wide as 
long (Fig. 217) while that of M. laticeps 
is narrow (Fig. 227). The median apoph- 
ysis of the palpus (at 4 hr in Fig. 221) has 
two tips. 

Distribution. Ecuador, Peru (Map 3B). 

Specimens Examined. ECUADOR Pichincha: 
Tinalandia, 12 km E Santo Domingo de los Colorados, 
750 m, beating vegetation, 11-17 May 1986, 1 imm., 
IS, 26 (G. B. Edwards, FSCA). PERU Ucayali: Co- 
lonia Calleria, Rio Calleria, 15 km from Ucayali, 10- 
30 Sept. 1961, 12 (B. Malkin, AMNH). 

Metazygia uma new species 
Figures 222-225; Map 3B 

Holotype. Female holotype from Puesto de Vigilan- 
cia Pakitza, Zona Reservada de Manu, Depto. Ma- 
dre de Dios, 11°58'S, 7ri8'W, Peru, inundated for- 
est, 30 Sept. 1987 (D. Silva D., J. Coddington), in 
MUSM. The specific name is an arbitrary combi- 
nation of letters. 

Description. Female holotype. Cara- 
pace orange. Chelicerae, labium, endites 
orange. Sternum orange. Coxae orange; 
legs orange but distal tips of tibiae, meta- 
tarsi and tarsi black. Dorsum of abdomen 
with longitudinal gray lines (Fig. 224); 
venter with a black rectangle (Fig. 225). 



Posterior median eyes 0.7 diameter of an- 
terior medians, laterals 0.7 diameter. An- 
terior median eyes 0.5 diameter apart, 3 
diameters from laterals. Posterior median 
eyes 0.4 diameter apart. Height of clypeus 
equals 0.6 diameter of anterior median eye. 
Total length 8.2 mm. Carapace 3.8 mm 
long, 2.5 wide, 2.3 behind lateral eyes. First 
femur 2.6 mm, patella and tibia 3.4, meta- 
tarsus 2.3, tarsus 1.1. Second patella and 
tibia 2.9 mm, third 1.7, fourth 2.7. 

Male. Coloration as in female, but ven- 
ter of abdomen all black. Posterior median 
eyes 0.8 diameter of anterior medians, lat- 
erals 0.5. Anterior median eyes 0.8 diam- 
eter apart. Posterior median eyes 0.3 di- 
ameter apart. Coxal hook very small. Sec- 
ond tibia thicker than first with two long 
macrosetae in a line. Height of clypeus 
equals 1 diameter of anterior median eye. 
Total length 3.9 mm. Carapace 2.0 mm 
long, 1.4 diameters wide, 0.9 wide behind 
lateral eyes. First femur 1.4 mm, patella 
and tibia 1.8, metatarsus 1.2, tarsus 0.7. 
Second patella and tibia 1.5 mm, third 0.8, 
fourth 1.1. 

Note. Male was matched with the fe- 
male because of similar coloration and the 
relatively wide cephalic region of the car- 
apace. 

Diagnosis. This species is distinct in that 
the eye region of the carapace is almost as 
wide as the thoracic region (Fig. 224). The 
male differs from M. laticeps by the curved 
embolus lamella with a short curved em- 
bolus "above" it (Fig. 225). This strange- 
looking species is apparently a Metazygia 
and closest to M. laticeps, which has also 
the carapace relatively wide anteriorly 
(Fig. 228). 

Natural History. The male was collect- 
ed in forest interior. 

Distribution. Amazon region, Peru to 
Brazil (Map 3B). 



Specimens Examined. PERU Madre de Dios: Zona 
Reservada Tambopata, 290 m, 8 July 1987, 19 (D. 
Silva D., MUSM). BRAZIL Amazonias: Reserva Flo- 
restal, 80 km from Manaus, 19 Feb. 1991, IS (H. 
Fowler, E. Venticinque, R. S. Vieira, MCZ). 



Metazygia • Levi 



117 



Metazygia laticeps (O. P.-Cambridge), 
new combination 
Figures 226-230; Map 3F 

Epeira laticeps O. P.-Cambridge, 1889: 18, pi. 4, fig. 

16, 2. Female holotype from Bugaba, Panama, in 

BMNH no. 1890.7.1.5020, examined. Keyserling, 

1892: 175, pi. 8, fig. 129, 2. 
Aranea laticeps: — F. P.-Cambridge, 1904: 516, pi. 

49, fig. 21, 2. Roewer, 1942: 845. 
Araneus laticeps: — Bonnet, 1955: 527. 

Note. The type specimen belonged to 
Keyserling. The vial containing the type 
also contains Keyserling's original, toothed, 
blue-bordered label, which is still faintly 
legible when dried. The first line reads 
Guatemala and not Bugaba. 

Description. Female from Pipeline 
Road, Panama. Carapace dark orange with 
median double line, black on each side of 
thoracic region. Chelicerae, labium, en- 
dites orange. Sternum dusky orange-brown. 
Coxae light orange, legs dusky orange- 
brown. Dorsum of abdomen with pairs of 
brackets (Fig. 228); venter with median 
dark gray patch (Fig. 229). Posterior me- 
dian eyes 0.8 diameter of anterior medi- 
ans, laterals 0.6 diameter. Anterior median 
eyes 0.8 diameter apart, 2 diameters from 
laterals. Posterior median eyes 0.6 diam- 
eter apart. Height of clypeus equals 0.6 
diameter of anterior median eye. Total 
length 10.0 mm. Carapace 3.7 mm long, 
3.1 wide, 2.2 behind lateral eyes. First fe- 
mur 3.4 mm, patella and tibia 4.4, meta- 
tarsus 2.7, tarsus 1.3. Second patella and 
tibia 3.7 mm, third 2.2, fourth 3.2. 

Male from Barro Colorado Island, Pan- 
ama. Color as in female. Posterior median 
eyes 0.7 diameter of anterior medians, lat- 
erals 0.7 diameter. Anterior median eyes 
0.8 diameter apart, 2.5 diameters from lat- 
erals. Posterior median eyes 0.3 diameter 
apart. Height of clypeus equals 0.4 di- 
ameter of anterior median eye. Abdomen 
as in female, but smaller. Total length 6.3 
mm. Carapace 3.4 mm long, 2.5 wide, 1.7 
behind lateral eyes. First femur 2.9 mm, 
patella and tibia 3.8, metatarsus 2.7, tarsus 
1.2. Second patella and tibia 3.1 mm, third 
1.7, fourth 2.5. 



Note. Males and females were matched 
because both have a dusky patch on the 
venter of the abdomen (Fig. 229) and be- 
cause both have the same wide distribu- 
tion. Males are much less common in col- 
lections than females. 

Variation. Total length of females 8.0 
to 11.0 mm. Females and male from Mato 
Grosso, Brazil, differed from those of other 
regions: the female epigynum is more 
rounded posteriorly and in posterior view 
there is a round depression ventrally, but 
the lateral and median sclerites are of the 
same width as the one illustrated (Fig. 227). 
The male from Mato Grosso has a more 
elongate median apophysis, and the two 
parallel prongs of the embolus are of sim- 
ilar width. 

Illustrations were made from a female 
from Pipeline Road, Panama, and a male 
from Barro Colorado Island, Panama. 

Diagnosis. The abdomen is elongate oval 
(Fig. 228). Metazygia laticeps female has 
a pointed, wide, triangular epigynum (Fig. 
226) with the posterior median plate 
slightly narrower than the lateral plates on 
each side (Fig. 227). The male has a large, 
semicircular median apophysis (at 5 hr in 
Fig. 230). 

Natural History. Females are collected 
in tropical forest by unrolling rolled-up 
leaves, their retreat, at a height of about 
150 cm. Specimens from Mato Grosso all 
came from gallery forest; Guyanas from 
forest savanna and swamp forest; and near 
Iquitos, Peru, from rain forest. 

Distribution. Panama to Rio de Janeiro 
and northern Bolivia (Map 3F). 

Specimens Examined. PANAMA Colon: Fort 
Sherman, 12 (MCZ). Panama: nr. Gamboa, edge of 
Canal, 1948, 12 (W. Eberhard, MCZ); Soberania Natl. 
Park, Pipeline Road, 8 km NW Gamboa, 12 (MCZ); 
Barro Colorado Isl,, Lago Gatun, 2S, 22, MCZ. 

TRINIDAD Port of Spain, 13 (MCZ). GUYANA 
Canje Ikuruwa River 05°70'N, 57°50'W, 12 (AMNH). 
FRENCH GUIANA nr. Placer Tresor, Roura Mtns., 
12 (MCZ); nr, Sautero, Matouri, 12 (MCZ). COLOM- 
BIA Satander: Rio Suarez, 800-1,000 m, 12 (AMNH). 
PERU hereto: Iquitos, 12 (AMNH); Explorama Lodge, 
80 km NE Iquitos, 12 (FSCA). San Martin: 32 km 
SE Moyabamba, \$ (AMNH). Junin: Amable Maria, 



118 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



12 (PAN). Madre de Dios: Zona Reservada Tambo- 
pata, 290 m, 19 (MUSM). BRAZIL Pard: Canind6, 
Rio Gurupi, 12 (AMNH). Amazonas: Tefe, 12 (S. Par- 
rish, MCZ); Rio Negro, Umarituba, 12 (NRMS); 
Maturaca, 12 (MCP). Espirito Santo: Linhares, Par- 
que Souterrana, 12 (MZSP). Rio de Janeiro: Rio de 
Janeiro, Jardim Botanico, 12 (MCZ); Parque da Ci- 
dade, 12 (MCZ). Mato Grosso: 260 km N Xavantina, 
400 m, 12''49'S, 51°46'W, 32, U (MCZ). BOLIVIA 
Beni: Chacobo Indian Village, Rio Benicito, 12 
(AMNH); Est. Biologica Beni, 12 (USNM). 

Metazygia mundulella Strand 
Figures 231-238; Map 3F 

Aranea (Metazygia) mundulella Strand, 1915: 114. 
Ten female, two male, and two immature syntypes 
from mud-dauber wasp nest, Joinville, Santa Ca- 
tarina State, Brazil, in SMF no. 4010, examined. 

Aranea mundulella: — Roewer, 1942: 848. 

Larinia mundula: — Bonnet, 1957: 2350. 

Note. Strand (1915: 114) wrote, "Since 
there is no certainty from the hterature as 
to which species the specimens belong, I 
make some descriptive remarks and pro- 
pose, if necessary, the name mundulella." 

Description. Female syntype. Carapace 
orange, cephalic region darker. Chelicer- 
ae, labium, endites dark orange. Sternum 
orange. Coxae, legs orange. Dorsum of ab- 
domen light with a pair of dark anterior 
patches and four pairs of dark lines (Fig. 
236). Venter light. Posterior median eyes 
same diameter as anterior medians, lat- 
erals 0.8 diameter. Anterior median eyes 
0.5 diameter apart, 0.9 diameter from lat- 
erals. Posterior median eyes 0.3 diameter 
apart, 2.1 diameters from laterals. Height 
of clypeus equals 0.5 diameter of anterior 
median eye. Total length 5.9 mm. Cara- 
pace 2.8 mm long, 2.1 wide, 1.4 behind 
lateral eyes. First femur 2.3 mm, patella 
and tibia 2.6, metatarsus 1.8, tarsus 0.8. 
Second patella and tibia 2.3 mm, third 1.4, 
fourth 2.0. 

Male syntype. Color, including marks 
on abdomen, as in female. Posterior me- 
dian eyes 0.9 diameter of anterior medi- 
ans, laterals 0.5 diameter, posterior laterals 
0.5. Anterior median eyes 0.7 diameter 
apart, 0.7 diameter from laterals. Posterior 
median eyes 0.2 diameter apart, 1.2 di- 
ameters from laterals. Height of clypeus 



equals 0.7 diameter of anterior median eye. 
Second tibia as thick as first, with macro- 
setae. Total length 4.5 mm. Carapace 2.4 
mm long, 2.0 wide, 0.9 wide behind lateral 
eyes. First femur 2.1 mm, patella and tibia 
2.7, metatarsus 1.9, tarsus 0.7. Second pa- 
tella and tibia 2.3 mm, third 1.3, fourth 
1.8. 

Note. The syntypes have lost all white 
pigment and also the silver pigment of the 
eyes, perhaps from having been in a buf- 
fered formaldehyde solution (Levi, 1989). 
All except for one specimen (Figs. 231, 
232) have the epigynum broken. Each side 
is broken off (Figs. 234, 235), apparently 
the result of mating. Males and females 
were collected together. 

Diagnosis. The epigynum, unlike that 
of M. genialis (Fig. 239), has a concave 
margin on each side and a flat scape (Figs. 
231, 233). In posterior view, it has a ventral 
pocket on each side (at 11 hr and 2 hr in 
Fig. 232). The male palpus, like that of M. 
genialis (Figs. 243, 244), has a lobe on the 
tegulum (at 12 hr in Figs. 237, 238) but 
differs in the shape of the embolus lamella 
and median apophysis (center and at 5 hr 
in Fig. 237). 

Natural History. All specimens came 
from a mud-dauber wasp nest. 

Metazygia genialis (Keyserling) 
Figures 239-246; Map 3F 

Epeira genialis Keyserling, 1892; 156, pi. 8, fig. 114, 
2. Two female syntypes from Rio Grande do Sul, 
Brazil, one has the epigynum broken, the other 
covered by secretions, in BMNH, examined. 

Epeira mundula: — Keyserling, 1892: 179, pi. 9, fig. 
132, S (not female lectotype). 

Aranea genialis: — Roewer, 1942: 843. 

Araneus genialis: — Bonnet, 1955: 507. 

Metazygia genialis: — Levi, 1991a: 179. 

Note. The male paralectotype of E. 
mundula belongs with the female of M. 
genialis. 

Description. Female from Santa Vitoria 
do Palmar, Rio Grande do Sul. Cephalic 
region of carapace dark brown, thoracic 
region yellowish. Chelicerae dark brown. 
Labium, endites dark brown. Sternum light 
brown. Coxae yellowish; legs with proxi- 



METAZYCIA'Levi 



119 



239 




Figures 239-246. Metazygia genialis (Keyserling). 239-242, female. 239-241, epigynum. 239, ventral. 240, posterior. 241, 
lateral. 242, dorsal. 243-246, left male palpus. 243, mesal. 244, ventral. 245, 246, palpus pulled apart. 

Figures 247-249. M. amalla n. sp., female. 247, 248, epigynum. 247, ventral. 248, posterior. 249, dorsal. 

Figures 250-252. M. ikuruwa n. sp., male. 250, 251, palpus. 250, mesal. 251, ventral. 252, dorsal. 

Abbreviations. C, conductor; H, hematodocha; E, embolus; L. emtwius lamella; M, median apophysis; P, paracymbium; R, 
radix; T, teguium; Y, cymbium. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



mal articles yellowish, distal brown. Dor- 
sum of abdomen with dusky outline of fo- 
lium (Fig. 242); venter gray, without 
marks. Posterior median eyes 0.6 diameter 
of anterior medians, laterals 0.6 diameter. 
Anterior median eyes 0.7 diameter apart, 
1 diameter from laterals. Posterior median 
eyes 0.3 diameter apart, 3 diameters from 
laterals. Height of clypeus equals 0.3 di- 



ameter of anterior median eye. Total length 
6.3 mm. Carapace 3.0 mm long, 2.3 wide, 
1.5 wide behind lateral eyes. First femur 
2.2 mm, patella and tibia 2.7, metatarsus 
1.8, tarsus 0.8. Second patella and tibia 2.5 
mm, third 1.7, fourth 2.1. 

Male from Santa Vitoria do Palmar, Rio 
Grande do Sul. Color as in female, but 
cephalic region yellowish. Posterior me- 



120 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



dian eyes 0.5 diameter of anterior medi- 
ans, laterals 0.5 diameter. Anterior median 
eyes 0.4 diameter apart, 0.5 diameter from 
laterals. Posterior medians 0.2 their di- 
ameter apart, 2.2 diameters from laterals. 
Height of clypeus equals 0.4 diameter of 
anterior median eye. Third femur with 
three short macrosetae; each only three 
times as long as wide. Total length 4.8 mm. 
Carapace 2.6 mm long, 2.2 wide, 1.1 be- 
hind lateral eyes. First femur 2.2 mm, pa- 
tella and tibia 2.7, metatarsus 2.2, tarsus 
0.7. Second patella and tibia 2.4 mm, third 
1.4, fourth 1.7. 

Note. Males and females were collected 
together. 

Variation. Total length of females 6.3 
to 7.7 mm, males 4.8 to 5.8. Illustrations 
were made from Santa Vitoria, Rio Grande 
do Sul, Brazil. 

Diagnosis. The epigynum has a knob 
(Figs. 239, 240) rather than a flat scape as 
in M. mundulella (Figs. 231-235). The 
male, like M. mundulella (Figs. 237, 238), 
has a tegulum (T in Fig. 245) with a lobe 
(at 12 hr in Figs. 243, 245) and a distinctive 
shape of the embolus lamella (L in Fig. 
245) and median apophysis (at 5 hr in Fig. 
243). 

Distribution. Bahia to Rio Grande do 
Sul States, Brazil (Map 3F). 

Specimens Examined. BRAZIL Bahia: Mucuri, Fa- 
zenda Farol, 11 Apr. 1979, 1<? (A. C. Viella, MCN 
11108). Rio Grande do Sul: Santa Vitoria do Palmar, 
Estayao Ecologica do Taim, 9 Apr. 1986, 4$, 16 (M. 
Rosenau, MCN 14821); Rio Grande, 4 Dec. 1986, 1.5 
(A. D. Brescovit, MCN 16287); Guaiba, 9 Jan. 1980, 
16 (M. H. Galileo, MCN 09182); Viamao, Fazenda 
Sanga da Porteira, 11-14 Apr. 1983, 22 (A. A. Lise, 
MCN 11565); Viamao, Lagoa do Casamento, 2 Apr. 
1975, 15 (A. A. Lise, MCN 02644a); Viamao, Esta^ao 
Exper. Fitotecnica de Aguas Belas, 30 Mar. 1977, \$ 
(E. H. Buckup, MCN 05567); Pelotas, Mav 1959, 1<3 
(C. Biezanko, AMNH). 

Metazygia amalla new species 
Figures 247-249; Map 3D 

Holotype. Female holotype from Pinhal, Est. Santa 
Catarina, Brazil, Jan. i948 (A. Mailer), in AMNH. 
The specific name is an arbitrary combination of 
letters. 



Description. Female holotype. Cara- 
pace orange, cephalic region orange- 
brown. Chelicerae dark orange-brown. 
Labium, endites brown. Sternum orange. 
Coxae light orange, legs orange-brown. 
Dorsum of abdomen with faint dark mark- 
ings forming outline of a folium (Fig. 249); 
venter light dusky. Posterior median eyes 
0.8 diameter of anterior medians, laterals 
0.8 diameter. Anterior median eyes 0.8 di- 
ameter apart, 1.1 diameters from laterals. 
Posterior median eyes 0.2 diameter apart. 
Height of clypeus equals 0.6 diameter of 
anterior median eye. Total length 6.3 mm. 
Carapace 2.7 mm long, 2.1 wide, 1.1 wide 
behind lateral eyes. First femur 2.4 mm, 
patella and tibia 2.7, metatarsus 1.8, tarsus 
0.8. Second patella and tibia 2.3 mm, third 
1.5, fourth 2.0. 

Variation. Total length of females 6.3 
to 6.5 mm. Illustrations were made from 
the female holotype. The holotype and 
paratype have the median area of the epi- 
gynum broken (Figs. 247, 248). 

Diagnosis. Metazygia amalla differs 
from other species by the wide posterior 
median plate of the epigynum (Fig. 248). 

Paratype. BRAZIL Santa Catarina: 
Pinhal, Jan. 1948, 29 (A. Mailer, AMNH). 

Metazygia ikuruwa new species 
Figures 250-252; Map 3D 

Holotype. Male holotype from Canje Ikuruwa River, 
05°70'N, 57°50'W,' Guyana, Aug.-Dec. 1961 (G. 
Bentley), in AMNH. The specific name is a noun 
in apposition after the type locality. 

Description. Male holotype. Carapace 
orange. Chelicerae, labium, endites or- 
ange; sternum, coxae orange. Legs orange, 
distal articles darker. Dorsum of abdomen 
with dark outline of folium (Fig. 252); ven- 
ter dusky. Posterior median eyes 0.8 di- 
ameter of anterior medians, laterals 0.7 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 0.5 diameter from laterals. 
Posterior median eyes 0.3 diameter apart, 
1.5 diameters from laterals. Height of 
clypeus equals 0.6 diameter of anterior 
median eye. Total length 4.4 mm. Cara- 
pace 2.4 mm long, 1.7 wide, 0.8 wide be- 



Metazycia • Levi 



121 



hind lateral eyes. First femur 2.3 mm, pa- 
tella and tibia 2.7, metatarsus 2.0, tarsus 
0.9. Second patella and tibia 2.3 mm, third 
1.3, fourth 1.9. 

Diagnosis. This species differs from M. 
gregalis by the large anchor-shaped me- 
dian apophysis (at 5 hr in Fig. 250). 

Paratypes. GUYANA Bartica: Kartabo, 
1920, 33 (CUC). 

Metazygia gregalis (O. P. -Cambridge) 
Figures 253-262; Map 3E 

Epeira gregalis O. P. -Cambridge, 1889: 22, pi. 5, fig. 
3, 2. Ten female syntypes from Veragua [Prov. 
Veraguas], Panama in BMNH, examined. Keyser- 
ling, 1892: 177, pi. 9, fig. 131, 2, <?. 

Metazygia gregalis: — F. P.-Cambridge, 1904: 501, 
pi. 47, fig. 24, 2, (5. Petrunkevitch, 1930: 327, figs. 
208-210, 2, 6. Roewer, 1942: 868. Bonnet, 1957: 
2819. 

Eustala tuceps Chamberlin, 1925: 217. Female ho- 
lotype from Barro Colorado Island [Lago Gatiin], 
Panama, in MCZ, examined. Roewer, 1942: 767. 
First synonymized by Banks, 1929: 95. 

Metazygia manni Bryant, 1945: 377, figs. 12, 13, 23, 
2, S. Male holotype from Cap Haitien, Haiti, in 
MCZ, examined. Brignoli, 1983: 274. NEW SYN- 
ONYMY. 

Metazygia similis Caporiacco, 1947: 25; 1948: 660, 
fig. 70, 2. Female holotype from Mackenzie, 
[06°00'N, 58°17'W], Guyana, in MZUF, examined. 
Brignoli, 1983: 274. NEW SYNONYMY. 

Synonymy. The genitalia of Metazygia 
manni and M. similis are similar to those 
of M. gregalis. No differences could be 
found. 

Description. Female from Panama. 
Carapace orange, cephalic region brown. 
Chelicerae brown. Labium, endites, ster- 
num orange. Coxae orange; legs orange. 
Dorsum of abdomen with dusky pattern 
over tiny white pigment spots (Fig. 257); 
venter light dusky without marks. Poste- 
rior median eyes 0.8 diameter of anterior 
medians, laterals 0.8 diameter. Anterior 
median eyes 0.8 diameter apart, 1.5 di- 
ameters from laterals. Posterior median 
eyes 0.5 diameter apart. Height of clypeus 
equals 0.8 diameter of anterior median eye. 
Total length 8.0 mm. Carapace 3.5 mm 
long, 2.7 wide, 1.5 behind lateral eyes. First 



femur 2.7 mm, patella and tibia 3.2, meta- 
tarsus 2.4, tarsus 0.9. Second patella and 
tibia 2.9 mm, third 1.8, fourth 2.5. 

Male from Panama. Color as in female. 
Posterior median eyes 0.7 diameter of an- 
terior medians, anterior laterals 0.7 di- 
ameter, posterior laterals 0.6. Anterior me- 
dian eyes 0.5 diameter apart, 0.7 diameter 
from laterals. Posterior median eyes 0.3 
diameter apart. Height of clypeus equals 
0.6 diameter of anterior median eye. Fangs 
modified (Figs. 261, 262). Endite without 
tooth, palpal femur without facing tuber- 
cle. First coxa with very small hook. Sec- 
ond tibia thicker than first, without special 
macrosetae. Total length 5.0 mm. Cara- 
pace 2.9 mm long, 2.1 wide, 1.3 wide be- 
hind lateral eyes. First femur 2.4 mm, pa- 
tella and tibia 2.8, metatarsus 2.1, tarsus 
0.9. Second patella and tibia 2.3 mm, third 
1.5, fourth 2.0. 

Note. Males and females are commonly 
collected together. 

Variation. Total length of females 6.2 
to 9.6 mm, males 4.0 to 6.0. The largest 
male and female both came from Pelotas, 
Rio Grande do Sul State, Brazil. The epi- 
gynum is quite variable: sometimes the 
posterior two bulges are absent, sometimes 
there is a median bulge. The illustrations 
were made from specimens from Barro 
Colorado Island, Panama. 

Diagnosis. The female epigynum in 
ventral view is wider than long (Figs. 253, 
254); that of M. benella and M. yobena is 
longer than wide (Figs. 263, 270). The me- 
dian apophysis (M) of the male palpus is 
a small hook (Figs. 258, 260); it is longer 
in M. benella (Fig. 267) and has a black 
wall in M. yobena (Fig. 274) 

Natural History. Specimens have been 
collected from the following places: brush 
along fences, on houses, and eaves of a 
building in Costa Rica; on a building at 
night in Panama; on walls under light at 
night in Paraguay; in sweeping river veg- 
etation in Bolivia; from a wasp nest in Su- 
rinam; and from rolled leaves in savanna, 
Depto. Beni, Bolivia. 

Distribution. Nicaragua, Greater Antil- 



122 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



les (except Jamaica), Tobago, south to Ar- 
gentina (Map 3E). 

Specimens Examined. NICARAGUA Bonanza 
(AMNH). COSTA RICA Heredia: Serapiqui (MCZ); 
La Selva (MCZ, USNM). Cartago: Turrialba (CAS). 
Puntarenas: Finca Selva Verde (DU). PANAMA Chi- 
riqui: Puerto Armuelles (FSCA); David (AMNH, 
MCZ). Veraguas: NE Puerto Mutis (MIUP). Herrera: 
Paris (MCZ). Code: Nata (AMNH). Colon: Santa Rosa 
(AMNH); Fort Gulick (AMNH); Puente Sobre Re- 
presa Madden (MIUP); Madden Dam (AMNH, MCZ). 
Panama: Reserva Forestal (MIUP); Barro Colorado 
Isl., Lago Gatun (CAS, MIUP); Frijoles (MCZ); Pedro 
Miguel (MCZ); Red Tank (MCZ). 

CUBA Santiago de Cuba: Cuabitas (AMNH). HAI- 
TI Cap Haitien (MCZ). DOMINICAN REPUBLIC 
Santo Domingo, Jardin Botanico (MCZ); Cruce de 
Jima Abajo, La Vega (MNSD). PUERTO RICO Aguas 
Buenas (JEC); Caguas (AMNH); Adjuntas (AMNH); 
Humacao (MCZ); Laguna Cartagena, Valle de Leras 
(MCZ); Loma Tinaja, Laguna Cartagena (AMNH); 
Mayaguez (AMNH); Toa Baja (AMNH). TOBAGO 
Bucco Bay (AMNH). 

VENEZUELA Sucre: 7 km E San Antonio del Gol- 
fo (USNM). Apure: Mantecal (MCZ), Distrito Fed- 
eral: San Jose del Avila, Caracas (AMNH). GUYANA 
Kartabo (AMNH). SURINAM Brokopondo Lake 
(AMNH). FRENCH GUIANA St. Laurent de Maroni 
(PAN). COLORADO Magdalena: Pozo Colorado, 10 
km W Santa Marta (AMNH). Santander: Rio Suarez 
(AMNH). Meta: Carimagua (MCZ); El Porvenir, 140 
m (MCZ); Finca Chenevo, 20 km N Rio Muco, 20 
km S El Porvenir (MCZ); Lomalinda, Puerto Lleras 
(CAS, MCZ); 15 km SW Puerto Lopez, Hda. Mozam- 
bique, 200 m (MCZ). Valle: Cali (AMNH, MCZ); 
Centr. Hidroelectr. Anchicaya (MCZ); Lago Calima, 
1,300 m (MCZ); Palmira (CAS); Rio Jamundi entre 
Cali y Jamundi (MCZ); Rio Para, below Buenos Aires 
(MCZ); Rio Tulud, 1,100 m (MCZ); Sevilla (AMNH). 
ECUADOR Sucumbios: Res. Fauna Cuyabeno, La- 
guna Grande (MCZ). Manabi: road betw. Crucita and 
Charapoto (MCZ). PERU Loreto: Estiron, Rio Am- 
piyacu (AMNH); Alto Amazonas, Pastaza (MCZ); 
Iquitos airport (FSCA); Jenaro Herrera (MUSM); Rio 
Putamayo (AMNH). Cajamarca: Jean (AMNH). 
Tumbes: Lechugal (PAN). Piura: San Lorenzo (MCZ); 
Guayaquil (CAS); Mallares, Rio Chira (CAS); Sullana 
(CAS). Lambayeque: pampa NW Oyotun (MCZ); 
UCA Yarina-Coche (IRSNB). Ucayali: Pucallpa 
(MUSM). Hudnuco: Higueras, Las Lomas (CAS); Tin- 
go Maria (AMNH); Monzon Valley, Tingo Maria 
(CAS). Ancash: Quillabamba (AMNH). BRAZIL 
Amazonas: Tefe (MCZ); Guajara, Rio Negro (AMNH); 
Lower Rio Negro (AMNH); Santo Antonio do I^a 
(MCN); Rio Xingu (MNRJ). Acre: mouth of Rio Em- 
bira, Rio Jurua (AMNH). Rondonia: Fazenda Rancho 
Grande, NE Cacaulandia (FSCA). Mato Grosso: Ba- 
rra do Tapirape (AMNH); Porto Velho, Rio Tapirape 
(AMNH); Juan Pinheiros, Rio Tapirape (AMNH). 
Mato Grosso do Sul: Corumba (AMNH). Minas Ger- 



ais: Lavras (MCZ); Minas de Serinha, Diamantina 
(AMNH); Uba (AMNH). Rio de Janeiro: Rio de Ja- 
neiro (MCZ). Sao Paulo: Botucatu (MZSP). Parana: 
Cataratas de Iguagu (MCZ); Ponta Grossa (AMNH). 
Santa Catarina: Blumenau (AMNH); Joa^aba (MZSP). 
Rio Grande do Sul: very common (MCN). URU- 
GUAY Maldonado: Punta del Este (MCZ). Colonia: 
Punta Gorda (CAS). PARAGUAY Central: Aregua 
(CAS); Villeta (MCZ); Asuncion (FSCA). Itapua: An- 
tidia Natianda (MCZ). BOLIVIA Pando: Abuna 
(MCZ). Beni: Chacobo Indian Village, Rio Benicito 
(AMNH); Estacion Biologica Beni, savanna, 50 km E 
San Borja (USNM); Espiritu, Yacuma (ZSM). AR- 
GENTINA Misiones: San Javier (MLP). Chaco: Selva 
del Rio de Oro (MEG). Formosa: Pto. Santos (MACN). 
Santiago del Estero: Santiago del Estero (MCZ). 

Metazygia benetla new species 
Figures 263-269; Map 3C 

Holotype. Female holotype, male paratype from near 
Cali, Valle, Colombia, ?1983 (W. Eberhard), in 
MCZ. The specific name is an arbitrary combina- 
tion of letters. 

Description. Female holotype. Cara- 
pace light orange, cephalic region darker 
orange. Chelicerae orange-brown. Labi- 
um, endites dark orange. Sternum light 
orange. Coxae, legs light orange. Dorsum 
of abdomen white (Fig. 266); venter light 
dusky. Posterior median eyes 0.8 diameter 
of anterior medians, anterior laterals 0.8 
diameter, posterior laterals 0.7. Anterior 
median eyes 0.7 diameter apart, 1.3 di- 
ameters from laterals. Posterior median 
eyes 0.2 diameter apart, 2.8 diameters from 
laterals. Height of clypeus equals 0.6 di- 
ameter of anterior median eye. Total length 
5.6 mm. Carapace 2.9 mm long, 2.3 wide, 
1.5 wide behind lateral eyes. First femur 
2.5 mm, patella and tibia 2.9, metatarsus 
2.1, tarsus 1.0. Second patella and tibia 2.7 
mm, third 1.3, fourth 2.3. 

Male paratype. Color as in female. Pos- 
terior median eyes 0.7 diameter of anterior 
medians, laterals 0.6 diameter. Anterior 
median eyes 0.7 diameter apart, 0.7 di- 
ameter from laterals. Posterior median eyes 
0.3 diameter apart, 1.7 diameters from lat- 
erals. Height of clypeus equals 0.4 diam- 
eter of anterior median eye. Fangs mod- 
ified (Fig. 269). Endite lacks tooth. Total 
length 5.0 mm. Carapace 2.5 mm long, 2.1 
wide, 1.0 behind lateral eyes 1.0 wide. First 



Metazygia 'Levi 



123 




Figures 253-262. Metazygia gregalis (O. P. -Cambridge). 253-257, female. 253-256, epigynum. 253, ventral. 254, subposterior. 
255, posterior. 256, lateral. 257, dorsal. 258-262, male. 258-260, left male palpus. 258, mesal. 259, ventral. 260, pulled apart. 
261, eye region, chelicerae, and right palpus. 262, fangs from below. 

Figures 263-269. M. benella n. sp. 263-266, female. 263-265, epigynum. 263, ventral. 264, posterior. 265, lateral. 266, dorsal. 
267-269, male. 267, 268, palpus. 267, mesal. 268, ventral. 269, eye region, chelicerae, and right palpus. 

Figures 270-276. M. yobena n. sp. 270-273, female. 270-272, epigynum. 270, ventral. 271 , posterior. 272, lateral. 273, dorsal. 
274-276, male. 274, 275, palpus. 274, mesal. 275, ventral. 276, fangs from below/. 

Abbreviations. C, conductor; E, embolus; M, median apophysis; R, radix. 



Scale lines. 1.0 mm, genitalia 0.1 mm. 



124 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



femur 2.1 mm, patella and tibia 2.5, meta- 
tarsus 2.0, tarsus 0.9. Second patella and 
tibia 2.3 mm, third 1.3, fourth 1.8. 

Note. Males and females were collected 
together. 

Variation. Most individuals have a fo- 
lium pattern on the abdomen as in Figure 

257. Total length of females 6.3 to 8.4 mm, 
males 4.2 to 5.1. Illustrations were made 
of the female holotype and male paratype 
collected with it. 

Diagnosis. The female epigynum (Fig. 
263) is longer than wide, lacks the posterior 
notch present in M. yobena (Fig. 270), and 
has a pair of lobes on the scape (at 10 hr 
and at 2 hr in Fig. 263). The male has a 
round tubercle on the tegulum (at 1 hr in 
Fig. 267), has a longer median apophysis 
(at 4 hr in Fig. 267) than M. gregalis (Fig. 

258, M in Fig. 260), and is without the 
black posterior wall (Fig. 267) present in 
M. yobena (at 4 hr in Fig. 274). 

Natural History. A male and female 
were collected in roadside shrubs at night 
near Cali, Colombia. 

Distribution. Panama and Colombia 
(Map 3C). 

Paratypes. COLOMBIA Valle: nr. Cali, 
1,000 m, no date, 22 (W. Eberhard 759, 
807, MCZ), 8 May 1973, 19 (W. Eberhard 
513, MCZ), Feb. 1975, 19 (W. Eberhard 
937, MCZ), 1973-1974, U (W. Eberhard, 
MCZ), 3 Mar. 1973, 19, 1<5 (H. Levi, W. 
Eberhard, MCZ); above Barrio Siloe, SW 
CaU, 3 July 1972, 16 (M. Corn, MCZ). 

Specimens Examined. PANAMA Panama: Barro 
Colorado Island, Lago Gatun, 25 Apr. 1946, 1$ (T. 
C. Schneirla, AMNH), July 1950, IS; 19 July 1954, 
13, Aug. 1954, 12, 14-18 Jan. 1958, 29, 6 Feb. 1958, 
19, 18 Feb, 1958, IS (A. M, Chickering, MCZ). 

Metazygia yobena new species 
Figures 270-276; Map 3C 

Holotype. Female holotype, one female, and two male 
paratypes from Mitu, 188 m, Depto. Vaupes, Co- 
lombia, at night in bamboo, 20 Apr. 1979 (M. Bar- 
reto), in MCZ. The specific name is an arbitrary 
combination of letters. 

Description. Female from Cuyabeno, 
Sucumbios Prov., Ecuador. Carapace 



brownish black, sides of thoracic region 
light orange. Chelicerae black. Labium, 
endites brown. Sternum yellowish, darker 
on each side. Coxae, legs dusty orange. 
Dorsum of abdomen with indistinct me- 
dian lighter band (Fig. 273); venter dark 
dusky. Posterior median eyes 0.9 diameter 
of anterior medians, laterals 0.7 diameter. 
Anterior median eyes 0.8 diameter apart, 
1.3 diameters from laterals. Posterior me- 
dian eyes 0.2 diameter apart, 2.5 diameters 
from laterals. Height of clypeus equals 0.7 
diameter of anterior median eye. Total 
length 6.0 mm. Carapace 3.1 mm long, 2.2 
wide, 1.3 wide behind lateral eyes. First 
femur 2.5 mm, patella and tibia 3.2, meta- 
tarsus 2.3, tarsus 1.1. Second patella and 
tibia 2.7 mm, third 1.6, fourth 2.3. 

Male paratype. Color as in female. Pos- 
terior median eyes 0.7 diameter of anterior 
medians, laterals 0.7 diameter. Anterior 
median eyes 0.7 diameter apart, 0.7 di- 
ameter from laterals. Posterior median eyes 
0.3 diameter apart, 1.9 diameters from lat- 
erals. Height of clypeus equals 0.6 diam- 
eter of anterior median eye. Fangs mod- 
ified (Fig. 276). Endite without tooth, pal- 
pal femur without facing tubercle. First 
coxa with small hook on its side. All legs 
with relatively long macrosetae. Total 
length 5.2 mm. Carapace 2.7 mm long, 2.1 
wide, 1.1 wide behind lateral eyes. First 
femur 2.4 mm, patella and tibia 2.9, meta- 
tarsus 2.1, tarsus 0.9. Second patella and 
tibia 2.4 mm, third 1.5, fourth 2.0. 

Note. Males and females were collected 
together. 

Variation. Total length of females 4.9 
to 7.5 mm, males 4.1 to 5.6. Most females 
have the folium pattern made up of pairs 
of brackets on the abdomen, which is typ- 
ical of the genus (Fig. 257). Some male 
palpi have only a small tubercle or none 
on the tegulum of the palpus (at 1 hr in 
Fig. 274). Illustrations were made from a 
specimen from the Cuyabeno Reserve, 
Sucumbios Prov., Ecuador, and from a pal- 
pus from a male collected in Depto. Vau- 
pes, Colombia. Figure 276 was made from 
a specimen from near Manaus, Brazil. 



Metazygia 'Levi 



125 



Diagnosis. The epigynum (Fig. 270) dif- 
fers from that of M. gregalis (Fig. 253) by 
being longer than wide and from that of 
M. benella (Fig. 263) by having a notch 
on the posterior margin (at 6 hr in Fig. 
270). The palpus differs from that of the 
two similar species by having a black wall 
on the hook-shaped median apophysis of 
the palpus (at 4 hr in Fig. 274). 

Natural History. Specimens came from 
bamboo in Colombia; from trees in a lake 
in the Cuyabeno Reserve, Ecuador; from 
trees in rain forest; from swamp plants in 
Peru; from forest savanna in Guyana; from 
falling into a canoe from overhanging veg- 
etation in Venezuela; and from cerrado 
shrub in Mato Grosso, Brazil. 

Distribution. Guyana and Amazon 
drainage (Map 3C). 

Paratypes. COLOMBIA Vaupes: Mitu, 
at night in bamboo, 20 Apr. 1971, 19, IS 
(M. Barreto, MCZ). 

Specimens Examined. VENEZUELA Amazonas: 
middle Rio Baria, 100 m (AMNH). GUYANA Ku- 
yuvvini River, from landing to Essequibo (AMNH); 
Upper Essequibo River (AMNH); Canje Ikuruwa Riv- 
er (AMNH); Kartabo (AMNH); Tumatumari 
(AMNH). FRENCH GUIANA Uassa [Uaga, Brazil] 
(PAN). COLOMBIA Meta: Finca Chenevo, 20 km S 
El Porvenir (MCZ); Hacienda Mozambique, 15 km 
SW Puerto Lopez (MCZ); Lomalinda, 3°18'N, 73°22'W 
(CAS); Carimagua, 175 m (MCZ), Amazonas: Arar- 
acuara (CV); Rio Pira, Apaporis, 0°25'S, 70°15'W 
(CAS). ECUADOR Napo: Coca, Rio Napo (L. Pena, 
MCZ). Sucumbios: Cuyabeno, common (MCZ, 
MECN). Pastaza: El Puyo, Rio Pastaza, 900 m (CAS). 
PERU Loreto: Explorama Lodge, 25 km NE Iquitos 
(FSCA); Estiron Rio Ampiacu (AMNH); Prov. Alto 
Amazonas, Pastaza (MCZ); Aquaitia (AMNH). Ama- 
zonas: Alto Rio Comainas, Puesto de Vigilancia (D. 
Silva D., MUSM). Hudnuco: Divisoria (AMNH); 
Monzon Valley, Tingo Maria (AMNH, CAS). Junin: 
Amable Maria (PAN). Cuzco: Chanchosmayo Valley 
(AMNH, CAS). Madre de Dios: Puerto Maldonado 
(AMNH); Zona Reserv. Tambopata, 12°50'S, 69°17'W 
(USNM); Zona Reserv. de Manu (MUSM); Alto Rio 
Madre de Dios (D. Silva D,, MUSM). BRAZIL Ama- 
zonas: Manaus, igapo Taruma-Mirim (INPA); Rio 
Autaz, Santa Amelia (NRMS). Pard: Belem (MCZ); 
Aldeia Ara9U, 20 km E Caninde (AMNH). Rondonia: 
Abuna (MCZ); Fazenda Rancho Grande, NE Cacau- 
landia (FSCA). Sao Paulo: Barueri (MZSP). BOLIVIA 
Beni: Espiritu, Yacuma (ZSM); Estacion Biol. Beni, 
14°47'S, 66°15'W (USNM); 19.5 km S Rurrenabaque 
(USNM). 



Metazygia voluptifica (Keyserling) 
Figures 277-284; Map 3D 

Epeira voluptifica Keyserling, 1892; 152, pi. 7, fig. 
112, 9, $. Female and male syntypes from Rio Gran- 
de [do Sul], Brazil, in BMNH, no. 1890.7. 1. 5041- 
5042, examined. 

Epeira mundula Keyserling, 1892: 179, pi. 9, fig. 132, 
2, not S. Female lectotype, here designated, from 
Rio Grande do Sul, Brazil, in BMNH, no. 
1890.7.1.5067, 5068, examined. NEW SYNONY- 
MY. 

Zilla punctata Keyserling, 1893: 305, pi. 15, fig. 225, 
2. Female holotype from Nova Friburgo, Brazil, 
lost. Not in BMNH, HECO, MCZ, NMW, USNM, 
ZMB. NEW SYNONYMY. 

Larinia mundula: — Simon, 1905: 10. Roewer, 1942: 
771. Bonnet, 1957: 2350. 

Aranea voluptifica: — Roewer, 1942: 856. 

Araneus voluptificus: — Bonnet, 1955: 631. 

Metazygia mundula: — Harrod, Levi, and Leiben- 
sperger, 1991: 246. 

Metazygia voluptifica: — Levi, 1991a: 180. 

Note. Keyserling described this species 
several times, first as Epeira voluptifica. 
Keyserling's female has a bracket folium 
as in M. gregalis (Fig. 257) and the epi- 
gynum lacks a scape. The type vial of 
Epeira mundula has a toothed, blue-bor- 
dered, 23-by-30-mm label of Keyserling 
reading, "Rio Grande do Sul, Epeira mun- 
dula Keys." The female is chosen as the 
lectotype because Keyserling's illustration 
of the female is recognizable while that of 
the male is not. The female lectotype has 
the scape of the epigynum torn off, as is 
that of most specimens. The male in the 
type vial is one that I associated with M. 
genialis. A second vial of E. mundula with 
a female syntype has a similar label, 20 by 
30 mm in size, and has also a different 
label with the number 1889.2.17. It also 
contains my typed label reading "2, 6 syn- 
types," added in 1974, when I examined 
the specimens and illustrated them. The 
type of Zilla punctata is lost, but the il- 
lustration of the epigynum matches this 
species. Keyserling gives the total length 
as 9.0 mm, larger than specimens I have 
examined. 

Description. Female from Guaiba, Rio 
Grande do Sul. Carapace light orange, ce- 
phalic area darker. Chelicerae brown. La- 



126 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



bium, endites orange. Sternum light or- 
ange. Coxae light orange; legs light orange, 
distal articles darker, dusky. Dorsum of 
abdomen with a pattern of paired lines or 
brackets (Fig. 282); venter with some white 
pigment behind epigynum. Posterior me- 
dian eyes 0.8 diameter of anterior medi- 
ans, laterals 0.7 diameter. Anterior median 
eyes 0.3 diameter apart, 1 diameter from 
laterals. Posterior median eyes 0.2 diam- 
eter apart, 1.6 diameters from laterals. 
Height of clypeus equals 0.5 diameter of 
anterior median eye. Total length 5.0 mm. 
Carapace 2.1 mm long, 1.6 wide, 0.9 wide 
behind lateral eyes. First femur 1.9 mm, 
patella and tibia 2.1, metatarsus 1.4, tarsus 
0.7. Second patella and tibia 1.9 mm, third 
1.1, fourth 1.6. 

Male from Santa Vitoria do Palmar. Col- 
or as in female, but less gray pigment on 
abdomen. Posterior median eyes same di- 
ameter as anterior medians, laterals 0.8 di- 
ameter. Anterior median eyes 0.7 their di- 
ameter apart, 0.7 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
1.3 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Second tibia thinner than first. 
Total length 4.5 mm. Carapace 2.1 mm 
long, 1.6 wide, 0.9 wide behind lateral eyes. 
First femur 2.2 mm, patella and tibia 2.7, 
metatarsus 2.0, tarsus 0.8. Second patella 
and tibia 2.4 mm, third 1.2, fourth 1.5. 

Note. Males and females were matched 
because collections came from the same 
locality (Minas Serinha, Minas Gerais) and 
because the carapace was of similar length. 
The female from Mato Grosso was col- 
lected with a male M. corumba; the E. 
mundula lectotype was with a male M. 
genialis. However, Keyserling described 
the female of E. voluptifica together with 
a male that I consider here the correct 
match. 

Variation. Most females have the scape 
torn off (Figs. 280, 281). Total length of 
females 4.8 to 7.3 mm, males 4.0 to 5.5. 
Illustrations of females were made from 
specimens from Guaiba, Rio Grande do 
Sul; the male from Santa Vitoria do Pal- 
mar, Rio Grande do Sul. 



Diagnosis. The female can be separated 
from others by the two almost round cir- 
cles of the epigynum in ventral view (Figs. 
277, 280). The female is distinguished from 
that of M. viriosa by the posterior median 
plate of the epigynum, which is slightly 
wider than each lateral plate (Figs. 278, 
281). The male is separated from others 
by the black fold of the tegulum of the 
palpus, which is distally serrate (T at 12 
hr in Fig. 283). 

Natural History. A female was collected 
from grass and brush along a fence in Co- 
lombia. 

Distribution. Colombia to Argentina 
(Map 3D). 

Specimens Examined. COLOMBIA Meta: Cari- 
magua, Oct. 1973, 19 (W. Eberhard, MCZ). PERU 
Madre de Dios: Alto Rio Madre de Dios, Playa Ma- 
ronal campsite, 24 Sept. 1987, 1<5 (D. Silva D., MUSM). 
BRAZIL Minas Gerais: Diamantina, Minas de Se- 
rrinha, 1945, 99, 13 (E. Cohn, AMNH). Mato Grosso 
do Sul: Corumba, 28-29 May 1960, 19 (B. Malkin, 
AMNH). Rio Grande do Sul: Encantado, 24 May 
1986, 15 (A, D. Brescovit, MCN 15125); Santa Vitoria 
do Palmar, Esta9ao Ecologica do Taim, 26 Nov. 1985, 
19 (M. Rosenau, MCN 14050), 9 Apr. 1986, IS (M. 
Rosenau, MCN 14824); Guaiba, Granja Carola, 23 
July 1986, 159 (M. A. L. Marques, MCN 15419); 
Triunfo 25 Jan, 1990, IS (A. B. Bonaldo, MCN 19387); 
Porto Alegre, Vila Assungao, 27 July 1988, 19 (R. 
Richter, MCN 18003). ARGENTINA Chaco: Resis- 
tencia, 29 (MACN). Corrientes: Corrientes, 19 (Z. von 
Beneden, ZMK). Entre Rios: Salto Grande, NE Con- 
cordia, Mar. 1964, 15 (M. E. Galiano, MEG). 

Metazygia viriosa (Keyserling), 
new combination 
Figures 285-288; IVIap 3A 

Epeira viriosa Keyserling, 1892: 165, pi. 8, fig. 122, 
9. Female holotype from Rio Grande do Sul, Brazil, 
in BMNH no. 1890.7.1.506, examined. 

Aranea viriosa: — Roewer, 1942: 856. 

Araneus viriosus: — Bonnet, 1955: 630. 

Description. Female holotype. Cara- 
pace orange-brown, cephalic region 
darkest. Chelicerae, labium, endites brown. 
Sternum dusky orange. Legs light orange. 
Dorsum of abdomen with anterior trans- 
verse black band (Fig. 288). Venter with- 
out marks or pigment. Posterior median 
eyes same diameter as anterior medians, 
laterals 0.8 diameter. Anterior median eyes 



METAZYGIA'Levi 



111 



277 



280 




292 



294 



."JD^ 





297 





295 



Figures 277-284. Metazygia voluptifica (Keyserling). 277-282, female. 277-281 , epigynum. 277, ventral. 278, posterior. 279, 
lateral. 280, ventral, scape torn off. 281 , posterior, scape torn off. 282, dorsal. 283, 284, left male palpus. 283, mesal. 284, 
ventral. 

Figures 285-288. M. viriosa (Keyserling), female. 285-287, epigynum. 285, ventral. 286, posterior. 287, lateral. 288, dorsal. 

Figures 289-291 . M. ituari n. sp., female. 289, 290, epigynum. 289, ventral. 290, posterior. 291 , dorsal. 

Figures 292-294. M. limonaln. sp., female. 292, 293, epigynum. 292, ventral. 293, posterior. 294, dorsal. 

Figures 295-297. M. tanica n. sp., male. 295, 296, left palpus. 295, mesal. 296, ventral. 297, dorsal. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



0.7 diameter apart, 0.7 diameter from lat- 
erals. Posterior median eyes 0.3 diameter 
apart, 1.5 diameters from laterals. Height 
of clypeus equals 0.7 diameter of anterior 
median eye. Abdomen spherical (Fig. 288). 
Total length 4.8 mm. Carapace 1.7 mm 
long, 1.3 wide. First femur 1.7 mm, patella 
and tibia 2.1, metatarsus 1.4, tarsus 0.6. 



Second patella and tibia 1.8 mm, third 1.1, 
fourth 1.7. 

Variation. All females had the scape of 
the epigynum torn off. Total length of fe- 
males 4.6 to 5.4 mm. Illustrations were 
made from the female holotype; Figure 
287 was made from a female from Juru- 
batuba. 



128 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Diagnosis. Metazygia viriosa differs 
from M. voluptifica by the markings of 
the abdomen (Fig. 288) and by the median 
posterior depression of the epigynum (Fig. 
286). 

Specimens Examined. BRAZIL Sao Paulo: Cocaia, 
Sept, 1950, 19 (H. Urban, MZSP 9662); Jurubatuba, 
6 July 1941, 2S (P. F. S. Pereira, MZSP 9619). Santa 
Catarina: Pinhal, Jan. 1948, Dec. 1948, 62 (A. Mailer, 
AMNH). 

Metazygia ituari new species 
Figures 289-291 ; Map 3D 

Holotype. Female holotype from Utiarity (Utiariti), 
Est. Mato Grosso, Brazil, 1961 (H. Lenko) in MZSP 
no. 4155. The specific name is an arbitrary com- 
bination of letters. 

Description. Female holotype. Cepha- 
lothorax light orange, only eyes with some 
black pigment. Dorsum of abdomen with 
three longitudinal white pigment bands 
(Fig. 291); venter with white pigment spots. 
Posterior median eyes 0.8 diameter of an- 
terior medians, laterals 0.8 diameter. An- 
terior median eyes 0.6 diameter apart, 0.7 
diameter from laterals. Posterior median 
eyes 0.3 diameter apart, 1.8 diameters from 
laterals. Ocular quadrangle narrower be- 
hind than in front. Height of clypeus equals 
0.4 diameter of anterior median eye. Ab- 
domen (Fig. 291). Total length 3.1 mm. 
Carapace 1.3 mm long, 1.1 wide, 0.6 wide 
behind lateral eyes. First femur 1.1 mm, 
patella and tibia 1.2, metatarsus 0.7, tarsus 
0.5. Second patella and tibia 1.1 mm, third 
0.7, fourth 0.9. 

Diagnosis. The female, which appar- 
ently has lost the scape of her epigynum, 
is separated from M. voluptifica and M. 
viriosa by the short, wide posterior median 
plate of the epigynum (Fig. 290). 

Metazygia limonal new species 
Figures 292-294; Map 4A 

Holotype. Female holotype from El Limonal, Alto 
Rio Madre de Dios, Depto. Madre de Dios, night 
collecting, 21 June 1988 (P. Lozada), in MUSM. 
The specific name is a noun in apposition after the 
type locality. 

Description. Female holotype. Cara- 
pace very light orange, black only between 



eyes. Chelicerae, labium, endites light or- 
ange. Sternum light orange. Legs light or- 
ange. Dorsum of abdomen white with an- 
terior dumbbell-shaped black marks (Fig. 
294); venter with a white square. Posterior 
median eyes same diameter as anterior 
medians, anterior laterals 0.7 diameter, 
posterior 0.6. Anterior median eyes 0.7 di- 
ameter apart, 0.3 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
1 diameter from laterals. Ocular quadran- 
gle narrower behind than in front. Height 
of clypeus equals 0.4 diameter of anterior 
median eye. Total length 4.2 mm. Cara- 
pace 1.7 mm long, 1.3 wide, 0.8 behind 
lateral eyes. First femur 1.9 mm, patella 
and tibia 2.4, metatarsus 1.7, tarsus 0.7. 
Second patella and tibia 2.1 mm, third 1.2, 
fourth 1.8. 

Variation. All females examined had the 
scape of the epigynum torn off (Figs. 292, 
293). Total length of females 3.0 to 4.2 
mm. Illustrations were made from the fe- 
male holotype. 

Diagnosis. Metazygia limonal differs 
from others by the Y-shaped posterior me- 
dian plate of the epigynum (Fig. 293). 

Distribution. From Depto. Madre de 
Dios, Peru, to northern Argentina (Map 
4A). 

Specimens Examined. BRAZIL Rio Grande do Sul: 
Santa Rosa, May 1955, 22 (C. Biezanko, AMNH). 
ARGENTINA Misiones: Puerto Rico, Dec. 1943, 12 

(MACN). 

Metazygia tanica new species 
Figures 295-297; Map 4A 

Holotype. Male holotype from Botanical Gardens, 
Georgetown, Guyana, 22 Feb. 1959 (A. Nadler), in 
AMNH. The specific name is an arbitrary combi- 
nation of letters. 

Description. Male holotype. Cephalo- 
thorax orange, sternum lighter. Dorsum of 
abdomen light with faint dusky outline of 
fohum (Fig. 297); venter light. Posterior 
median eyes 0.8 diameter of anterior me- 
dians, laterals 0.7 diameter. Anterior me- 
dian eyes 0.6 diameter apart, 0.5 diameter 
from laterals. Posterior median eyes 0.2 
diameter apart, 1.3 diameters from later- 



Metazygia • Levi 



129 



als. Height of clypeus equals 0.8 diameter 
of anterior median eye. Endite with mi- 
nute tooth. Total length 4.2 mm. Carapace 
2.5 mm long, 1.9 wide, 0.9 behind lateral 
eyes. First femur 2.3 mm, patella and tibia 
2.7, metatarsus 2.0, tarsus 0.9. Second pa- 
tella and tibia 2.4 mm, third 1.5, fourth 
2.0. 

Diagnosis. This male differs from M. 
voluptifica by having a structure (embolus 
or embolus lamella) of the palpus in a di- 
agonal position with its sides almost par- 
allel (Fig. 295). 

Metazygia vaupes new species 
Figures 298-302; Map 4A 

Holotype. Female holotype from Mitu, Depto. Vau- 
pes, 200 m, Colombia, Feb. 1975 (P. A. Schneble), 
in MCZ. The specific name is a noun in apposition 
after the type locality. 

Description. Female holotype. Cara- 
pace yellowish. Chelicerae, labium, en- 
dites yellowish. Sternum yellowish. Legs 
yellowish. Dorsum of abdomen without 
color but with white pigment spots around 
anterior and sides (Fig. 301); venter with 
white pigment spots. Posterior median eyes 
1.2 diameters of anterior medians, laterals 
0.8 diameter. Anterior median eyes 0.8 di- 
ameter apart, 0.3 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
0.8 diameter from laterals. Ocular quad- 
rangle square. Height of clypeus equals 0.3 
diameter of anterior median eye. Total 
length 3.1 mm. Carapace 1.17 mm long, 
1.04 wide, 0.54 behind lateral eyes. First 
femur 1.52 mm, patella and tibia 1.82, 
metatarsus 1.35, tarsus 0.48. Second patella 
and tibia 1.53 mm, third 0.87, fourth 1.26. 

Male from Depto. Loreto, Peru. Color 
as in female but anterior white band of 
abdomen less distinct. Posterior median 
eyes 0.9 diameter of anterior medians, lat- 
erals 0.7 diameter. Anterior median eyes 
0.7 diameter apart, 0.2 diameter from lat- 
erals. Posterior median eyes 0.2 diameter 
apart, 1 diameter from laterals. Ocular 
quadrangle narrower behind than in front. 
Height of clypeus equals 0.6 diameter of 
anterior median eve. First coxa with small 



hook. Second tibia thicker than first, with 
one long macroseta and several short ones 
distally, all in one line. Total length 2.0 
mm. Carapace 1.2 mm long, 0.9 wide, 0.4 
wide behind lateral eyes. First femur 1.4 
mm, patella and tibia 1.7, metatarsus 1.2, 
tarsus 0.5. Second patella and tibia 1.3 mm, 
third 0.7, fourth 1.0. 

Note. Males and females were collected 
in the same location in Rondonia, and both 
lack dark pigment dorsally on the abdo- 
men. 

Variation. Total length of females 3.1 
to 3.2 mm. Illustrations were made from 
the female holotype and a male from Ron- 
donia. 

Diagnosis. The abdomen appears heart- 
shaped (Fig. 301). The epigynum is very 
distinct with a short rounded scape (Figs. 
298, 299). The male has a shorter, smaller 
embolus lamella (Fig. 302) than M. cas- 
taneoscutata (Fig. 308). 

Natural History. Specimens have been 
collected in grass and shrubs in Loreto, 
Peru, and rain forest in Rondonia, Brazil. 

Distribution. Amazon region (Map 4A). 

Specimens Examined. PERU Loreto: Rio Manatee, 
18 July 1989, 13 (G. B. Edwards, FSCA). Hudnuco: 
Monzon Valley, Tingo Maria, 23 Sept. 1954, IS (E. 
I. Schlinger, E. S. Ross, CAS). BRAZIL Rondonia: 
Fazenda Rancho Grande, NE Cacaulandia, 6-15 Dec. 
1990, 19, 13 (G. B. Edwards, J, E. Eger, FSCA). 

Metazygia castaneoscutata (Simon) 
Figures 303-308; Map 4B 

Araneus castaneoscutatus Simon, 1895: 806. Female 
holotype from Amazonas [specimen labeled as 
coming from Iquitos to Pebas, Peru], in MNHN, 
examined. Bonnet, 1955: 452. 

Aranea castaneoscutata: — Roewer, 1942: 838. 

Metazygia castaneoscutata: — Levi, 1991a: 177. 

Description. Female holotype. Cara- 
pace light orange with a median black 
band. Chelicerae black. Labium, endites, 
sternum light orange. Legs dusky orange. 
Dorsum of abdomen with black band 
around anterior, continuing around sides 
of abdomen and meeting black ring around 
spinnerets (Figs. 306, 307); a narrow band 
of white pigment spots parallel to band on 
dorsum and a median dorsal dusky patch. 



130 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Venter with a dusky T-shaped mark and 
a black ring around spinnerets (Fig. 307). 
Posterior median eyes 0.7 diameter of an- 
terior medians, laterals 0.5 diameter. An- 
terior median eyes 0.3 diameter apart, 0.2 
diameter from laterals. Posterior median 
eyes 0.2 diameter apart, 0.8 diameter from 
laterals. Height of clypeus equals 0.3 di- 
ameter of anterior median eye. Total length 
3.4 mm. Carapace 1.2 mm long, 1.0 wide, 
0.5 wide behind lateral eyes. First femur 
1.3 mm, patella and tibia 1.5, metatarsus 
1.0, tarsus 0.4. Second patella and tibia 1.3 
mm, third 0.7, fourth 1.1. 

Male from Alto Rio Comaina. Color as 
in female. Posterior median eyes 0.9 di- 
ameter of anterior medians, laterals 0.6 
diameter. Anterior median eyes 0.2 di- 
ameter apart, almost touching laterals. 
Posterior median eyes 0.2 diameter apart, 
0.8 diameter from laterals. Height of clyp- 
eus equals 0.3 diameter of anterior median 
eye. Fourth coxa with a tiny macroseta on 
left side only. Second tibia thicker than 
first, both first and second with macrose- 
tae. Total length 2.1 mm. Carapace 1.2 
mm long, 0.9 wide, 0.5 behind lateral eyes. 
First femur 1.1 mm, patella and tibia 1.3, 
metatarsus 0.8, tarsus 0.4. Second patella 
and tibia 1.1 mm, third 0.6, fourth 0.8. 

Note. Males and females were matched 
because of similar color and markings and 
because they were collected together. 

Variation. The scape of the epigynum 
may be broken off (Fig. 304). Total length 
of females 3.1 to 4.5 mm, males 2.1 to 2.3. 
Illustrations were made from a female from 
the Tambopata Reservation and a male 
from Alto Rio Comaina. 

Diagnosis. Metazygia castaneoscutata 
can be separated from others by the spher- 
ical abdomen (Fig. 306) and its coloration. 
The epigynum has a longer scape (Fig. 



303) than M. vaupes (Fig. 298) and a dif- 
ferently shaped posterior median plate 
(Fig. 305). The male has a longer embolus 
lamella (at 10 hr in Fig. 308) and more 
distinct conductor (center of Fig. 308) than 
M. vaupes (Fig. 302). 

Natural History. Specimens came from 
forest interior near Manaus. 

Distribution. Amazon region (Map 4B). 

Specimens Examined. PERU Amazonas: Alto Rio 
Comaina, Puesto da Vigilancia 22, Falso Paquisha, 
850-1,150 m, Cordillera del Condor, 24 Oct. 1987, 
IS, 28 Oct. 1987, 19, 1,5 (D. Silva D., MUSM). Madre 
de Dios: Zona Reservada Tambopata, trocha princi- 
pal, 290 m, 12°50'S, 69°17'W, 13-29 May 1988, 22 
(D. Silva D., MUSM). BRAZIL Am,azonas: Reserva 
Ducke, Manaus, 26 Mar. 1974, 19 (L. P. Albuquerque, 
MCN 20047); Reserva Campina, 22 Jan. 1973, 19 
(MCN 20053); Reserva Cabo Frio, 80 km from Ma- 
naus, 23 Jan. 1991, 19; Colosso Reserve, 80 km from 
Manaus, 19 Mar. 1991, U, 19, 4 June 1991, 19; Reserva 
Dimona, 80 km from Manaus, 26, 27 Mar. 1991, 3$, 
14, 15 May 1991, 29; km 41 Reserve, 80 km from 
Manaus, 17 Apr. 1991, 19 (all H. Fowler, E. Ventic- 
inque, R. S. Vieira, MCZ). Mato Grosso: Sinop, Oct. 
1975, 19 (M. Alvarenga, AMNH); Jacare, Xingu, Nov. 
1961, 16 (Werner, AMNH). 

Metazygia octama new species 
Figures 309-313; Map 4B 

Holotype. Female holotype from near Cali, 1,000 m 
elev., Depto. Valle, Colombia (W. Eberhard, no. 
821), in MCZ. The specific name is an arbitrary 
combination of letters. 

Description. Female holotype. Cara- 
pace orange, eye region black, thoracic re- 
gion lightest. Chelicerae, labium, endites 
orange. Sternum orange. Legs orange. Ab- 
domen white with a black band around 
anterior (Fig. 312), venter with a white 
square between epigynum and spinnerets. 
Posterior median eyes 0.9 diameter of an- 
terior medians, laterals 0.5 diameter. An- 
terior median eyes 0.6 diameter apart, 0.4 
diameter from laterals. Posterior median 



Figures 298-302. Metazygia vaupes n. sp. 298-301, female. 298-300, epigynum. 298, ventral. 299, posterior. 300, lateral. 
301 , dorsal. 302, left male palpus. 



Figures 303-308. M. castaneoscutata (Simon). 303-307, female. 303-305, epigynum. 303, ventral. 304, ventral, scape torn. 
305, posterior. 306, dorsal. 307, abdomen, ventral. 308, male palpus. 



METAZYGlA'Levi 131 




Figures 309-313. M. octama n. sp. 309-312, female. 309-312, epigynum. 309, ventral. 310, posterior. 31 1 , lateral. 312, dorsal. 
313, male palpus. 

Figures 314-317. M. floresta n. sp., female. 314-316, epigynum. 314, ventral. 315, posterior. 316, lateral. 317, dorsal. 

Figures 31 8-321 . M. mariahelenae n. sp., male. 31 8, 31 9, palpus. 31 8, mesal. 31 9, ventral. 320, dorsal. 321 , abdomen, ventral. 

Scale lines. 1 .0 mm, genitalia 0.1 mm. 



132 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



eyes 0.3 diameter apart, 1 diameter from 
laterals. Height of clypeus equals 0.5 di- 
ameter of anterior median eye. Total length 
5.5 mm. Carapace 2.1 mm long, 1.6 wide, 
0.8 behind lateral eyes. First femur 2.1 
mm, patella and tibia 2.5, metatarsus 2.0, 
tarsus 0.7. Second patella and tibia 2.3 mm, 
third 1.4, fourth 2.0. 

Male from type locality. Color as in fe- 
male. Carapace with slight lobe. Posterior 
median eyes same diameter as anterior 
medians, anterior laterals 0.8 diameter. 
Anterior median eyes 0.8 diameter apart, 
0.3 diameter from laterals. Posterior me- 
dian eyes 0.2 diameter apart, 1 . 1 diameters 
from laterals. Height of clypeus equals 0.6 
diameter of anterior median eye. Third 
and fourth coxa each with one long mac- 
roseta. Second tibia thicker than first, with 
several large macrosetae. Total length 3.7 
mm. Carapace 1.6 mm long, 1.3 wide, 0.5 
wide behind lateral eyes. First femur 1.7 
mm, patella and tibia 2.2, metatarsus 1.7, 
tarsus 0.6. Second patella and tibia 1.5 mm, 
third 1.0, fourth 1.5. 

Note. Males and females were collected 
together. 

Variation. Total length of females 4.1 
to 5.1 mm, males 3.1 to 3.7. Illustrations 
were made from the female holotype and 
a male paratype. 

Diagnosis. Metazygia octama females 
have a thicker epigynal scape (Figs. 309, 
310) than M. vaupes (Fig. 298) and M. 
castaneoscutata (Fig. 303). In males the 
palpus has a hair-like, S-shaped embolus 
that lies between the transparent lamella 
and conductor (between center and at 11 
hr in Fig. 313). 

Natural History. A female and imma- 
tures were collected at night on a roadside 
shrub near Cali, Colombia. When living 
they were dark green with reddish first 
pair of legs. 

Distribution. From Panama to Depto. 
Madre de Dios, Peru (Map 4B). 

Paratypes. COLOMBIA Valle: nr. Cali, 
3 Mar. 1973, 2 imm., 19 (H. Levi, W. Eber- 
hard, MCZ), 1973-1974, 12, 5<5, 1976, 1<3, 
1977, 16, 1983, SS (W. Eberhard, MCZ), 



no dates, 39, 1<5 (W. Eberhard no. 660, 667, 
956, MCZ). 

Specimens Examined. PANAMA Panama: Forest 
Reserve, Aug. 1936, U (A. M. Chickering, MCZ). 
PERU Madre de Dios: 15 km E Puerto Maldonado, 
26 Feb. 1989, 19 (D. Silva D., MUSM). 

Metazygia floresta new species 
Figures 314-317; Map 4B 

Holotype. Female holotype from Floresta dos Ma- 
cacos, Est. Guanabara [Est. Rio de Janeiro], Brazil, 
Feb. 1961 (M. Alvarenga), in AMNH. The specific 
name is a noun in apposition after the type locality. 

Description. Female holotype. Cara- 
pace orange, eye region black. Chelicerae 
brown. Labium, endites dusky orange. 
Sternum orange. Legs orange. Dorsum of 
abdomen white with a dark band around 
the anterior (Fig. 317). Venter with a large 
white square between epigynum and spin- 
nerets. Eyes large. Posterior median eyes 
0.8 diameter of anterior medians, laterals 
0.6 diameter. Anterior median eyes 0.7 di- 
ameter apart, 0.6 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
1 diameter from laterals. Height of clypeus 
equals 0.3 diameter of anterior median eye. 
Total length 3.8 mm. Carapace 1.5 mm 
long, 1.0 wide, 0.5 behind lateral eyes. First 
femur 1.6 mm, patella and tibia 1.9, meta- 
tarsus 1.3, tarsus 0.5. Second patella and 
tibia 1.6 mm, third 0.9, fourth 1.3. 

Diagnosis. Abdomen subspherical, flat- 
ter anteriorly, and slightly pointed poste- 
riorly (Fig. 317). This female differs from 
all others in the smooth, sclerotized scape 
(Figs. 314-316) with fused plates in pos- 
terior view (Fig. 315). 

Specimens Examined. BRAZIL Rio de Janeiro: 
Nova Igua^u, Miguel Couto, July 1961, 19 (M. Al- 
varenga, AMNH). 

Metazygia mariahetenae new species 
Figures 318-321 ; Map 4C 

Holotype. Male holotype from Reserva Ducke, Ma- 
naus, Est. Amazonas, Brazil, Aug. 1971 (M. E. Ga- 
liano), in MACN. The species is named after the 
collector. 

Description. Male holotype. Carapace 
light orange, eye region dusky. Chelicerae 



Metazygm 'Levi 



133 



dusky. Labium, endites dusky. Sternum 
dusky light orange. Legs light orange, dis- 
tally dusky. Dorsum of abdomen white 
with black band around anterior (Fig. 320). 
Venter with white longitudinal band on 
each side of genital furrow (Fig. 321). Pos- 
terior median eyes 0.8 diameter of anterior 
medians, anterior laterals 0.7 diameter, 
posterior 0.5. Anterior median eyes their 
diameter apart, 0.6 diameter from laterals. 
Posterior median eyes 0.4 diameter apart, 
their diameter from laterals. Ocular quad- 
rangle narrower behind than in front. 
Height of clypeus equals 0.6 diameter of 
anterior median eye. Fourth coxa with 
short macroseta. Second tibia thicker than 
first, with three to four strong macrosetae 
on distal quarter. Total length 2.8 mm. 
Carapace 1.4 mm long, 1.1 wide, 0.5 be- 
hind lateral eyes. First femur 1.5 mm, pa- 
tella and tibia 2.0, metatarsus 1.5, tarsus 
0.5. Second patella and tibia 1.4 mm, third 
0.9, fourth 1.3. 

Diagnosis. The male is distinguished by 
an embolus that points clockwise (center 
of Fig. 318) and a median apophysis that 
is rectangular in ventral view (at 3 hr in 
Fig. 318). 

Metazygia nigrocincta (F. P.-Cambridge), 
new combination 
Figures 322-327; Map 4G 

Aranea nigrocincta F. P.-Cambridge, 1904: 513, pi. 
49, figs. 11, 12, 9, S. Female and male syntypes 
from Bugaba, Panama, in BMNH, lost. Roewer, 
1942: 848. 

Araneus nigrocinctus: — Bonnet, 1955: 550. 

The types are lost. F. P. -Cambridge's 
illustration fits this species. 

Description. Female from Fortin de los 
Flores, Veracruz, Mexico. Carapace or- 
ange, eye region black. Chelicerae dark 
brown. Labium, endites dark dusky or- 
ange. Sternum dusky orange. Coxae or- 
ange; legs dark dusky on orange. Dorsum 
of abdomen white with a black band across 
anterior (Fig. 525); venter with a white 
square (Fig. 326). Posterior median eyes 
0.8 diameter of anterior medians, laterals 



0.6 diameter. Anterior median eyes 0.7 di- 
ameter apart, 0.7 diameter from laterals. 
Posterior median eyes 0.3 diameter apart, 
1.6 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Total length 4.0 mm. Cara- 
pace 1.9 mm long, 1.4 wide, 0.7 behind 
lateral eyes. First femur 1.8 mm, patella 
and tibia 2.3, metatarsus 1.5, tarsus 0.6. 
Second patella and tibia 2.0 mm, third 1.2, 
fourth 1.7. 

Male from Veracruz, Mexico. Color as 
in female. Carapace with small lobes. Pos- 
terior median eyes same diameter as an- 
terior medians, laterals 0.8 diameter. An- 
terior median eyes 0.7 diameter apart, 0.7 
diameter from laterals. Posterior median 
eyes 0.2 diameter apart, 1.2 diameters from 
laterals. Height of clypeus equals 0.5 di- 
ameter of anterior median eye. First coxa 
with large hook, third and fourth each with 
a short macroseta. Total length 2.7 mm. 
Carapace 1.4 mm long, 1.2 wide, 0.6 be- 
hind lateral eyes. First femur 1.6 mm, pa- 
tella and tibia 1.8, metatarsus 1.2, tarsus 
0.5. Second patella and tibia 1.5 mm, third 
0.9, fourth 1.3. 

Note. Males and females were collected 
together. 

Variation. Total length of females 3.8 
to 4.7 mm, males 2.5 to 2.7. Some males 
have macrosetae on the fourth coxae only, 
not on the third. Specimens from Jalisco, 
Mexico, have two white spots on the un- 
derside of the abdomen. Illustrations were 
made from specimens from Veracruz, 
Mexico. 

Diagnosis. The female's abdomen is 
subspherical, widest anteriorly (Fig. 325). 
Females of this species are distinguished 
from those of M. lagiana (Figs. 328, 329) 
by having a thin, straight epigynal scape 
(Fig. 322) and a Y-shaped posterior me- 
dian plate with openings at the end of the 
arms (Figs. 323, 328, 329). The male's pal- 
pus has an undulating median apophysis 
(Fig. 327) that is similar to that of M. cas- 
taneoscutata (Fig. 308) but is narrower at 
its base. The embolus lamella of this spe- 
cies (at 11 hr in Fig. 327) does not extend 



134 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



beyond the cymbium tip as it does in M. 
castaneoscutata. 

Distribution. Mexico to Panama (Map 

4G). 

Specimens Examined. MEXICO San Luis Potosi: 
Tamanzunchale, 27 Sept. 1939, 12 (C. M. Bogert, H. 
E. Yokes, AMNH). Jalisco: 16.3 km NE La Huerta, 
6 Aug. 1967, 12 (R. E. Leech, REL); Esta. Biol. Cha- 
mela, 100 m, Sept. 1988, 22, 3<5, Sept. 1990, 12 (W. 
Eberhard, MCZ). Veracruz: Fortin de las Flores, 
18°53'N, 96°53'W, 25 Apr. 1963, 12 (W. Gertsch, W. 
Ivie, AMNH); Tlapacoyan, 300 m, 7, 8 July 1946, 2$ 
(H. Wagner, AMNH); Los Tuxtlas Biol. Sta., 9-29 
July 1990, 12 (B. Traw, MCZ); Canyon of Rio Metlac, 
nr. El Fortin [?], 17 Dec. 1948, 12 (H. B. Leech, CAS). 
HONDURAS Tela, 1-17 Apr., 1<5 (F. Dybas, AMNH). 
PANAMA Chiriqui: Boquete, July 1939, 12 (A. M. 
Chickering, MCZ). 



Metazygia lagiana new species 
Figures 328-332; Map 4C 

Holotijpe. Female holotype from Cataratas de Igua- 
9u, Misiones Prov., Argentina, 5 Oct. 1963 (M. E. 
Galiano), in MACN. The species name is an arbi- 
trary combination of letters. 

Description. Female. Carapace shiny 
with head black grading into the orange 
sides of the thoracic region. Chelicerae, 
labium, endites dark brown. Sternum 
black. Coxae yellow; legs yellow with black 
ring around end of first tibiae. Dorsum of 
abdomen white, with black band around 
anterior and black spots posteriorly (Fig. 
330); venter with black median band from 
pedicel to and enclosing spinnerets (Fig. 
331). Posterior median eyes 1 diameter of 
anterior medians, anterior laterals 1 di- 
ameter, posterior 0.8. Anterior median eyes 
their diameter apart, 0.7 from laterals. 
Posterior median eyes 0.5 their diameter 
apart, 1.7 from laterals. Height of clypeus 
equals 0.5 diameter of anterior median eye. 
Total length 4.0 mm. Carapace 1.5 mm 



long, 1.3 wide, 0.7 behind lateral eyes. First 
femur 1.5 mm, patella and tibia 1.9, meta- 
tarsus 1.2, tarsus 0.6. Second patella and 
tibia 1.6 mm, third 1.0, fourth 1.4. 

Male paratype. Color differs from that 
of female: carapace orange with eye re- 
gion black, sternum orange. Abdomen with 
anterior black band, venter with a wider 
than long white rectangle. Posterior me- 
dian eyes 0.8 diameter of anterior medi- 
ans, anterior laterals 0.5 diameter, poste- 
rior laterals 0.4. Anterior median eyes 0.8 
diameter apart, 0.8 from laterals. Posterior 
median eyes 0.3 diameter apart, slightly 
more than one from laterals. First coxae 
with large hook on venter, third and fourth 
each with a macroseta. Second tibiae 
thicker than first with long macroseta al- 
most in middle on venter. Total length 2.9 
mm. Carapace 1.7 mm long, 1.4 wide. First 
femur 1.8 mm, patella and tibia 2.2, meta- 
tarsus 1.6, tarsus 0.6. Second patella and 
tibia 1.5 mm, third 0.9, fourth 1.4. 

Note. Male and female were collected 
at the same locality. 

Variation. Total length of females 3.7 
to 4.0 mm. 

Diagnosis. The female differs from that 
of M. nigrocincta by having an epigynal 
scape that is thin, bent, and transparent 
(Fig. 328) and a posterior median plate 
upside-down heart-shaped (Fig. 329). The 
male palpus features a knob-shaped me- 
dian apophysis (at 4 hr in Fig. 332). 

Natural History. A female has been col- 
lected in cerrado scrub in Mato Grosso, 
Brazil. 

Distribution. Depto. Madre de Dios, 
Peru, to northern Argentina (Map 4C). 

Paratypes. ARGENTINA Misiones: 
Gral. Belgrano, Jan. 1966, 19; Dec. 1972, 
IS (M. E. Galiano, MACN). 



Figures 322-327. Metazygia nigrocincta (F. P.-Cambridge). 322-326, female. 322-324, epigynum. 322, ventral. 323, posterior. 
324, lateral. 325, dorsal. 326, atxJomen, ventral. 327, left male palpus. 

Figures 328-332. M. lagiana n. sp. 328-331, female. 328, 329, epigynum. 328, ventral. 329, posterior. 330, dorsal. 331, 
atxlomen, ventral. 332, male palpus. 



Figures 333-336. M. carimagua n. sp., female. 333-335, epigynum. 333, ventral. 334, posterior. 335, lateral. 336, dorsal. 



M £ TAZYGIA • Levi 1 35 




Figures 337-341. M. toque n. sp., female. 337-339, epigynum. 337, ventral. 338, posterior. 339, lateral. 340, dorsal. 341, 
abdomen, ventral. 

Figures 342-345. M. cienaga n. sp., female. 342-344, epigynum. 342, ventral. 343, posterior. 344, lateral. 345, dorsal. 

Figures 346-350. M. souza n. sp., female. 346-348, epigynum. 346, ventral. 347, posterior. 348, lateral. 349, dorsal. 350, 
abdomen, ventral. 



Scale lines. 1.0 mm, genitalia 0.1 mm. 



136 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



Specimens Examined. PERU Madre de Dios: Re- 
serva de Manu, Puesta de Vigilancia Pakitza, 11°58'S, 
71°18"W, 6 Oct. 1987, 19 (D. Silva D., J. Coddington, 
USNM). BRAZIL Mato Grosso: 260 km N Xavantina, 
12°49'S, 51°46'W, Feb., Apr. 1969, 12 (Xavantina 
Cachimbo Exped., MCZ). BOLIVIA Beni: Est. Biol. 
Beni, 14°47'S, 66°15'W, ca. 225 m, 8-14 Nov. 1989, 
19 (J. Coddington, USNM). 

Metazygia carimagua new species 
Figures 333-336; Map 4C 

Holotype. Female holotype and one female paratype 
from Carimagua, 100 m, Depto. Meta, Colombia, 
Oct. 1973, grass and bushes along fence (W. Eber- 
hard), in MCZ. The specific name is a noun in 
apposition after the type locality. 

Description. Female holotype. Cara- 
pace orange. Chelicerae, labium, endites 
orange. Sternum, legs orange. Dorsum of 
abdomen white, with an indistinct pair of 
dusky patches anteriorly (Fig. 336). Venter 
with white transverse bar of white pig- 
ment spots behind epigynum. Posterior 
median eyes 0.8 diameter of anterior me- 
dians, laterals 0.7 diameter. Anterior me- 
dian eyes 0.3 diameter apart, 0.9 diameter 
from laterals. Posterior median eyes 0.2 
diameter apart, 1.5 diameters from later- 
als. Height of clypeus equals 0.6 diameter 
of anterior median eye. Total length 3.4 
mm. Carapace 1.6 mm long, 1.2 wide, 0.7 
behind lateral eyes. First femur 1.7 mm, 
patella and tibia 2.1, metatarsus 1.7, tarsus 
0.6. Second patella and tibia 1.6 mm, third 
0.9, fourth 1.4. 

Diagnosis. The species lacks black pig- 
ment, even in the eye region (Fig. 336). 
The epigynum differs from that of other 
species by having a pentagonal shape in 
ventral view (Fig. 333) and a narrow pos- 
terior median plate (Fig. 334). 

Specimen Examined. COLOMBIA Meta: Cari- 
magua, 100 m, 19 (W. Eberhard 633, MCZ). 

Metazygia loque new species 
Figures 337-341 ; Map 4C 

Holotype. Female holotype from Rurrenabaque, Beni, 
Bolivia, Oct.-Nov. 1956 (L. Pena), in IRSNB. The 
specific name is an arbitrary combination of letters. 

Description. Female holotype. Cara- 
pace orange-yellow , cephalic region dusky. 



Chelicerae brown. Labium, endites brown. 
Sternum dusky orange, borders darkest. 
Coxae, legs orange-yellow with distal ar- 
ticles darkest. Dorsum of abdomen with 
dense white pigment spots, anterior with 
black transverse band (Fig. 340); venter 
with a pair of white longitudinal bands and 
a pair of white spots on light gray (Fig. 
341). Posterior median eyes same diameter 
as anterior medians, laterals 0.8 diameter. 
Anterior median eyes 0.8 diameter apart, 
0.6 diameter from laterals. Posterior me- 
dian eyes 0.2 diameter apart, 0.9 diameter 
from laterals. Height of clypeus equals 0.7 
diameter of anterior median eye. Total 
length 4.5 mm. Carapace 1.7 mm long, 1.2 
wide, 0.7 behind lateral eyes. First femur 
1.9 mm, patella and tibia 2.3, metatarsus 
1.6, tarsus 0.5. Second patella and tibia 2.0 
mm, third 1.1, fourth 1.5. 

Diagnosis. The abdomen is oval, widest 
in middle, and slightly flattened anteriorly 
(Fig. 340). The female is distinguished by 
having an epigynum that is flat, ventrally 
projecting, and longer than wide (Figs. 
337-339). 

Metazygia cienaga new species 
Figures 342-345; Map 4D 

Holotype. Female holotype from along Arroyo Frio, 
La Cienaga, Prov. La Vega, 19°04'N, 70°51"W, Do- 
minican Republic, 8 Jan. 1986 (S. Larcher), in 
USNM. The specific name is a noun in apposition 
after the type locality. 

Description. Female holotype. Cara- 
pace orange, cephalic region black. Che- 
licerae, labium, endites orange. Sternum 
orange. Coxae orange, legs dusky orange. 
Dorsum of abdomen white with two an- 
terior black marks that are fused ventrally 
above carapace (Fig. 345). Venter white, 
with large white square between epigyn- 
um and spinnerets. Posterior median eyes 
0.7 diameter of anterior medians, laterals 
0.6 diameter. Anterior median eyes 0.7 di- 
ameter apart, 0.8 diameter from laterals. 
Posterior median eyes 0.3 diameter apart, 
1.5 diameters from laterals. Height of 
clypeus equals 0.3 diameter of anterior 
median eye. Total length 5.2 mm. Cara- 



METAZYGIA»Levi 



137 



pace 2.3 mm long, 1.7 wide, 0.9 behind 
lateral eyes. First femur 2.2 mm, patella 
and tibia 2.7, metatarsus 1.8, tarsus 0.8. 
Second patella and tibia 2.2 mm, third 1.4, 
fourth 1.9. 

Diagnosis. The abdomen is oval, widest 
in middle, anteriorly flattened slightly with 
median protuberance (Fig. 345). The epi- 
gynum has a median, wide lobe (Fig. 342) 
and a triangular, posterior median plate 
(Fig. 343). 

Metazygia souza new species 
Figures 346-350; Map 4D 

Holotype. Female holotype from Ilha de Maraca, Rio 
Uraricoera, Roraima State, Brazil, 25 Sept. 1987 
(M. E. L. Souza), in MCN no. 20059. The speciBc 
name is a noun in apposition after the collector. 

Description. Female holotype. Cara- 
pace yellow, eye region black. Chelicerae 
dusky yellow. Labium, endites, sternum 
yellow. Coxae, legs yellow. Dorsum of ab- 
domen with dense white pigment and a 
black band around the anterior (Fig. 349), 
sides and venter without pigment (Fig. 
350). Posterior median eyes 0.8 diameter 
of anterior medians, laterals 0.7 diameter. 
Anterior median eyes 0.6 diameter apart, 
0.6 diameter from laterals. Posterior me- 
dian eyes 0.4 diameter apart, 1.3 diameters 
from laterals. Height of clypeus equals 0.2 
diameter of anterior median eye. Total 
length 4.0 mm. Carapace 1.5 mm long, 1.2 
wide, 0.6 behind lateral eyes. First femur 
1.7 mm, patella and tibia 2.1, metatarsus 
1.5, tarsus 0.6. Second patella and tibia 1.7 
mm, third 1.0, fourth 1.5. 

Diagnosis. Unlike that of other species, 
the epigynum of M. souza has a posterior 
median notch (Figs. 346, 347). 

Metazygia lopez new species 
Figures 351-359; Map 4D 

Holotype. Female holotype and one female and six 
male paratypes from Hacienda Mozambique, 15 
km SW Puerto Lopez, 500 m elev., Depto. Meta, 
Colombia (W. Eberhard), in MCZ. The specific 
name is a noun in apposition after the type locality. 

Description. Female holotype. Cara- 
pace, sternum, legs yellow; legs with tips 



of tarsi black. Dorsum of abdomen with 
white pigment and a dusky band that is 
broken in middle around anterior (Fig. 
355). Venter white behind epigynum, with 
transverse dark mark in front of spinnerets 
(Fig. 356). Posterior median eyes 0.8 di- 
ameter of anterior medians, laterals 0.7 
diameter. Anterior median eyes a little less 
than their diameter apart, a little less than 
their diameter from laterals. Posterior me- 
dian eyes 0.3 their diameter apart, 1.7 from 
laterals. Height of clypeus equals 0.4 di- 
ameter of anterior median eye. Total length 
4.2 mm. Carapace 1.7 mm long, 1.2 wide, 
0.6 behind lateral eyes. First femur 1.8 
mm, patella and tibia 2.1, metatarsus 1.7, 
tarsus 0.7. Second patella and tibia 1.7 mm, 
third 1.0, fourth 1.4. 

Male paratype. Coloration as in female. 
Carapace shiny with indistinct round tho- 
racic depression enclosing a median lon- 
gitudinal mark. Posterior median eyes 1 
diameter of anterior medians, laterals 0.8 
diameter. Anterior median eyes their di- 
ameters apart, 0.5 from laterals. Posterior 
median eyes almost touching, 1.7 diame- 
ters from laterals. Height of clypeus equals 
0.4 diameter of anterior median eye. First 
coxae with small hook. Fourth coxae with 
a pointed tubercle. Abdomen oval, longer 
than wide. Total length 2.8 mm. Carapace 
1.5 mm long, 1.1 wide, 0.5 behind lateral 
eyes. First femur 1.7 mm, patella and tibia 
2.2, metatarsus 1.7, tarsus 0.7. Second pa- 
tella and tibia 1.6 mm, third 0.9, fourth 
1.2. 

Note. Males and females have been col- 
lected together. The epigynum of some 
specimens is completely covered by amor- 
phous hard material, probably placed there 
by the male after mating. 

Variation. Total length of females 3.2 
to 4.5 mm. Eberhard (personal commu- 
nication) reports the species to be green 
when alive. Illustrations were made from 
the holotype and paratypes. 

Diagnosis. The abdomen is spherical 
(Figs. 355, 356) and, unlike many species, 
the eye region of the carapace is light in 
color (Fig. 355) and the posterior median 



138 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



plate of the epigynum is dumbbell-shaped 
(Fig. 352). The male has a tubercle on the 
fourth coxa and a complex-shaped con- 
ductor (C in Fig. 359) and a long, thorn- 
shaped embolus (E in Figs. 357-359). 

Natural History. Specimens have been 
collected in grass and brush along a fence 
in Carimagua, Colombia; in grassland jun- 
gle at Puerto Lleras, Colombia; and in sa- 
vanna, fogging trees, in Venezuela. 

Distribution. Venezuela, Amazon re- 
gion (Map 4D). 

Specimens Examined. VENEZUELA Gudrico: 
Mato Masaquaral, 45 km S Calabozo, 19 Apr. 1980, 
12 (K. Rabenold, MCZ). COLOMBIA Meta: Cari- 
magua, 100 m, Oct. 1973, 19, IS (W. Eberhard, MCZ); 
Hacienda Mozambique, 15 km SW Puerto Lopez, 
119, 16 (W. Eberhard, MCZ); Lomalinda, Puerto 
Lleras, 3°18'N, 73°22'W, 12 Jan. 1986, 19 (B. T. Car- 
roll, MCZ), Sept. 1987, 29 (B. T. Carroll, CAS). PERU 
Ucayali: Laguna Cashibococha, 25 km nr. Pucallpa, 
30 Dec. 1987, 19 (M. Remo, MUSM). BRAZIL Ama- 
zonas: Lago do Jose, Manaus, 9 Aug. 1979, 19 (J. Adis, 
MCN 20057); Ilha de Curari, Manaus, 3 Aug. 1987, 
19 (J. Adis, MCN 20056); Ilha de Marchantaria, Rio 
SoHmoes, 59°58'W, 3n5'S, 2 Sept. 1992, 43 (J. Adis 
et al., INPA). 

Metazygia samiria new species 
Figures 360-364; Map 4D 

Holotype. Female holotype and three female para- 
types from Rio Samiria, fogging and night collect- 
ing, Depto. Loreto, Peru, 8-31 May 1990 (T. Erwin, 
D. Silva D. ), in MUSM, one paratype in MCZ. The 
specific name is a noun in apposition after the type 
locality. 

Description. Female holotype. Cara- 
pace light orange, cephalic region darker, 
eye area black. Chelicerae brown-black. 
Labium, endites dusky orange. Sternum 
orange with sides dusky. Legs orange with 
indistinct darker dusky rings. Dorsum of 
abdomen black around anterior, otherwise 



light (Fig. 363). Venter gray to black with 
a pair of white patches between epigynum 
and spinnerets and with white anteriorly 
on sides of pedicel (Fig. 364). Posterior 
median eyes 0.9 diameter of anterior me- 
dians, laterals 0.9 diameter. Anterior me- 
dian eyes 0.7 diameter apart, 0.6 diameter 
from laterals. Posterior median eyes 0.3 
diameter apart, L4 diameters from later- 
als. Height of clypeus equals 0.7 diameter 
of anterior median eye. Total length 5.6 
mm. Carapace 2.1 mm long, L6 wide, 0.8 
behind lateral eyes. First femur 2.1 mm, 
patella and tibia 2.7, metatarsus 1.9, tarsus 
0.7. Second patella and tibia 2.5 mm, third 
1.3, fourth 2.1. 

Variation. Total length of females 4.5 
to 5.7 mm. Illustrations were made from 
the female holotype. 

Diagnosis. Metazygia samiria differs 
from M. ducke in the shape of the lateral 
plates in posterior view of the epigynum 
(Fig. 361). 

Distribution. Western Amazon region 
(Map 4D). 

Specimens Examined. PERU Hudnuco: Dantas- 
La Molina, SW Puerto Inca, 09°28'S, 75°00"W, 22 May 
1987, 19 (D. Silva D., MUSM); Cucharas, Huallaga 
Valley, Feb.-Apr. 1954, 19 (F. Woytkowski, CAS). 
Madre de Dios. 15 km E Puerto Maldonado, 12°33'S, 
69°03'W, 26 Feb. 1989, 19 (D. Silva D., MUSM); Zona 
Reservada Tambopata, trocha principal, 290 m, 
12°50'S, 69°17'W, 69 (D, Silva D., MUSM). 

Metazygia ducke new species 
Figures 365-369; Map 4D 

Holotype. Female holotype from Reserva Ducke, 
Manaus, Est. Amazonas, Brazil, Aug. 1971 (M. E. 
Galiano), in MACN. The specific name is a noun 
in apposition after the type locality. 

Description. Female holotype. Cara- 
pace orange. Chelicerae brown. Labium, 



Figures 351-359. Metazygia lopez n. sp. 351-356, female. 351-354, epigynum. 351 , 354, ventral. 352, posterior. 353, lateral. 
351-353, (Colombia). 354, (Brazil). 355, dorsal. 356, abdomen, ventral. 357-359, left male palpus. 357, mesal. 358, ventral. 359, 
mesal, pulled apart. 

Figures 360-364. M. samiria n. sp., female. 360-362, epigynum. 360, ventral. 361, posterior. 362, lateral. 363, dorsal. 364, 
abdomen, ventral. 



Figures 365-369. M. ducke n. sp., female. 365-367, epigynum. 365, ventral. 366, posterior. 367, lateral. 368, dorsal. 369, 
abdomen, ventral. 



METAZYGiA'Levi 139 




Figures 370-373. M. erratica (Keyserling), female. 370-372, epigynum. 370, 372, ventral. 371 , posterior. 371 , (Mato Grosso). 
372, (Holotype). 373, dorsal. 

Figures 374-377. M. manu n. sp, male. 374, 375, palpus. 374, mesal. 375, ventral. 376, dorsal. 377, abdomen, ventral. 
Abbreviations. A, terminal apophysis; C, conductor; E, embolus; I, stipes; M, median apophysis; R, radix. 
Scale lines. 1.0 mm, genitalia 0.1 mm. 



140 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



endites brown. Sternum orange, dusky on 
each side. Coxae and legs orange with dis- 
tal articles of legs darker. Dorsum of ab- 
domen with dense white pigment spots and 
an anterior transverse black band (Fig. 
368). Venter with a pair of white patches 
on gray (Fig. 369). Posterior median eyes 
same diameter as anterior medians, lat- 
erals 0.8 diameter. Anterior median eyes 
0.6 diameter apart, 0.5 diameter from lat- 
erals. Posterior median eyes 0.2 diameter 
apart, 1 diameter from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Total length 5.0 mm. Cara- 
pace 2.1 mm long, 1.5 wide, 0.8 behind 
lateral eyes. First femur 2.1 mm, patella 
and tibia 2.7, metatarsus 1.8, tarsus 0.6. 
Second patella and tibia 2.3 mm, third 1.2, 
fourth 1.9. 

Variation. The specimen from Bolivia 
has a black sternum and other minor dif- 
ferences. 

Diagnosis. Metazygia ducke differs from 
M. samiria (Fig. 361) by the shape of the 
lateral plates in posterior view of the epi- 
gynum and the square median plate, which 
has a textured area at its ventral end (Fig. 
366). 

Distribution. Amazon region (Map 4D). 

Specimens Examined. BOLIVIA Beni: Est. Biol. 
Beni, 14°47'S, eG'lS'W, 225 m, 8-14 Nov. 1989, 19 
(J. Coddington et at., USNM). 

Metazygia erratica (Keyserling), 
new combination 
Figures 370-373; Map 4E 

Epeira erratica Keyserling, 1883: 197, pi. 15, fig. 3, 
9. Female holotype from "Provinz Amazonas," Bra- 
zil, in HECO, examined. Keyserling, 1892: 161, pi. 
8, fig. 119, 9. 

Aranea errans Roewer, 1942: 841. New name for 
erratica, since thought preoccupied by Aranea er- 
ratica Olivier, 1789. 

Araneus erraticus: — Bonnet, 1955: 501. 

Description. Female holotype. Cara- 
pace orange. Chelicerae, labium, endites 
orange. Sternum, coxae, legs light orange. 
Abdomen whitish (Fig. 373). Posterior me- 
dian eyes 0.8 diameter of anterior medi- 
ans, laterals 0.7 diameter. Anterior median 



eyes 0.3 diameter apart, 0.7 from laterals. 
Posterior median eyes 0.6 diameter apart, 
1.4 diameters from laterals. Height of 
clypeus 0.5 diameter of anterior median 
eye. Legs without macrosetae. Total length 
2.9 mm. Carapace 1.4 mm long, 0.9 wide, 
0.6 behind lateral eyes. First femur 1.5 
mm, patella and tibia 1.7, metatarsus 1.2, 
tarsus 0.6. Second patella and tibia 1 .3 mm, 
third 0.7, fourth 1.1. 

Variation. Keyserling records a silvery 
coloration of the abdomen; the specimens 
available have lost all pigment. Total length 
2.9 to 3.9 mm. Figure 372 was made from 
the holotype; Figures 370, 371, and 373 
are from a specimen from Mato Grosso. 

Diagnosis. The absence of black in the 
eye region (Fig. 373) and the thick folded 
lips of the epigynum in ventral view (Figs. 
370, 372) separate M. erratica from M. 
samiria and similar species. All specimens 
had black amorphous material covering 
the openings of the epigynum (on the left 
of Figs. 370-372) which was not found in 
related species. 

Specimens Examined. BRAZIL Mato Grosso: Ba- 
rra do Tapirape, 1-5 Jan, 1961, 19 (B. Malkin, AMNH); 
Utiariti, 25 Oct. 1966, 19 (F. Lenko, Pereira, MZSP 
6064). 

Metazygia manu new species 
Figures 374-377; IVIap 4E 

Holotype. Male holotype from Puesto de Vigilancia 
Pakitza, Zona Reservada de Manu, Depto. Madre 
de Dios, ll''58'S, 71''18'W, Peru, night collecting, 
30 Sept. 1987 (D. Silva D., J. Coddington), in MUSM. 
The specific name is a noun in apposition after the 
type locality. 

Description. Male holotype. Carapace 
orange, cephalic area gray, black between 
eyes. Chelicerae, labium, endites black. 
Sternum black. Coxae light orange; legs 
orange except for black ring distally on 
first tibia. Dorsum of abdomen white with 
anterior transverse black band, and pos- 
terior pair of black patches that fuse to a 
median band above spinnerets (Fig. 376). 
Venter with distinct black band covering 
both genital area and spinnerets and con- 
tinuing into dorsal black patches (Fig. 377). 



Metazycia 'Levi 



141 



Carapace with double border above tirst 
coxae. Posterior median eyes 0.8 diameter 
of anterior medians, laterals 0.7 diameter. 
Anterior median eyes 0.5 diameter apart, 
0.2 diameter from laterals. Posterior me- 
dian eyes 0.3 diameter apart, their diam- 
eter from laterals. Height of clypeus equals 
0.4 diameter of anterior median eye. Sec- 
ond tibia thicker than first, with a mac- 
roseta on swollen area. Total length 2.7 
mm. Carapace 1.24 mm long, 1.04 wide, 
0.54 behind lateral eyes. First femur 1.03 
mm, patella and tibia 1.18, metatarsus 0.75, 
tarsus 0.38. Second patella and tibia 1.14 
mm, third 0.81, fourth 1.12. 

Diagnosis. The male palpus differs from 
others by having a long, fine, "diagonal," 
embolus (Fig. 374) and by the complex 
shape of the median apophysis (at 4 hr in 
Fig. 374, below center of Fig. 375). 

Metazygia genaro new species 
Figures 378-384; Map 4E 

Holotype. Female holotype from Genaro Herrera, 
04°55'S, 73''45'W, Depto. Loreto, Peru, 26 Aug, 
1988 (D. Silva D.), in MUSM. The specific name 
is a noun in apposition after the type locahty. 

Description. Female holotype. Cara- 
pace light orange, cephalic region brown, 
eye area black. Chelicerae dark brown. 
Labium, endites, sternum brown. Coxae 
orange; first two legs dark brown, last two 
orange. Dorsum of abdomen white with a 
black band around anterior (Fig. 381); 
venter with a pair of white patches (Fig. 
382). Posterior median eyes same diameter 
as anterior medians, laterals 0.7 diameter. 
Anterior median eyes 0.8 diameter apart, 
0.6 diameter from laterals. Posterior me- 
dian eyes 0.4 diameter apart, 1.1 diameters 
from laterals. Height of clypeus equals 0.4 
diameter of anterior median eye. Total 
length 4.3 mm. Carapace 1.7 mm long, 1.3 
wide, 0.7 behind lateral eyes. First femur 
1.9 mm, patella and tibia 2.1, metatarsus 
1.5, tarsus 0.5. Second patella and tibia 1.8 
mm, third 1.1, fourth 1.6. 

Male. As in female, but orange areas 
more yellowish (Fig. 384); venter with a 
pair of white pigment patches on gray (Fig. 



382). Carapace with lobes above first coxae 
(Fig. 384). Posterior median eyes same di- 
ameter as anterior medians, laterals 0.7 di- 
ameter. Anterior median eyes 0.8 diame- 
ter apart, 0.2 diameter from laterals. Pos- 
terior median eyes 0.3 diameter apart, 0.8 
diameter from laterals. Ocular quadrangle 
narrower behind than in front. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. First coxa with hook, third 
and fourth each with macroseta on a soft 
tubercle. Second tibia thicker than first with 
macrosetae, one of them long. Total length 
2.5 mm. Carapace 1.30 mm long, 1.04 
wide, 0.48 behind lateral eyes. First femur 
1.49 mm, patella and tibia 1.91, metatarsus 
1.40, tarsus 0.52. Second patella and tibia 
1.31 mm, third 0.80, fourth 1.14. 

Note. Male and female were matched 
because in both the anterior dorsal bands 
of the abdomen are intense black, with 
white adjacent to the black both anteriorly 
and posteriorly. Both sexes have a pair of 
white patches on black on the underside 
of the abdomen (Fig. 382). Male and fe- 
male did not come from the same locality. 

Diagnosis. A broad, triangular epigynal 
scape as seen in ventral view (Fig. 378) 
distinguishes the female. Males are distin- 
guished from M. voxanta by the shape of 
the palpal sclerites (Fig. 383). 

Distribution. Depto. Loreto, Peru (Map 
4E). 

Specimen Examined. PERU Loreto: Rio Manatee 
[a tributary of the Amazon between Explorers Lodge 
and Rio Napo], 18 July 1989, grass and shrubs, 16 (G. 
B. Edwards, FSCA). 

Metazygia voxanta new species 
Figures 385-390; Map 4E 

Holotype. Female holotype, female paratype, and 
two male paratypes from 260 km N Xavantina, 
12°49'S, 51°46'W, 400 m, Mato Grosso State, Brazil, 
campo-grassland, Feb. -Apr. 1969 (Xavantina- 
Cachimbo Expedition), 12 holotype and 13 para- 
type in MCN, others in MCZ. The specific name 
is an arbitrary combination of letters. 

Description. Female holotype. Cara- 
pace orange, eye region black. Chelicerae, 
labium, endites brownish. Sternum or- 
ange. Coxae, legs orange. Dorsum of ab- 



142 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



domen white, with anterior and posterior 
black marks (Fig. 388). Venter dusky with 
few irregularly spaced white pigment spots. 
Posterior median eyes 0.8 diameter of an- 
terior medians, laterals 0.6 diameter. An- 
terior median eyes 0.8 diameter apart, 0.5 
diameter from laterals. Posterior median 
eyes 0.2 diameter apart, 1 diameter from 
laterals. Height of clypeus equals 0.5 di- 
ameter of anterior median eye. Total length 
2.5 mm. Carapace 0.9 mm long, 0.7 wide, 
0.4 behind lateral eyes. First femur 0.8 
mm, patella and tibia 1.0, metatarsus 0.6, 
tarsus 0.4. Second patella and tibia 0.9 mm, 
third 0.5, fourth 0.7. 

Male paratype. Color as in female. The 
carapace has a lobe above the first coxa 
(Fig. 390). Posterior median eyes 0.7 di- 
ameter of anterior medians, laterals 0.5 
diameter. Anterior median eyes 0.3 di- 
ameter apart, 0.2 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
1 diameter from laterals. Height of clypeus 
equals 0.8 diameter of anterior median eye. 
First coxa with hook, fourth with one short 
macroseta. Second tibia thicker than first, 
with four macrosetae on distal half. Total 
length 2.0 mm. Carapace 1.0 mm long, 0.8 
wide, 0.4 behind lateral eyes. First femur 
0.8 mm, patella and tibia 1.1, metatarsus 
0.6, tarsus 0.4. Second patella and tibia 0.9 
mm, third 0.5, fourth lost. 

Note. Males and females were collected 
together. 

Diagnosis. The epigynum has two lobes 
in ventral view (Fig. 385) and an hour- 
glass-shaped median plate in posterior view 
(Fig. 386). The male is similar to that of 
M. genaro but has an embolus lamella and 



a median apophysis of different shape (Fig. 
389). 



Metazygia peckorum new species 
Figures 391-400; IVIap 4F 

Holotype. Female from La Chiquita, 5 m elev., 11 
km SE San Lorenzo, Esmeraldes Prov., Ecuador, 
3-10 June 1975 (S. and J. Peck), in MCZ. The 
species is named after the collectors. 

Description. Female holotype. Cara- 
pace light orange, cephalic region dusky. 
Chelicerae, labium, endites brown. Ster- 
num black. Coxae light orange. First two 
pairs of legs brown, except for proximal 
end of femora light orange, last two pairs 
of legs light orange. Dorsum of abdomen 
white with anterior dark band (Fig. 399). 
Venter black with a pair of white patches 
(Fig. 400). Posterior median eyes 0.8 di- 
ameter of anterior medians, laterals 0.6 
diameter. Anterior median eyes 0.7 di- 
ameter apart, 0.7 diameter from laterals. 
Posterior median eyes 0.3 diameter apart, 
1.1 diameters from laterals. Height of 
clypeus equals 0.8 diameter of anterior 
median eye. Total length 4.6 mm. Cara- 
pace 1.9 mm long, 1.5 wide, 0.8 behind 
lateral eyes. First femur 2.0 mm, patella 
and tibia 2.3, metatarsus 1.7, tarsus 0.6. 
Second patella and tibia 2.0 mm, third 1.2, 
fourth 1.8. 

Variation. Total length of females 4.0 
to 5.1 mm. 

Diagnosis. The oval to triangular me- 
dian area of the epigynum has a longitu- 
dinal groove and a minute flat scape at its 
tip (Figs. 391, 394, 397), and the posterior 
median plate is constricted dorsally in pos- 



Figures 378-384. Metazygia genaro n. sp. 378-382, female. 378-380, epigynum. 378, ventral. 379, posterior. 380, lateral. 
381, dorsal. 382, abdomen, ventral. 383, 384, male. 383, left palpus. 384, dorsal. 

Figures 385-390. M. voxanta n. sp. 385-388, female. 385-387, epigynum. 385, ventral. 386, posterior. 387, lateral. 388, dorsal. 
389, 390, male. 389, palpus. 390, carapace. 

Figures 391^00. M. peckorum n. sp., female. 391-398, epigynum. 391 . 394. 397, ventral. 392, 395, posterior. 393, 396, 398, 
lateral. 391-393, (Peru). 394-396, (holotype. Ecuador). 397, 398 (Para, Brazil). 399, dorsal. 400, abdomen, ventral. 

Figures 401^04. M. moldira n. sp., female. 401-403, epigynum. 401, ventral. 402, posterior. 403, lateral. 404, dorsal. 



METAZYGiA'Levi 143 




Figures 405-408. M. valentim n. sp., female. 405-407, epigynum. 405, ventral. 406, posterior. 407, lateral. 408, abdomen, 

lateral. 

Figures 409-413. M. bahia n. sp., female. 409-411, epigynum. 409, ventral. 410, posterior. 411, lateral. 412, dorsal. 413, 
abdomen, ventral. 



Scale lines. 1.0 mm, genitalia 0.1 mm. 



144 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



terior view (Figs. 392-395): these char- 
acters separate M. peckorum from M. mol- 
dira (Figs. 401, 402). 

Distribution. From Colombia to Bahia 
State, Brazil (Map 4F). 

Specimens Examined. COLOMBIA Valle: Central 
Hidroelectrica Anchicaya, 1978, 12 (W. Eberhard, 
MCZ). ECUADOR Los Rios: Juan Montalvo, Mar. 
1938, 19 (W. Clarke-Macintyre, AMNH). PERU 
Htidntico: Monzon Valley, Tingo Maria, 10 Nov. 1954, 
19 (E. I. Schlinger, E. S. Ross, CAS). BRAZIL Rorai- 
ma: Ilha de Maraca, Rio Uraricoera, 22 July 1987, 19 
(A. A. Lise, MCN 20061). Pard: Belem, Aug. 1971, 
19 (M. E. Galiano, MEG). Rahia: Camaca, Fazenda 
Matiapa, 16 Oct. 1978, 19 (J. S. Santos, MCN 11117); 
Uru^uca, Fazenda Almada, 26 Nov. 1977. 19 (J. S. 
Santos, MCN 10321). 

Metazygia motdira new species 
Figures 401-404; Map 4F 

Holotype. Female holotype from 15 km E Puerto 
Maldonado, 12°33'S, 69°03"W, 200 m, Depto. Ma- 
dre de Dios, Peru, 26 Feb. 1989 (D. Silva D.), in 
MUSM. The specific name is an arbitrary combi- 
nation of letters. 

Description. Female holotype. Cara- 
pace orange, eye region black. Chelicerae 
orange with a proximal dusky patch. La- 
bium, endites dusky orange. Sternum hght 
orange. Legs light orange. Dorsum of ab- 
domen white with a black band around 
anterior (Fig. 404). Venter with a white 
square between epigynum and spinnerets. 
Eyes subequal in size. Anterior median eyes 
their diameter apart, 0.8 diameter from 
laterals. Posterior median eyes 0.5 diam- 
eter apart, L2 diameters from laterals. 
Height of clypeus equals 0.4 diameter of 
anterior median eye. Total length 3.8 mm. 
Carapace L9 mm long, 1.4 wide, 0.8 be- 
hind lateral eyes. First femur 2.1 mm, pa- 
tella and tibia 2.5, metatarsus 1.8, tarsus 
0.7. Second patella and tibia 2.3 mm, third 
1.3, fourth 1.8. 

Variation. Total length of females 3.8 
to 5.6 mm. Illustrations were made from 
the female holotype. 

Diagnosis. Metazygia moldira is similar 
to M. peckorum, but the posterior median 
plate of the epigynum is flask-shaped and 
widest dorsally (bottom of Fig. 402). 



Distribution. Western Amazon region 
(Map 4F). 

Specimens Examined. ECUADOR Sucumhnos: 
bridge over Rio Cuyabeno, O.OFS, 76°18'W, 8, 9 Aug. 
1988, 19 (W. Maddison, MCZ). PERU Loreto: Rio 
Samiria, 8-31 May 1990, 19 (T. Erwin, D. Silva D., 
MUSM). 

Metazygia valentim new species 
Figures 405-408; Map 4F 

Holotype. Female holotype and two immatures from 
Sao Valentim, Est. Rio Grande do Sul, Brazil, 16 
Oct. 1976 (R. Scherer), in MCN no. 04782. The 
specific name is a noun in apposition after the type 
locality. 

Description. Female holotype. Cara- 
pace dusky brown, eye region black. Che- 
licerae, labium, endites dark brown. Ster- 
num dark brown. Coxae light yellowish; 
legs light yellowish with tips of tarsi darker. 
Dorsum of abdomen white with anterior 
transverse black band; sides with a black 
patch (Fig. 408). Venter with a black band 
starting anteriorly from the transverse band 
and posteriorly enclosing spinnerets. Pos- 
terior median eyes same diameter as an- 
terior medians, laterals 0.8 diameter. An- 
terior median eyes 0.8 diameter apart, 0.7 
diameter from laterals. Posterior median 
eyes 0.3 diameter apart, 1.6 diameters from 
laterals. Laterals 0.5 their diameter apart. 
Height of clypeus equals 0.8 diameter of 
anterior median eye. Total length 3.1 mm. 
Carapace 1.4 mm long, 1.1 wide, 0.6 be- 
hind lateral eyes. First femur 1.6 mm, pa- 
tella and tibia 1.9, metatarsus 1.2, tarsus 
0.5. Second patella and tibia 1.6 mm, third 
0.8, fourth 1.3. 

Diagnosis. The epigynum of this species 
differs from that of other species by having 
a transverse posterior lip in ventral view 
(Fig. 405) and a T-shaped posterior me- 
dian plate in posterior view (Fig. 406). 

Metazygia bahia new species 
Figures 409-413; Map 4F 

Holottjpe. Female holotype from Fazenda Jacaranda, 
Itamaraju, Bahia State, Brazil, 9 Dec. 1977 (J. S. 
Santos), in MCN no. 11030. The specific name is a 
noun in apposition after the type locality. 



METAZYCIA'Levi 



145 



Description. Female holotype. Cara- 
pace orange, eye region black. Chelicerae 
brown. Labium, endites dusky orange. 
Sternum dusky orange. Legs orange. Dor- 
sum of abdomen with dense white pig- 
ment spots and with a black band around 
anterior (Fig. 412). Venter with indistinct 
pair of white patches, dusky around spin- 
nerets (Fig. 413). Posterior median eyes 
0.9 diameter of anterior medians, anterior 
laterals 0.7 diameter, posterior 0.8. Ante- 
rior median eyes 0.4 diameter apart, 0.6 
diameter from laterals. Posterior median 
eyes 0.4 diameter apart, one diameter from 
laterals. Height of clypeus equals 0.4 di- 
ameter of anterior median eye. Total length 
4.2 mm. Carapace 2.0 mm long, 1.4 wide, 
0.8 behind lateral eyes. First femur 1.9 
mm, patella and tibia 2.3, metatarsus 1.6, 
tarsus 0.6. Second patella and tibia 2.0 mm, 
third 1.2, fourth 1.6. 

Variation. Total length of females 4.0 
to 4.3 mm. Illustrations were made from 
the holotype. 

Diagnosis. Unlike the epigynum of M. 
peckorum (Figs. 391, 392), the two cones 
on the epigynum of M. bahia are medially 
separated (Figs. 409, 410). 

Distribution. Bahia State to Sao Paulo 
State, Brazil (Map 4F). 

Specimens Examined. BRAZIL Sao Paulo: Alto da 
Serra Barreira das Camellas, Nov. 1941, 19 (J. Dom- 
igo, MZSP 9641); Caraguatatuba, 14 July 1964, 12 
(Exped. Depto. Zool., MZSP). 

Metazygia rothi new species 
Figures 414, 415; Map 4F 

Holotype. Male holotype from Lomalinda, nr. Puerto 
Lleras, Depto. Meta, 300 m, 3°18'S, 73°22'W, Co- 
lombia, Mar. 1988 (V. Roth), in MCZ. The species 
is named after the collector. 

Description. Male holotype. Carapace 
olive-white, black between eyes. Chelic- 
erae, labium, endites, sternum, legs olive- 
white. Dorsum of abdomen white with 
black band around anterior (Fig. 415). 
Venter dusky with an indistinct pair of 
white patches. Posterior median eyes 0.7 
diameter of anterior medians, laterals 0.7 



diameter. Anterior median eyes 0.5 di- 
ameter apart, 0.3 diameter from laterals. 
Posterior median eyes 0.2 diameter apart, 
1 diameter from laterals. Height of clypeus 
equals 0.8 diameter of anterior median eye. 
Second tibia barely thicker than first, with- 
out macrosetae. Total length 2.1 mm. Car- 
apace 1.04 mm long, 0.78 wide, 0.41 be- 
hind lateral eyes. First femur 0.96 mm, 
patella and tibia 1.18, metatarsus 0.78, tar- 
sus 0.37. Second patella and tibia 0.94 mm, 
third 0.58, fourth 0.78. 

Note. This delicate, small male might 
be that of M. carimagua. A recent molt 
may have given the specimen its olive- 
white color. 

Diagnosis. Metazygia rothi differs from 
others by having the sickle-shaped em- 
bolus positioned on the conductor and by 
the shape of the median apophysis and 
conductor (Fig. 414). 

Metazygia cazeaca new species 
Figures 416, 417; Map 41 

Holotype. Male holotype from Jacareacanga, Est. Para, 
Brazil, Oct. 1959 (M. Alvarenga), in AMNH. The 
specific name is an arbitrary combination of letters. 

Description. Male holotype. Carapace 
dark brown, except for light orange me- 
dian thoracic region. Chelicerae orange. 
Labium, endites dusky orange. Sternum 
orange. Coxae light orange, legs orange. 
Dorsum of abdomen black with a trans- 
verse light band (Fig. 417), sides light. 
Venter with black trapezoid between gen- 
ital groove and spinnerets. Posterior me- 
dian eyes 0.8 diameter of anterior medi- 
ans, laterals 0.5 diameter. Anterior median 
eyes 0.5 diameter apart, 0.3 diameter from 
laterals. Posterior median eyes 0.3 diam- 
eter apart, their diameter from laterals. 
Height of clypeus equals 0.6 diameter of 
anterior median eye. Endite without tooth. 
First coxa with minute hook. (Distal leg 
articles broken off.) Total length 2.1 mm. 
Carapace 1.1 mm long, 0.9 wide, 0.5 be- 
hind lateral eyes. Third patella and tibia 
0.7 mm. 

Diagnosis. The coloration of the body. 



146 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



the elongate shape of the abdomen, widest 
in posterior half (Fig. 417), and the shape 
of the sclerites of the palpus (Fig. 416) are 
unlike any other species of Metazygia. This 
species may not belong to the genus. 

Metazygia cunha new species 
Figures 418, 419; Map 4H 

Holotype. Male holotype from Jaboticabal, Sao Paulo 
State, Brazil, 4 Oct. 1986 (H. F. do Cunha), in MCN 
no. 17820. The specific name is a noun in apposition 
after the name of the collector. 

Description. Male holotype. Carapace 
yellow, eye region black. Chelicerae, la- 
bium, endites yellow. Sternum yellow. Legs 
yellow. Dorsum of abdomen white, with 
a band around the anterior and an indis- 
tinct, transverse, white patch adjacent and 
posterior to it (Fig. 419). Venter with a 
pair of white patches on light gray, with 
another posterior pair of white spots. Car- 
apace with small lobes (Fig. 419). Median 
eyes large. Posterior median eyes same di- 
ameter as anterior medians, laterals 0.6 di- 
ameter. Anterior median eyes 0.4 diame- 
ter apart, 0.2 diameter from laterals. Pos- 
terior median 0.2 diameter apart, 0.8 di- 
ameter from laterals. Height of clypeus 
equals 0.6 diameter of anterior median eye. 
First coxa with large hook, third and fourth 
with a macroseta; the macroseta on the 
fourth coxa on a tubercle. Total length 3.0 
mm. Carapace 1.4 mm long, 1.0 wide, be- 
hind lateral eyes 0.5 wide. First femur 1.7 
mm, patella and tibia 2.2, metatarsus 1.7, 
tarsus 0.6. Second patella and tibia 1.5 mm, 
third 0.9, fourth 1.3. 

Diagnosis. Metazygia cunha differs 
from others by the shape of the embolus 
lamella, conductor, and median apophysis 
(Fig. 418). 

Metazygia aldela new species 
Figures 420-422; Map 41 

Holotype. Male holotype from Adeia Ara9u, Igarape 
Gurupi-Uma, 50 km E of Caninde, Rio Gurupi, 
Est. Para, Brazil, 2-30 May 1963 (B. Malkin), in 
AMNH. The specific name is an arbitrary combi- 
nation of letters. 

Description. Male holotype. Carapace 
orange. Chelicerae dusky orange. Labium, 



endites dusky orange. Sternum light or- 
ange with black on each side. Legs orange. 
Dorsum of abdomen whitish with trans- 
verse black band around anterior (Fig. 
421); venter black, with a pair of round 
pigmentless patches side by side (Fig. 422). 
Posterior median eyes same diameter as 
anterior medians, laterals 0.6 diameter. 
Anterior median eyes 0.6 diameter apart, 
0.5 diameter from laterals. Posterior me- 
dian eyes 0.3 diameter apart, their diam- 
eter from laterals. Height of clypeus equals 
0.6 diameter of anterior median eye. Car- 
apace with small lobes (Fig. 421). First 
coxa with large hook. Total length 3.5 mm. 
Carapace 1.8 mm long, 1.5 wide, 0.6 be- 
hind lateral eyes. First femur 2.0 mm, pa- 
tella and tibia 2.5, metatarsus 1.8, tarsus 
0.6. Second patella and tibia 1.9 mm, third 
1.1, fourth 1.7. 

Diagnosis. Metazygia aldela differs 
from others by the large sclerotized em- 
bolus lamella (at 11 hr) and the pointed 
tooth (at 4 hr) on the median apophysis 
(Fig. 420). 

Metazygia atama new species 
Figures 423, 424; Map 4H 

Holotype. Male holotype from Fazenda Matiapa, Ca- 
maca, Bahia State, Brazil, 14 Oct. 1978 (J. S. San- 
tos), in MCN no. 11078. The specific name is an 
arbitrary combination of letters. 

Description. Male holotype. Carapace 
light orange, eye region black. Chelicerae, 
labium, endites brown. Sternum brown. 
Coxae light orange, legs light orange. Dor- 
sum of abdomen white with black band 
around anterior (Fig. 424); venter black. 
Posterior median eyes 0.8 diameter of an- 
terior medians, laterals 0.7 diameter. An- 
terior median eyes 0.7 diameter apart, 0.3 
diameter from laterals. Posterior median 
eyes 0.2 diameter apart, 1 from laterals. 
Height of clypeus equals 0.8 diameter of 
anterior median eye. Carapace with small 
lobes (Fig. 424). First coxa with large hook, 
third with one long macroseta and several 
smaller ones, fourth with one macroseta. 
Second tibia thicker than first, swollen in 
middle with two long macrosetae and some 
others. Total length 2.9 mm. Carapace 1.6 



METAZYGIA'Levi 147 



417 




Figures 414, 415. Metazygia rothin. sp., male. 414, left palpus. 415, dorsal. 

Figures 416, 417. M. cazeaca n. sp., male. 416, palpus. 417, dorsal. 

Figures 418, 419. M. cunha n. sp., male. 418, palpus. 419, dorsal. 

Figures 420-422. M. aldela n. sp., male. 420, palpus. 421 , dorsal. 422, abdomen, ventral. 

Figures 423, 424. M. atama n. sp., male. 423, palpus. 424, dorsal. 

Figures 425^28. M. oro n. sp., male. 425, 426, palpus. 425, mesal. 426, ventral. 427, dorsal. 428, abdomen, ventral. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



mm long, 1.3 wide, 0.6 behind lateral eyes. 
First femur 1.7 mm, patella and tibia 2.3, 
metatarsus 1.7, tarsus 0.6. Second patella 
and tibia 1.6 mm, third 1.0, fourth 1.5. 

Diagnosis. The palpus of Metazygia 
atama differs from that of other males by 
the large median apophysis that faces the 
cymbium (Fig. 423). 



Metazygia oro new species 
Figures 425-428; Map 4! 

Holotype. Male holotype from Rio Colorado, El Oro 
Prov., Ecuador, 4 Nov. 1942 (R. Walls), in CAS. 
The specific name is a noun in apposition after the 
type locality. 

Description. Male holotype. Carapace 
dusky orange. Chelicerae orange. Labium, 



148 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



endites, sternum brown. Legs orange. Ab- 
domen white with a black band around 
anterior (Fig. 427). Venter black with a 
pair of white patches (Fig. 428). Carapace 
with small lobes (Fig. 427). Posterior me- 
dian eyes same diameter as anterior me- 
dians, laterals 0.6 diameter. Carapace with 
small lobes (Fig. 427). Anterior median eyes 
0.8 diameter apart, 0.6 diameter from lat- 
erals. Posterior median eyes 0.3 diameter 
apart, 1.1 diameters from laterals. Height 
of clypeus equals 1 diameter of anterior 
median eye. First coxa with large hook, 
fourth with a short macroseta. Total length 
3.1 mm. Carapace 1.5 mm long, 1.2 wide, 
0.6 behind lateral eyes. First femur 1.6 
mm, patella and tibia 2.1, metatarsus 1.4, 
tarsus 0.5. Second patella and tibia 1.5 mm, 
third 0.9, fourth 1.4. 

Diagnosis. This species is distinguished 
by the unique shape of the embolus la- 
mella and the median apophysis of the 
palpus (above center and at 3 hr, respec- 
tively, in Fig. 425, at 10 hr and center in 
Fig. 426). 

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150 Bulletin Museum of Comparative Zoology, Vol. 154, No. 2 



INDEX 



Valid names are printed in italics. Page numbers refer to main references, starred page numbers to 
illustrations. 



adisi, Metazygia, 104, 105* 

albonigra, Metazygia, 88 

aldela, Metazygia, 146, 147* 

amalla, Metazygia, 119*, 120 

arnoi, Metazygia, 104, 105* 

atalaya, Metazygia, 109*, 110 

atama, Metazygia, 146, 147* 

bahama, Metazygia, 81, 83* 

bahia, Metazygia, 143*, 144 

harueri, Metazygia, 97*, 98 

benella, Metazygia, 122, 123* 

bolivia, Metazygia, 105*, 106 

calix, Metazygia, 72 

carimagua. Metazygia, 135*, 136 

carolinalis, Metazygia, 72 

carrizal, Metazygia, 101*, 102 

castaneoscutata, Aranea, 129 

castaneoscutata, Metazygia, 129, 131'' 

castaneoscutatus, Araneus, 129 

cazeaca, Metazygia, 145, 147* 

chenevo, Metazygia, 109*, 110 

chicanna, Metazygia, 90, 91* 

cienaga, Metazygia, 135*, 136 

coamensis, Larinia, 99 

corima, Metazygia, 109*, 111 

coriimba, Metazygia, 113*, 114 

crabroniphila, Aranea, 112 

crahroniphila, Metazygia, 112, 113* 

crewi, Aranea, 99 

crewi, Araneus, 99 

crewi, Metazygia, 99, 101* 

crewi, Singa, 99 

cunha, Metazygia, 146, 147* 

curari, Metazygia, 105*, 106 

dilatata, Aranea, 88 

dilatatus, Araneus, 88 

dubia, Aranea, 82 

dubia, Epeira, 82 

dubia, Metazygia, 82, 83* 

ducke, Metazygia, 138, 139* 

enabla, Metazygia, 95, 97* 

errans, Aranea, 140 

erratica, Epeira, 140 

erratica, Metazygia, 139*, 140 

erraticus, Araneus, 140 

fecunda, Epeira, 93 

floresta, Metazygia, 131*, 132 

genaro, Metazygia, 141, 143* 

genialis, Aranea, 118 

genialis, Epeira, 118 

genialis, Metazygia, 118, 119* 

genialis, Araneus, 118 

goeldii, Metazygia, 101*, 104 

gregalis, Epeira, 121 

gregalis, Metazygia, 121, 123* 



ikuruwa, Metazygia, 119*, 120 
incerta, Aranea, 93 
incerta, Epeira, 93 
incerta, Metazygia, 91*, 93 
incertus, Araneus, 93 
ipago, Metazygia, 86, 87* 
ipanga, Metazygia, 107, 109* 
isabelae, Metazygia, 96, 97* 
ituari, Metazygia, 127*, 128 
jamari, Metazygia, 97*, 99 
keyserlingi, Metazygia, 87*, 89 
lagiana, Metazygia, 134, 135* 
laticeps, Aranea, 117 
laticeps, Araneus, 117 
laticeps, Epeira, 117 
laticeps, Metazygia, 115*, 117 
lazepa, Metazygia, 109*, 110 
limonal, Metazygia, 127*, 128 
livida, Metazygia, 72 
lopez, Metazygia, 137, 139* 
loque, Metazygia, 135*, 136 
maculata, Epeira, 93 
manni, Metazygia, 121 
manu, Metazygia, 139*, 140 
mariahelenae, Metazygia, 131*, 132 
matanzas, Metazygia, 113, 113* 
Metazygia, 66 

moldira, Metazygia, 143*, 144 
mollybyrnae, Araneus, 94 
mollybyrnae, Singa, 94 
moraballicus, Araneus, 82 
moraballii, Aranea, 82 
moraballii, Epeira, 82 
mundula, Epeira, 118, 125 
mundula, Larinia, 118, 125 
mundula, Metazygia, 125 
mundulella, Aranea, 118 
mundulella, Metazygia, 115*, 118 
nigrocincta, Aranea, 133 
nigrocincta, Metazygia, 133, 135* 
nigrocinctus, Araneus, 133 
nobas, Metazygia, 101, 103* 
octama, Metazygia, 130, 131* 
oro, Metazygia, 147, 147* 
pallidula, Aranea, 94 
pallidula, Epeira, 94 
pallidula, Metazygia, 91*, 94 
pallidulus, Araneus, 88, 94 
palloides, Araneus, 94 
paquisha, Metazygia, 101*, 103 
pastaza, Metazygia, 91*, 92 
patiama, Metazygia, 85, 87* 
peckorum, Metazygia, 142, 143* 
pimentel, Metazygia, 83*, 85 
punctata, Zilla, 125 



Metazygia 'Levi 



151 



redfordi, Metazygia, 96, 97* 
rogenhoferi, Metazygia, 97*, 98 
rogenhoferi, Zilla, 98 
rogenhoferi, Zygiella, 98 
rothi, Metazygia, 145, 147* 
samiria, Metazygia, 138, 139* 
saturnino, Metazygia, 111, 113* 
sendero, Metazygia, 114, 115* 
serian, Metazygia, 108, 109* 
similis, Metazygia, 121 
simplicissima, Aranea, 94 
simplicissima, Epeira, 94 
simplicissimus, Araneus, 94 
souza, Metazygia, 135*, 137 
taman, Metazygia, 101*, 102 
tanica, Metazygia, 127*, 128 
tapa, Metazygia, 91*, 92 
tuceps, Eustala, 121 
uma, Metazygia, 115*, 116 
unguiformis, Metazygia, 72 
uraricoera, Metazygia, 108, 109* 



uratron, Metazygia, 109*, 111 
valentim, Metazygia, 143*, 144 
vaupes, Metazygia, 129, 131* 
vaurieorum, Metazygia, 101*, 102 
viriosa, Aranea, 126 
viriosa, Epeira, 126 
viriosa, Metazygia, 126, 127* 
viriosus, Araneus, 126 
voluptifica, Aranea, 125 
voluptifica, Epeira, 125 
voluptifica, Metazygia, 125, 127* 
voluptificus, Araneus, 125 
voxanta, Metazygia, 141, 143* 
wittfeldae, Epeira, 81 
wittfeldae, Metazygia, 81, 83* 
yobena, Metazygia, 123*, 124 
yucumo, Metazygia, 105*, 106 
zilloides, Aranea, 88 
zilloides, Araneus, 88 
zilloides, Epeira, 86 
zilloides, Metazygia, 86, 87* 



(US ISSN 0027-4100) 



aulletln of the 

Museum of 

Comparative 

Zoology 



Orb-Weaving Spiders 
Actinosoma, Spilasma, Micrepeira, Pronous, 
and Four New Genera (Araneae: Araneidae) 



HERBERT W. LEVI 



HARVARD UNIVERSITY 

CAMBRIDGE, MASSACHUSETTS, U.S.A. 



VOLUME 154, NUMBER 3 
31 JULY 1995 



(US ISSN 0027-4100) 



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dinidae (Mollusca: Bivalvia). 265 pp. 

3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- 
derms. 284 pp. 

4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the 
Present Day. 236 pp. 

5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology 
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6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. 

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Insects. {Bulletin of the M.C.Z., Vol. 108.) Reprinted 1971. 

Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. 

Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First Inter- 
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ORB-WEAVING SPIDERS ACTINOSOMA, SPILASMA, MICREPEIRA, 
PRONOUS, AND FOUR NEW GENERA (ARANEAE: ARANEIDAE) 



HERBERT W. LEVP 

Abstract. Actinosoma contains one species, A. 
pentacanihum. Spilasma has three species, two of 
which are new. Micrepeira has seven species, of which 
five are new. Pronous has 14 species, of which 11 are 
new. The new genera are Spinepeira, with one new 
species known only from a female; Hingstepeira, with 
four species, three of them new; Madrepeira, with 
one new species; and Tatepeira, with four species, 
three of which are new and of doubtful generic place- 
ment. There is one new generic synonymy and 11 
new synonymies of species names. Species of all these 
genera are known only from the Americas. 

The species of several of these genera make unusual 
webs. Actinosoma pentacanthum lives over water on 
emergent vegetation. Hingstepeira, Spilasma, and 
Micrepeira build retreats into the web. Madrepeira 
makes a ladderweb. Most of these spiders are rarely 
collected. The names used by R. W. G. Kingston in 
his observations on unusual webs are identified. 

INTRODUCTION 

This is one of a series of papers revising 
Neotropical araneid orb weavers. Previous 
papers are listed in Levi (1993a). Since 
1993, revisions of Lewisepeira Levi 
(1993b), Kaira O. P. -Cambridge (Levi, 
1993c), and Acacesia Simon (Glueck, 1994) 
have been published. 

This paper is dedicated to those collec- 
tors who contributed specimens with pho- 
tographs of their webs. They are W. Eber- 
hard, J. Coddington, H. Hofer, and R. L. 
C. Baptista. Without their photographs and 
notes on web structure, this revision would 
lack important data. The photographs 
made it possible to place many of the spe- 
cies named by Kingston (1932) in his book 
on Guyana spiders. Kingston did not in- 



' Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge, Massachusetts, 02138. 



tend to describe new species and did so 
only because no keys were available and 
nobody could name the spiders whose un- 
usual webs he had observed and illustrat- 
ed. (See Note, p. 209.) 

Among the species described here are 
many that probably are quite common but 
are rarely collected. Actinosoma occurs on 
emergent vegetation and is known to dive 
and run over water if disturbed. Spider 
collectors are usually not equipped with 
waders or boats, and the knowledge that 
schistosomiasis occurs in areas in the Lesser 
Antilles, northern Venezuela, coastal Su- 
rinam, and eastern Brazil (I AM AT, 1992) 
further deters arachnologists from splash- 
ing in ponds. 

Pronous may be equally difficult to col- 
lect. More than half of the available spec- 
imens were picked up by A. M. Chicker- 
ing. Pronous makes a small web in leaf 
litter and disappears into litter at the 
slightest disturbance. Presumably by 
spending much time sifting leaf litter, 
Chickering got more specimens than other 
collectors. 

The genera in this revision are not close- 
ly related. Actinosoma has a paramedian 
apophysis, the conductor is in the middle 
of the tegulum area, there are no spines 
on the median apophysis, and there is no 
distal hematodocha (Fig. 9); in the female 
the epigynum is a broad lobe (Figs. 1-3). 
The characters cited are synapomorphies 
with Alpaida O. P. -Cambridge (Levi, 
1988). Tatepeira, the other extreme, lacks 
a paramedian apophysis and has the con- 
ductor on the side of the tegulum, and the 
median apophysis has spines and well-de- 



Bull. Mus. Comp. Zool., 154(3): 153-213, July, 1995 153 



154 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



Table 1. 



Some characters of species belonging to genera revised here, with other genera 
included for comparison. 



ACT ALP SPI HIN MCA PRO MEP SPL ACA MAD AM TAT ARA NEC 

PME - - -[ + ]-[ + ]---- - - - - 

head - - --[ + ]-- + + - - - -- 

IV >i - - -- + +---- - - -- 

abd.shape [ + ] - [ + ] [ + ] [ + ] [ + ] [ + ] - - [ + ] [ + ] [ + ] 

abd.pttrn. + - - - - --[ + ][ + ]- - - - - 

abd. spins. + — *— - — _____ _ _ __ 

epig.lobe + + ++ + -[ + ]+ + - - - -- 

epig.scp. - - — — — ~[ + ]~~+ + + + + 

scape pnt. — — — — — ~[ + ]~~~" + + ~~ 

scape pck. ________ — —_ — + + 

3 dwarf - - ?- +*+- + -- -*- -*- 

3abdom - - ?- + +-[ + ]-- " - -- 

patella 1 1* ? 1 1 1 1 1 1 2 2* 2 2* 2 

M tooth -- ?--- + --+ + + + + 

Oagella __?-_- + --- + + -- 

PM + + ? + +*?-? + ----- 

C on edge - - ?- _*__-_+ + + + + 

DH __?____-_+ + + + + 

A + + ? + + --- + + + + + + 

A cplx. - + ?_-_---+ + + +- 

Genera: ACA, Acacesia; ACT, Actinosoma; ALP, Alpaida; AM, Amazonepeira; ARA, Araneus; HIN, 
Hingstepeira, MAD, Madrepeira, MCA, Micrathena; MEP, Micrepeira; NEO, Neoscona; PRO, Pronous; 
SPI, Spinepeira; SPL, Spilasma, TAT, Tatepeira. Revised genera are set in italic. 

Abbreviations: PME, posterior median eyes modified; head, cephalic region modified; IV > I, leg 4 longer 
than leg 1; abd.shape, abdomen shape modified; abd.pttrn., abdomen pattern modified; abd. spins., median 
posterior spines or tubercles present; epig.lobe, epigynum a lobe; epig.scp., epigynum with scape; scape pnt., 
scape tip pointed; scape pck., scape tip with distal pocket; 6 dwarf, male dwarfed and may lack endite tooth, 
coxal hook; S abdom, male abdomen modified with ventral scutum; patella, palpal patella with 1 or 2 setae; 
M tooth, median apophysis with tooth; flagella, median apophysis with flagella; PM, paramedian apophysis; 
C on edge, conductor on edge of tegulum; DH, distal hematodocha present; A, terminal apophysis present; 
A cplx., terminal apophysis branched. +, present; [ + ], an autapomorphy of genus; -, absent; *, variable in 
some species. 



veloped distal hematodocha (Fig. 224); the 
female has an epigynum with an annulat- 
ed scape (Figs. 217-219) and a pair of 
humps on the abdomen (Fig. 220). These 
characters all are synapomorphies with Ar- 
aneus and Aculepeira. Other genera here, 
Pronous, Micrepeira, and Spilasma, have 
some intermediate characters (Table 1). 

Readers of manuscripts often suggest 
that more information on apomorphies 
should be provided for construction of 
cladograms. But while some help can be 
given, the judgment about what is a good 
apomorphy is subject to reconsideration 
and relies on knowledge of other genera, 
some not yet revised. A study of relation- 
ships and useful apomorphies will be in 
order at the end of the revisions. Some 



examples of recent changes follow: Earlier 
it was thought that the presence of one or 
two macrosetae on the male palpal patella 
was useless in phylogeny and was not used. 
Now, however, despite many exceptions, 
it appears that the Araneus group of gen- 
era generally has two macrosetae, while 
those close to Alpaida have one (Table 1). 
In another example, the usual armature of 
araneid males, such as an endite tooth, a 
hook on the first coxa, or the modified sec- 
ond tibia seems to be lost in males that are 
very small compared to the female [with 
many exceptions, e.g., Acanthepeira Marx 
(Levi, 1976)]. At present, such armature is 
of little use in phylogeny. Another case of 
changed value concerns the pointed scape 
of the epigynum and the paired flagella of 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 155 



the median apophysis in the male palpus, 
formerly considered a synapomorphy for 
about five genera close to Araneus. But 
now these features are also noted in Mi- 
crepeira (Fig. 162, M in Fig. 179), which 
otherwise is close to Alpaida O. P. -Cam- 
bridge (Levi, 1988). At this stage the best 
information that can be provided is limited 
to the autapomorphies of individual re- 
vised genera. 

MATERIALS AND 
ACKNOWLEDGMENTS 

Specimens of the following collections 
were used, and I thank their curators for 
making the spiders available: 

AD A. Dean, College Station, Texas, 

United States 

AMNH American Museum of Natural 
History, New York, United States; 
N. Platnick, L. Sorkin 

BMNH Natural History Museum, Lon- 
don, England; P. Hillyard, F. 
Wanless 

CAS California Academy of Sciences, 
San Francisco, California, United 
States; W. J. Pulawski, D. Ubick, 
C. Griswold 

DU D. Ubick, San Francisco, Califor- 

nia, United States 

FMLT Fundacion Miguel Lillo, Tucu- 
man, Argentina; J. A. Corronca 

FSCA Florida State Collection of Ar- 
thropods, Gainesville, Florida, 
United States; G. B. Edwards 

HECO Hope Entomology Collections, 
Oxford University, Oxford, En- 
gland; I. Lansbury, M. Atkinson 

INPA Instituto Nacional de Pesquisas da 
Amazonia, Manaus, Est. Amazo- 
nas, Brazil; C. Magalhaes 

IRSNB Institut Royal des Sciences Natu- 
relles de Belgique, Brussels, Bel- 
gium; L. Baert 

JAK J. A. Kochalka, Ciudad Universi- 
taria, Paraguay 

MACN Museo Argentino de Ciencias Na- 
turales, Buenos Aires, Argentina; 
E. A. Maury 

MCN Museu de Ciencias Naturais, Fun- 



da^ao Zoobotanica do Rio Grande 
do Sul, Porto Alegre, Rio Grande 
do Sul, Brazil; E. H. Buckup 

MCP Museu de Ciencias, Pontificia 
Universidade Catolica do Rio 
Grande do Sul, Porto Alegre, Rio 
Grande do Sul, Brazil; A. A. Lise 

MCZ Museum of Comparative Zoolo- 
gy, Cambridge, Massachusetts, 
United States 

MECN Museo Ecuatoriano de Ciencias 
Naturales, Quito, Ecuador; L. 
Aviles 

MEG M. E. Galiano; Buenos Aires, Ar- 
gentina 

MHNG Museum d'Histoire Naturelle, Ge- 
neve, Switzerland; V. Mahnert 

MIUP Museo de Invertebrados, Univer- 
sidad de Panama, Panama; D. 
Quintero A. 

MNHN Museum National d'Histoire Na- 
turelle, Paris, France; J. Heur- 
tault, C. Bollard 

MUSM Museo de Historia Natural, Univ- 
ersidad Nacional Mayor de San 
Marcos, Lima, Peru; D. Silva D. 

MZCR Museo Zoologico de Universidad 
de Costa Rica, San Jose, Costa 
Rica; C. E. Valerio 

MZSP Museu de Zoologia, Universidade 
de Sao Paulo, Sao Paulo, SP, Bra- 
zil; P. Vanzolini, J. L. Leme 

MZUF Museum Zoologico de La "Spe- 
cola," Universita di Firenze, Flor- 
ence, Italy, L. Bartolozzi, S. Mas- 
cherini 

NMB Naturhistorisches Museum, Basel, 
Switzerland; E. Sutter, A. Hanggi 

NRMS Naturhistoriska Riksmuseet, Stock- 
holm, Sweden; T. Kronestedt 

PAN Polska Akademia Nauk, Warsza- 
wa, Poland; J. Proszynski, A. Slo- 
jewska, E. Kierych 

REL R. E. Leech, Edmonton, Alberta, 
Canada 

RLCB R. L. C. Baptista, Rio de Janeiro, 

Brazil 
SMF Forschungsinstitut Senckenberg, 
Frankfurt am Main, Germany; M. 
Grasshoff 



156 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



UCVC Universidad Central de Venezue- 
la, Caracas, Venzuela; J. Racenis 

USNM National Museum of Natural His- 
tory, Smithsonian Institution, 
Washington, D.C., United States; 
J. Coddington, S. F. Larcher 

ZMK Zoologisk Museum, Copenhagen, 
Denmark; H. Enghoff 

ZSM Zoologische Staatssammlung, Mu- 
nich, Germany 

In addition, I am grateful to J. A. Cor- 
ronca, who sent specimens and photo- 
graphs of Actinosoma habitat, W. Eber- 
hard for habitat and behavior information, 
I. Lansbury for information on O. P. -Cam- 
bridge types, T. Kronestedt for type in- 
formation on Actinosoma species, P. Van- 
zolini and S. F. Larcher for locality infor- 
mation, H. G. Fowler and his students for 
making their large Amazonian collections 
available, and R. L. C. Baptista, J. A. Cod- 
dington, W. Eberhard, and H. Hofer for 
photographs of webs. A. Johnston, L. Lei- 
bensperger, L. R. Levi, and W. Piel read 
the paper and made many improvements 
in the writing, and two anonymous readers 
also suggested changes. Especially helpful 
were H. Hofer, E. H. Buckup, and M. A. 
L. Marques, who read a draft of the manu- 
script meticulously and made me aware 
of many misprints. National Institutes of 
Health and the National Science Foun- 
dation grants supported the start of this 
research. Publications costs were covered, 
in part, by the Wetmore-Colles Fund. 

METHODS 

Most methods used in these studies are 
described in Levi (1993a). 

Internal genitalia of Pronous specimens 
were examined by removing the epigyn- 
um using minutennadeln (used otherwise 
for mounting small insects) mounted on a 
wooden stick. The epigynum was placed 
in Hoyer's medium between two cover- 
slips so that either side could be examined. 
An outline drawing was made with a dis- 
secting microscope, at the same magnifi- 
cation as the illustration of the epigynum. 
Details were added later. Finally, the epi- 



gynum was removed from the mount and 
placed in a 2-by-4-mm glass vial stoppered 
with cotton and kept with the female spec- 
imen. 

Palpi were spread out by pulling apart 
rather than by expanding, because pulling 
apart keeps all sclerites in similar positions 
and makes it easier to match and illustrate 
the labeled sclerites with those of the con- 
tracted palpus. This is of special impor- 
tance in genera related to Araneus that 
have well-developed distal hematodocha. 

The sizes of the eyes are expressed as 
diameters of the anterior median eyes; dis- 
tances between eyes of the anterior row 
are expressed as diameters of the anterior 
median eyes (in profile), and distances be- 
tween eyes of the posterior row are given 
as diameters of the posterior median eyes 
(in profile). The height of the clypeus, the 
distance between the anterior median eyes 
and the edge of the carapace, is expressed 
as diameter of the anterior median eyes 
(Levi, 1993a, fig. 28f). These measure- 
ments are approximate. 

To examine for the presence of a ta- 
petum, the eyes of Pronous specimens were 
first checked with reflected light. When 
no tapetum was found in the posterior me- 
dian eyes, the eye region of the carapace 
(of a female P. quintana) was cut off, 
cleared with methyl benzoate, and ex- 
amined more carefully. 



Actinosoma Holmberg 

Actinosoma Holmberg, 1883: 239. Type species Ac- 
tinosoma pentacanthum (Walckenaer) designated 
by Holmberg and by monotypy. The gender of the 
name is neuter (Bonnet, 1955: 156). 

Diagnosis. Actinosoma is readily sep- 
arated from all other Neotropical araneid 
genera by the five large sclerotized, orange 
spines on the abdomen of both female and 
male (Figs. 4-6, 10), an autapomorphic 
character. The genitalia are very similar 
to those of Alpaida, but the terminal 
apophysis of the male palpus is a simple 
structure (A in Fig. 9); in Alpaida it is 
subdivided. 



AcTiNosoMA, Spilasma, MiCREPEiRA, Pronous • Levi 157 




pentacanthum 



Map 1 . Distribution of Actinosoma pentacanthum. 



Actinosoma differs from Micrathena, 
with which it is commonly confused, by 
having the first pair of legs longer than the 
fourth, by lacking stridulating structures 
on the book-lungs (present in most Mi- 
crathena), and by lacking a square to rect- 
angular abdomen of the male. The abdo- 
men of male Actinosoma has spines (Fig. 
10) just as in the female; male Micrathena 
lack them. 

Description. Female. Abundant black 
pigment between median eyes (Fig. 7). 
Width of cephalic region behind eyes 
slightly more than half thoracic width. 
Epigynum with projecting, rounded shelf, 
its tip a rounded lobe (Figs. 1-3) as in 
Alpaida. 

Male. Width of eye region behind eyes 
about half the width of the thoracic region 
(Fig. 11). Palpus with conductor in middle 
of tegulum (Fig. 8, C in Fig. 9); tegulum 
without sclerites on distal, ventral side 
(right, in left palpus at 3 hr in Fig. 8, T 
in Fig. 9). Paramedian apophysis (PM in 
Fig. 9) is a separate, lightly sclerotized, 
L-shaped sclerite attached by soft stalk to 
conductor. Median apophysis cup-shaped 



(M in Fig. 9; Levi, 1988, fig. 10), as it often 
is in Alpaida. Embolus supported by para- 
median apophysis; its tip on conductor (E 
in Fig. 9). Terminal apophysis a narrow 
lobe above embolus (A in Fig. 9). 

Relationship. The lobe of the epigyn- 
um of the female resembles that of Al- 
paida (see Levi, 1988) and Micrathena (see 
Levi, 1985); it is a synapomorphy. The 
palpus has the conductor in the center of 
the tegulum (C in Fig. 9) as in Alpaida 
and Micrathena, and there is a parame- 
dian apophysis (PM in Fig. 9) as in these 
two genera [it may be missing in some 
Micrathena (Levi, 1985)], another syna- 
pomorphy of various genera. Unlike Ara- 
neus, Neoscona, Madrepeira, Tatepeira, 
and others (Figs. 213, 224), the median 
apophysis lacks sharp spines or flagella-like 
extensions. 

Natural History. Actinosoma penta- 
canthum lives on emergent aquatic veg- 
etation above the water surface in the mid- 
dle of ponds and puddles. 

Distribution. Only one species is wide- 
spread in South America (Map 1). 

Misplaced. Actinosoma heteracantha 



158 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



Mello-Leitao, 1943, is Wagneriana het- 
eracantha (Mello-Leitao) (Levi, 1991c). 

Actinosoma riscoi Archer (1971: 158, 
figs. 3, 4, 9) is Rubrepeira rubronigra (Mel- 
lo-Leitao) (Levi, 1991a). 



Actinosoma pentacanthum (Walckenaer) 
Figures 1-12; Map 1 

Plectana pentacantha Walckenaer, 1841: 170. Lo- 
cality of specimens not known but may have been 
from Cayenne (French Guiana); specimens lost. 

Acrosoma stelligeriim Thorell, 1860: 301; 1868: 26. 
Origin of specimen unknown and type specimens 
not in NRMS, lost. Specimens determined by Tho- 
rell from Corumba, Mato Grosso [Mato Grosso do 
Sul], Brazil, in NRMS, examined. Similarity of de- 
scription of stelligerum and pentacantha first no- 
ticed by Butler (1873). Name first synonymized by 
Holmberg (1883). 

Acrosoma pentacanthum: — Butler, 1873: 428. 
Holmberg, 1883: 237. 

Acrosoma pulcherrima Holmberg, 1876: 143 (p. 21 
in a separate publication, not seen). Specimens from 
Puerto Obligada [Vuelta de Obligado, Prov. Buenos 
Aires, 33°35'S, 59°49'W (USDI, 1968)], Argentina, 
lost. First synonymized by Holmberg (1883). 

Actinosoma pentacanthum: — Holmberg, 1883: 239. 
Roewer, 1942: 778. Bonnet, 1955: 156. 

Cyrtarachne quinquespinosa Keyserling, 1892: 55, 
pi. 3, fig. 44, 9, S. Five immature and five female 
syntypes from Espirito Santo, Brazil, in USNM, 
examined. Goldi, 1892: 227. First synonymized by 
Simon (1895). 

Araneus pentacanthus: — Simon, 1895: 819, fig. 869, 
2. Badcock, 1932: 27. 

Araneus pentacantha: — Zapfe, 1957: 29, fig. 3, 9, 5. 

Islote. Acrosoma Perty, 1833, with the 
type species A. swainsoni, is a subjective 
synonym of Micrathena (Levi, 1985). Cyr- 
tarachne Thorell (1868: 10) was a name to 
replace Cyrtogaster Keyserling (1864: 80) 
[the name preoccupied by Walckenaer 
(1833) for a hymenopteran]. The genus has 
the type Cyrtogaster grubei, by mono- 
typy, from Mauritius, Indian Ocean, for 
an araneid with the carapace high in tho- 
racic region and abdomen triangular, wid- 
er than long, pointed behind, and having 
four dorsal tubercles. 

According to Holmberg (1883: 227), the 
information of 1876 was published on page 
143, 1876 (date from Holmberg); it is page 
21 on a photocopy of the journal that was 
available. 



The syntypes of Cyrtarachne quin- 
quespinosa were unexpectedly found in 
unsorted collections of the USNM. The 
USNM specimens were recognized as Key- 
serling specimens because they were la- 
beled "Espirito Santo, Cyrtarachne 
5-spinosa Keys.", on a label, 24 by 30 mm, 
having toothed perforations on all four sides 
and a blue frame, a short distance inside 
from the white teeth. This is a typical Key- 
serling label. A second plain old label in 
the vial read "Espirito Santo, Rio Minas". 

Description. Female from Puerto Napo, 
Ecuador. Carapace orange, eye region 
black. Chelicerae orange, distally black. 
Labium, endites black. Sternum, coxae or- 
ange; legs without rings, dusky black, fem- 
ora darkest. Dorsum of abdomen with or- 
ange and black, the black including paired 
white patches (Fig. 4); venter black with 
a pair of white lines (Fig. 6). Eyes sub- 
equal. Anterior median eyes their diam- 
eter apart, 2 diameters from laterals. Pos- 
terior median eyes 0.8 diameter apart, 2.5 
diameters from laterals. Ocular quadran- 
gle slightly narrower behind than in front, 
slightly longer than wide. Height of clyp- 
eus equals 1.1 diameters of anterior me- 
dian eye. Abdomen with five prominent 
spines (Figs. 4-6). Total length 8.0 mm. 
Carapace 3.2 mm long, 2.7 wide, 1.5 be- 
hind lateral eyes. First femur 3.4 mm, pa- 
tella and tibia 3.9, metatarsus 2.7, tarsus 
1.5. Second patella and tibia 3.3 mm, third 
2.3. Fourth femur 3.4, patella and tibia 
3.7, metatarsus 2.5, tarsus 1.1. 

Male from Puerto Napa, Ecuador. Col- 
oration as in female (Fig. 10). Posterior 
median eyes 0.8 diameter of anterior me- 
dians, laterals 0.8 diameter. Anterior me- 
dian eyes 0.7 diameter apart, 1.4 diameters 
from laterals. Posterior median eyes 0.6 
diameter apart, 2 diameters from laterals. 
Ocular quadrangle narrower behind, 
slightly longer than wide. Height of clyp- 
eus equals 1.2 diameters of anterior me- 
dian eye. Endite with tooth facing a tu- 
bercle on the coxa and a tooth on the prox- 
imal end of the palpal femur. Palpal pa- 
tella with one macroseta. First coxa with 
hook. Second tibia thicker than first but 



ACTINOSOMA, Spilasma, MiCREPEiRA, Pronous • Levi 159 




Figures 1-12. Actinosoma pentacanthum (V\Ja\ckenaer). 1-7, female. 1-3, epigynum. 1 , ventral. 2, posterior. 3, lateral. 4, dorsal. 
5, lateral. 6, abdomen ventral. 7, eye region and chelicerae. 8-12, male. 8, left palpus, mesal view. 9, palpus pulled apart. 10, 
dorsal. 11, carapace. 12, eye region, chelicerae and right palpus. 

Figures 13-20. Spinepeira schlingeri n. sp., female. 13-15, epigynum. 13, ventral. 14, posterior. 15, lateral. 16, dorsal. 17, 
lateral. 18, carapace. 19, carapace and chelicera. 20, eye region and chelicerae. 



Abbreviations. A, terminal apophysis. C, conductor. E, embolus. M, median apophysis. PM, paramedian apophysis. R, radix. T, 
tegulum. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



160 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



without macrosetae. Abdomen as in fe- 
male (Fig. 10). Total length 6.7 mm. Car- 
apace 3.1 mm long, 2.3 wide; 1.1 behind 
lateral eyes. First femur 3.4 mm, patella 
and tibia 3.8, metatarsus 2.4, tarsus 1.1. 
Second patella and tibia 3.1 mm, third 2.0. 
Fourth femur 3.3 mm, patella and tibia 
3.4, metatarsus 2.1, tarsus 1.0. 

Note. Males and females were collect- 
ed together. 

Variation. Total length of females 7.0 
to 9.7 mm, males 6.0 to 7.0 mm. The bases 
of the five spines vary in width in different 
specimens. Specimens from the Napo Riv- 
er, Ecuador, were used for the illustrations, 
except for the dorsal view of the female 
(Fig. 4), which was made from a specimen 
from Mato Grosso, Brazil (NRMS). Re- 
cently collected specimens had bright or- 
ange, black, and white coloration. 

Diagnosis. The abdomen with five 
sclerotized spines in both males and fe- 
males (Figs. 4, 5, 10), and the orange, white, 
and black coloration, separates the species 
from all other araneids. 

Natural History. Holmberg (1876) 
captured the spiders along the sides of a 
brook. Goldi (1892) found this red spider 
in earthen organ-pipe tubes of wasps, 
which were attached to walls and window 
frames in Santo Antonio de Padua, Est. 
Rio de Janeiro, Brazil. The Colombian 
specimens were taken from a Sceliphron 
wasp nest. All others were from ponds or 
streams. Specimens from the Napo River, 
Ecuador, were collected by sweeping be- 
low water and water surface in middle of 
a swamp. A female from Manaus, Brazil, 
came from Paspalum grass, which forms 
vast floating meadows in Amazonian riv- 
ers. Badcock (1932: 27) recorded the spe- 
cies as "common everywhere on surface 
of swamps" in Paraguayan Chaco. Bad- 
cock's locality is described by Carter (1928: 
97), the collector, as an island surrounded 
by swamps with daily maximum temper- 
atures of 90° F (42° C), mean 85° F (30° 
C), with surface layer of water at 108° F 
(43° C). A specimen from Paraguay 
(MHNG) was collected in a swamp. Cor- 



ronca (personal correspondence) reported 
webs in flooded grasslands, in tropical Ban- 
ado areas. When the web is disturbed, the 
spider dives or walks on the water surface. 
This species is probably just as common in 
rain forest ponds as in wet areas of dry 
regions, but collectors are more likely to 
be attracted to marshes and ponds in dry 
areas. Galiano reported in collecting data 
from Santa Fe Province, Argentina, that 
the spider makes a horizontal web on float- 
ing plants. 

Distribution. Amazon area to Buenos 
Aires Province, Argentina (Map 1). 

Specimens Examined. COLOMBIA Amazonas: 

25 km N Leticia, 29 (MCZ). ECUADOR Napo: 10 
km E Puerto Napo, S side Rio Napo, 69, 6(5, 14 imm. 
(MCZ). PERU "northern PERU", Reanrer [local. ?], 
49 (AMNH). Loreto: Parinari Canyon, Rio Samiria, 
19, 1(5, imm. (AMNH). Hudnuco: Monzon Valley, 
Tingo Maria, 19, 13 (CAS). BRAZIL Amazonas: Porto 
America [05°10'S on Rio Madeira or OySQ'S on Rio 
Purus, Vanzolini, personal communication], 19 
(AMNH); Rio Autaz, Santa Amelia, 19 (NRMS); Rio 
Autaz, Cururuzinho, \$ (NRMS); Manaus, Bilhares, 

26 (NRMS); Manaus, Flores, 19 (NRMS); Rio Negro, 
19 (MZSP); Manaus, Igarape, km 30, M-Caracarai, 19 
(INPA); Manaus, 19 (INPA). Mato Grosso: Pocone, 
km 11 Transpantaneira, imm. (MCZ). Rio de Janeiro: 
Santo Antonia de Padua (Goldi, 1892). Sao Paulo: 
Reprisa, Sao Jose de Rio Preto, 19, U (MZSP). BO- 
LIVIA El Beni: Yacuma, Espirito, 19 (ZSM); Rosario, 
29 (USNM); 19 (M. R. Lopez, USNM). La Paz: Reves, 
14°03'S, 68°01'W (Zapfe, 1957); upper lake on c'ha- 
caltaya, 19 (ZSM). PARAGUAY San Pedro: 2 km NW 
Lima, imm. (MHNG). Presidente Hayes: Transcha- 
co, km 320, 29 (IRSNB); Makthlawaiya (Badcock, 
1932). Paraguari: nr. Ybytymi, 119, 5(5 (MCZ). Gtiaird: 
As. Tacuara. Central: Esterus del San Lorenzo, 19, 
1(5 (CAS); Asunci6n, 19 (MCZ); Villa del Maestro, San 
Lorenzo, 29, 1(5, 2 imm. (MHNG). Caaguazu: 20 km 
N Cnl. Oviedo, 19 (MHNG). ARGENTINA Formosa: 
Reserva Ecologica El Bagual, 59, 1,5 (FMLT, MCZ). 
Chaco: Selva del Rio de Oro, 19, 1(5 (MEG). Santa 
Fe: Arroyo El Toba, 19, 13 (MEG); Tortagal, 59, 1(5 
(FMLT). Buenos Aires: San Nicolas, Baradero, Pilar 
(Holmberg, 1883); Buenos Aires, 19 (MCZ); Delta de 
Parana, Arroyo Espera, 13 (MEG); Escobar, 1(5 
(MACN); Campana, 29 (CAS); Tigre, 29 (BMNH). 



Spinepeira new genus 

Type species. Spinepeira schlingeri. The name is an 
arbitrary combination of letters attached to "epeira". 
The name of the genus is feminine. 



AcTiNOSOMA, Spilasma, MicREPEiRA, Pronous • Levi 



161 



Diagnosis. This genus, which is close 
to Alpaida, differs by having a pair of long 
projections on the abdomen, one anterior 
median tubercle, and two posterior me- 
dian tubercles (Figs. 16, 17). 

Relationship. The two posterior me- 
dian tubercles represent a synapomorphy 
found in numerous genera including Al- 
paida, Acanthepeira, Eriophora, Eustala, 
and Wagneriana. With the exception of 
Eustala, these genera have males with a 
paramedian apophysis in the palpus. Ad- 
ditional characters placing the genus close 
to Alpaida are the black pigmentation be- 
tween the median eyes (Figs. 18-20) and 
lack of setae on the carapace. 



schli 




Map 2. Distribution of Spinepeira schlingeri. 



Spinepeira schlingeri new species 
Figures 13-20; Map 2 

Holotype. Female holotype from Monzon Valley, 
Tingo Mana, Depto. Huanuco, Peru, 18 Dec. 1954 
(E. I. Schlinger, E. S. Ross), in CAS. The species is 
named after the collector, entomologist E. I. Schlin- 
ger. 

Description. Female holotype. Cara- 
pace yellow-white, with a longitudinal 
black line through the middle, area be- 
tween median eyes black (Fig. 18). Che- 
licerae, labium, endites, sternum black. 
Legs yellow-white. Dorsum of abdomen 
with pairs of black and white patches (Figs. 
16, 17); venter with genital area and spin- 
nerets black and a large black patch be- 
tween (Fig. 17). Posterior median eyes 1.5 
diameters of anterior medians, anterior 
laterals 1 diameter, posterior laterals 1.5 
diameters. Anterior median eyes 1.3 di- 
ameters apart, 2 diameters from laterals. 
Posterior median eyes their diameter apart, 
1.5 diameters from laterals. Ocular quad- 
rangle wider behind than in front. Height 
of clypeus equals 1 diameter of anterior 
median eye. Abdomen with a pair of long 
projections and one pair of lateral tuber- 
cles, an anterior median tubercle and two 
posterior median tubercles (Figs. 16, 17). 
Total length 5.5 mm. Carapace 1.7 mm 
long, 1.5 wide, 0.9 wide behind lateral eyes. 
First femur 2.2 mm, patella and tibia 2.7, 



metatarsus 2.0, tarsus 0.7. Second patella 
and tibia 2.0 mm, third 1.3, fourth 1.8. 

No additional specimens have been 
found. 

Hingstepeira new genus 

Type species. Epeira folisecens Hingston, 1932. The 
name is an arbitrary combination of letters linking 
part of explorer Hingston's name to "epeira". The 
gender of the name Hingstepeira is feminine. 

Diagnosis. Hingstepeira is separated 
from most other araneid genera by the 
shape of the abdomen: elongate, oval, usu- 
ally widest at the posterior half (Fig. 24), 
with the spinnerets overhung by an ante- 
rior to the posterior tip of the abdomen 
(Figs. 25, 39, 44, 52). It differs from Singa 
by having less black coloration in the me- 
dian eye region (Figs. 26-28) and by the 
genitalia, especially the palpus, which in 
Hingstepeira has a prominent paramedian 
apophysis (PM in Fig. 30). In the close 
spacing of the posterior median eyes (Fig. 
38) and the shape and color of the abdo- 
men, Hingstepeira is similar to Metazy- 
gia, but the paramedian apophysis of the 
male palpus (PM in Fig. 30) separates the 
genera. Unlike other male araneids, except 
the males of Metazygia gregalis (O. P.- 
Cambridge), M. benella Levi, and M. yob- 
ena Levi, the Hingstepeira male has the 



162 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



distal end of the chelicerae modified and 
with a frontal tooth (Fig. 33). The Meta- 
zygia species have a widened fang faced 
by a wide anterior tooth (Levi, 1994, fig. 
261). 

There are other genera in which the 
females have an elongate abdomen ex- 
tending beyond the spinnerets. Hingste- 
peira lacks the thick first pairs of legs of 
the southeast Asian Perilla Thorell, 1895, 
and Hingstepeira's abdomen is wider. 
[Perilla has been placed by Brignoli (1983) 
in Nephilinae, which belongs in the family 
Tetragnathidae.] Hingstepeira differs from 
the Mediterranean and African Nemos- 
colus Simon, 1895, by having a parame- 
dian apophysis in the palpus. 

Hingstepeira differs from Arachnura 
Vinson, 1863, which has a similar web, by 
lacking the narrow tail; also, the male 
Hingstepeira lacks a sclerotized shield on 
the abdomen. Hingstepeira differs from 
the Indopacific Milonia Thorell, 1890, in 
lacking Milonia's silver coloration. (Mil- 
onia may be a tetragnathid.) 

Description. Cephalothorax glabrous, 
orange to orange-brown, median eye re- 
gion black, without marks; cephalic area 
darker than thorax. Cephalic region in both 
sexes wide behind lateral eyes, more than 
half width of thoracic region (Figs. 26, 32). 
Median ocular quadrangle narrower be- 
hind than in front. Abdomen oval, longer 
than wide, widest in middle or posterior 
(Fig. 24). Abdomen coloration variable; all 
but one species (H. isherton) with trans- 
verse black rectangle between epigynum 
and spinnerets (Figs. 25, 44, 52). Epigyna 
variable, often with a smooth, bulbous pro- 
jection (Figs. 21, 35). 

The male of only one species (H. foli- 
secens) is known. Male carapace slightly 
smaller than that of female (Figs. 32-34). 
Endite with tooth (Fig. 34) facing tubercle 



on palpal femur. First coxa with hook on 
distal margin (Fig. 34). Palpal patella with 
one macroseta (Fig. 34). First and second 
legs with macrosetae. 

Relationship. The presence of the 
paramedian apophysis of the male palpus 
(PM in Fig. 30), the absence of a distal 
hematodocha, the presence of the conduc- 
tor (C) in the middle of the tegulum, and 
the free section of the tegulum on the up- 
per right in the left palpus (T in Fig. 30) 
place the genus close to Alpaida. 

Natural History. The Hingstepeira 
web resembles that of Arachnura by hav- 
ing a coiled up leaf as retreat in the upper 
vertical radius (Pi. 1). 

Distribution. There are four species, all 
South American, found in the Amazon area 
and Guyanas. 

Separating Species. The species differ 
in the markings on the abdomen. Males 
and females of the same species have sim- 
ilar markings. 



Key to Species of Hingstepeira 

The only male known is that of H. foHsecens (Figs. 
29-34), 

1. Epigynum with median knob-shaped 

structure (Figs. 21-23, 35-37) 2 

Epigynum otherwise, without median knob- 
shaped structure 3 

2(1). In ventral view of epigynum, round struc- 
ture facing posteriorly (Fig. 21); abdo- 
men with a dorsal, median, posterior 

black patch (Fig. 24) folisecens 

In posterior view of epigynum, round 
structure facing anteriorly (Fig. 35); ab- 
dominal pattern is a series of facing 
brackets (Fig. 38) isherton 

3(1). In ventral view of epigynum, the median 
area, anterior of a transverse bar, is con- 
cave (Fig. 40); abdomen with black bands 

on sides (Fig. 43) arnolisei 

In ventral view epigynum convex (Figs. 45, 
48); abdomen with two dorsal black bands 
(Fig. 51 ) dimona 



Plate 1 . Hingstepeira folisecens (Hingston). Upper web with 35 cm diameter, hub about 140 cm above forest floor. Lower web 
about 35 cm diameter, hub 160 cm above forest floor (photos, H. Hofer). 



AcTiNOSOMA, Spilasma, Micrepeira, Pronuus • Levi 163 




164 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



folisecens 

SURINAM 




'>\>\^r^^^'^'^ ' '-^GUYANA 
'XJ~X'^~ '' VENEZUELA y ) 




O arnolisei 
# dimona 
A isherton 



Map 3. Distribution of Hingstepeira species. 



Hingstepeira folisecens (Hingston) 
Plate 1; Figures 21-34; Map 3 

Epeira folisecens Hingston, 1932; 364, figs. 51, 52, 
webs. Female from Moraballi Creek, 2 mi [3.2 km] 
east of Essequibo River, 12 mi [19 km] south of 
Bartica, Guyana, lost. Not in BMNH, HECO. 

Larinia bristowei Mello-Leitao, 1940: 180, figs. 6, 7, 
S. Male holotype from Moraballi Creek, Essequibo 
River, Guyana, in BMNH, examined. Brignoli, 1983: 
272. NEW SYNONYMY. 

Aranea folisecens: — Roewer, 1942: 842. 

Araneus folisecens: — Bonnet, 1955: 504. 

Note. Hingston's (1932, figs. 51, 52) il- 
lustration of the web of this species resem- 
bles the web recently photographed by H. 
Hofer (Pi. 1). Also, Hingston described the 
"abdomen . . . grey-brown darkening to 



apex which is nearly black, . . . with spin- 
nerets at about the centre, area behind 
spinnerets brown, a quadrate black patch 
in front of spinnerets. . . . Total length 5 
mm" (p. 364). 

The type localities of Epeira folisecens 
and Larinia bristowei are the same. 

Description. Female from Taruma 
Mirim, Amazonas State, Brazil. Carapace 
orange, lightest posteriorly, area between 
eyes black. Chelicerae, labium, endites or- 
ange. Sternum, coxae light orange. Legs 
dusky orange. Dorsum of abdomen light 
gray, posterior end with a black patch (Fig. 
24); venter with a dark, dusky trapezoidal 
patch between spinnerets and epigynum 
(Fig. 25). Posterior median eyes 0.8 di- 
ameter of anterior medians, laterals 0.5 
diameter. Anterior median eyes 0.7 di- 
ameter apart, 1.3 diameters from laterals. 
Posterior median eyes 0.2 diameter apart, 
2.5 diameters from laterals. Height of 
clypeus equals 0.4 diameter of anterior 
median eye. Abdomen elongate, widest 
behind middle (Fig. 24). Total length 5.7 
mm. Carapace 2.1 mm long, 1.4 wide, 1.1 
behind lateral eyes. First femur 1.3 mm, 
patella and tibia 1.8, metatarsus 0.7, tarsus 
0.5. Second patella and tibia 1.6 mm, third 
1.4, fourth 1.6. 

Male holotype of Larinia bristowei. 
Carapace orange, black around eyes. Che- 
licerae, labium, endites, sternum, legs or- 
ange. Dorsum of abdomen whitish with 
three pairs of dusky patches and a black 
patch above spinnerets (Fig. 31); venter 
light gray. Posterior median eyes 0.6 di- 
ameter of anterior medians, laterals 0.5 
diameter. Anterior median eyes 0.6 di- 
ameter apart, slightly more than their di- 
ameter from laterals. Posterior median eyes 
0.6 diameter apart, 3 diameters from lat- 
erals. Height of clypeus slightly less than 
diameter of anterior median eye. Abdo- 
men oval (Fig. 31). First and second legs 
armed with macrosetae; second thicker 
than first. Total length 3.7 mm. Carapace 
1.9 mm long, 1.4 wide. First femur 1.4 
mm, patella and tibia 1.9, metatarsus 1.2, 
tarsus 0.6. Second patella and tibia 1 .6 mm, 
third 1.1, fourth 1.5. 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 165 



Note. Males and females have been col- 
lected together and share similar markings 
on the abdomen, although males are darker 
than females. 

Variation. Total length of females 4.7 
to 7.4 mm, males 3.1 to 4.6. The sides of 
the abdomen in ventral view may be dark. 
The posterior patch on the abdomen and 
ventral markings may be almost absent, or 
gray to black. Illustrations were made from 
a female from near Manaus, Brazil (Figs. 
21-28), from the male holotype of Larinia 
bristowei (Fig. 29), and from specimens 
from near Manaus, Brazil (Figs. 30-34). 

Diagnosis. The elongate abdomen and 
the characteristic markings, a black patch 
on the posterior end (Figs. 24, 25, 31), sep- 
arate this species from all other araneids. 

Natural History. Hingstepeira folise- 
cens makes a vertical web with a curled 
leaf retreat attached to an upper radius 
(Kingston, 1932, figs. 51, 52; Pi. 1). Webs 
have been found in the interior and border 
of forests. They were found in igapo (pe- 
riodically flooded forests) near Manaus and 
on the Manaus-Itacotiara in campina for- 
est highway and in terra firma forest near 
Manaus. Adult males were common only 
during February and March. 

Distribution. Guy anas and Brazilian 
Amazonian area (Map 3). 

Specimens Examined. SURINAM Saramacca: 
Voltzberg-Raleighvallen Nature Reserve, 04°32'N, 
56°32'W, 8 Feb. 1982, 19 (D. Smith Trail, MCZ). 
Merowijne: Anapaike Village, Lawa River, 8-29 Nov. 
1963, 12 (E. Malkin, AMNH). Brokopondo: Browns 
Berg, 5°N, 55''27'W, 20 Feb. 1982, 12 (D, Smith Trail, 
MCZ). FRENCH GUIANA Cayenne: Montagnes 
Kaw, nr. Camp Caiman, ca. 27 km SE Paura, 04°33'N, 
52°09'W, 100-300 m, 25 Aug. 1988, 12; 1 Sept. 1988, 
1(5 (S. Marshall, USNM). BRAZIL Amazonas: Taruma 
Mirim, igapo, Manaus, 24 Apr. 1984, 12 (J. Adis, 
INPA); 25 Feb. 1987, 12 (H. Hofer, INPA); 5 Dec. 
1987, 12 (H. Hofer, INPA); km 192, highway Manaus 
to Itacotiara, 18 Apr. 1987, 12 (H, Hofer, INPA); 80 
km N Manaus, 9 Mar. 1989, 16 (H. Fowler, MCZ); 
Colosso Reserve, 80 km N Manaus, 18 Jan. 1989, 12, 
1(3; Jan. 1989-June 1991, 632, 12(5; Reserva Cabo Frio, 
80 km N Manaus, Oct. 1989-June 1991, 132, 3(5; 13 
May 1992, 12; Reserva Dimona, 80 km N Manaus, 
1989-1992, 32; Mar.-June 1991, 182; Reserva Km 41, 
80 km N Manaus, Mar.-May 1991, 72; Reserva C. de 
Powell, 80 km N Manaus, 32; Reserva Florestal, 80 
km N Manaus, 19 Feb. 1991, 32; Reserva Gaviao, 21 
Feb.-21 Oct. 1991, 32; (all H. Fowler, R. S. Vieira, 



E. Venticinque, MCZ); Reserva Porto Alegre, 80 km 
N Manaus, 1989-1992, 12; 27 May 1992, 12 (H. G. 
Fowler, MCZ). 



Hingstepeira isherton new species 
Figures 35-39; Map 3 

Holotype. Female holotype from Isherton, Guyana, 
10 Nov. 1937 (W. G. Hassler), in AMNH. The spe- 
cific name is a noun in apposition after the locality. 

Description. Female holotype. Cara- 
pace orange, cephalic region darkest. Che- 
licerae orange-brown. Labium, endites 
dark brown. Sternum orange, sides dusky. 
Legs orange. Dorsum of abdomen with 
white pigment spots and five pairs of 
brackets (Fig. 38); venter black around 
spinnerets fading into surrounding area, 
except for clear border posteriorly (Fig. 
39). Posterior median eyes 0.8 diameter of 
anterior medians, laterals 0.6 diameter. 
Anterior median eyes 0.8 their diameter 
apart, 1.8 diameters from laterals. Poste- 
rior median eyes 0.2 diameter apart, 4 di- 
ameters from laterals. Height of clypeus 
equals 0.2 diameter of anterior median eye. 
Abdomen elongate oval, widest posteriorly 
(Fig. 38). Total length 7.0 mm. Carapace 
3.4 mm long, 2.1 wide, 1.5 behind lateral 
eyes. First femur 2.3 mm, patella and tibia 
3.0, metatarsus 2.0, tarsus 0.8. Second pa- 
tella and tibia 2.7 mm, third 1.6, fourth 
2.3. 

Note. This female has a dorsal abdom- 
inal pattern similar to Metazygia, and its 
eyes resemble those of Metazygia (Levi, 
1994). The epigynum also resembles that 
of Metazygia goeldii Levi. The main 
Hingstepeira character is the elongate ab- 
domen (Fig. 38). Only finding a male will 
ascertain the placement. 

Diagnosis. The shape of the epigynum 
(Figs. 35-37) and the dorsal abdominal 
pattern (Fig. 38) separate this species from 
other Hingstepeira. 



Hingstepeira arnolisei new species 
Figures 40-44; Map 3 

Holotype. Female holotype from Ilha de Maraca, 
Rio Uraricoera, Roraima Stae, Brazil, 5 Dec. 1987 



166 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



(A. A. Lise), in MCN no. 20062. The species is 
named after the collector. 

Description. Female holotype. Cara- 
pace light orange-yellow. Chelicerae, la- 
bium, endites, sternum, legs yellowish 
white. Dorsum of abdomen with four lon- 
gitudinal white bands, sides black anteri- 
orly and posteriorly (Fig. 43); venter with 
a tranverse black rectangle between epi- 
gynum and spinnerets, sides black (Fig. 
44). Posterior median eyes 0.8 diameter of 
anterior medians, anterior laterals 1.2 di- 
ameters, posterior laterals 0.8 diameter. 
Posterior median eyes oval. Anterior me- 
dian eyes their diameter apart, 2.3 diam- 
eters from laterals. Posterior median eyes 
their narrow diameter apart, 5 diameters 
from laterals. Height of clypeus equals di- 
ameter of anterior median eye. Abdomen 
elongate, 1.8 times as long as wide (Fig. 
43). Total length 8.0 mm. Carapace 3.1 
mm long, 2.3 wide, 1.8 behind lateral eyes. 
First femur 1.9 mm, patella and tibia 3.0, 
metatarsus 1.8, tarsus 0.9. Second patella 
and tibia 2.5 mm, third 1.7, fourth 2.5. 

Diagnosis. Hingstepeira arnolisei dif- 
fers from other Hingstepeira species by 
having a concave area on the anterior of 
the epigynum (Fig. 40) and by the lateral, 
black banding of the abdomen (Figs. 43, 
44). 



Hingstepeira dimona new species 
Figures 45-52; Map 3 

Holotype. Female holotype from Reserva Dimona, 
80 km north of Manaus, Amazonas State, Brazil, in 
forest, 15 May 1991 (H. Fowler, R. S. Vieira, E. 
Venticinque), in MCN no. 25543. The specific name 
is a noun in apposition after the type locality. 



Description. Female holotype. Cara- 
pace orange, darkest in midline. Chelic- 
erae dark orange. Labium, endites, ster- 
num orange. Legs orange with indistinct 
dark orange rings. Dorsum of abdomen 
with two black bands, midline and sides 
of bands white (Fig. 51); venter with a 
black transverse rectangle between epi- 
gynum and spinnerets, sides and posterior 
black (Fig. 52). Posterior median eyes 0.8 
diameter of anterior medians, laterals 0.6 
diameter. Anterior median eyes 0.6 di- 
ameter apart, 2 diameters from laterals. 
Posterior median eyes 0.3 diameter apart, 

2.7 diameters from laterals. Height of 
clypeus equals 0.5 diameter of anterior 
median eye. Abdomen oval (Fig. 51). Total 
length 5.2 mm. Carapace 2.5 mm long, 1.8 
wide, 1.3 behind lateral eyes. First femur 

1.8 mm, patella and tibia 2.3, metatarsus 
1.4, tarsus 0.7. Second patella and tibia 1.9 
mm, third 1.1, fourth 1.8. 

Variation. Total length of females 4.7 
to 6.0 mm. The borders of the posterior 
lateral plates of the epigynum (Figs. 46, 
49) are variable. Figures 45-47, 51, and 
52 were made from the female holotype, 
and Figures 48-50 from the specimen from 
Reserva Ducke. 

Diagnosis. The domed structure of the 
epigynum (Figs. 45-50) and the dorsal 
black bands of the abdomen (Fig. 51) sep- 
arate this species from other Hingstepeira. 
The species may belong to Metazygia and 
be close to M. laticeps (Kevserling) (Levi, 
1995, figs. 226-230). 

Natural History. All females were col- 
lected in forest. 

Paratypes. BRAZIL Amazonas: Reserva Dimona, 
80 km N Manaus, 26 Mar. 1991, 22; 17 Mav 1991, 



Figures 21-34. Hingstepeira folisecens (Hingston). 21-28, female. 21-23, epigynum. 21 , ventral. 22, posterior. 23, lateral. 24, 
dorsal. 25, abdomen ventral. 26, carapace. 27, carapace and chelicera. 28, eye region and chelicerae. 29-34, male. 29, left 
palpus. 30, palpus pulled apart. 31 , dorsal. 32, carapace. 33, carapace and chelicera. 34, eye region, chelicerae, and right palpus. 

Figures 35-39. H. istierton n. sp., female, 35-37, epigynum. 35, ventral. 36, posterior. 37, lateral. 38, dorsal. 39, abdomen, 
ventral. 



Figures 40-44. 
ventral. 



H. arnolisei n. sp., female. 40-42, epigynum. 40, ventral. 41, posterior. 42, lateral. 43, dorsal. 44, abdomen. 



ACTINOSOMA, Spilasma, Micrepeira, Pronous • Levi 167 




Figures 45-52. H. dimona n. sp., female. 45-50, epigynum. 45, 48, ventral. 46, 49, posterior. 47, 50. lateral. 45^7, (holotype). 
48-50, (paratype). 51, dorsal. 52, abdomen, ventral. 

Abbreviations. A, terminal apophysis. C, conductor. E, embolus. M, median apophysis. PM, paramedian apophysis. 
Scale lines. 1.0 mm, genitalia 0.1 mm. 



168 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



19; 25 June 1991, 22 (H. Fowler, R. S. Vieira, E. 
Venticinque, MCZ). 

Specimens Examined. BRAZIL Amazonas: Re- 
serva Ducke, Manaus, 26 July 1973, 19 (L. P. Albu- 
querque, MCN 20050); 27 Feb. 1974, 29 (L. P. Al- 
buquerque, MCN 23561); Reserva Km 41, 80 km N 
Manaus, 21 May 1991, 19 (H. Fowler, R. S. Vieira, 
E. Venticinque, MCZ); Cabo Frio, 80 km N Manaus, 
13 Mar. 1989, 19; 27 May 1989, 19 (H. Fowler, R. S. 
Vieira, E. Venticinque, MCZ). 



Pronous Keyserling 

Pronous Keyserling, 1881: 547. Type species Pronous 
tuberculifer Keyserling, 1881, by monotypy. The 
gender of the name is masculine (Bonnet, 1958: 
3778). 

Paphlagon O. P.-Cambridge, 1893: 117. Type species 
Paphlagon beatus O. P.-Cambridge, 1893, by 
monotypy. Species (erroneously) and genus syn- 
onymized with P. tuberculifer by O. P.-Cambridge 
(1898: 281). 

Zigana Chamberlin and Ivie, 1936: 53. Type species 
Zigana wixoides Chamberlin and Ivie, 1936, by 
monotypy. NEW SYNONYMY. 

Note. The synonymy of Zigana with 
Pronous was recognized by Chickering, 
judging from specimen labels, but was 
never published. 

Diagnosis. The large posterior median 
eyes, which have lost their tapeta, are about 
twice the diameter of the anterior median 
eyes, face laterally, and are closer to the 
lateral eyes than to each other (Figs. 53- 
56, 58-62). This apomorphy separates 
Pronous from all other araneid genera in- 
cluding Micrathena. Micrathena and 
Pronous share two apomorphies: the fourth 
legs are longer than the first, and the ab- 
domen of the male is almost rectangular 
(Figs. 61,62). 

Pronous differs from Hypsosinga by 
having the clypeus height equal to the di- 
ameter of the anterior median eye (in 
Hypsosinga the clypeus is higher) and by 
having the fourth legs longer than the first. 

Pronous may be confused with liny- 
phiid genera (e.g., Dubiaranea Mello-Lei- 
tao, 1943 = Paranesticus Mello-Leitao, 
1944: 333 = Hormembolus Millidge, 1991), 
which have similar large posterior median 
eyes. The linyphiids, however, have a high 



clypeus and thinner legs and may have a 
sclerotized stridulating ridge laterally on 
the chelicerae. 

Description. Female. Species all with 
similar coloration and pattern: cephalo- 
thorax orange to orange-brown, sides of 
thoracic region sometimes darker. Chelic- 
erae, labium, and endites orange. Sternum 
light orange-red. Coxae and legs dusky or- 
ange to brown, rarely black. Abdomen light 
orange with six black patches, two on mid- 
dle tubercles and two pairs on posterior 
tubercles (Figs. 56, 57). Some specimens 
have scattered white pigment spots, es- 
pecially on anterior median tubercle, and 
gray to black shading on posterior and sides 
(Fig. 55). Venter usually without pigment, 
sometimes with a median black streak. 
Living specimens are all bright orange-red 
with black marks. 

Carapace punctate. Anterior median 
eyes always slightly larger than laterals and 
posterior medians about 2 diameters of an- 
terior medians (Figs. 53-55). Median eye 
quadrangle wider behind than in front 
(Fig. 53). Height of clypeus about equal 
to diameter of anterior median eyes. Legs 
with femur almost as long as patella and 
tibia. Unlike Micrathena, Pronous lacks a 
stridulating area on the book-lung covers. 
Carapace width behind lateral eyes about 
half maximum diameter of carapace (Fig. 
53). Posterior median eyes about 2 diam- 
eters of anterior medians. Anterior eyes 
their diameter apart, about 2 diameters 
from laterals. Posterior median eyes 1 to 
1.5 diameters apart, 0.5 to 0.8 diameter 
from laterals. Abdomen with a small an- 
terior median tubercle and three pairs of 
tubercles, one pair in middle, two pairs 
posterior (Pi. 2, Figs. 56, 57). Anterior me- 
dian tubercle usually absent in females 
from South America (Fig. 57). Epigynum 
(Fig. 63) without scape, sclerotic areas 
transparent and difficult to delineate. 

Male. Slightly smaller than female. Car- 
apace punctate. Width of carapace behind 
lateral eyes narrower than half of width 
in thoracic region (Fig. 58). Posterior me- 
dian eyes 1.5 to 2 diameters of anterior 



AcTiNOSOMA, Spilasma, MiCREFEiRA, Fronovs • Levi 169 



median eyes. Anterior median eyes 0.6 to 
1 diameter apart, 1 to almost 2 diameters 
from laterals. Posterior median eyes 1.1 to 
1.5 their diameter apart, 0.5 to 1 diameter 
from laterals. Difference in length be- 
tween first legs and longer fourth legs is 
always less in males than in females. Ab- 
domen with a rectangular, lightly sclero- 
tized, dorsal scutum, widest in anterior half, 
with only faint indications of tubercles 
(Figs. 60-62). 

Male endite with tooth (Fig. 59), palpal 
femur with facing tubercle. Palpal patella 
with one macroseta (Fig. 59). First coxa 
with a small hook posteriorly on distal rim 
of margin, second femur with opposing 
small groove (Fig. 59). Second tibia with 
macrosetae on anterior face. Palpus with 
median apophysis within frame of tegul- 
um in median view (M in Figs. 64, 65). 
Conductor positioned centrally on the te- 
gulum (C in Figs. 64, 65). A lobe of te- 
gulum overhangs conductor (at 12 hr in 
Figs. 64, 65). Embolus a curved structure, 
variable in thickness and shape in different 
species (E in Figs. 64, 65). Palpus without 
paramedian apophysis. 

Variation. All species are remarkably 
similar in coloration, shape, and size (Pi. 
2, Figs. 56, 57). Also, the variation of spec- 
imens of different species collected in the 
same area is similar: most South American 
females have smaller tubercles on the ab- 
domen (Fig. 57) than those from Central 
America (Fig. 56), and the two specimens 
collected in Guyana (P. nigripes and P. 
intus) have black legs. 

Relationship. The long fourth legs and 
nearly rectangular male abdomen of Pron- 
ous are characters shared with Micrath- 
ena; both are believed to be synapomor- 
phies. However, Micrathena uses the long 
fourth legs to hold its abdomen upside 
down, horizontally, at a right angle to the 
web, making the sky and leaves the back- 
ground for the bright dorsal coloration and 
the ground the background for the dull 
ventral view. Pronous, on the other hand, 
adopts a resting position more like that of 
other araneids (Eberhard, personal com- 



munication). Thus, the homology of the 
long legs in the two genera is uncertain. 

Natural History. The species makes 
small vertical webs, about 7 to 10 cm di- 
ameter, above leaf litter in forests (per- 
sonal observations in Panama; Pi. 2). The 
web has few supporting threads and may 
catch walking insects and insects flying 
above ground level (Lubin, 1978). At the 
slightest disturbance, the spider falls from 
the web and disappears in leaf litter. Spec- 
imens are thus difficult to collect. About 
half of the available Pronous specimens 
were collected by A. M. Chickering, who 
laboriously sifted leaf litter. 

Distribution. Fourteen species are 
American, most coming from Mexico and 
Central America (Map 4). The four species 
described by Simon (Roewer, 1942: 967) 
from Asia and Madagascar are probably 
all misplaced. The holotypes of only two 
of the four were found in the MNHN. 
Pronous laevisternis Simon, 1908, from 
Tonkin, the area around Hanoi, Vietnam, 
and P. taprobanicus Simon, 1895, from 
Ceylon (Sri Lanka), were examined and 
found to belong to Hypsosinga. The first 
is H. pygmaea (Sundevall, 1831), NEW 
SYNONYMY (see Levi, 1975). Another 
two, more recently described, Pronous 
minutus Saito, 1939 (in Yaginuma, 1986), 
and Pronous tetraspinulus Yin, Wang, Xie 
and Peng, 1990 (Platnick, 1993: 462) from 
China and Japan are probably also mis- 
placed. The fourth legs are much shorter 
than the first in P. minutus (Song, personal 
communication); P. minutus may also be 
Hypsosinga. Pronous chelifer Hasselt 
(1882: 24), from Sumatra, is Gea spinipes 
C. L. Koch, 1843 (Levi, 1983). 

Separating Species. Males are readily 
separated by the structure of their palpi, 
the shape of the embolus, the median 
apophysis, conductor, and tegular lobe. 
Females are difficult to separate: the epi- 
gyna of females of different species are 
similar. The borders of structures in the 
epigynum are transparent and difficult to 
delineate. The cleared epigynum showing 
ducts (cleared in Hoyer's medium) is help- 



170 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 




ACTINOSOMA, Spilasma, Micrepeira, Pronous • Levi 171 



ful, but the dorsal view of the epigynum 
(vulva) is less so. 



Key to Female Pronous 
The terms used in this key are illustrated in Figure 



63. 
1. 



2(1). 



3(2). 



4(3). 



5(4). 



6(5). 



7(1). 



Females from South America (Maps 4B, 
D, F, G) 2 

Females from Mexico or Central Amer- 
ica (Maps 4A, B, C, E, F, G) 7 

Epigynum with V-shaped dusky patch 
(Figs. 63, 131, 135); lateral and pos- 
terior overhang of epigynum wide 
(Figs. 131, 135); in posterior view long 
a.xis of opening longitudinal (Figs. 134, 

138); South America (Map 4D) 

tuberculifer 



Epigynum otherwise (Figs. 98, 107, 117, 
1 22, 1 27 ) 3 

Epigynum with posterior median plate 
narrow ventrally, wider dorsally (at 
center of Fig. 120); lateral overhang 
with a pointed projection in ventral 
view (Fig. 117); Colombia (Map 4G) 



Sides of posterior median plate straight 
dorsally (Figs. 101, 106, 125, 130) 4 

Anterior lip of epigynum visible in ven- 
tral view (Figs. 103, 107, 122, 127) 5 

Anterior lip not visible in ventral view 
(Fig. 98); Colombia (Map 4F) wixoides 

Epigynum with anterior lip small (Fig. 
127); dusky patches with a pair of 
round spots (Fig. 127); Guyana (Map 
4G ) n igripes 

Epigynum with anterior lip larger (Figs, 
103, 107, 122); dusky patches other- 
wise 

Anterior lip swollen in median (Figs. 103, 
107); posterior lip curved anteriorly in 
median (Figs. 103, 107); Costa Rica to 
Ecuador, Venezuela, northern Brazil 
(Map 4B) intus 

Anterior lip not swollen in median (Fig. 
122); Colombia (Map 4G) pance 

Mexico (Maps 4A, C, E) 8 

Honduras to Panama (Maps 4A, B, C, E, 
F, G) _ _ 10 

Epigynum with a median notch (Fig. 71) 
quintana 

Epigynum without notch (Figs. 66, 75) 



9(8). 



10(7). 



11(10). 



12(11) 



Median area of epigynum with anterior 
margin of epigynum V-shaped (Fig. 
66) beatus 

Median area of epigynum with margins 
parallel forming a small bordered lon- 
gitudinal groove (Fig. 75) felipe 

Epigynum in ventroposterior view with 
a transverse, oval median plate having 
a small median longitudinal ridge (Fig. 
114); margin of epigynum rebordered 
(Fig. 112); Panama (Map 4G) shanus 

Epigynum otherwise (Figs. 79, 81, 83 
85, 88, 90) 11 

In posterior view, epigynum median 
plate with wide median split (Fig. 96); 
Costa Rica (Map 4A) golfito 

Epigynum otherwise (Figs. 69, 82, 101) 
12 



Posterior lips of epigynum almost tri- 
angular in shape with a posterior me- 
dian bulge (Fig. 79); Honduras (Map 

4E) lancetilla 

Epigynum otherwise (Figs. 66, 83, 88 

98) 13 

13(12). No distinct anterior lip visible in ventral 

illg view (Figs. 66, 98) 14 

Anterior lip visible in ventral view, 
4 sometimes only in median area (Figs. 

83, 88) 15 

14(13). Posterior median plate wider dorsally 
than ventrally and only barely notched 
ventrally (at 12 hr in Figs. 68, 69); 

Costa Rica (Map 4A) beatus 

- Sides of posterior median plate about 

parallel and with a long median di- 
vision (Fig. 101); in cleared view with 
a pair of median wide duct curves (Fig. 

99); Panama (Map 4F) wixoides 

Q 15(13). Margin of epigynum in ventral view with 
a median longitudinal bordered groove 

(Fig, 83); Costa Rica (Map 4C) peje 

Epigynum otherwise (Figs. 88, 103, 107) 
16 



16(15). Ventroposterior view of epigynum with 
two curved openings, lateral plates 
constricting median plate; median 
plate with long groove (Fig. 90); Costa 

Rica (Map 4 A) colon 

- Epigynum otherwise, lateral plates only 
slightly constricting median plate (Figs. 
105, 109); median plate with short 
ventral split (Figs. 106, 1 10); Costa Rica 
to Colombia (Map 4B) _ intus 



Plate 2. Pronous tuberculifer, bright orange-red coloration with black patches and dusky legs, total length 5 mm. Web of 
Pronous wixoides about 10 cm wide (lower photo. W. Eberhard). 



172 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



Key to Male Pronous 

The males of P. colon, P. lancetilla, and P. nigripes 
are unknown. 

1. Outer, upper edge of median apophysis 
(M in Figs. 64, 65) with a square notch, 
transverse spine of median apophysis 
thick (Fig. 97); Costa Rica (Map 4A) ... 

golfito 

Median apophysis without square corner 
notch (Figs. 64, 70, 74, 116) 2 

2(1). Distal, outer edge of median apophysis 
with an outer spine and an elongate 
notch between it and edge (Figs. 70, 74, 
116, at 3 hr in Fig. 70); Mexico to Pan- 
ama (Map 4) 3 

Outer edge of median apophysis entire 
(Figs. 64, 78, 87, 121) 5 

3(2). Embolus (E in Figs. 64, 65) a semicircular 
disk (Fig. 70); Mexico, Costa Rica (Map 

4A) beatus 

Embolus filamentous (Figs. 74, 116) 4 

4(3). Transverse spine of median apophysis close 
to its "upper" edge (Fig. 74); Mexico 
(Map 4C) quintana 

- Transverse spine of median apophysis in 

middle of median apophysis (Fig. 116); 
Panama (Map 4G) shanus 

5(2). Embolus a long fine thread, whose length 
(if stretched) is longer than diameter of 
palpus (Fig. 102); Panama to Colombia, 

Ecuador (Map 4F) wixoides 

Embolus otherwise (Figs. 87, 121, 139) 6 

6(5). Embolus a semicircular disk (partly trans- 
parent) (Fig. 139); South America (Map 

4D) _ tuberculifer 

Embolus more or less filamentous (Figs. 
78, 111, 121, 126) 7 

7(6). Spine of median apophysis annulate (Fig. 

121); Colombia (Map 4G) valle 

- Spine of median apophysis smooth (Figs. 

87, 97, 126) 8 

8(7). Median apophysis pointed distally "above" 
spine (at 3 hr in Fig. 78); Mexico (Map 
4E) felipe 

- Median apophysis rounded "above" spine 

(Figs. 87, 111, 126) 9 

9(8). Embolus thick, wider than opening of 
notch, the "upper" area enclosed by 
embolus (Fig. 87); Costa Rica (Map 4C) 
peje 



Embolus a finer thread (Figs. Ill, 116) 



10 



10(9). Embolus a fine thread (Fig. 126); tegulum 
lobe truncate (at 12 hr in Fig. 126); 

Colombia (Map 4G) pence 

Embolus thick, tegulum lobe rounded 
(Fig. Ill); Costa Rica to Ecuador (Map 
4B) intus 



Pronous beatus (0. P.-Cambridge) 
Figures 66-70; Map 4A 

Paphlagon beatus O. P.-Cambridge, 1893: 117, pi. 
14, fig. 10, 2. Female from Teapa [Tabasco State], 
Mexico, in BMNH, lost. Erroneously synonymized 
with P. tuberculifer by O. P.-Cambridge (1898: 
281). 

Note. The holotype female, without 
male of P. beatus, could not be found in 
the BMNH. Cambridge later received 
more specimens and cited P. beatus as a 
synonym of P. tuberculatus (in error). 
These later females, together with males, 
were found in the BMNH to be P. quin- 
tana. There are two specimens in the 
HECO collection, one male and one fe- 
male. The only label in the vial was made 
out for my loan, with my loan number, 
the Paphlagon beatus name, and locality, 
Teapa. However, the specimens also be- 
longed to P. quintana. From O. P. -Cam- 
bridge's illustration, it is clear that the Hope 
Collection specimens cannot be the types. 
This was verified by a letter from I. Lans- 
bury, stating that the holotype should be 
in the Natural History Museum in London. 
I suspect that when O. P.-Cambridge syn- 
onymized Paphlagon beatus (erroneously) 
the type label of P. beatus was changed, 
although it will always remain the type of 
this name, whether synonymized with an- 
other name or not. 

Description. Female from Huehuetan. 



Figures 53-65. Morphology of Pronous. 53-57, 63, female. 53, carapace. 54, eye region and chelicerae. 55, lateral. 56, 57, 
dorsal. 63, epigynum, diagrammatic. 58-62, 64, 65, male. 58, carapace. 59, eye region, chelicerae, right palpus, left first coxa 
and second trochanter, and proximal part of femur. 60, lateral. 61, 62, dorsal. 63, epigynum, diagrammatic. 64, 65, left male 
palpus. 64, palpus pulled apart. 65, ventral. 53, 54, 56, 58-61 , 64, 65, P. intus. 55, P. wixoides. 57, 62, P. tuberculifer 



Figures 66-70. Pronous beatus (O. P.-Cambridge). 66-69, female epigynum. 66, ventral. 67, ventral, cleared. 68, ventroposterior. 
69, posterior. 70, male left palpus, mesal. 



AcTiNOSOMA, Spilasma, Micrepeira, Prunous • Levi 173 





U$^ j /jl^'H*^" 



Figures 71-74. P. quintana n. sp. 71-73, female epigynum. 71, ventral. 72, ventral, cleared. 73, posterior. 74, male palpus. 
Figures 75-78. P. felipe n. sp. 75-77, female epigynum. 75, ventral. 76, ventral, cleared. 77, posterior. 78, male palpus. 
Abbreviations. C, conductor. E, embolus. M, median apophysis. R, radix. T, tegulum. 
Scale lines. 1.0 mm, genitalia 0.1 mm. 



174 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



Total length 4.5 mm. Carapace 2.0 mm 
long, 1.7 wide, 1.1 behind lateral eyes. First 
femur 2.2 mm, patella and tibia 2.2, meta- 
tarsus 1.7, tarsus 0.9. Second patella and 
tibia 1.9 mm, third 1.5. Fourth femur 2.9 
mm, patella and tibia 2.8, metatarsus 2.1, 
tarsus 0.7. 

Male from Huehuetan. Abdomen elon- 
gate, intermediate between P. intus (Fig. 
61) and P. tuberculifer (Fig. 62). Total 
length 4.3 mm. Carapace 1.9 mm long, 1.6 
wide, 0.8 behind lateral eyes. First femur 
2.1 mm, patella and tibia 2.0, metatarsus 
1.8, tarsus 0.9. Second patella and tibia 1.7 
mm, third 1.3. Fourth femur 2.7 mm, pa- 
tella and tibia 2.4, metatarsus 2.2, tarsus 
0.9. 

Note. Males and females were collect- 
ed together. 

Variation. Total length of females 4.5 
to 4.8 mm, males 4.3 to 4.7. The specimens 
illustrated came from near Huehuetan, 
Chiapas. 

Diagnosis. The epigynum of the fe- 
male differs from P. quintana (Figs. 71- 
73) in lacking a median notch in ventral 
view (Fig. 66). In posterior view the epi- 
gynum differs from that of P. felipe (Fig. 
77) by having a median plate that is nar- 
row ventrally and widest dorsally (Fig. 69). 
The male has a round semicircular em- 
bolus (Fig. 70), as does the South American 
P. tuberculifer (Fig. 139). The outer edge 
of the median apophysis has a notch sep- 
arating off an outer spine (Fig. 70); the 
transverse spine has nearly parallel sides 
and, unlike P. quintana (Fig. 74), a large 
lobe above (Fig. 70). 

Distribution. Mexico, perhaps to Costa 
Rica (Map 4A). 

Specimens Examined. MEXICO Nayarit: San Bias, 
4-5 Aug. 1947, IS (B. Malkin, C, M. Goodnight, 
AMNH). Jalisco: Rio Pitillal, Playa Grande, 5 km E 
Puerto Mallarta, 12 July 1989, 19; 8 July 1992, 39 (R. 
West, MCZ); Chamela, Sept, 1989, 19 (W. Eberhard, 
MCZ). Oaxaca: 3.2 km SE Niltepec, 16°32'N, 94°33'W, 
16 Aug. 1966, 19 (J., W. Ivie, AMNH); 8 km E Ta- 
panatepec, 230 m, 28 Aug. 1967, 19 (R. E. Leech, 
REL). Chiapas: Esquintla, 19 (Crawford, MCZ); 10 
km N Arriaga, 305 m, 23 Aug. 1972, 19 (C. Mullinex, 
K. Lucas, CAS); Finca Santa Marta, nr. Huehuetan, 
31 July-1 Aug. 1950, 59, 2$ (C, M. Goodnight, 



AMNH); 16 km SE Tierra y Libertad [?], 1,000 m, 
23 Aug. 1972, 13 (C. MulHnex, K. Lucas, CAS). COS- 
TA RICA Cartago: El Cedral, Navarro Orosi-Car- 
tago, 29 Nov. 1979, 19, doubtful determination (C. 
E. Valeric, MZCR). 



Pronous quintana new species 
Figures 71-74; Map 4C 

Pronous tuberculatus: — O. P. -Cambridge, 1898: 281, 
pi. 36, fig. 13, S (misidentification). 

Holotype. Male holotype, male paratype, and two 
female paratypes from 31 km NE of Felipe Carrillo 
Puerto, on Highway 307 toward Tulum, ca. 19°48'N, 
87°52'W, Quintana Roo, Mexico, 17 July 1983 (W, 
Maddison, R. S. Anderson), in MCZ. The specific 
name is a noun in apposition after the locality. 

Note. Cambridge specimens (1898, but 
not 1893), both females and males, are all 
P. quintana. 

Description. Female paratype. Color- 
ation as in other species, legs dark orange, 
dusky marks on sides of carapace. Total 
length 5.4 mm. Carapace 2.1 mm long, 1.9 
wide, 1.1 wide behind lateral eyes. First 
femur 2.0 mm, patella and tibia 1.9, meta- 
tarsus 1.7, tarsus 0.7. Second patella and 
tibia 1.8 mm, third 1.5. Fourth femur 2.8 
mm, patella and tibia 2.7, metatarsus 2.1, 
tarsus 0.9. 

Male holotype. Abdomen relatively 
short, intermediate between P. intus (Fig. 
61) and P. tuberculifer (Fig. 62), with soft, 
bulging sides. Black patches small. Total 
length 4.3 mm. Carapace 1.9 mm long, 1.5 
wide, 0.7 behind lateral eyes. First femur 
1.2 mm, patella and tibia 1.9, metatarsus 
1.3, tarsus 0.7. Second patella and tibia 1.5 
mm, third 1.1. Fourth femur 2.1 mm, pa- 
tella and tibia 2.0, metatarsus 1.6, tarsus 
0.6. 

Note. Males and females were matched 
on the basis of two collections that includ- 
ed both sexes. 

Variation. Total length of females 3.6 
to 5.4 mm, males 3.6 to 4.3. The illustration 
of the male palpus (Fig. 74) was made 
from the holotype; the female (Figs. 71- 
73) was drawn from a specimen from near 
La Palma, Veracruz. 

Diagnosis. The female differs from 
other Mexican species by having a deep 



AcTiNOSOMA, Spilasma, Micrepeira, Fronovs • Levi 175 




Map 4. Distribution of Pronous species. 



176 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



median notch on the posterior margin of 
the epigynum (Fig. 71). The male differs 
by having a short, thread-hke embolus and 
by the median apophysis having a lateral 
notch, separating off an outer spine (Fig. 
74). 

Natural History. The male holotype 
was found hanging by a thread from veg- 
etation. The female was found close to the 
ground in forest, along the edge of a road 
in a small orb web with its hub open. 

Distribution. Southern Mexico (Map 
4C). 

Specimens Examined. MEXICO Veracruz: Es- 
tacion de Biologi'a Tropical "Los Tuxtlas", nr. La 
Palma, 15 km N Catemaco, ca. 18°36'N, 95°07"W, 1, 
2 Aug. 1983, 19, 16 (W. Maddison, AMNH ex MCZ); 
Aug. 1986, 2$ (W. Eberhard 3363, 3366, MCZ). Chia- 
pas: Pichucalco, 17 Julv 1947, 12 (C, M. Goodnight, 
AMNH); Palenque, July 1981, 12 (C. Gold, CAS). 
Tabasco: Teapa, 22, 2$ (BMNH, HECO). 



Pronous felipe new species 
Figures 75-78; Map 4E 

Holotype. Male holotype from 31 km NE of Felipe 
Carrillo Puerto on Highway 307 to Tulum, 19°48'N, 
87°52'W, Quintana Roo, Mexico, 17 July 1983 (W. 
Maddison, R. S. Anderson), in MCZ. The specific 
name is a noun in apposition after the locality. 

Description. Female from Coba Ruins, 
Quintana Roo. Total length 4.9 mm. Car- 
apace 2.2 mm long, 1.9 wide, behind lat- 
eral eyes 0.9 wide. First femur 2.1 mm, 
patella and tibia 2.1, metatarsus 1.6, tarsus 
0.7. Second patella and tibia 2.0 mm, third 
1.6. Fourth femur 2.9 mm, patella and 
tibia 2.6, metatarsus 2.3, tarsus 0.9. 

Male holotype. Coloration light, abdo- 
men intermediate in length between P. 
intus (Fig. 61) and P. tuberculifer (Fig. 
62). Posterior dorsal black patches small. 
Posterior dorsal tubercles distinct. Total 
length 4.3 mm. Carapace 1.8 mm long, 1.5 
wide, 0.7 wide behind lateral eyes. First 
femur 1.7 mm, patella and tibia 1.7, meta- 
tarsus 1.3, tarsus 0.7. Second patella and 
tibia 1.4 mm, third 1.1. Fourth femur 2.1 
mm, patella and tibia 2.0, metatarsus 1.7, 
tarsus 0.7. 

Note. Males and females were matched 



by elimination, both being the third Mex- 
ican species after males and females of the 
other two species were associated. 

Variation. Total length of females 4.9 
mm, males 3.8 to 4.3. The illustrations were 
made from specimens from the Coba Ru- 
ins. 

Diagnosis. The female differs from P. 
beatus (Figs. 66-69) and P. quintana (Figs. 
71-73) by having a median longitudinal 
bordered groove in ventral view of the 
epigynum (Fig. 75). In the male, the pal- 
pus has a short embolus enclosing a shallow 
notch, and the median apophysis has a dis- 
tal, "upper" point and no outer spine (Fig. 
78). 

Distribution. Eastern Mexico (Map 4E). 

Specimens Examined. MEXICO San Luis Potosi: 
Tamazunchale, 6, 7 Julv 1941, 12 (L. I. Davis, AMNH); 
19 April 1963, 12 (W. J. Gertsch, W. Ivie, AMNH); 
26 June 1947, 12 (B. Malkin, AMNH). Veracruz: Cor- 
doba, 20-27 July 1976, 12 (C. H., H. Seevers, AMNH); 
Tecolutla, 13 Oct. 1947, 16 (H. M. Wagner, AMNH). 
Quintana Roo: Coba Ruins, ca. 20°30'N, 87°42'W, 18 
July 1983, 12 (W. Maddison, MCZ). 



Pronous lancetilla new species 
Figures 79-82; Map 4E 

Holotype. Female holotype from Lancetilla, Depto. 
Atlantida, Honduras, July 1929 (A. M. Chickering), 
in MCZ. The specific name is a noun in apposition 
after the locality. 

Description. Female holotype. Total 
length 4.4 mm. Carapace 2.0 mm long, 1.6 
wide, 0.9 behind lateral eyes. First femur 
1.9 mm, patella and tibia 1.9, metatarsus 
1.3, tarsus 0.7. Second patella and tibia 1.7 
mm, third 1.2. Fourth femur 2.5 mm, pa- 
tella and tibia 2.1, metatarsus 1.8, tarsus 
0.8. 

Diagnosis. Pronous lancetilla differs 
from P. golfito (Figs. 93-96) and other 
Pronous by having in ventral view of the 
epigynum a semicircular anterior margin, 
no anterior lips visible, and a transverse 
posterior lip forming a triangle on each 
side (Fig. 79). In posterior view, the me- 
dian plate lacks the deep split (Fig. 82) 
present in P. golfito (Fig. 96). The internal 



AcTiNOSOMA, Spilasma, MiCREPEiRA, Pronuus • Levi 177 



genitalia are simple (Fig. 80) as in P. gol- 
fito (Fig. 94). 



Pronous peje new species 
Figures 83-87; Map 4C 

Holotype. Male holotype from La Selva, near Puerto 
Viejo, Heredia Prov.', Costa Rica, 1-3 Dec. 1981 (J. 
Coddington), in MCZ. The specific name is a noun 
in apposition after the name of a stream near the 
type locaHty. 

Description. Female paratype. Color- 
ation as in other species but legs brown. 
Total length 4.3 mm. Carapace 1.9 mm 
long, 1.5 wide, 0.9 behind lateral eyes. First 
femur 1.9 mm, patella and tibia 1.9, meta- 
tarsus 1.3, tarsus 0.7. Second patella and 
tibia 1.6 mm, third 1.3. Fourth femur 2.4 
mm, patella and tibia 2.1, metatarsus 1.8, 
tarsus 0.7. 

Male holotype. Coloration as in other 
species but legs black. Abdomen elongate 
as in P. intus (Fig. 61). Total length 3.2 
mm. Carapace 1.5 mm long, 1.1 wide, 0.6 
behind lateral eyes. First femur 1.4 mm, 
patella and tibia 1.3, metatarsus 1.0, tarsus 
0.6. Second patella and tibia 1.1 mm, third 
0.8. Fourth femur 1.6 mm, patella and 
tibia 1.4, metatarsus 1.2, tarsus 0.6. 

Note. Males and females were collect- 
ed at the same locality. 

Variation. Total length of females 4.1 
to 4.4 mm. The illustrations (Figs. 83-87) 
were made from the holotype and para- 
type. 

Diagnosis. Pronous peje differs from 
other species found in Costa Rica by the 
large circular margins of the epigynum 
and the presence of a bordered median 
longitudinal groove with parallel sides (Fig. 
83). The embolus of the male encloses a 
small space with a diameter smaller than 
the diameter of the embolus at its widest 
(Fig. 87). 

Natural History. A female from Limon 
Province was collected in a wet forest. An- 
other from La Selva was collected with six 
unwrapped eggs. 

Specimens Examined. COSTA RICA Heredia: La 
Selva, nr. Puerto Viejo, Feb. 1986, 29 paratypes (W. 



Eberhard, MCZ). Liynoii: Cerro Tortuguero, Tortu- 
guero, 110 m, 6 Jan. 1986, 19 (J. Coddington, USNM). 
PANAMA Chiriqui: La Fortuna, 5 Apr. 1984, 29 (W. 
Eberhard, MCZ). 



Pronous colon new species 
Figures 88-92; Map 4A 

Holotype. Female holotype from near Villa Colon, 
San Jose Prov., Costa Rica, Nov. 1990 (W. Eber- 
hard), in MCZ. The specific name is a noun in 
apposition after the locality. 

Description. Female holotype. Total 
length 4.8 mm. Carapace 2.1 mm long, 1.7 
wide, 1.1 wide behind lateral eyes. First 
femur 2.3 mm, patella and tibia 2.3, meta- 
tarsus 1.8, tarsus 0.9. Second patella and 
tibia 2.1 mm, third 1.6. Fourth femur 2.7 
mm, patella and tibia 2.7, metatarsus 2.1, 
tarsus 1.0. 

Diagnosis. Pronous colon differs from 
P. golfito (Fig. 93) and P. peje (Fig. 83) 
by the wide overhangs and almost circular 
openings on each side of the anterior lip 
as seen in the ventral view of the epigynum 
(Fig. 88). In posterior view, the epigynum 
of this species differs from others by hav- 
ing the median plate constricted by a shal- 
low lobe of the lateral plates (Figs. 90, 91). 

Pronous golfito new species 
Figures 93-97; Map 4A 

Holotype. Male from 3 km northeast of Golfito, 100 
m, collected on and under logs, Puntarenas Prov., 
Costa Rica, 22, 23 May 1987 (D. Ubick), in CAS. 
The specific name is a noun in apposition after the 
locality. 

Description. Female paratype. Total 
length 5.0 mm. Carapace 2.1 mm long, 1.7 
wide, 0.8 behind lateral eyes. First femur 
2.1 mm, patella and tibia 2.1, metatarsus 
1.5, tarsus 0.7. Second patella and tibia 1.8 
mm, third 1.4. Fourth femur 2.7 mm, pa- 
tella and tibia 2.5, metatarsus 2.0, tarsus 
0.8. 

Male holotype. Abdomen elongate as in 
P. intus (Fig. 61). Total length 3.8 mm. 
Carapace 1.7 mm long, 1.3 wide, 0.6 be- 
hind lateral eyes. First femur 1.9 mm, pa- 
tella and tibia 1.8, metatarsus 1.3, tarsus 



178 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



0.7. Second patella and tibia 1.4 mm, third 
1.1. Fourth femur 2.0 mm, patella and 
tibia 1.9, metatarsus 1.6, tarsus 0.7. 

Note. Males and females were collect- 
ed in the same area. 

Variation. Total length of females 4.8 
to 5.0 mm. The illustrations (Figs. 93-97) 
were made from the holotype and para- 
type. 

Diagnosis. The female differs from P. 
wixoides by having only one pair of dark 
patches between seminal receptacles and 
anterior margin of the epigynum (Fig. 93) 
and by having the ventroposterior open- 
ings narrowly oval (Fig. 95). In posterior 
view it differs by the wide median notch 
(Fig. 96). The male differs from all others 
by the small nick on the "upper right" 
corner of the median apophysis of the left 
palpus (Fig. 97). 

Natural History. Specimens were col- 
lected in forest vegetation. 

Paratypes. COSTA RICA Puntarenas: Golfito, 24 
July 1981, 2$ paratypes (G. B. Edwards, FSCA); 24 
July 1981, 19 (B. K.' Dozier, FSCA). 

Specimens Examined. COSTA RICA Puntaren- 
as: Osa Peninsula, 4 km SW Rincon, 8-12 Mar. 1967, 
19 (Organiz. for Tropical Studies, MCZ). 

Pronous wixoides (Chamberlin and Ivie) 

new combination 

Figures 55, 98-102; Map 4F 

Pronous beatus: — Banks, 1929: 96 (misidentifica- 
tion). 

Zigana wixoides Chamberlin and Ivie, 1936: 53, pi. 
16, figs. 137, 138, 9. A female lectotype, here des- 
ignated, and four female and one immature male 



paralectotypes (all type specimens in poor condi- 
tion), from Barro Colorado Island [Lago Gatun, 
Panama Prov.], Panama, in AMNH, exannined. 
Roewer, 1942: 883. Bonnet, 1959: 4962. 



Description. Female from Barro Col- 
orado Island, Panama. Total length 3.7 
mm. Carapace 1.6 mm long, 1.4 wide, 0.9 
behind lateral eyes. First femur 1.7 mm, 
patella and tibia 1.8, metatarsus 1.3, tarsus 
0.6. Second patella and tibia 1.5 mm, third 
1.1. Fourth femur 2.1 mm, patella and 
tibia 2.0, metatarsus 1.7, tarsus 0.7. 

Male from Barro Colorado Island, Pan- 
ama. Abdomen relatively short (Fig. 62). 
Total length 3.6 mm. Carapace 1.6 mm 
long, 1 .3 wide, eyes 0.6 behind lateral eyes. 
First femur 1.7 mm, patella and tibia 1.6, 
metatarsus 1.4, tarsus 0.7. Second patella 
and tibia 1.3 mm, third 1.0. Fourth femur 
1.8 mm, patella and tibia 1.8, metatarsus 
1.5, tarsus 0.6. 

Note. The sexes were matched on the 
basis of the connecting ducts in the epi- 
gynum of the female (Fig. 99), which cor- 
respond to the long embolus of the male. 
These ducts are longer in P. wixoides than 
in P. intus and P. shanus. The distribution 
of male specimens from Panama to Ec- 
uador matches that of the females (Map 
4F). 

Variation. One female individual of 
doubtful determination from Serra Nueva 
Granada, northern Colombia, has the ab- 
domen almost completely black. Total 
length of females 3.6 to 5.0 mm, males 3.4 
to 4.1. The illustrations (Figs. 98-102) were 



Figures 79-82. Pronous lancetilla n. sp., female epigynum. 79, ventral. 80, ventral, cleared. 81, ventroposterior. 82, posterior. 

Figures 83-87. P. pe/'e n. sp. 83-86, female epigynum. 83, ventral. 84, ventral, cleared. 85, posterior. 86, lateral. 87, male left 
palpus. 

Figures 88-92. P. colon n. sp. 88-91 , female epigynum. 88, ventral. 89, ventral, cleared. 90, ventroposterior. 91 , posterior. 92, 
lateral. 

Figures 93-97. P. golfito n. sp. 93-96, female epigynum. 93, ventral. 94, ventral, cleared. 95, ventroposterior. 96, posterior. 
97, male palpus. 



Figures 98-102. P. wixoides (Cfiamberlin and Ivie). 98-101, female epigynum. 98, ventral. 99, ventral, cleared. 100, ventro- 
posterior. 101, posterior. 102, male palpus. 



AcTiNOSOMA, Spilasma, Micrepeira, Pronovs • Levi 179 




Figures 103-111. P. intus n. sp. 103-110, female epigynum. 103, 107, ventral. 104, 108, ventral, cleared. 105, 109, ventro- 
posterior. 106, 110, posterior. Ill, male palpus. 103-106, 111, (Panama). 107-110, (Ecuador). 



Scale lines. 0.1 mm. 



180 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



made from specimens from Barro Colo- 
rado Island, Panama. 

Diagnosis. This species is slightly 
smaller than P. intus and P. shanus. The 
female Pronous wixoides differs from re- 
lated Central American species by having 
four (rather than two) dark patches on the 
anterior of the epigynum. The anterior pair 
of patches are in the shape of a tranverse 
hne (Fig. 98). The internal genitalia differ 
by having the duct loops facing each other 
in the median (Fig. 99). The female is dif- 
ficult to separate from P. intus, which has 
an anterior lip showing in the median (Figs. 
103, 107); P. wixoides does not (Fig. 98). 
The epigynum may have to be cleared for 
diagnosis. In contrast, the male differs from 
all other species by having a much longer 
filamentous embolus (Fig. 102). 

Natural History. Specimens were col- 
lected in second growth forest near Bue- 
naventura, Colombia. 

Distribution. Panama, Colombia to 
southern Ecuador (Map 4F). 

Specimens Examined. PANAMA Code: El Valle, 
Jan. 1936, 29 (J. A. Griswold, MCZ); July 1936, U (A. 
M. Chickering, MCZ); Oct. 1954, 19 (W. E. Lundy, 
AMNH). Colon: Fort Davis, 14 Aug. 1936, 12 (A. M. 
Chickering, MCZ); Porto Bello [Portobelo], 12 Aug. 
1936, 19 (A. M. Chickering, MCZ), Panama: Barro 
Colorado Island, very common. COLOMBIA Mag- 
dalena: Serra Nueva Granada, S. N. de Santa Marta, 
1,311 m, 24 May 1975, 19 (doubtful determ., J. A. 
Kochalka, JAK). Meta: ca. 15 km SW Puerto Lopez, 
Hacienda Mozambique, 200 m, 19 (W. Eberhard 1748, 
MCZ). Valle: Hydroelectric Dam on Rio Anchicaya, 
400 m, 1978, 15 (W. Eberhard, MCZ); no date, 19 
(W. Eberhard, 851, MCZ); 1975, 19 (W. Eberhard, 
MCZ); 28 km E Buenaventura, 20 Jan, 1970, 19 (W. 
Eberhard 247, MCZ), Narino: nr, Barbacoas, 20 m, 
20 Mar, 1974, 19 (W, Eberhard 750, MCZ), ECUA- 
DOR "Peru, Palmal" [Palmal, Ecuador, see Palmales, 
03°41'S, 80°00'W, El Oro Prov,, 93 m (Stephens and 
Traylor, 1983)], \6 (K, Jelski, J. Sztolcman, PAN), 

Pronous intus new species 

Figures 53, 54, 56, 58-61, 64, 65, 1 0S- 
Ill; Map4B 

Holotype. Male holotype and three female para- 
types from Barro Colorado Island, Lago Gatun, 
Prov, Panama, Panama, June 1950 (A. M. Chick- 
ering), in MCZ. The specific name is an arbitrary 
combination of letters. 



Description. Female paratype. Total 
length 4.7 mm. Carapace 1.9 mm long, 1.6 
wide, 1.1 behind lateral eyes. First femur 
1.9 mm, patella and tibia 2.1, metatarsus 
1.5, tarsus 0.7. Second patella and tibia 1.8 
mm, third 1.3. Fourth femur 2.5 mm, pa- 
tella and tibia 2.5, metatarsus 1.8, tarsus 
0.7. 

Male holotype. Abdomen elongate (Fig. 
61). Total length 4.0 mm. Carapace 1.9 
mm long, 1.5 wide, 0.7 behind lateral eyes. 
First femur 1.8 mm, patella and tibia 1.9, 
metatarsus 1.4, tarsus 0.7. Second patella 
and tibia 1.4 mm, third 1.2. Fourth femur 
2.0, patella and tibia 1.9, metatarsus 1.7, 
tarsus 0.7. 

Note. Males and females were matched 
on the basis of their distributions: both have 
been found from Costa Rica to northern 
South America. Also, clearing the epigyn- 
um of P. intus reveals a shorter duct than 
in P. wixoides, corresponding to a shorter 
embolus in the male palpus. 

Variation. The abdomen of a male from 
Trinidad is shorter than that of other spec- 
imens. Total length of females 4.6 to 5.6 
mm, males 3.5 to 4.7. Figures 53, 54, 56, 
58-61, 103-106, and 111 were made from 
specimens from Barro Colorado Island, 
Panama, and Figures 107-110 from Ecu- 
adorean specimens. 

Diagnosis. In ventral view the open- 
ings of the epigynum have a wider anterior 
sclerotized lip at the median (Figs. 103, 
107) than has P. wixoides (Fig. 98), and 
when cleared, the connecting ducts appear 
shorter (Figs. 104, 108) than those of P. 
wixoides (Fig. 99). The male palpus has a 
median apophysis (Fig. Ill) as in P. wix- 
oides but, unlike P. wixoides (Fig. 102), 
has a shorter, thicker embolus, lightly col- 
ored on the inside (Fig. 111). 

Natural History. Specimens have been 
found in premontane forest and by sweep- 
ing meadows in Curumani, Colombia. A 
male was collected in a pitfall trap near 
Fortuna, Panama. A male from Luepa, 
Venezuela, came from a Malaise trap in 
cloud forest. 

Distribution. Costa Rica to Venezuela 



ACTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 181 



and Ecuador, northern Amazonas State, 
Brazil (Map 4B). 

Paratypes. PANAMA Panama: Barro Colorado 
Island, 16-20 July 1924, 15 (N. Banks); July 1934, 69; 
Jan. 1936. 49; June 1936; 1 9; July 1936; \$ (USNM 
ex MCZ); June 1939, 19, 1<5; July 1939, 1<5; Aug. 1939, 
59; Sept. 1939, 29, 26; July 1950, 179, 2<5; Aug. 1950, 
39; Julv 1954, 99, 2(5; Aug. 1954, 13; Jan. 1958, 89, 53; 
Feb. 1958. 163, 369 (MCZ, 13 in AMNH ex MCZ) (all 
A. M. Chickering, MCZ); 29 (K. W. Cooper, AMNH); 
Nov. -Dec. 1939, 19 (G. C. Wood, AMNH). 

Specimens Examined. COSTA RICA Limon: Ba- 
taan [Batan], 16 June 1951, 19 (O. L. Cartwright, 
USNM). Puntarenas: Golfito, 16 July 1981, 13 (B. K. 
Dozier, FSCA); Osa Peninsula, Sirena, 10 m, Feb. 
1984, 13 (W. Eberhard, MCZ). PANAMA Bocas del 
Toro: Fortuna, Chiriqui Grande Road, 8°47'N, 
82''11'W, 16-18 July 1987, 13 (D. Olson, MCZ). Chi- 
riqui: David, 26 Nov. 1975, 19 (D. Quintero, MCZ). 
Code: Valle, July 1936, 13 (A. M. Chickering, MCZ). 
Panama: Summit, Sept. 1946, 19 (N. L. H. Krauss, 
AMNH); Gamboa, Jan. 1958, 19, 13 (A. M. Chick- 
ering, MCZ); Forest Reserve, 28 July 1954, 13 (A. M. 
Chickering, MCZ). Darien: Villa Darien, 12-18 Feb. 
1984, 13 (M. N. Garcia, MIUP). WEST INDIES Trin- 
idad: nr. Port of Spain, 1913, 23 (R. Thaxter, MCZ). 
VENEZUELA Monagas: Caripito, Mar. 1942, 19 (W. 
Beebe, AMNH). Bolivar: 10 km N Luepa, Gran Sa- 
bana, 26 June-11 July 1987, 1,500 m, 13 (S., J. Peck, 
AMNH). Distrito Federal: "La Moka" [probably a 
villa near Caracas; H. Enghoff, personal communi- 
cation], 1891, 19 (ZMK). COLOMBIA Bolivar: Car- 
tagena, 21 Dec. 1964, 19 (P. R. Craig, CAS). Cesar: 
Curumani, 22 July 1968, 13 (B. Malkin, AMNH). 
Meta: ca. 20 km N Rio Muco, ca. 20 km S El Porvenir, 
Finca Chenevo, 170 m, no date, 13 (W. Eberhard, 
MCZ); 15 km SW Puerto Lopez, 200 m, 1972, 19 (W. 
Eberhard, MCZ). Antioquia: Cancheras, Mutata, July 
1963, 13 (P. B. Schneble, MCZ). Valle: 50 km S Bue- 
naventura, Mar. 1973, 29 (W. Eberhard, MCZ). EC- 
UADOR Sucumbios: bridge over Rio Cuyabeno, on 
road betw. Tarapoa & Tipishca, 00°01'S, 76°18'W, 
25-30 June 1988, 19, 13 (W. Maddison, MCZ). BRA- 
ZIL Amazonas: Parque Nacional do Pico da Neblina, 
5 Oct. 1990, 13 (A. A. Lise, MCP). 



Pronous shanus new species 
Figures 112-116; IVIap 4G 

Holotype. Male holotype and three female para- 
types from Barro Colorado Island, Gatun Lake, 
Panama Prov., Panama, June 1950 (A. M. Chick- 
ering), in MCZ. The specific name is an arbitrary 
combination of letters. 

Description. Female paratype. Total 
length 5.0 mm. Carapace 2.0 mm long, 1.8 
wide, 0.9 behind lateral eyes. First femur 



2.1 mm, patella and tibia 2.2, metatarsus 
1.7, tarsus 0.8. Second patella and tibia 2.0 
mm, third 1.4. Fourth femur 2.8 mm, pa- 
tella and tibia 2.6, metatarsus 2.1, tarsus 
0.9. 

Male holotype. Abdomen elongate as in 
P. intus (Fig. 61). Total length 4.1 mm. 
Carapace 1.8 mm long, 1.4 wide, 0.6 be- 
hind lateral eyes. First femur 2.1 mm, pa- 
tella and tibia 2.0, metatarsus 1.5, tarsus 
0.7. Second patella and tibia 1.5 mm, third 
1.3. Fourth femur 2.2 mm, patella and 
tibia 2.1, metatarsus 1.8, tarsus 0.7. 

Note. Males and females were matched 
on the basis of similar geographical distri- 
bution and because both male and female 
genitalia differed strongly from genitalia 
of other species of Pronous. (Females have 
been collected with males of the other two 
Panamanian species, P. wixoides and P. 
intus.) 

Variation. Some females have the ven- 
ter and sides of the abdomen black. Total 
length of females 4.3 to 5.6 mm, males 3.8 
to 4.8. The illustrations (Figs. 112-116) 
were made from the holotype and para- 
types. 

Diagnosis. Females are easily separat- 
ed from other species by the rebordered 
margin of the epigynum (Fig. 112) and, 
in ventroposterior view, by the transverse, 
oval median plate having a slight median, 
longitudinal ridge (Fig. 114). The male 
differs from the other two sympatric spe- 
cies, P. intus and P. wixoides, by having 
a short, thin embolus and the median 
apophysis with a narrow transverse spine 
as well as an outer spine at a right angle 
to the first (at 4 hr in Fig. 116). There is 
a narrow notch between the outer spine 
and the median apophysis, best seen in a 
more median view of the palpus. 

Distribution. Panama Canal area (Map 
4G). 

Paratypes. PANAMA Panama: Barro Colorado 
Island, Aug. 1936, 13 (AMNH ex MCZ); June, 1 Aug. 
1939, 29, 33; July 1950, 19, 13; Aug. 1950, 19 (USNM 
ex MCZ); Jan. 1958, 13 (all A. M. Chickering, MCZ 
or ex MCZ). 

Specimens Examined. PANAMA Code: El Valle, 



182 Bulletin Museum of Comparative Zoologij, Vol. 154, No. 3 



July 1936, 13 (A. M. Chickering, MCZ). Panama: 
Arraijan, Aug. 1936, IS (A. M. Chickering, MCZ); 
Forest Preserve, 29 Jan. 1958, 1<5 (A. M. Chickering, 
MCZ); Summit, Aug. 1950, 29, IS (A. M. Chickering, 
MCZ); nr. Gamboa, July 1981, IS (W. Eberhard, 
USNM ex MCZ); Julv 1984, 12 (W. Eberhard 2669, 
MCZ); Cocoli, 24 Nov. 1954, 12 (W, Lundy, AMNH); 
Farfan, 9 Jan. 1958, 12 (A. M. Chickering, USNM e.x 
MCZ); Fort Kobbe, 3 Aug, 1983, 1<5 (H., L. Levi, H. 
Stockwell, MCZ); Miraflores Locks, 3 Jan. 1958, 12 
(A. M. Chickering, MCZ). 



Pronous valle new species 
Figures 117-121; Map 4G 

Holotype. Male holotype from near Saladito, 1,700 
m, Depto. Valle, Colombia, Mar. 1976 (W. Eber- 
hard 1051), in MCZ. The specific name is a noun 
in apposition after the locality. 

Description. Female paratype. Abdo- 
men lacks an anterior median tubercle, 
other tubercles small. Total length 5.6 mm. 
Carapace 2.3 mm long, 1.9 wide, 1.1 be- 
hind lateral eyes. First femur 1.9 mm, pa- 
tella and tibia 2.0, metatarsus 1.5, tarsus 
0.8. Second patella and tibia 1.8 mm, third 
1.3. Fourth femur 2.5 mm, patella and 
tibia 2.4, metatarsus 1.9, tarsus 1.0. 

Male holotype. Abdomen length be- 
tween P. intus (Fig. 61) and P. tubercu- 
lifer (Fig. 62). Total length 3.8 mm. Car- 
apace 1.7 mm long, 1.4 wide, 0.7 behind 
lateral eyes. First femur 1.8 mm, patella 
and tibia 1.5, metatarsus 1.2, tarsus 0.7. 
Second patella and tibia 1.4 mm, third 1.1. 
Fourth femur 1.9 mm, patella and tibia 
1.8, metatarsus 1.4, tarsus 0.7. 

Note. Males and females were matched 
on the basis of having been collected from 
the same locality. 

Variation. The male holotype is newly 
molted and the palpus was preserved when 
still soft. An older male might have other 



areas of the palpus sclerotized and darker 
than in the one illustrated (Fig. 121). 

Diagnosis. The female of P. valle dif- 
fers from females of P. pance and P. tub- 
erculifer species by having the angle of 
the epigynum margin on each side more 
obtuse (Fig. 117) and, in posterior view, 
the median plate widens dorsally (at 6 hr 
in Fig. 120). The male differs from other 
species by having a lightly colored lobe on 
the outside curvature of the embolus (in 
center of Fig. 121) and by having an an- 
nulate transverse spine on the median 
apophysis (Fig. 121). 

Paratype. COLOMBIA Valle: Saladito, 1,700 m, 
Apr. 1977, 12 (W. Eberhard 1162, MCZ). 

Pronous pance new species 
Figures 122-126; Map 4G 

Holotype. Male holotype and one male and five 
female paratypes from Rio Pance, near Cali, 1,100 
m, Colombia, 23 Mar. 1970 (W. Eberhard 1-290), 
in MCZ. The specific name is a noun in apposition 
after the locality. 

Description. Female paratype. The sides 
of the carapace are dusky, and there is a 
median, dorsal gray line on the abdomen. 
Total length 4.5 mm. Carapace 2.1 mm 
long, 1.7 wide, 0.9 behind lateral eyes. First 
femur 1.9 mm, patella and tibia 1.8, meta- 
tarsus 1.3, tarsus 0.7. Second patella and 
tibia 1.7 mm, third 1.3. Fourth femur 2.1 
mm, patella and tibia 2.1, metatarsus 1.7, 
tarsus 0.7. 

Male paratype. Abdomen shape be- 
tween P. intus (Fig. 61) and P. tubercu- 
lifer (Fig. 62). Total length 3.8 mm. Car- 
apace 1.7 mm long, 1.4 wide, 0.7 behind 
lateral eyes. First femur 1.7 mm, patella 
and tibia 1.7, metatarsus 1.3, tarsus 0.7. 



Figures 112-116. Pronous stianus n. sp. 112-115, female epigynum. 112, ventral. 113, ventral, cleared. 114, ventroposterior. 
115, posterior. 116, male left palpus. 

Figures 117-121. P. valle n. sp. 117-120, female epigynum. 117, ventral. 118, ventral, cleared. 119, ventroposterior. 120, 
posterior. 121, male palpus. 



Figures 122-126. P. pance n. sp. 122-125, female epigynum. 122, ventral. 123, ventral, cleared. 124, ventroposterior. 125, 
posterior. 126, male palpus. 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 183 




Figures 127-130. P. nighpes Capohacco, female epigynum. 127, ventral. 128, ventral, cleared. 129, ventroposterior. 130, 
posterior. 

Figures 131-139. P. tuberculifer n. sp. 131-138, female epigynum. 131, 135, ventral. 132-136, ventral, cleared. 133, 137, 
ventroposterior. 134-138, posterior. 139, male palpus. 131-134, 139, (Peru). 135-138, (Guyana). 

Scale lines. 0.1 mm. 



184 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



Second patella and tibia 1.5 mm, third 1.1. 
Fourth femur 1.8 mm, patella and tibia 
1.8, metatarsus 1.5, tarsus 0.5. 

Note. Males and females were collect- 
ed together. 

Variation. Total length of females 4.5 
to 4.8 mm. 

Diagnosis. The female differs from 
other species by the semicircular openings 
and the wide anterior lip of the epigynum 
(Fig. 122). The male has a short filamen- 
tous embolus (Fig. 126), narrower than that 
of P. intus (Fig. Ill) and P. valle (Fig. 
121) and a larger transverse spine on the 
median apophysis (Fig. 126) than in P. 
shanus (Fig. 116) and P. valle (Fig. 121). 

Paratype. COLOMBIA Valle: Rio Pance nr. Call, 
1,100 m, 22 June 1970, 19 (W. Eberhard 290, MCZ). 

Specimens Examined. COLOMBIA Cauca: Rio 
Palace, Totoro, 1,800 m, oak forest, 24 Mar. 1967, 19, 
uncertain determination (R. Root, W. L. Brown, MCZ). 

Pronous nigripes Caporiacco 
Figures 127-130; Map 4G 

Pronous nigripes Caporiacco, 1947: 25; 1948: 662, 
figs. 71, 72, 9, S. Female lectotype, immature par- 
alectotype, designated by Levi (1985: 542), from 
Port Diamond (perhaps Great Diamond, or Dia- 
mond Plantation, both 6°43'N, 58°11'W), Guyana, 
in MZUF, examined. Male paralectotype is Mi- 
crathena acuta (Walckenaer) (Levi, 1985). 

Description. Female lectotype. Color- 
ation as in other species, except sternum 
light; coxae dark or brown, legs black. Car- 
apace slightly rugose. Total length 4.8 mm. 
Carapace 1.9 mm long, 1.6 wide, 0.9 be- 
hind lateral eyes. First femur 2.0 mm, pa- 
tella and tibia 2.0, metatarsus 1.5, tarsus 
0.7. Second patella and tibia 1.8 mm, third 
1.3. Fourth femur 2.7 mm, patella and 
tibia 2.4, metatarsus 2.1, tarsus 0.8. 

Variation. Total length of females 4.7 
to 4.8 mm. The illustrations (Figs. 127- 
130) were made from the lectotype. 

Diagnosis. The pair of black patches 
within the dusky patch of the epigynum 
(Fig. 127) separate this species from P. 
tuberculifer, P. intus, and other South 
American species. Unlike P. tuberculifer, 
P. nigripes has the openings in posterior 



view of the epigynum with their long axis 
transverse (Fig. 130). The epigynum lacks 
overhang above the openings (Fig. 127). 

Specimens Examined. GUYANA Tumatumari, 
21 July 1936, 19 (MZUF). 



Pronous tuberculifer Keyserling 

Plate 2, Figures 57, 62, 131-139; Map 
4D 

Pronous tuberculifer Keyserling, 1881: 548, pi. 16, 
fig. 1, 9, S. Male lectotype and two female para- 
lectotypes from Amable Maria [Depto. Junin], Peru, 
in PAN, examined; Keyserling, 1892: 35, pi. 2, fig. 

31, 9, 6. 

Note. Simon (1895: 863, figs. 922-924), 
Bonnet (1958: 3778) and Roewer (1942: 
967) erroneously considered all American 
specimens, and all citations of the genus 
Pronous, to refer to P. tuberculifer. 

Description. Female from Tingo Mar- 
ia, Peru. Abdomen lacking anterior me- 
dian tubercle; other tubercles small (Fig. 
57). Total length 5.4 mm. Carapace 2.2 
mm long, 1.8 wide, 1 .0 behind lateral eyes. 
First femur 1.9 mm, patella and tibia 2.0, 
metatarsus 1.5, tarsus 0.7. Second patella 
and tibia 1.8 mm, third 1.3. Fourth femur 
2.5 mm, patella and tibia 2.3, metatarsus 
1.9, tarsus 0.8. 

Male from Tingo Maria, Peru. Abdo- 
men as in Figure 62. Total length 3.7 mm. 
Carapace 1.8 mm long, 1.5 wide, 0.7 be- 
hind lateral eyes. First femur 1.7 mm, pa- 
tella and tibia 1.7, metatarsus 1.3, tarsus 
0.7. Second patella and tibia 1.3 mm, third 
1.1. Fourth femur 2.0 mm, patella and 
tibia 1.8, metatarsus 1.7, tarsus 0.7. 

Note. Males and females were paired 
in the original description, and both be- 
long to the only species of Pronous col- 
lected so far in Peru. 

Variation. Total length of females 3.9 
to 5.4 mm, males 3.3 to 4.6. The female 
illustrated (Figs. 131-134) came from Tin- 
go Maria, Peru, the male (Fig. 139) from 
the Depto. Amazonas, Peru, and Figures 
135-138 from Guyana. 

Diagnosis. The epigynum differs from 
that of other Pronous by having a V-shaped 



I 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 185 



dusky patch and by the lateral overhang 
covering the openings (Figs. 131, 135). In 
posterior view the longest diameter of the 
opening is longitudinal (Figs. 131, 135); in 
others it is transverse (Fig. 130). The em- 
bolus is almost semicircular (it may be 
transparent; Fig. 139); a similar embolus 
is found only in the Mexican P. beatus. 
Also, unlike all other species, P. tubercu- 
lifer has the conductor with a small cylin- 
drical projection directed to the right (in 
a left palpus pointing toward 3 hr in Fig. 
139) and a short tegulum lobe (Fig. 139). 

Natural History. The male from Alto 
Rio Comaina came from secondary veg- 
etation at the border of a swamp. 

Distribution. Guyana, Amazon area to 
Misiones Prov., Argentina. Male specimens 
(readily determined) have been found in 
Minas Gerais, Brazil, and in Paraguay and 
Bolivia (Map 4D). 

Specimens Examined. GUYANA Mackenzie 
[06°00'N, 58°ITW], 10 km below Three Friends on 
Demerara River, 55 m el. (Stephens and Traylor, 
1985), Sept. 1931, 12 (MZUF). COLOMBIA Putu- 
mayo: Rio Putumayo, nr. Puerto Asis, no date, 19 (W. 
Eberhard 434, MCZ). ECUADOR Sucumbios: bridge 
over Rio Cuyabeno on road betw. Tarapoa & Tipish- 
ca, O'Ol'S, TenS'W, 8-9 Aug. 1988, 19 (W. Maddison, 
MCZ); Reserva Faunistica Cuyabeno, Laguna Gran- 
de, Sendero La Hormiga, 00°00', 76°10'W, 2-5 Aug. 
1988, 29 (W. Maddison, MCZ). Napo: Pampeya, Rio 
Napo, May 1965, 29 (L. Pena, MCZ). Pichincha: Cay- 
ambe, 2,300 m, June 1965, 1<5 (L. Pefia, MCZ). Za- 
mora Chinchipe: Rio Jumbue, 1 June 1965, 1<5 (L. 
Pena, MCZ). PERU Amazonas: Alto Rio Comaina, 
Puesto de Vigilancia, 22, Falso Paquisha, 850-1,150 
m, 21 Oct.-3 Nov. 1987, 1<5 (D. Silva D. MUSM). 
Hudnuco: Cucharas, Huallaga Valley, Feb.-Apr. 1954, 
19 (F. Woytkowski, CAS); Tingo Maria, 26 May 1947, 
19, 1$ (J, C. Pallister, AMNH), 2 June 1967, 19 (A. F. 
Archer, S. Risco, AMNH); Monzon Valley nr. Tingo 
Maria, 10 Nov. 1954, 29 (E. S. Ross, E. I. Schlinger, 
CAS). BRAZIL Amazonas: Reserva Porto Alegre, 80 
km N Manaus, 27 May 1992, 19 (H. G. Fowler, MCZ). 
Paraiba: Independencia, 1911, 19 (W. M. Mann, MCZ). 
Pernambuco: Pernambuco [Recife], (SMF). Minas 
Gerais: Mina Serinha, Diamantina, Dec. 1944, 16 (E. 
Cohn, AMNH). Parana: Cataratas do Igua?u, 18-25 
Mar. 1985, 29 (H., L. Levi, MCZ). PARAUGAY Con- 
cepcion: Apa, Jan.-Feb. 1909, U (AMNH). Cordil- 
lera: San Bernardino, 19 (E. Reimoser, MCZ). BO- 
LIVIA El Beni: Estac. Biol. Beni, 14°47'S, 65°15"W, 
225 m, 8-14 Nov. 1989, 15 (J. Coddington et al, 
USNM). ARGENTINA Misiones: Montecarlo, Jan. 
1966, 19 (M. E. Galiano, MEG). 



Spilasma Simon 

Spilasma Simon, 1895; 794, fig. 856, 3 abdomen, fig. 
857 9 eye region. Type species Spilasma artifex 
Simon, 1895 (=S. duodecimguttata (Keyserling, 
1880)] by original designation and monotypy. The 
gender of the name is feminine (Bonnet, 1958: 4121). 

Diagnosis. Spilasma has a narrow ce- 
phalic region of the carapace (Figs. 146, 
154) as in Cyclosa and Acacesia. Spilasma 
differs from Cyclosa by having few setae 
on the carapace and lacking the dark rings 
on the legs (Fig. 145). Spilasma differs from 
both Cyclosa and Acacesia by having an 
oval abdomen, widest in the middle, with 
six dorsal pairs of white patches, and dif- 
fers from many araneids by having no 
markings in the midline of the abdomen 
and no folium pattern (Fig. 145). 

Description. Cephalothorax glabrous, 
orange to orange-brown without marks. 
Abdomen oval, widest in middle, with dor- 
sal paired patches of white (Fig. 145) and 
no ventral markings. Posterior median eyes 
subequal to anterior medians and their di- 
ameter or less apart. The epigynum has a 
ventrally extending lobe (Figs. 140, 142) 
with posterior sculpturing (Fig. 143). 

Male smaller than female, with ventral 
sclerotized area from sides of pedicel to 
genital groove (Fig. 153). Endite without 
tooth, no coxal hook. Palpal patella with 
two weak setae. Second tibia thinner than 
first. 

Relationship. Spilasma can be placed 
with the Alpaida group of species, on the 
basis of the following characters: the epi- 
gynum is lobe-shaped (Figs. 140-144); the 
tegulum is free in the upper right corner 
of the left palpus (T in Fig. 151); the con- 
ductor is in the middle of the tegulum, not 
the upper right in the left palpus (C in Fig. 
151); the median apophysis is without 
spines or flagella (except distally) and pro- 
jects beyond bulb of palpus (M in Fig. 151); 
in the upper left of the conductor (in the 
left palpus) there is a pocket (C in Fig. 
161), which may be a homolog with the 
paramedian apophysis. The narrow ce- 
phalic region of the carapace is probably 
a synapomorphy with Acacesia Simon 



186 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



^^^^^^^ 


^'fV 




llfi 


^EHH^t^^^^^S^^^^'^" i^^n^ 


IS 


wSmi 


B 


t^i^i^BSriWmmm 


^^^^5c^ ' 


1^ ■ •^'■'la ■ 






Plate 3. Spilasma duodecimguttata. Web diameter of upper web about 25 cm, lower one 10.4 cm {upper photos, R. L. C. 
Baptista; lower photo, J. A. Coddington). 



(Glueck, 1994) and perhaps also with Cy- 
closa. 

Natural History. The web is horizon- 
tal, pulled up at the hub, with sticky threads 
(W. Eberhard, personal communication) 
and, above the hub, a median granular 
retreat, with sand grains, having a lateral 
flap (Pi. 2). The flap pulls in when dis- 
turbed. The retreat is also used to house 
the eggsacs (Lubin, 1978). 

Distribution. There are three Ameri- 
can species, with only one specimen each 
available for two of them, from Honduras 
to Rio de Janeiro State, Brazil. Spilasma 
africana Simon, 1903, from Spanish Guin- 
ea, is the only described species not from 
America. The specimens of S. africana are 
lost, and the species was synonymized with 



Prasonica seriata Simon, 1895, by Gras- 
shoff (1971). Prasonica is a genus related 
to M angora. 

Key to Spilasma Species 

The female of S. utaca and the male of S. baptistai 
are not known. 



Males 

Females 



2(1). 



3(1). 



A pair of deep longitudinal grooves on car- 
apace (Fig. 161); base of embolus with 
duct making two loops (Fig. 160); Peru 
(Map 5) utaca 

Carapace without grooves (Fig. 152); em- 
bolus duct with one loop (Figs. 149-151); 
from Honduras to Bolivia and southern 
Brazil (Map 5) duodecimguttata 

In posterior view of epigynum, slit-shaped 
openings face laterally (Fig. 157); Per- 
nambuco, Brazil (Map 5) baptistai 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 187 




D baptistai 

# duodecimguttata 

A utaca 



Map 5. Distribution of Spilasma species. 



In posterior view of epigynum, slit-shaped 
openings face anteromedially (Figs. 141, 
143); Honduras to Bolivia and south- 
eastern Brazil duodecimguttata 



Spilasma duodecimguttata (Keyserling) 
Plate 3; Figures 140-155; Map 5 

Singa duodecimguttata Keyserling, 1880: 302, pi. 4, 
fig. 6, 9. Female holotype from New Granada [old 
name for Colombia], in BMNH, examined. Key- 
serling, 1893: 286, pi. 14, fig. 211, 9. Roewer, 1942: 
877. 

Epeira lamentaria Keyserling, 1883: 199, pi. 15, (?fig. 
5); 1892: 174, pi. 8, (?fig. 128). Female holotype 
from Amazon Prov., Brazil at HECO, examined. 
NEW SYNONYMY. 

Spilasma tredecimguttata Simon, 1895: 789, figs. 856, 
857, 6 abdomen, 9 eyes; 1897: 476. Many male and 
female syntypes from the Amazon in MNHN no. 
1010, examined. Roewer, 1942: 776. Bonnet, 1958: 
4121. NEW SYNONYMY. 

Spilasma artifex Simon, 1895: 794; 1897: 477, pi. 12, 
figs. 1-5, pi. 13, fig. 1, web. Female holotype from 
San Esteban, Venezuela, MNHN no. 10127, ex- 
amined. Roewer, 1942: 776. Bonnet, 1958: 4121. 
NEW SYNONYMY. 

Epeira davisi Hingston, 1932: 365, fig. 40, web. Spec- 



imens from Guyana, in BMNH, lost. NEW SYN- 
ONYMY. 

Aranea lamentaria: — Roewer, 1942: 845. 

Araneus duodecimguttatus: — Bonnet, 1955: 449. 

Araneus lamentarius: — Bonnet: 1955: 526. 

Spilasma coccineum Caporiacco, 1955: 344, fig. 29, 
(3. Male holotype from Rancho Grande, Venezuela, 
in UCVC, not examined. Brignoli, 1983: 280. NEW 
SYNONYMY. 

Note. The small illustration of Keyser- 
ling (1883, fig. 5; and the same in 1892, 
fig. 128) does not look like the epigynum 
of this species. Keyserling may have illus- 
trated an artifact or debris. Caporiacco's 
(1955) illustration of the male palpus of S. 
coccineum is good. 

Lopez (1985: 86) described a new spe- 
cies, Spilasma richei, collected along the 
path from Roura to Kaw, southeast of Cay- 
enne, French Guyana. He placed it in the 
Museum de Beziers, France, but it has been 
lost (Lopez, 1986, personal communica- 
tion). Few diagnostic characters are pro- 
vided for S. richei except for noting the 
difference in "la spinulation des pattes et 



188 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



la form precise de I'epigyne", but a de- 
tailed description is forthcoming (Lopez, 
1985: 87). This is considered a nomen nu- 
dum. It is also not listed in Platnick's cat- 
alogs (1989, 1993). 

The genitalia of S. duodecimguttata are 
quite variable. When many specimens 
from different locations are available, one 
readily sees the reason for the many syn- 
onymies cited here. 

Epeira davisi was erroneously placed in 
Cyrtophora by Levi (1991b: 179). 

Description. Female from Depto. Meta, 
Colombia. Carapace, chelicerae, labium, 
endites, sternum, legs, bright orange. Dor- 
sum of abdomen orange, with a gray cast, 
with six pairs of symmetrical white patches 
(Fig. 145); venter darker gray. Posterior 
median eyes same diameter as anterior 
medians, laterals 0.7 diameter. Anterior 
median eyes 0.3 diameter apart, 0.8 di- 
ameter from laterals. Posterior median eyes 
0.5 diameter apart, 1.1 diameters from lat- 
erals. Ocular rectangle, slightly longer than 
wide. Height of clypeus equals 0.4 diam- 
eter of anterior median eye. Sternum 
slightly wider than long. Abdomen oval, 
widest anterior of middle (Fig. 145). Total 
length 4.5 mm. Carapace 2.0 mm long, 1.9 
wide, 0.8 behind lateral eyes. First femur 
2.3 mm, patella and tibia 2.3, metatarsus 
1.3, tarsus 1.0. Second patella and tibia 2.0 
mm, third 1.2, fourth 2.0. 

Male from Depto. Meta, Colombia. Col- 
or as in female. Distal leg articles darkest. 
Posterior median eyes 0.7 diameter of an- 
terior medians, laterals 0.5 diameter. An- 
terior median eyes 0.2 diameter apart, 0.6 
diameter from laterals. Posterior median 
eyes 0.7 diameter apart, 1.2 diameters from 
laterals. Height of clypeus equals the di- 
ameter of an anterior median eye. Abdo- 
men widest in middle, with a soft, but 



sclerotized shield in the area anterior to 
the genital groove (Fig. 153). Total length 
3.2 mm. Carapace 1.6 mm long, 1.3 wide, 
0.5 behind lateral eyes. First femur 1.6 
mm, patella and tibia 1.6, metatarsus 1.0, 
tarsus 0.7. Second patella and tibia 1.4 mm, 
third 0.9, fourth 1.3. 

Note. Males and females have been col- 
lected together. 

Variation. The genitalia of no two 
specimens look quite alike, and each new 
specimen makes one think they have a new 
species. Most variation is in the protuber- 
ance of the epigynum: some having a semi- 
circular keel (Figs. 140-142), some not 
(Figs. 143, 144). The male palpus also var- 
ies in structure. The tip of the median 
apophysis and the curvature of the em- 
bolus are unusually variable (Figs. 149, 
150). Some individuals lack white patches 
on the abdomen. Total length of females 
3.0 to 7.0 mm, males 2.2 to 3.6. Most il- 
lustrations were made from specimens 
from Depto. Meta, Colombia, but Figures 
143-145 and 150 were made from speci- 
mens from 80 km north of Manaus, Brazil. 

Diagnosis. The first femur and patella- 
tibia are of about the same length, unlike 
those of most other araneids. The epigyn- 
um differs from S. baptistai by having the 
slit openings face anteriomedially (Figs. 
141, 143); the male differs from S. utaca 
by lacking the pair of grooves on the car- 
apace (Fig. 161). The male palpus super- 
ficially resembles that of Ocrepeira yaelae 
Levi (1993a, fig. 359). 

Natural History. Spilasma duodecim- 
guttata is found in foliage of wet forest in 
Guapiles, Costa Rica; in lowland forest. El 
Dorado, Venezuela; in rain forest near Le- 
ticia, Colombia; in secondary forest by a 
lake 15 km southwest of Puerto Lopez, 
Colombia; in forest interior in reserves 



Figures 140-155. Spilasma duodecimguttata (KeyserWng). 140-148, female. 140-144, epigynum. 140, 143, ventral. 141, pos- 
terior. 142, 144, lateral. 145, dorsal. 146, carapace. 147, carapace and cheliera. 148, eye region and chelicerae. 149-155, male. 
149-151, left palpus. 149, 15C, mesal. 151, pulled apart. 149, 151, (Depto. Meta, Colombia). 151, (N of Manaus, Brazil). 152, 
dorsal. 153, abdomen ventral. 154, carapace. 155, eye region, chelicerae and right palpus. 



Figures 156-159. S. baptistai, female. 156-158, epigynum. 156, ventral. 157, posterior. 158, lateral. 159, dorsal. 



ACTINOSOMA, Spilasma, Micrepeira, Pronous • Levi 189 




Figures 160, 161. S. utaca, male. 160, palpus. 161, dorsal. 

Abbreviations. C, conductor. E, embolus. M, median apophysis. R, radix. T, tegulum. 

Scale lines. 1 mm, genitalia 0.1 mm, except Figures 146-148, 153-155, 0.5 mm. 



190 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



north of Manaus, Brazil. In Rio de Janeiro, 
Brazil, a web was found to be 40 cm above 
ground. 

Distribution. Honduras to Rio de Ja- 
neiro State, Brazil, and Bolivia (Map 5). 

Specimens Examined. HONDURAS Atlantida: 
Lancetilla, 9 (MCZ). COSTA RICA Limon: 5.5 km 
E Guapiles, $ (DU). Heredia: Finca La Selva, 9, S 
(CAS, MCZ). Puntarenas: Reserva Carara nr. Tar- 
coles, 9 (MCZ). Cartago: Turrialba, S (AD). PANAMA 
Panama: Barro Colorado Island, 9, S (MCZ); Cerro 
Galero, S (MCZ), WEST INDIES Trinidad: Old Gold 
Mine, Arima Ward, 9 (USNM). VENEZUELA Ara- 
gua: Rancho Grande, <5 (AMNH), 9 (MCZ), 9, S 
(USNM). Bolivar: 22 km S El Dorado, S (AMNH). 
Falcon: 3 km S El Hondo de Uria, 15 km E Curi- 
magua, 9 (MCZ). GUYANA Shudikar River, Upper 
Essequibo River, $ (AMNH); Upper Essequibo River, 
Onoro Region, 9 (AMNH); Essequibo River opposite 
Twasinki Mtns., imm. (AMNH); Upper Essequibo 
River, nr. Akaramukra Rapids, 3 (AMNH). SURI- 
NAM Saramacca: Voltzberg Raleighvallen Nature 
Reserve, 9 (MCZ). FRENCH GUIANA nr. Camp Cai- 
man, Montagnes Kaw, 9 (USNM). COLOMBIA San- 
tander: Carare, Opon Capote 6°38'N, 73°55'W, 6 
(MCZ). Meta: 15 km SW Puerto Lopez, 9, <5 (MCZ). 
Valle: 28 km E Buenaventura, <3 (MCZ); Central An- 
chicaya, 9 (MCZ). Putumayo: Rio Putumayo, nr. Pto. 
Asis, 9 (MCZ). Amazonas: Amacayacu, 48 km NW 
Leticia, S (MCZ); 18 km N Leticia, S (AMNH); Le- 
ticia, 9, 6 (CAS). ECUADOR Sucumbios: Reserva 
Faunistica Cuyabeno, 9 (MCZ, MECN). Napo: Rio 
Coca and Napo, <5 (MCZ); 48.6 km NE Baeza, S (MCZ). 
PERU Hudnuco: Panguana, Rio Pachitea, 9°37'S, 
74°56'W, 9 (MCZ); Monzon Valley, Tingo Maria 9, <5 
(CAS); Dantas La Molina, Quebrada Sapote, SW 
Puerto Inca, 270 m, 9°38'S, 75°00'W, 19 (MUSM). 
Madre de Dios: Zona Reserva de Manu, 9 (USNM); 
15 km E Puerto Maldonado, 6 (CAS); Res. Tambo- 
pata, 12°50'S, 69°17'W, 9, S (USNM). BRAZIL Ro- 
raima: Ilha de Maraca, 9, $ (INPA). Amazonas: Cabo 
Frio Reserve, 80 km N Manaus, 9, $ (INPA, MCN, 
MCZ); Colosso Reserve, 80 km N Manaus, 9, 3 (INPA, 
MCN, MCZ); Dimona Reserve, 80 km N Manaus, 9, 
& (MCZ); Gaviao Reserve, 80 km N Manaus, 3 (MCZ); 
Km 41 Reserve, 80 km N Manaus, 9, 6 (MCZ); Fa- 
zenda Esteio, Manaus, $ (INPA); Belem, periodically 
flooded forest, nr. confluence with Rio Solimoes, S 
(AMNH); Manaus, 9, $ (MACN); Reserva Porto Ale- 
gre, 80 km N Manaus, 9 (MCZ); Reserva Ducke nr. 
Manaus, 9 (MCN, INPA); Parque Nacional do Pico 
da Neblina, Maturaca, 9 (MCP). Pard: Caninde, 9 
(AMNH). Bahia: Encruzilhada, $ (AMNH); Fazenda 
Matiapa, Camacan, $ (MCN). Mato Grosso: Barra de 
Tapirape, 3 (AMNH, MCZ); Chapada dos Guimaraes, 
9 (AMNH); Sinop, S (AMNH); Villa Vera, 55°30'S, 
12°46'W, 6 (AMNH). Rio de Janeiro: Mangaratiba, S 
(AMNH); Barra da Tijuca, 9 (RLCB); Morro da Urea, 
9 (RLCB). BOLIVIA El Bent: 27 km SW Yucumo, 
500 m, 15°50'S, eg-H'W, 6 (USNM). 



Spitasma baptistai new species 
Figures 156-159; Map 5 

Holotype. Female from Dois Irmaos Reservation, 
Recife, Pernambuco State, Brazil, 25 Jan. 1989 (R. 
L. C. Baptista, A. P. Chaves 2654) in MZSP. The 
species is named after the collector. 

Description. Female holotype. Cara- 
pace, chelicerae, sternum, labium, and en- 
dites orange. Legs orange, distal articles 
darkest. Dorsum of abdomen with pairs of 
white patches (Fig. 159); venter gray. Eyes 
small. Posterior median eyes 0.8 diameter 
of anterior medians, laterals 0.8 diameter. 
Anterior median eyes 0.2 diameter apart, 
1 diameter from laterals. Posterior median 
eyes 1 diameter apart, 1.3 diameters from 
laterals. Ocular quadrangle slightly longer 
than wide, slightly wider behind. Height 
of clypeus equals 0.5 diameter of the an- 
terior median eye. Abdomen oval (Fig. 
159). Total length 3.0 mm. Carapace 1.40 
mm long, 1.19 wide, 0.57 behind lateral 
eyes. First femur 1.30 mm, patella and 
tibia 1.32, metatarsus 0.67, tarsus 0.52. Sec- 
ond patella and tibia 1.19 mm, third 0.71, 
fourth 1.10. 

Diagnosis. The eyes are smaller than 
those of S. duodecimguttata (Fig. 159) and 
in posterior view of the epigynum, the me- 
dian plate (Fig. 157) is relatively wider 
than that of S. duodecimguttata (Figs. 141, 
143), and the openings face laterally (Fig. 
157). 

Natural History. The specimen was 
collected with its retreat. Vestiges of the 
web were projecting from refuge between 
leaves of a bush, 1.5 m above ground in 
secondary forest. 

No other specimens were found. 

Spitasma utaca new species 
Figures 160, 161; Map 5 

Holotype. Male holotype from 1,600 to 2,200 m, 
Utcuyacu [above Merced, 1,465 m, 11°12'S, 
75°28'W; Stephens and Traylor, 1983], Depto. Ju- 
nin, Peru, 4 Apr. 1948 (F. Woytkowski), in AMNH. 
The specific name is an arbitrary combination of 
letters. 

Description. Male holotype. Carapace 
dark orange-brown. Chelicerae, labium. 



AcTiNOSOMA, Spilasma, MiCREi'EiRA, Pronovs • Levi 191 



endites, sternum dark brown. Coxae brown, 
except posterior half of fourth coxae, which 
are hght yellowish; legs dark brown. Dor- 
sum of abdomen black with two pairs of 
white spots, the first round, the second di- 
amond-shaped, and with transverse pos- 
terior dark bands (Fig. 161); venter black. 
Posterior median eyes 0.7 diameter of an- 
terior medians, anterior laterals 0.7 di- 
ameter, posterior 0.6. Anterior median eyes 
0.2 diameter apart, 0.4 diameter from lat- 
erals. Posterior median eyes 0.4 diameter 
apart, 1.2 diameters from laterals. Ocular 
quadrangle narrower behind than in front. 
Height of clypeus equals 0.7 diameter of 
anterior median eye. Sternum corniculate. 
Abdomen oval, widest behind middle (Fig. 
161). Total length 3.0 mm. Carapace 1.59 
mm long, 1.19 wide, 0.53 wide behind lat- 
eral eyes. First femur 1.00 mm, patella and 
tibia 1.72, metatarsus 0.87, tarsus 0.69. Sec- 
ond patella and tibia 1.56 mm, third 0.93, 
fourth 1.45. 

Diagnosis. The carapace of S. utaca 
has a pair of grooves (Fig. 161) not present 
in S. duodecimguttata (Fig. 152), and the 
duct loops twice in the proximal end of 
the embolus (Fig. 160). 

No other specimens were found. 

Micrepeira Schenkel 

Micrepeira Schenkel, 1953: 26. Type species M. al- 
bomaculata Schenkel by monotypy. The gender of 
the name Micrepeira is feminine. 

Diagnosis. Micrepeira differs from 
most other araneid genera, including Ar- 
aneus and Singa by the domed sternum 
(Figs. 186, 191) and a subspherical abdo- 
men with bold, white dorsal markings: of- 
ten a wide median, longitudinal band and 
a wide transverse band or a few pairs of 
transverse patches, but no white markings 
on the venter (Figs. 165, 176, 185, 190, 
195, 204). Also in Micrepeira, the femora 
are the same length as the adjacent patella 
and tibia; in other genera, the femora are 
shorter. Micrepeira resembles the Indo- 
pacific Anepsion Strand, 1929 (Chrysan- 
thus, 1969), but has a narrower cephalic 
region. It resembles the African Pherenice 



Thorell, 1899, but has a low clypeus, less 
than the diameter of the anterior median 
eye. Micrepeira differs from all similar 
genera by the following: the epigynum has 
a wide shelf, resembling that of Alpaida, 
with a tiny, soft, distal, pointed scape, eas- 
ily overlooked (Figs. 162, 192, 200), by the 
male palpus, which has a large median 
apophysis bearing two flagella (Figs. 168, 
177, M in Fig. 179), and a conductor either 
on the edge or inward on the tegulum, and 
no terminal apophysis or embolus lamella 
(Figs. 179, 180). The median apophysis 
flagella are, as usual, associated with a 
pointed scape of the female epigynum. 

Description. Female. Cephalothorax 
glabrous, orange to brown without distinct 
marks, cephalic region wide (Figs. 166, 
167). Eyes subequal. Median ocular quad- 
rangle almost square. Height of clypeus 
0.6 to 0.8 diameter of the anterior median 
eye. Legs relatively short and thick (Fig. 
204). 

Males much smaller than females (Fig. 
181), width behind eye region less than 
half width of carapace. Endite with small 
tooth (M. fowleri) or none (M. hoeferi), 
no tubercle or tooth on palpal femur. No 
hook on first coxa, palpal patella with one 
macroseta. Second tibia as thick as or thin- 
ner than first, without any large macro- 
setae. Median apophysis of palpus has two 
proximal flagella associated with a pointed 
scape of the epigynum. 

Relationship. The broad lobe of the 
epigynum, the palpus with a bare tegulum 
(at 1-2 hr in Fig. 168), the lack of distal 
hematodocha, the lack of terminal apoph- 
ysis or embolus lamella, and the single 
macroseta on the palpal patella place the 
genus close to Alpaida. But the tiny, point- 
ed scape of the epigynum (Figs. 162-164) 
and the two flagella of the median apoph- 
ysis (Fig. 168, M in Fig. 179) suggest re- 
lationship with Kaira, Aculepeira, Ama- 
zonepeira, and Metepeira, all genera clos- 
er to Araneus. Although the pointed scape 
and flagella on the median apophysis were 
previously considered synapomorphies, 
evidence now indicates that this may be a 
homoplasious character, as the make-up of 



192 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 




Plate 4. Micrepeira hoeferi n. sp. Diameter of web 20 cm, hub of web 150 cm above forest floor (photo, H. H6fer). 



the genitalia of Micrepeira resembles so 
much more that of Alpaida. The position 
of the conductor (C in Figs. 179, 180, at 
1 hr in Fig. 177 and at 11 hr in Fig. 178) 
is on the edge of the tegulum but is at the 



distal end and not above the median 
apophysis as in Araneus. 

Natural History. Webs are known of 
three species (M. hoeferi, M. tubulofa- 
ciens, and M. velso). All have the shape 



ACTiNOSOMA, Spilasma, MiCREPEiRA, Pronous • Levi 193 



f \GUYANA 
(•k>'-rT>-~ FR. GUIANA 




D f o w I e r i \'>{^. I. 
▲ pachitea \ PERU 
# tubulofaciensV^^ '[' 
V velso ^-c^^/f^ 



SURINAM 

FR. GUIANA 




n albomaculata 
# hoeferi 
Vsmithae 



BOLIVIA 'r-- ---\\ \ l'^ 




Map 6. Distribution of Micrepeira species. 



of the lower half of a circle hanging on a 
line with a retreat above the hub (Pi. 4). 
The retreat is made of detritus and silk 
and has a lid that can be closed (Hofer 
note on label with specimen of M. hoeferi). 
At its pointed end, the retreat has two 
strong silk support lines covered by some 
detritus. 

Distribution. All seven species are Cen- 
tral and South American (Map 6). 



Key to Micrepeira Species 

Males of M. albomaculata, M. pachitea, M. smi- 
thae, M. tubulofaciens, and M. velso are unknown. 

1 . Male 2 

Female 3 

2(1). Middle of median apophysis with a lobe 
(Fig. 168); Amazonian Ecuador to Ma- 

naus region of Brazil (Map 6) fowleri 

Middle of median apophysis without lobe 
(M in Figs. 177-180); Guianas to Peru, 
Mato Grosso, Brazil (Map 6) hoeferi 



194 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



3(1). Sternum with an anterior longitudinal 
groove (Fig. 191); Peruvian Amazon (Map 

6) pachitea 

Sternum without such groove (Fig. 186) 4 

4(3). Epigynum in ventral view a convex lobe 
(Fig. 200); Falcon State, Venezuela (Map 

6) _ _ albomaculata 

Epigvnum flat (Figs. 162, 171, 182, 187, 
192, 196) 5 

5(4). Epigynum posterior median plate over- 
hanging laterals as in Figure 197; Costa 

Rica (Map 6) velso 

Posterior median plate otherwise; South 
America 6 

6(5). Epigynum with posterior margin straight 
in ventral view and with narrow scape 
(Figs. 171, 174); Guianas, to Peru, and 
Mato Grosso, Brazil (Map 6) hoeferi 

- Epigynum more or less triangular in ven- 
tral view or a triangular scape (Figs. 162, 
182, 1 92 ) 7 

7(6). Triangle formed by epigynum taking up 
most of posterior margin of epigynum 

(Figs. 162, 182) 8 

Triangle formed by epigynum making up 
less than half of posterior margin of epi- 
gynum (Fig. 192); Surinam (Map 6) 

smithae 

8(7). Epigynal lobe marked as in Figure 182; 

Guianas (Map 6) tuhulojaciens 

Epigynum without dark marks (Fig. 162); 
Amazonian Ecuador to Manaus region of 
Brazil (Map 6) _ fowleri 



Micrepeira fowleri new species 
Figures 162-170; Map 6 

Holotype. Female holotype from 80 km north of 
Manaus, Km 41 Reserve, forest interior, Amazonas 
State, Brazil, 14 Apr. 1991, 200 m (H. G. Fowler, 
E. V'enticinque, R. S. Vieira), in MCN no. 25536. 
The species is named after the collector. 

Description. Female holotype. Cara- 
pace orange. Chelicerae, labium, endites 
orange. Sternum dusky orange. Coxae or- 
ange; legs dusky orange, darker distally. 
Dorsum of abdomen dark dusky with a 
longitudinal, median band of white pig- 
ment patches and a pair of white patches; 
posterior black with a pair of small pos- 
terior patches (Fig. 165); venter dusky with 
a median darker band from epigynum to 
spinnerets. Posterior median eyes 0.8 di- 
ameter of anterior medians, laterals 0.6 
diameter. Anterior median eyes 0.4 di- 
ameter apart, 1 diameter from laterals. 



Posterior median eyes 0.7 diameter apart, 
1.5 diameters from laterals. Total length 
3.0 mm. Carapace 1.36 mm long, 1.24 
wide, 0.75 behind lateral eyes. First femur 
1.18 mm, patella and tibia 1.18, metatarsus 
0.62, tarsus 0.54. Second patella and tibia 
1.09 mm, third 0.65, fourth 1.03. First tibia 
0.22 mm thick. 

Male paratype. Color as in female ex- 
cept lacking anterior abdominal white pig- 
ment patches, having only the posterior 
pair. Posterior median eyes 0.8 diameter 
of anterior medians, laterals 0.6 diameter. 
Anterior median eyes 0.7 diameter apart, 
0.7 diameter from laterals. Posterior me- 
dian eyes 0.8 diameter apart, 1.2 diameters 
from laterals. Endite with small tooth. To- 
tal length 1.8 mm. Carapace 1.07 mm long, 
0.88 wide, 0.44 behind lateral eyes. First 
femur 0.91 mm, patella and tibia 0.81, 
metatarsus 0.49, tarsus 0.40. Second patella 
and tibia 0.72 mm, third 0.45, fourth 0.65. 

Note. Males and females were collect- 
ed together; both have the posterior of the 
abdomen black. 

Variation. Total length of females 2.4 
to 3.5 mm, males 1.8 to 2.1. The anterior, 
median, transverse groove of the epigyn- 
um (center of Fig. 162) may be absent or 
different in width from that illustrated. 
Illustrations were made from specimens 
collected 80 km north of Manaus. 

Diagnosis. Micrepeira fowleri, unlike 
other species, has its abdomen black pos- 
teriorly, contrasting with the lighter an- 
terior three-quarters (Fig. 165). Females 
are smaller than M. hoeferi and are sep- 
arated from them by having a large tri- 
angular lobe of the epigynum with a cen- 
tral transverse groove or dark mark be- 
tween two darker areas (Fig. 162) and by 
having concave sides of the posterior me- 
dian plate (Fig. 163). The male differs from 
that of M. hoeferi by having a lobe in the 
middle of the median apophysis (Fig. 168). 
It is possible that all specimens considered 
M. fowleri are variants of M. tubulofa- 
ciens. 

Natural History. Specimens of M. fow- 
leri were collected in forest border and 



ACTINOSOMA, Spilasma, Micrepeira, Pronovs • Levi 195 




Figures 1 62-1 70. Micrepeira fowleri n. sp. 1 62-1 67, female. 1 62-1 64, epigynum. 1 62, ventral. 1 63, posterior. 1 64, lateral. 1 65, 
dorsal. 1 66, carapace. 1 67, eye region and chelicerae. 1 68-1 70, male. 1 68, left palpus. 1 69, carapace. 1 70, eye region, chielicerae, 
and righit palpus. 

Figures 171-181. Micrepeira hoeferin. sp. 171-176, female. 171-175, epigynum. 171, 174, ventral. 172, 175, posterior. 173, 
lateral. 171-173, (Mato Grosso State). 174, 175, (Amazonas State). 176, dorsal. 177-181, male. 177-180, left palpus. 177, 
mesal. 178, ventral. 179, 180 pulled apart. 181, dorsal. 

Abbreviations. C, conductor. E, embolus. M, median apopfiysis. R, radix. T, tegulum. 

Scale lines. 1.0 mm, genitalia 0.1 mm, except Figures 166, 167, 169, 170, 0.5 mm. 



196 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



forest interior. The retreat has parallel 
sides, about 4.0 to 4.5 mm wide and 10 to 
20 mm long. 

Distribution. Amazonian Ecuador, 
Peru to Manaus Brazil (Map 6). 

Paratypes. BRAZIL Amazonas: Km 41 Reserve, 25 
May 1991, 19 (H. G. Fowler, E. Venticinque, R. S. 
Vieira). 

Specimens Examined. ECUADOR Sucumhnos: 
Cuyabeno Reserve, Laguna Grande, 13 Feb. 1984, 
12 (L. Aviles, MECN). PERU Madre de Dios: Zona 
Reservada Tambopata, 290 m, 12°50'S, 69''17"W, 14 
June 1988, 12 (D. Silva D., MUSM). BRAZIL Ama- 
zonas: 80 km N Manaus, 2''24'S, 59'"52'W, 26 Feb. 
1989, 6, 9 Nov. 1989, 1<5 (H. G. Fowler, MCZ); Cabo 
Frio Reserve, 80 km N Manaus, 1989, 1990, 52, 26; 
Colosso Reserve, 80 km N Manaus, 1989, 1990, 1012, 
16<? (H. G. Fowler, E. Venticinque, R. S. Vieira, MCZ); 
Dimona Reserve, 1989-1992, 52, IS (H. G. Fowler, 
MCZ); Reserva Porto Alegre, 80 km N Manaus, 1989- 
1992, 22 (H. G. Fowler, MCZ); Reservacida de Powel, 
20 Apr. 1991, 12 (H. G. Fowler, MCZ); Reserva Ducke, 
4 Oct. 1990, 12 (H. Hofer, INPA). 



Micrepeira hoeferi new species 
Plate 4; Figures 171-181; Map 6 

Holotype. Female holotype from igapo Taruma 
Mirim, near Manaus, Amazonas State, Brazil, 11 
Feb. 1988 (H. Hofer), in INPA. The species is named 
after H. Hofer, who collected it and photographed 
its web. 

Spilasma tubulofaciens: — Quintero, 1974: 307, figs. 
1-6, 2, web. 

Description. Female holotype. Cara- 
pace dusky orange, sides of thorax darkest. 
Chelicerae light brown. Labium, endites, 
sternum dark orange-brown. Coxae, legs 
brown with indistinct darker rings. Dor- 
sum of abdomen with single and paired 
white patches on black (Fig. 176); sides, 
venter black. Posterior median eyes 1 di- 
ameter of anterior medians, laterals 0.8 
diameter. Anterior median eyes 0.8 di- 
ameter apart, 2.2 diameters from laterals. 
Posterior median eyes 1 diameter apart, 3 
diameters from laterals. Total length 5.0 
mm. Carapace 2.5 mm long, 2.1 wide, 1.5 
behind lateral eyes. First femur 1.8 mm, 
patella and tibia 1.8, metatarsus 1.1, tarsus 
0.8. Second patella and tibia 1.7 mm, third 
1.3, fourth 1.5. 



Male from Mato Grosso. Color darker 
than female. Carapace, sternum brownish 
black. Only proximal portion of femur, 
metatarsi, and tarsi light; others contrast- 
ingly ringed. Dorsum of abdomen with 
white cross and a pair of white posterior 
spots (Fig. 181), venter with white band 
around anterior above carapace. Posterior 
median eyes 0.8 diameter of anterior me- 
dians, laterals 0.7 diameter. Anterior me- 
dian eyes 0.8 diameter apart, 1.5 diameters 
from laterals. Posterior median eyes their 
diameter apart, 2.1 diameters from later- 
als. Total length 2.4 mm. Carapace 1.3 mm 
long, 1.3 wide, 0.7 behind lateral eyes. First 
femur 1.2 mm, patella and tibia 1.2, meta- 
tarsus 0.7, tarsus 0.6. Second patella and 
tibia 1.1 mm, third 0.7, fourth 0.9. 

Note. Males and females have been col- 
lected together; they have a similar color 
pattern on the abdomen (Figs. 176, 181). 

Variation. Total length of females 5.0 
to 6.3 mm, males 1.5 to 2.4. Figures 171- 
173 and 176-181 were made from speci- 
mens from Mato Grosso and Figures 174 
and 175 from the female holotype. 

Diagnosis. Females are larger than 
those of M. tubulofaciens, and the epi- 
gynum, in ventral view, has a straight pos- 
terior border (Figs. 171, 174). In both M. 
fowleri and M. tubulofaciens, the epigyn- 
um has a triangular median lobe in ventral 
view (Figs. 162, 182). 

Natural History. With the type spec- 
imen is a note saying that it came from a 
vertical web, the orb truncate above, with 
a cone-shaped retreat with cover; web di- 
ameter 20 cm, built about 1.5 m above 
ground, mesh width 2 to 3 mm (Pi. 4). The 
retreat is made of silk and detritus particles 
and is about 8 mm wide and 13 mm long. 
Another measured 7 by 30 mm. Two spec- 
imens collected by R. L. C. Baptista were 
1.8 and 2.0 m above ground, in forest. The 
female from French Guiana was collected 
with the female of M. tubulofaciens in 
forest clearing. 

Distribution. French Guiana to Peru, 
to Mato Grosso, Brazil (Map 6). It may 
occur in Paraguay, on the basis of a dia- 



AcTiNOSOMA, Spilasma, MiCREPEiRA, Pronous • Levi 197 



gram of a web, sent by J. Kochalka, with 
an inquiry about the genus of the builder. 

Specimens Examined. FRENCH GUIANA Cri- 
que Limonade, 3-4 km SW Saul, 21-22 Dec. 1972, 
19 (D. Quintero, MCZ); Montagnes Kaw nr. Camp 
Caiman, 27 km SE Roura, 4°33'N, 52''09"W, 25 Aug. 
1988, 9 (S. Marshall, USNM), PERU Madre de Dios: 
Zona Reservada Tambopata, 290 m, 12°50'S, 69°17'W, 
4 June 1988, 19 (J. Coddington, USNM), BRAZIL 
Roraima: Ilha de Maraca, 20 July 1987, 29 (A. A. Lise, 
MCN 20065); 19 (F. P. Benton, MCN 20066). Ama- 
zonas: Parq. Nacion. Pico da Neblina, Maturaca, 11 
Oct. 1990, 19 (A. A. Lise, MCP); Colosso Reserve, 80 
km N Manaus, 18 Jan. 1989, 29 (H. G, Fowler, MCZ); 
2 March 1990, 19 (H. G. Fowler, R. S. Vieira, E. 
Venticinque, MCZ); 80 km N Manaus, 17 Jan. 1989, 
19 (H. Fowler, MCZ); Reserva Campina, Manaus, 22 
Jan. 1973, U (L. P. Albuquerque, MCN 21179); Re- 
serva Ducke, Manaus, 4 Aug. 1987, 19 (A. A. Lise, 
MCN 23357); 15 Aug. 1991, 29 (A. D. Brescovit, MCN 
21392). Pernambuco: Dois Irmaos Forest Reserve, 25 
Jan. 1989, 19 (R. L. C. Baptista, A, P. Chaves, RLCB). 
Mato Grosso: Chapada dos Guimaraes, 15-26 July 
1992, 49, 26, 2 imm. (A. A. Lise, A. Braul, MCP, 2358, 
2361); Chavantina, 22 Jan. 1947, 19, doubtful det. (H. 
Sick, MZSP). Espirito Santo: Dois Bocas Forest Res- 
ervation, 15 Nov. 1988, 29 (R. L. C. Baptista, A. P. 
Chaves, RLCB). 



Micrepeira tubulofaciens (Hingston) 
new combination 
Figures 182-186; Map 6 

Epeira tubulofaciens Hingston, 1932: 366, figs. 40, 
41 (webs). Female holotype from Moraballi Creek, 
2 mi [3.2 km] east of Essequibo River, 12 mi [38.5 
km] south of Bartica, Guyana, in BMNH, lost. 

Aranea tubulifaciens: — Roewer, 1942; 854. 

Epeirella tubulofaciens: — Mello-Leitao, 1948: 164. 

Araneus tubulifaciens: — Bonnet, 1955: 620. 

Note. Kingston's illustrations (1932, 
figs. 40, 41) of Epeira tubulofaciens match 
the web of M. hoeferi (Pi. 4). Hingston 
described this species as having "colour 
black with a white area in front where base 
overhangs cephalothorax and a broad white 
T-shaped mark on the dorsum, the hori- 
zontal limb of the T crossing the mid-dor- 
sum transversely and the vertical limb run- 
ning forward to the center of the base." 
This description may fit all Micrepeira 
species; however, this is the species found 
in Guyana (although others may occur 
there), and Hingston gives the size of 3.1 



mm, while the similar, widespread M. hoe- 
feri is about 5 mm total length. 

A female was examined, which was cit- 
ed by Mello-Leitao (1948), and is depos- 
ited in the BMNH. 

I am not following Bonnet (1955) and 
Roewer (1942) in changing the spelling of 
"tubulofaciens", because Hingston was 
consistent in spelling the name with an 
"o". 

Description. Female from French Gui- 
ana. Cephalothorax light orange, legs 
slightly dusky, distal ends darkest. Dorsum 
of abdomen black with gray and white 
pigment patches (Fig. 185); venter with 
black band from genital groove to spin- 
nerets and around spinnerets; sides dusky 
orange. Posterior median eyes same di- 
ameter as anterior medians, laterals 0.9 di- 
ameter. Anterior median eyes 0.5 diame- 
ter apart, 1 diameter from laterals. Pos- 
terior median eyes 0.5 diameter apart, 1.1 
diameters from laterals. Total length 3.1 
mm. Carapace 1.3 mm long, 1.3 wide, 0.7 
behind lateral eyes. First femur 1.2 mm, 
patella and tibia 1.2, metatarsus 0.7, tarsus 
0.5. Second patella and tibia 1.1 mm, third 
0.6, fourth 1.1. 

Variation. Total length of females 3.1 
to 3.6 mm. The illustrations were made 
from a female from French Guiana. 

Diagnosis. This species is smaller than 
M. hoeferi, and its epigynum is a large 
triangle with distinct markings (Fig. 182). 
It differs from M. fowleri by having the 
black cap on the posterior of the abdomen 
less distinct (Fig. 185). 

Natural History. A retreat measured 
5.4 mm wide and 13 mm long. Judging 
by Hingston's illustrations, the web is sim- 
ilar to that of M. hoeferi (PI. 1). The spec- 
imen from French Guiana came with a 
female of the larger M. hoeferi and was 
collected in forest clearing; the specimen 
from Canje Ikuruwa River came from for- 
est savanna. 

Distribution. Guianas (Map 6). 

Specimens Examined. GUYANA Canje Ikuruwa 
River, 05''30'N, 57°30"W, Aug.-Dec. 1961, 19 (G. 
Bentley, AMNH); Higher Potaro River Distr. 19 (R. 



198 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



Lloyd, BMNH); Kartabo, 1924, 12 (AMNH). 
FRENCH GUIANA Crique Limonade, 3-4 km SW 
Saul, 21, 22 Dec. 1972, 9 (D. Quintero, MCZ); Mon- 
tagnes Kaw, nr. Camp Caiman, ca. 27 km SE Roura, 
4°33'N, 52°09'W, 5, 7 Aug. 1988, 29 (S. Marshall, 
USNM). 



Micrepeira pachitea new species 
Figures 187-191; Map 6 

Holotype. Female holotype from Panguana, Rio 
Pachitea, Depto. Huanuco, Peru, 9°37'S, 74°56'W, 
1987 (C. Manhart), in MCZ. The specific name is 
a noun in apposition after the locality. 

Description. Female holotype. Cara- 
pace orange-brown. Chelicerae, labium, 
endites, sternum brown. Legs brown. Dor- 
sum of abdomen black with white patches 
(Fig. 190); venter black with a white trans- 
verse band anterior of pedicel. Posterior 
median eyes 0.8 diameter of anterior me- 
dians, anterior laterals 0.8 diameter, pos- 
terior 1 diameter. Anterior median eyes 
0.7 diameter apart, 2 diameters from lat- 
erals. Posterior median eyes 0.8 diameter 
apart, 2.4 diameters from laterals. Total 
length 4.8 mm. Carapace 2.3 mm long, 1.9 
wide, 1.3 behind lateral eyes. First femur 
2.0 mm, patella and tibia 1.9, metatarsus 
1.0, tarsus 0.8. Second patella and tibia 1.8 
mm, third 1.3, fourth 1.6. 

Variation. Total length of females 4.8 
to 5.6 mm. 

Diagnosis. The sternum of M. pachitea 
has an anterior median groove (Fig. 191) 
and the epigynum is a broad, wide triangle 
(Fig. 187). 

Natural History. The cone-shaped re- 
treat, preserved with the specimen, is about 
7 mm wide and 21 mm long, another is 
13 mm long. 

Distribution. Peruvian Amazon (Map 6). 

Specimens Examined. PERU Loreto: Genaro 
Herrera, 04°55'S, 73°45'W, 28 Aug. 1988, 39 (D. Silva 
D., MUSM). Madre de Dios: Zona Reservada Tam- 
bopata, 290 m, 14 June 1988, 19 (D. Silva D., MUSM). 

Micrepeira smithae new species 
Figures 192-195; Map 6 

Holotype. Female holotype from Voltzberg-Ral- 
eighvallen Reserve, 4°45'N, 56°10'W, Surinam, 



April, May 1984 (D. Smith Trail), in MCZ. The 
species is named after the collector, arachnologist 
D. Smith. 

Description. Female holotype. Cara- 
pace dark brown. Chelicerae, labium, en- 
dites, sternum, dark brown. Legs brown. 
Dorsum of abdomen black with white 
chevrons (Fig. 195); venter black, with 
paired anterior white spots, one lying above 
each third femur when the spider is in 
resting position. Posterior median eyes 
same diameter as anterior medians, ante- 
rior laterals 0.8 diameter, posterior laterals 
1 diameter. Anterior median eyes their di- 
ameter apart, 2 diameters from laterals. 
Posterior median eyes 0.8 diameter apart, 
2.2 wide diameters from laterals. Total 
length 3.5 mm. Carapace 1.8 mm long, 1.7 
wide, 1.1 behind lateral eyes. First femur 
1.4 mm, patella and tibia 1.3, metatarsus 
0.7, tarsus 0.6. Second patella and tibia 1.3 
mm, third 0.9, fourth 1.2. 

Diagnosis. The epigynum of M. smi- 
thae differs from others by having a small 
triangle at the posterior margin, tapering 
into a tiny scale (Fig. 192), and by having 
a wide depressed median plate in posterior 
view (Fig. 193). 



Micrepeira velso new species 
Figures 196-199; Map 6 

Holotype. Female holotype and immature paratype 
from Finca La Selva, near Puerto Viejo, Heredia, 
Costa Rica, Jan. 1978 (W. Eberhard no. 1280), in 
MCZ. The specific name is an arbitrary combina- 
tion of letters. 

Description. Female holotype. Cara- 
pace dark orange-brown. Chelicerae or- 
ange-brown. Labium, endites, sternum or- 
ange-brown. Legs brown. Dorsum of ab- 
domen black with white patches (Fig. 199); 
venter black with paired anterior white 
patches lying above fourth coxae. Eyes 
subequal in size. Anterior median eyes 0.8 
diameter apart, 2.5 diameters from later- 
als. Posterior median eyes their diameter 
apart, 2.7 diameters from laterals. Total 
length 4.7 mm. Carapace 2.1 mm long, 2.0 
wide, 1.1 behind lateral eyes. First femur 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 199 




200 





Figures 182-186. Micrepeira tubulofaciens (Hingston), female. 182-184, epigynum. 182, ventral. 183, posterior. 184, lateral. 
185, dorsal. 186, sternum. 

Figures 187-191. M. pachitea n. sp., female. 187-189, epigynum. 187, ventral. 188, posterior. 189, lateral. 190, dorsal. 191, 

sternum. 

Figures 192-195. M. smithae n. sp., female. 192-194, epigynum. 192, ventral. 193, posterior. 194, lateral. 195, dorsal. 

Figures 196-199. M. velso n. sp., female. 196-198, epigynum. 196, ventral. 197, posterior. 198, lateral. 199, dorsal. 

Figures 200-204. M. albomaculata Schenkel, female. 200-202, epigynum. 200, ventral. 201 , posterior. 202, lateral. 203, dorsal. 
204, lateral. 

Scale lines. 1.0 mm, genitalia 0.1 mm, except Figures 186, 191, 0.5 mm. 



200 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



1.8 mm, patella and tibia 1.7, metatarsus 
0.9, tarsus 0.7. Second patella and tibia 1.6 
mm, third 1.2, fourth 1.4. 

Diagnosis. This species differs by the 
triangular median piece of the epigynum 
overhanging the posterior margin (Fig. 
196) and by the raised median posterior 
plate that overhangs the lateral plates (Fig. 
197). 

Natural History. The web was at the 
edge of a clearing and was tattered in the 
evening, fresh in the morning, suggesting 
that the spider builds early in the morning. 
The spider was in a retreat and pulled the 
sides of the retreat together when dis- 
turbed. The web was 28 cm wide (Eber- 
hard, personal communication and pho- 
tographs); the retreat, collected with the 
specimen, was 6 mm wide and 11 mm 
long. 

Paratype. COSTA RICA Heredia: Sarapiqui nr. 
Puerto Viejo, Sept. 1981, 12 (C. E. Griswold, CAS). 



Micrepeira albomaculata Schenkel 
Figures 200-204; Map 6 

Micrepeira albomaculata Schenkel, 1953: 26, fig. 24, 
9. Female holotype from El Pozon, Depto. Acosta, 
Est. Falcon, Venezuela, in NMB, examined. Brig- 
noli, 1983: 276. 

Description. Female holotype. Cara- 
pace dark orange, sides of thoracic region 
dusky. Chelicerae, labium, endites, ster- 
num orange. Legs dark orange. Dorsum 
of abdomen black with median and paired 
white patches (Figs. 203, 204); venter 
dusky, without white pigment. Chelicerae 
with two teeth on anterior margin. Eyes 
subequal. Anterior median eyes 0.8 di- 
ameter apart, 1.5 diameters from laterals. 
Posterior median eyes their diameter apart, 
1.5 diameters from laterals. Total length 
3.5 mm. Carapace 1.43 mm long, 1.31 
wide. First femur 1.01 mm, patella and 
tibia 1 .03, metatarsus 0.57, tarsus 0.52. Sec- 
ond patella and tibia 1.04 mm, third 0.68, 
fourth 0.96. 

Diagnosis. The bulbous median lobe of 



the epigynum (Fig. 200) and the T-shaped 
posterior median plate (Fig. 201) separate 
this species from all other known Micre- 
peira species. 



Madrepeira new genus 

Type species. Madrepeira amazonica. The name is 
an arbitrary combination of letters prefixed to 
"epeira . The name is feminine. 

Diagnosis. The genus is close to Acu- 
lepeira, Amazonepeira, Metepeira, Kaira, 
and Tatepeira, all of which exhibit two 
known synapomorphies: the pointed scape 
of the epigynum and the pair of flagella 
on the median apophysis (M in Fig. 213) 
of the male palpus. Madrepeira differs by 
the shape of the epigynum, which has lat- 
eral and median plates fused into a base 
and, on each side of the attachment of the 
scape, a scale formed from the posterior 
median plate (Figs. 205-208). It also dif- 
fers from other genera by the diamond- 
shaped abdomen with paired lateral humps 
and by having spindly legs with only a few 
long setae (Fig. 209). The genus also differs 
in making a ladderweb (Pi. 5), similar to 
that of Scoloderus. 

Relationship. Characters that ally this 
genus to Araneus include placement of the 
conductor (C in Fig. 213), covering the 
tegulum at the upper right (in the left pal- 
pus, at 2-3 hr in Fig. 212), the presence 
of distal hematodocha (DH) between the 
embolus (E) and the terminal apophysis (A 
in Fig. 213), the median apophysis with a 
distal spine and a proximal pair of flagella 
(M in Fig. 213), and two palpal patellar 
macrosetae (at 10 hr in Fig. 216) (the last 
character sometimes missing in a few spe- 
cies belonging to this group). The female 
has an annulate scape as do most relatives 
of Araneus (but also Parawixia and Er- 
iophora, both Alpaida relatives). 

Natural History. Madrepeira makes a 
short ladderweb in forests (Pi. 5). 

Distribution. There is only one Neo- 
tropical species. 



AcTiNOSOMA, Spilasma, MiCREFEiRA, Pronous • Levi 201 




Plate 5. Madrepeira amazonica n. sp. Web of an immature, about 5.1 cm wide, 16.5 cm long, calculated from the size of the 
spider (photo, J. Coddington). 



Madrepeira amazonica new species 
Plate 5; Figures 205-216; Map 7 

Holotype. Male holotype from Albergue "Cuzco 
Amazonica," 200 m, 12°33'S, 69°03'W, Rio Madre 
de Dios, Depto. Madre de Dios, Peru, 9 Mar. 1990 
(D. Silva D.), in MUSM. The specific name is a 
noun in apposition after the type locahty. 

Description. Female paratype. Cara- 
pace yellowish white with pink on each 
side of cephalic region. Chelicerae, labi- 
um, endites, sternum, legs yellowish white. 
Dorsum of abdomen with black dots, ex- 
cept for two light patches, one on each 
side (Fig. 209). Centers of light patches 
with some pink dots. Sides with a black 



line bordering the dorsal black spots and 
the white side of venter; venter yellowish 
with white on each side. Posterior median 
eyes 0.5 diameter of anterior medians, an- 
terior laterals 0.6 diameter, posterior lat- 
erals 0.5 diameter. Anterior median eyes 
their diameter apart, their diameter from 
laterals. Posterior median eyes 1.5 diam- 
eters apart, 2.2 diameters from laterals. 
Ocular quadrangle narrower behind than 
in front. Height of clypeus equals 0.4 di- 
ameter of anterior median eye. One che- 
licera with five teeth on anterior margin, 
other with four teeth; four teeth on one 
posterior margin, three on other; denticles 



202 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 




amazonica 



Map 7. Distribution of Madrepeira amazonica. 



in grooves. Legs spindly with siiort, tine 
setae and a few very long, dark macrosetae 
(Fig. 214). Abdomen diamond-shaped (Fig. 
209). Total length 5.0 mm. Carapace 2.1 
mm long, 1.7 wide, 1.0 behind lateral eyes. 
First femur 3.7 mm, patella and tibia 4.2, 
metatarsus 3.6, tarsus 1.5. Second patella 
and tibia 3.4 mm, third 1.7, fourth 2.5. 

Male holotype. Color as in female. Pos- 
terior median eyes 0.5 diameter of anterior 
medians, anterior laterals 0.5 diameter, 
posterior laterals 0.4 diameter. Anterior 
median eyes their diameter apart, 0.3 di- 
ameter from laterals. Posterior median eyes 
their diameter apart, 1.5 diameters from 
laterals. Ocular quadrangle narrower be- 
hind than in front. Height of clypeus equals 
0.2 diameter of anterior median eye. En- 
dite with cone-shaped tooth facing a tu- 
bercle on palpal femur. Palpal patella with 
two macrosetae (Fig. 216). First coxa with 
a minute hook on distal margin. First tibia 
with a pair of macrosetae on a slight tu- 
bercle (Fig. 214). Second tibia as thick as 
first, not modified Abdomen diamond- 
shaped. Total length 3.0 mm. Carapace 
1.43 mm long, 1.30 wide, 0.68 behind lat- 
eral eyes. First femur 2.92 mm, patella and 



tibia 3.32, metatarsus 2.62, tarsus 1.17. Sec- 
ond patella and tibia 2.56 mm, third 1.17, 
fourth 1.87. 

Note. Males and females were matched 
because they have similar markings and 
abdomen shape and were collected to- 
gether. The black horseshoe-shaped ap- 
pendage of the embolus (Figs. 212, 213) 
breaks off and stays attached to the epi- 
gynum (Fig. 206), perhaps blocking in- 
semination by other males. 

Variation. Total length of females 4.7 
to 5.8 mm, males 3.0 to 3.5. Illustrations 
were made from a female paratype and 
male holotype, except for Figures 210, 211, 
215, and 216, which were made from Bo- 
livian specimens. One immature specimen 
had four white patches across the abdo- 
men, one anterior median patch, and five 
confluent posterior patches each with a 
pink center and white outside. Some in- 
dividuals have black rings on the patella. 
Others had a red line around the white 
patches on the abdomen. Eyes of Bolivian 
specimens are slightly farther apart than 
those from Peru. 

Natural History. Most specimens were 
obtained by night collecting. Bolivian 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 203 




.OCz-^O 






}/ 205 




Figures 205-21 6. Madrepeira amazonica n. sp. 205-21 1 , female. 205-208, epigynum. 205, ventral. 206, ventral, mated. 207, 
posterior. 208, lateral, 209, dorsal. 210, carapace. 211, eye region and chelicerae. 212-216, male. 212, 213, left palpus. 212, 
mesal, 213, pulled apart. 214, first left patella and tibia, prolateral. 215, carapace. 216, eye region, chelicerae, and right palpus. 

Abbreviations. A, terminal apophysis. C, conductor. DH, distal hematodocha. E, embolus. M, median apophysis. R, radix. T, 
tegulum. 

Scale lines. 1.0 mm, genitalia 0.1 mm, except Figures 210, 211, 215, 216, 0.5 mm. 



specimens were observed in a ladderweb 
(PI. 5). 

Distribution. Amazon region to south- 
ern Mato Grosso, southern Bahia States, 
Brazil (Map 7). 

Paratypes. PERU Madre de Dios: Albergue "Cuz- 
co Amaconica", 200 m, Rio Madre de Dios, 16 May- 
12 June 1989, 4$, IS (D. Silva D., MUSM, 12 in MCZ). 

Specimens Examined. PERU Madre de Dios: Zona 
Reservada Tambopata, 30 km SW Puerto Maldonado, 
6-14 Sept. 1984, IS (T. L. Erwin et al., USNM). BRA- 



ZIL Roraima: Ilha de Maraca, Rio Uraricoera, 25 
Mar. 1987, 19 (A. A. Lise, MCN 20070). Amazonas: 
Lago de Jose, Manaus, 9 Aug. 1987, 19 (J. Adis, MCN 
20071); Rio Taruma, Mirim, Manaus, 30 July 1979, 
1 imm. (J. Adis et al., MCN 20054). Bahia: Fazenda 
Nossa Senhora das Neves, Itamarajii, 9 Oct. 1978, 1 
S (J. S. Santos, MCN 11089); Fazenda Matiapa, Ca- 
macan, 16 Oct. 1978, 19 (J. S. Santos, MCN 11113). 
Mato Grosso: Pocone, Fazenda Santa Ines, Pantanal, 
5-11 Aug. 1992, 3 imm, IS (A. A. Lise, A. Braul, MCP 
2362). BOLIVIA Beni: Station Biologica Beni, on trail 
betw. Zones 1 & 2, 12 Sept. 1987, 19 (J. Coddington, 
USNM). 



204 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



Tatepeira new genus 

Type species. Aranea tatarendensis Tullgren, 1905. 
The name is an arbitrary combination of letters, 
prefixed to "epeira"; its gender is feminine. 

Diagnosis. Tatepeira is close to Acu- 
lepeira, Amazonepeira, Madrepeira, Me- 
tepeira, and Kaira, all of which have a 
pointed scape on the epigynum (Figs. 217- 
219) and, in the male, a pair of flagella on 
the median apophysis (M in Fig. 224). Ta- 
tepeira differs from these genera by hav- 
ing a pair of large dorsal humps on the 
abdomen (Figs. 220, 227, 234, 238). (Acu- 
lepeira vdsite Levi, 1991a, does have humps 
on the abdomen and may belong here.) 
The type species T. tatarendensis has the 
scape of the epigynum on a pedestal (Figs. 
217-219), making the epigynum longer 
than wide as seen from the side (Fig. 219). 
ISlote. Three of the four species are 
placed here tentatively: Tatepeira itu lacks 
a terminal apophysis and the distal he- 
matodocha in the palpus (Figs. 229, 230). 
Tatepeira stadelmani and T. carrolli are 
similar to each other, but until their males 
are known their generic placement is un- 
certain. Before I revised Kaira (Levi, 
1993c), I considered these two to belong 
to Kaira, but the distal articles of the legs 
are not as curved and spinose as in species 
of Kaira. 

Relationship. The pointed scape of the 
epigynum (Figs. 217, 225) and the two 
flagella on the median apophysis (M in Fig. 
224) relate Tatepeira to Aculepeira, Ama- 
zonepeira, Madrepeira, Metepeira, and 
Kaira. These genera, as others related to 
Araneus and Neoscona, has the palpus with 
distal hematodocha (DH in Fig. 224), the 
conductor is near the edge of the tegulum 
below the arrow-shaped tip of the terminal 
apophysis (Fig. 223, C in Fig. 224), and 
there are two setae on the palpal patella. 
Natural History. The web is not known 
for any of the species. 

Distribution. All four species are Neo- 
tropical. 



Key to Species Here Placed in Tatepeira 

The males of T. stadelmani and T. carrolli are not 
known. 

1 . Female _ 3 

Male _ 2 

2( 1 ). Palpal flagella attached to distal end of me- 
dian apophysis of palpus and median 
apophysis without row of teeth (Figs. 222, 
223, M in Fig. 224); widespread (Map 8) 

__ __ __ tatarendensis 

Flagella on proximal end of median apoph- 
ysis, distal end with teeth (Figs. 229, 230); 
southern Brazil (Map 8) _ itu 

3(1). Scape of epigynum on a pedestal (Figs. 217- 

219); widespread (Map 8) tatarendensis 

Epigynum flat (Figs. 225, 226, 231-233, 
235-237) _ _ - - 4 

4(3). Abdomen humps facing dorsally (Fig. 227); 
total length less than 4 mm; southern Bra- 
zil (Map 8) ._ - itu 

Large humps facing laterally (Figs. 234, 
238); total length more than 5 mm 5 

5(4). In posterior view of epigynum, only lateral 
plates sclerotized and lateral plates with 
a ventral concave notch (Fig. 232); Hon- 
duras (Map 8) stadelmani 

In posterior view of epigynum, median 
plates sclerotized and ventrally curled 
(Fig. 236); Colombia (Map 8) carrolli 



Tatepeira tatarendensis (Tullgren) 
new combination 
Figures 217-224; Map 8 

Aranea tatarendensis Tullgren, 1905: 34, pi. 5, fig. 
12, 9. Female holotype from Tatarenda [Depto. 
Tarija, 600 m, northeast of Aguairenda, 21°50'S, 
63"'37'W, on border between tropical forest and dry 
woods (Paynter, 1993)], Bolivia, in NRMS, exam- 
ined. Roewer, 1942: 853. 

Araneus holmi Caporiacco, 1955: 354, fig. 34, 2. Fe- 
male holotype from Maiquetia, Distr. Federal, 
Venezuela, in UCVC. NEW SYNONYMY. 

Araneus tatarendensis: — Bonnet, 1955: 609. 

Araneus akeholmi Brignoli, 1983: 252. New name for 
A. holmi, preoccupied. NEW SYNONYMY. 

Wixia tatarendensis: — Levi, 1991b: 177. 

Description. Female from northern 
Colombia. Carapace yellowish white, 
dusky on each side of head, and carapace 
underlain by a white pigment patch. Car- 
apace lightly sclerotized, without setae, not 
shiny. Chelicerae orange. Labium, endites 
dusky. Sternum black, lightest in center. 
Coxae yellowish white; legs yellowish. 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 



205 




Figures 217-224. Tatepeira tatarendensis (Tullgren). 217-221, female. 217-219, epigynum. 217, ventral. 218, posterior. 219, 
lateral. 220, dorsal. 221, abdomen, ventral. 222-224, left male palpus. 222, mesal. 223, ventral. 224, pulled apart. 

Figures 225-230. T. itu n. sp. 225-228, female. 225-226, epigynum. 225, ventral. 226, posterior. 227, dorsal. 228, abdomen, 
ventral. 229, 230, palpus. 229, mesal. 230, ventral. 

Figures 231-234. T. stadelmani n. sp., female. 231-233, epigynum. 231 , ventral. 232, posterior. 233, lateral. 234, dorsal. 

Figures 235-238. T. carrollin. sp., female. 235-237, epigynum. 235, ventral. 236, posterior. 237, lateral. 238, dorsal. 

Abbreviations. A, terminal apophysis. C, conductor. DH, distal hematodocha. E, embolus. M, median apophysis. R, radix. T, 
tegulum. 

Scale lines. 1.0 mm, genitalia 0.1 mm. 



206 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



HONDURAS 




V Carroll i 
Q itu 

A stadelmani 
• tatarendensis 










Mf^, 



Map 8. Distribution of Tatepeira species. 



Dorsum of abdomen white with pairs of 
curved black hnes approaching each other 
posteriorly, and dusky spotted areas (Fig. 
220); venter black between epigynum and 
spinnerets with a wide, white, longitudinal 
streak on each side and a white spot on 
each side of black spinnerets (Fig. 221). 
Posterior median eyes 1.3 diameters of an- 
terior medians, laterals same diameter as 
anterior medians. Anterior median eyes 1 
diameter apart, 1.5 diameters from later- 
als. Posterior median eyes 0.8 diameter 
apart, 2.2 diameters from laterals. Ocular 
quadrangle square. Height of clypeus 
equals 0.5 diameter of anterior median eye. 
Abdomen slightly longer than wide with 
anterolateral humps (Fig. 220). Total 
length 2.9 mm. Carapace 1.6 mm long, 1.2 
mm wide, 0.7 behind lateral eyes. First 
femur 1.5 mm, patella and tibia 1.8, meta- 
tarsus 1.3, tarsus 0.6. Second patella and 
tibia 1.7 mm, third 0.9, fourth 1.5. 

Male from northern Colombia. Color as 



in female. Carapace with longitudinal me- 
dian line in thoracic region. Posterior me- 
dian eyes same diameter as anterior me- 
dians, laterals 0.8 diameter. Anterior me- 
dian eyes their diameter apart, 1.5 diam- 
eters from laterals. Posterior median eyes 
0.8 diameter apart, 2 diameters from lat- 
erals. Ocular quadrangle narrower behind 
than in front. Height of clypeus equals 0.8 
diameter of anterior median eye. Endite 
with tooth, palpal femur with facing tu- 
bercle. Palpal patella with two macrosetae. 
First coxa with large hook. Second tibia 
thicker than first, swollen, and with pro- 
lateral macrosetae. Abdomen oval without 
humps. Total length 3.4 mm. Carapace 1.5 
mm long, 1.4 wide, 0.7 behind lateral eyes. 
First femur 2.2 mm, patella and tibia 2.4, 
metatarsus 1.6, tarsus 0.7. Second patella 
and tibia 1.6 mm, third 1.1, fourth 1.8. 

Note. Males and females were paired 
on the basis of matching genitalia, the epi- 
gynum having a pointed scape and the 



AcTiNOSOMA, Spilasma, Micrepeira, Pronous • Levi 207 



median apophysis of the male a pair of 
flagella. Also, both have a similar colora- 
tion and black rings around the eyes. They 
were collected near each other. 

Variation. Total length of males 2.9 to 
3.4 mm. Most males have a dorsal folium 
on the abdomen. The illustrations were 
made from individuals from northern Co- 
lombia. 

Diagnosis. Tatepeira tatarendensis is 
separated from other species placed in this 
genus by the epigynum, which has the 
scape on a pedestal (Figs. 217-219), and 
by the median apophysis of the male, which 
has a pair of flagella on its distal end (Figs. 
222-224). 

Natural History. Males were collected 
in the herb layer in Colombia, in campo- 
grassland in Mato Grosso, Brazil. 

Distribution. Northern Colombia to 
southern Bolivia and Mato Grosso, Brazil 
(Map 8). 

Specimens Examined. COLOMBIA Magdalena: 
Bahia Concha, Tayrona Park, 10 km E Santa Marta, 
from low vegetation, 23 June 1985, 19 (H.-G. Muller, 
SMF 36900); Villa Culebra nr. Bonda, 10 km E Santa 
Marta, Oct. 1985, 3(5, 1 imm. (H.-G. Muller, SMF, 
MCZ); 1-11 Nov. 1985, IS (H.-G. Muller, SMF). Ces- 
ar: Velledupar, 4-9 June 1968, \$ (B. Malkin, AMNH). 
BRAZIL Mato Grosso: 260 km N Xavantina, 12°49'S, 
5r46'W, 400 m, Feb.-Apr. 1969, IS (Xavantina- 
Cachimbo Exped., MCZ). 



Tatepeira itu new species 
Figures 225-230; Map 8 

Holotype. Female holotype from Fazenda Pau 
d'Alho, Municipio de Itu, Sao Paulo State, Brazil, 
2 Feb. 1959 (F. Lane), in AMNH. The specific 
name is a noun in apposition after the type locality. 

Description. Female holotype. Cara- 
pace yellowish with dusky patches, white 
pigment underneath center of thoracic area 
and black rings around eyes. Chelicerae 
yellowish, proximally dusky. Labium 
dusky, endites yellow-white. Sternum 
dusky orange. Coxae yellowish; legs yel- 
lowish with narrow brownish to black rings. 
Dorsum of abdomen white with black 
patches (Fig. 227); venter with a pair of 
black patches with some white pigment 



posteriorly and to sides (Fig. 228). Poste- 
rior median eyes 1.3 diameters of anterior 
medians, anterior laterals 0.9, posterior lat- 
erals 1.2 diameters. Anterior median eyes 
their diameter apart, 1 diameter from lat- 
erals. Posterior median eyes 1.5 diameters 
apart, 1.5 diameters from laterals. Ocular 
quadrangle wider behind than in front. 
Height of clypeus equals 0.2 diameter of 
the anterior median eye. Abdomen oval 
(shrivelled), with two humps (Fig. 227). 
Total length 3.4 mm. Carapace 1.41 mm 
long, 1.36 wide, 0.85 wide behind lateral 
eyes. First femur 1.62 mm, patella and 
tibia 2.31 , metatarsus 1 .39, tarsus 0.58. Sec- 
ond patella and tibia 1.82 mm, third 1.11, 
fourth 1.45. 

Male paratype. Color as in female. Pos- 
terior median eyes 1.3 diameters of ante- 
rior medians, laterals 0.9 diameter. Ante- 
rior median eyes 2 diameters apart, 2 di- 
ameters from laterals. Posterior median 
eyes 1.2 diameters apart, 1.5 diameters 
from laterals. Ocular quadrangle slightly 
wider behind than in front. Height of clyp- 
eus equals 0.2 diameter of the anterior me- 
dian eye. Endite without tooth. Palpal pa- 
tella without macroseta. First coxa without 
hook. Second tibia as thick as first. Ab- 
domen as in female. Total length 3.0 mm. 
Carapace 1.45 mm long, 1.27 wide, 0.75 
wide behind lateral eyes. First femur 1.78 
mm, patella and tibia 1.98, metatarsus 1.22, 
tarsus 0.52. Second patella and tibia 1.78 
mm, third 0.94, fourth 1.29. 

Note. Males and females were matched 
on the basis of their genitalia, the epigyn- 
um with a pointed scape, and the median 
apophysis with a pair of flagella. Also, they 
have a similar abdomen shape and similar 
markings. The species is tentatively placed 
in Tatepeira. 

Diagnosis. The heart-shaped frame 
around the epigynum (Figs. 225, 226) and 
the shape and armature of the median 
apophysis separate T. itu from other spe- 
cies with similar genitalia. Unlike similar 
species, the palpus lacks a terminal apoph- 
ysis and distal hematodocha (Figs. 229, 
230). 



208 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



Paratype. BRAZIL Sao Paulo-. Jabaquara, Cidade 
Sao Paulo, 21 Dec. 1945, 1<5 paratype (H. Sick, AMNH). 

Specimens Examined. BRAZIL Rio Grande do 
Sul: Montenegro, 15 Nov. 1977, 12 (H. Buckup, MCN 

7535). 



Tatepeira stadetmani new species 
Figures 231-234; Map 8 

Holotype. Female holotype from Subirana, Depto. 
Yoro, Honduras, no date (Stadelman), in MCZ. The 
species is named after the collector. 

Description. Female holotype. Cara- 
pace orange with darker orange spots and 
a median longitudinal gray line; clypeus 
black. Black band covering anterior me- 
dian eyes, branching on each side, with 
upper branch covering lateral eyes, lower 
covering edge of cephalic region. Chelic- 
erae orange, with darker spots and patches. 
Labium, endites dusky brown. Sternum 
brown, lighter in middle toward labium. 
Coxae orange, mottled brown; legs orange, 
with irregular narrow brown rings. Dor- 
sum of abdomen white with indistinct pos- 
terior gray folium; folium as wide in front 
as behind, bordered by pairs of reverse 
parentheses (Fig. 234). Sides with longi- 
tudinal black line, which stops abruptly 
toward dorsum and fades toward venter. 
Venter colorless with a pair of faint white 
lines. Dorsum of carapace with a circular 
thoracic depression. Posterior median eyes 
0.8 diameter of anterior medians, laterals 
0.6 diameter. Anterior median eyes 0.9 di- 
ameter apart, 1.5 diameters from laterals. 
Posterior median eyes their diameter apart. 
Ocular quadrangle narrower behind than 
in front. Height of clypeus equals 0.6 di- 
ameter of anterior median eye. Abdomen 
with pair of lateral humps (Fig. 234). Total 
length 7.5 mm. Carapace 3.4 mm long, 2.7 
wide, 1.5 behind lateral eyes. First femur 
4.5 mm, patella and tibia 5.6, metatarsus 
3.4, tarsus 1.4. Second patella and tibia 4.9 
mm, third 2.7, fourth 3.8. 

Note. The abdomen of the holotype is 
shrivelled and damaged. 

Diagnosis. Tatepeira stadetmani is 
similar to T. carrolli but differs in having 
a circular thoracic depression and an epi- 



gynum having sclerotized lateral plates 
with a wide ventral notch in posterior view 

(Fig. 232). 

Tatepeira carrolli new species 
Figures 235-238; IVlap 8 

Holotype. Female holotype from Lomalinda, ■! 
03°18'N, 73°22'W, near Puerto Lleras, 300 m, grass- 
lands "with patches of jungle and marsh", Depto. 
Meta, Colombia, 13 Apr. 1986 (B. T. Carroll), in 
MCZ. The species is named after the collector. 

Description. Female holotype. Cara- 
pace yellowish with irregular dusky marks, 
area between median eyes and clypeus 
gray, indistinctly branching on sides as in 
T. stadelmani. A dark mark in deep, lon- 
gitudinal thoracic groove. Labium, endites 
dusky yellow. Sternum dusky yellow with 
median white streak. Coxae, legs yellowish 
with indistinct narrow dark rings. Dorsum 
of abdomen whitish with dusky marks; fo- 
lium widest in front (Fig. 238); sides with 
a distinct longitudinal black line fading 
toward venter; venter with little pigment, 
a dusky white square between epigynum 
and spinnerets. Posterior median eyes 0.8 
diameter of anterior medians, laterals 0.7 
diameter. Anterior median eyes 1.3 di- 
ameters apart, 2.1 diameters from laterals. 
Posterior median eyes 1.2 diameters apart. 
Ocular quadrangle narrower behind than 
in front. Height of clypeus equals diameter 
of anterior median eye. Abdomen with pair 
of lateral humps and two more pairs of 
dorsal humps in a line between big humps, 
also pairs of smaller bulges on sides pos- 
teriorly (Fig. 238). Total length 10.5 mm. 
Carapace 5.2 mm long, 4.5 wide, 2.2 be- 
hind lateral eyes. First femur 7.0 mm, pa- 
tella and tibia 8.4, metatarsus 5.5, tarsus 
2.0. Second patella and tibia 8.0 mm, third 
4.0, fourth 6.2. 

Diagnosis. Tatepeira carrolli is similar 
to T. stadelmani but differs by having a 
longitudinal groove in the thoracic region 
(Fig. 238) and by having the epigynum 
with the posterior median plate sclerotized 
and elevated above the lateral plates, its 
ventral borders curling toward the sides 
(Fig. 236). 



AcTiNOSUMA, Spilasma, MicREPEiRA, Pronvus • Levi 



209 



NOTE 

In 1929, Major R. W. G. Kingston (who 
previously had pubhshed 10 books and pa- 
pers on spiders, as cited in Bonnet, 1945) 
led an expedition to British Guiana. He 
had previously been on expeditions to 
tropical Asia. The 3-month expedition to 
Guyana included 12 Europeans and 12 In- 
dians. They collected from treetops by 
climbing trees with spike ladders, spiked 
boots, pulleys, and observation chairs. A 
report on this expedition was published by 
Kingston in 1932 (A Naturalist in the Gui- 
ana Forest). 

Kingston's volume contains 84 pages on 
the expedition and 151 pages on spider 
webs and protective adaptations of spiders. 
Various chapters describe protective and 
warning devices of insects, especially ants, 
termites, and caterpillar cases, and nest 
suspensions of birds and spiders. A chapter 
on tree roofs reports collecting 2,000 in- 
sects. It is stated that all specimens are 
deposited in the British Museum, Natural 
History (Natural History Museum, Lon- 
don, England), but the spiders have not 
been found. 

Most important, Kingston includes an 
appendix of new spider species found. Un- 
able to find a specialist to help him deter- 
mine species, he gave new names to the 
spiders observed. Twenty-seven spiders are 
named and described, and these names are 
listed in the catalogs of Roewer (1942) and 
in Bonnet (1955, 1958, 1959). It has been 
difficult to interpret Kingston's half-page 
descriptions without specimens. He did not 
often describe genitalia or other species- 
specific characters; however, the spiders' 
webs are illustrated in earlier chapters. 
Some species and webs are small and ap- 
pear to be immature. Evidence for this 
occurs in the descriptions of three new spe- 
cies of Argiope, where he records the sta- 
bilimenta of immatures (Levi, 1968: 346). 
The symmetrical webs illustrated were 
drawn by Kingston with a ruler and may 
not depict species-specific characters. But 
some of the webs give information that 
(along with the description of coloration 



of the specimens) provides clues for 
matching Kingston's names with speci- 
mens more recently collected in the 
Guianas. A list of names and synonymies 
are given here: 

Epeira morahallii (pp. 171, 172, 363) is Metazygia 
dubia (Keyserling) (Levi, 1995). 

E. folisecens (pp. 167-170, 364) is Hingstepeira fol- 
isecens (see Description section of this species). 

£. sacculifaciens (pp. 153-155, 364) may be imm. 
Micrepeira tubulofaciens (see Description section 
of this species). 

E. lodiculafaciens (pp. 157-159, 365) is a Cyrtophora 
sp. 

E. davisi (pp. 146-149, 365) is Spilasma duodecim- 
guttata (Keyserhng) (see Description section of this 
species). 

E. tiibtdofaciens (pp. 150-153, 366) is Micrepeira 
tubulofaciens (see Description section of this spe- 
cies). 

E. essequibensis (pp. 182, 183, 366) is the male of 
Eustala sp. 

E. nidificans (pp. 173, 174, 367) is probably an im- 
mature Eriophora fuliginea (C. L. Koch). 

E. foliplicans (pp. 175, 176, 367) might be an Erio- 
phora. 

Turckheimia morahallii (pp. 171-175, 200, 368) is 
Parawixia kochi (Taczanowski). NEW SYNONY- 
MY. 

T. tuberculata (pp. 178, 369) is Parawixia kochi (Tac- 
zanowski). NEW SYNONYMY. 

Argiope filiargentata, A. cuyunii, and A. filiinfracta 
are A. argentata (Levi, 1968). 

Epeira nidificans is 8 mm long, the ab- 
domen a little longer than wide, with three 
pale yellow bands, the ventral area with a 
conspicuous quadrate black area. The ven- 
tral dark area suggests that the species is 
an Eriophora, probably E. fuliginea. 

Epeira foliplicans is a relatively large 
species, 12 mm total length. However, un- 
like species of Araneus and Eriophora it 
has a median light band on the underside 
of the abdomen, and because it has a re- 
treat it is probably not a Eustala. 

The species of the genus Turckheimia 
are misplaced, as this name is a synonym 
of Cyclosa. The description of the tuber- 
cles on the abdomen places the two species 
in Parawixia kochi, a common species in 
Guyana. 

I have not attempted to place Kingston's 
six species described in Cyclosa, because 
Neotropical species of that genus still have 
to be revised. 



210 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



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AcTiNOSOMA, Spilasma, MiCREPEiRA, Pronovs • Levi 211 



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212 Bulletin Museum of Comparative Zoology, Vol. 154, No. 3 



INDEX 

Valid names are printed in italics. Page numbers refer to main references, starred page numbers to illustrations. 



Actinosoma, 156 
africana, Spilasma, 186 
akeholmi, Araneus, 204 
albomaculata, Micrepeira, 199*, 200 
amazonica, Madrepeira, 201, 203* 
argentata, Argiope, 209 
arnolisei, Hingstepeira, 165, 167* 
artifex, Spilasma, 187 

baptistai, Spilasma, 189*, 190 
beatus, Paphlagon, 172 
beatus, Pronous, 172, 173* 
beatus, Pronous, 178 
bristowei, Larinia, 164 

carrolli, Tatepeira, 205*, 208 
chelifer, Pronous, 169 
coccineum, Spilasma, 187 
colon, Pronous, 177, 179* 
cuyunii, Argiope, 209 
Cyclosa, 209 
Cyrtophora, 209 

davisi, Cyrtophora, 188 
davisi, Epeira, 187, 209 
dimona, Hingstepeira, 166, 167* 
dubia, Metazygia, 209 
duodecimguttata, Singa, 187 
duodecimguttata, Spilasma, 187, 189* 
duodecimguttatus, Araneus, 187 

Eriophora, 209 
essequibensis, Epeira, 209 
Eustala, 209 

felipe, Pronous, 173*, 176 
filiargentata, Argiope, 209 
filiinfracta, Argiope, 209 
foliplicans, Epeira, 209 
folisecens, Aranea, 164 
folisecens, Araneus, 164 
folisecens, Epeira, 164, 209 
folisecens, Hingstepeira, 164, 167* 
fowleri, Micrepeira, 194, 195* 
fuliginea, Eriophora, 209 

golfito, Pronous, 177, 179* 

heteracantha, Actinosoma, 157 
heteracantha, Wagneriana, 158 
Hingstepeira, 161 
hoeferi, Micrepeira, 195*, 196 
holmi, Araneus, 204 

intus, Pronous, 179*, 180 
isherton, Hingstepeira, 165, 167* 
itu, Tatepeira, 205*, 207 



kochi, Parawixia, 209 

laevisternis, Pronous, 169 
lamentaria, Aranea, 187 
lamentaria, Epeira, 187 
lamentarius, Araneus, 187 
lancetilla, Pronous, 176, 179* 
lodiculafaciens, Epeira, 209 

Madrepeira, 200 
Micrepeira, 191 
minutus, Pronous, 169 
moraballii, Epeira, 209 
moraballii, Turckheimia, 209 

nidificans, Epeira, 209 
nigripes, Pronous, 183*, 184 

pachitea, Micrepeira, 198, 199* 

pance, Pronous, 182, 183* 

Paphlagon, 168 

peje, Pronous, 177, 179* 

pentacanthum, Acrosoma, 158 

pentacanthum, Actinosoma, 158, 159* 

pentacanthus, Araneus, 158 

Plectana, 158 

Pronous, 168 

pulcherrima, Acrosoma, 158 

pygmaea, Hypsosinga, 169 

quinquespinosa, Cyrtarachne, 158 
quintana, Pronous, 173*, 174 

riscoi, Actinosoma, 158 
rubronigra, Rubrepeira, 158 

sacculifaciens, Epeira, 209 
schlingeri, Spinepeira, 159*, 161 
seriata, Prasonica, 186 
shanus, Pronous, 181, 183* 
smithae, Micrepeira, 198, 199* 
Spilasma, 185 
Spinepeira, 160 
spinipes, Gea, 169 
stadelmani, Tatepeira, 205*, 208 
stelligerum, Acrosoma, 158 

taprobanicus, Pronous, 169 
tatarendensis, Aranea, 204 
tatarendensis, Araneus, 204 
tatarendensis, Tatepeira, 204, 205* 
tatarendensis, Wixia, 204 
Tatepeira, 204 
tetraspinulus, Pronous, 169 
tredecimguttata, Spilasma, 187 
tuberculata, Turckheimia, 209 
tuberculatus, Pronous, 174 



AcTiNOSOMA, Spilasma, MiCREFEiRA, Pronous • Levi 213 



tiiberculifer, Pronous, 183*, 184 
tubiilifaciens, Aranea, 197 
tubulifaciens, Araneus 197 
tubulofaciens, Epeira, 197, 209 
tubulofaciens, Epeirella, 197 
tubulofaciens. Micrepeira, 197, 199" 
tubulofaciens, Spilasma, 196 

utaca, Spilasma, 189*, 190 



valle, Pronous, 182, 183* 
velso, Micrepeira, 198, 199* 
visite, Aculepeira, 204 

wixoides, Pronous, 178, 179" 
wixoides, Zigana, 178 

Zigana, 168 



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© The President and Fellows of Harvard College 1995. 



PELEGRINA FRANGANILLO AND OTHER JUMPING SPIDERS 
FORMERLY PLACED IN THE GENUS METAPHIDIPPUS 
(ARANEAE: SALTICIDAE) 



WAYNE P. MADDISON' 



CONTENTS 

Abstract 216 

Introduction 217 

Acknowledgments 218 

Materials and Methods 219 

Explanation of Morphological and Behavioral 

Terms 222 

The Subfamily Dendryphantinae 226 

Relationships within the Dendryphantinae 231 

The Bagheera Group of Genera 232 

Two Genera That Have Exchanged Species 
with Metaphidippus: Dendryphantes 

and Beata 235 

The Proper Placements of Metaphidippus 

Species 237 

Phanias F. P.-Cambridge, 1901 238 

Terralonus new genus 239 

Ghelna new genus 239 

The Genus Pelegrina Franganillo, 1930 240 

Phylogeny within Pelegrina 245 

Identifying Species of Pelegrina and the 

Metaphidippus niannii Group 247 

Key to the Males of All Species of Pelegrina 
and Those Metaphidippus mannii 
Group Species Occurring in the United 

States 248 

Key to the Female Pelegrina of the Eastern 
United States and Canada (East of the 

Mississippi River and Manitoba) 257 

Key to the Female Pelegrina of the Great 
Plains (between the Rocky Mountains 

and the Mississippi River) 258 

Key to the Female Pelegrina of the Pacific 
Coast of the United States and Western 

Canada 258 

Key to the Female Pelegrina and mannii 

Group of Arizona 259 

Key to the Pelegrina and Nagaina females 

of Mexico and Central America 260 

Descriptions of the Species of Pelegrina 262 

1. Pelegrina galathea (Walckenaer, 1837) — 263 



' Department of Ecology and Evolutionary Biol- 
ogy, University of Arizona, Tucson, Arizona 85721. 



2. Pelegrina proxima (G. & E. Peckham, 
1901) 265 

3. Pelegrina dithalea new species 268 

4. Pelegrina edrilana new species 269 

5. Pelegrina proterva (Walckenaer, 1837) __ 270 

6. Pelegrina peckhamorum (Kaston, 

1973) 272 

7. Pelegrina neoleonis new species 273 

8. Pelegrina tristis new species 274 

9. Pelegrina sabinema new species 275 

10. Pelegrina pervaga (G. & E. Packham, 
1909) 276 

11. Pelegrina kastoni new species 277 

12. Pelegrina flavipedes (G. & E. Peckham, 
1888) 278 

13. Pelegrina flaviceps (Kaston, 1973) 279 

14. Pelegrina exigua (Banks, 1892) 281 

15. Pelegrina montana (Emerton, 1891) 283 

16. Pelegrina insignis (Banks, 1892) 284 

17. Pelegrina chaimona new species 286 

18. Pelegrina clemata (Levi & Levi, 1951) .. 287 

19. Pelegrina aeneola (Curtis, 1892) 288 

20. Pelegrina halia new species 290 

21. Pelegrina chalceola new species 291 

22. Pelegrina jurcata (F. P.-Cambridge, 

1901) 292 

23. Pelegrina volcana new species 294 

24. Pelegrina bicuspidata (F. P.-Cam- 
bridge, 1901) 295 

25. Pelegrina ochracea (F. P.-Cambridge, 

1 90 1 ) 295 

26. Pelegrina morelos new species 296 

27. Pelegrina huachuca new species 296 

28. Pelegrina arizonensis (G. & E. Peck- 
ham, 1901) 297 

29. Pelegrina helenae (Banks, 1921) 298 

30. Pelegrina verecunda (Chamberlin & 
Gertsch, 1930) 299 

31. Pelegrina clavator new species 300 

32. Pelegrina pallidata (F. P.-Cambridge, 
1901) 301 

33. Pelegrina variegata (F. P.-Cambridge, 

1 901 ) 302 

34. Pelegrina yucatecana new species 303 



Bull. Mus. Comp. Zool., 154(4): 215-368, January, 1996 215 



216 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



35. Pelegrina sandaracina new species 304 

36. Pelegrina tillandsiae (Kaston, 1973) 305 

37. Pelegrina bunites new species 306 

38. Pelegrina orestes new species 307 

The Genus Nagaina G. & E. Peckham, 1896 ... 308 

39. Nagaina incunda G. & E. Peckham, 

1896 308 

Species of the mannii Group of the United 

States and Canada 310 

40. Metaphidippus mannii (G. & E. Peck- 
ham, 1888) 310 

41. Metaphidippus diplacis (ChamberHn, 

1 924 ) 3 1 2 

42. Metaphidippus tricolor ChamberUn & 

Ivie, 1941 314 

43. Metaphidippus chera (ChamberHn, 

1 924 ) 3 1 5 

44. Metaphidippus carmenensis (Chamber- 
Hn, 1924) 316 

45. Metaphidippus emmiltus new species .... 317 

Literature Cited 318 

Index 322 



Abstract. The genus Pelegrina FranganiUo con- 
tains 38 species of dendryphantine jumping spiders 
from North and Central America that were formerly 
placed in the genus Metaphidippus F. O. Pickard- 
Cambridge. The close relatives of the Dendryphan- 
tinae may include the Europhryinae and several 
smaller groups, for they share an embolus that is 
coiled counterclockwise (left palp) and separated from 
the tegulum by a fully expandable hematodocha. The 
subfamily Dendryphantinae itself is delimited by the 
derived conditions of a carina on the underside of 
the male chelicera, the coil of the embolus folded 
back so as to be hidden behind the base of the em- 
bolus, and S-shaped epig\nal openings. 

Within the subfamily, generic relationships are 
poorly understood, but it is clear that the genus Me- 
taphidippus is polyphyletic. The genus should in- 
clude at most a few species closely related to the 
neotropical genera Messua G. & E. Peckham, Bag- 
heera G. & E. Peckham, and Gastromicans Mello- 
Leitao. Gastromicans is removed from synonymy with 
Beata G. & E. Peckham. The following new com- 
binations are established for species in this group: 
Bagheera prosper (G. & E. Peckham), Messua cen- 
tralis (G. & E. Peckham), Messua dentiger (F. P.- 
Cambridge), Messua donalda (Kraus), Messua lata 
(Chickering), Messua laxa (Chickering), Messua lim- 
bata (Banks), Messua moma (F. P. -Cambridge), Mes- 
sua octonotata (F. P. -Cambridge), Messua pura 
(Bryant), Messua tridentata (F. P. -Cambridge), Gas- 
tromicans albopilosa (G. & E. Peckham), Gastromi- 
cans hondurensis (G. & E. Peckham), Gastromicans 
levispina (F. P. -Cambridge), Gastromicans noxiosa 
(Simon), and Gastromicans vigens (G. & E. Peck- 
ham). The combination Messua desidiosa G. & E. 
Peckham is revived. The proper placement of various 
groups currently assigned to Metaphidippus is dis- 



cussed, and the harfordii group is transferred to the 
genus Phanias F. P. -Cambridge, which appears to be 
relatively distantly related to most other dendry- 
phantines. The following new combinations are es- 
tablished: Phanias albeolus (ChamberHn & Ivie), 
Phanias concoloratus (Chamberlin & Gertsch), 
Phanias dominatus (Chamberlin & Ivie), Phanias 
furcifer (Gertsch), Phanias furcillatus (F. P. -Cam- 
bridge), Phanias harfordii (G. & E. Peckham), Phan- 
ias monticola (Banks), Phanias neomexicanus (Banks), 
and Phanias ivatonus (Chamberlin & Ivie). Also re- 
moved from Metaphidippus are the mylothrus and 
castaneus groups, for which the new genera Terra- 
lonus and Ghelna are described, thus yielding the 
new combinations Terralonus californicus (G. & E. 
Peckham), Terralonus mylothrus (Chamberlin), Ter- 
ralonus unicus (Chamberlin & Gertsch), Terralonus 
shaferi (Gertsch & Riechert), Terralonus versicolor 
(G. & E. Peckham), Terralonus vittatus (Banks), Ter- 
ralonus fraternus (Banks), Ghelna castanea (Hentz), 
Ghelna barrotvsi (Kaston), Ghelna sexmaculata 
(Banks), and Ghelna canadensis (Banks). The new 
combination Sassacus paiutus (Gertsch) is estab- 
lished. The vitis group is retained in Metaphidippus. 
The limits of the genera Dendryphantes C. L. Koch 
and Beata G. & E. Peckham are also reconsidered. 
The combination Dendryphantes nigromaculatus 
Keyserling is revived and the following combinations 
established: Beata hispida (G. & E. Peckham), Beata 
inconcinna (G. & E. Peckham), Beata maccunii (G. 
& E. Peckham), and Beata rustica (G. & E. Peck- 
ham). Dryphias (G. & E. Peckham) is synonymized 
with Beata. 

The largest group removed from Metaphidippus 
is placed in the genus Pelegrina FranganiUo, 1930, 
whose species are, with some exceptions, distin- 
guished from other dendr\ phantines b\ the presence 
of two terminal rami retrolateral to the embolus open- 
ing, an embolic hematodocha that bulges distally, 
wrinkles on the anterior margin of the male cheliceral 
fang, a distinct band of pale scales on the side of the 
face, and male courtship with the first legs held low 
and forward. The following species are moved into 
Pelegrina: P. aeneola (Curtis), P. arizonensis (G. & 
E. Peckham), P. bicuspidata (F. P. -Cambridge), P. 
clemata (Levi & Levi), P. exigua (Banks), P. flaviceps 
(Kaston), P.flavipedes (G. & E. Peckham), P.furcata 
(F. P. -Cambridge), P. galathea (Walckenaer), P. he- 
lenae (Banks), P. insignis (Banks), P. montana 
(Emerton), P. ochracea (F. P. -Cambridge), P. palli- 
data (F. P. -Cambridge), P. peckhamorum (Kaston), 
P. pervaga (G. & E. Peckham), P. proterva (Wal- 
ckenaer), P. proxima (G. & E. Peckham), P. tilland- 
siae (Kaston), P. variegata (F. P. -Cambridge), and P. 
verecunda (Chamberlin & Gertsch). Pelegrina prox- 
ima is shown to be a senior synonym of Pelegrina 
geniciilata FranganiUo, the latter being the types spe- 
cies of Pelegrina. A neotype is designated for Attus 
galathea Walckenaer. Seventeen species are de- 
scribed as new: P. balia, P. bunites, P. chaimona, P. 
chalceola, P. clavator, P. dithalea, P. edrilana, P. 



Pelegrina Jumping Spidkrs • Maddison 217 



huaclntca. P. kastoni, P. morelos, P. neoleonis, P. 
orestes. P. sabinema, P. sandaracina, P. tristis, P. 
volcano, and P. yucatecana. Eitoplirys Icucophaea C 
L. Koch, Icius crassivcnfer Ke>serling, am) Meta- 
phidippus digitatus F. P. -Cambridge are newly syn- 
onymized with P. galathea; Dendrijphantes uteanus 
Chamberhn & Gertsch with P. aeneola; and Dendry- 
phantes mimus Chamberhn with P. furcata. Eu- 
ophrys concolor Banks is removed from synonymy 
with P. proterva and considered a senior synonym of 
Sittaciis cursor Barrows, yielding the new combina- 
tion Sitticus concolor. Identification keys are pre- 
sented for all Pelegrina males and for females from 
restricted geographical regions. All species are de- 
scribed and illustrated. Male/female associations were 
achieved for all species north of Mexico. Courtship 
behavior is described for 22 species of Pelegrina, 
karyotypes for 10 species, and habitat information 
for most species. 

The genus Pelegrina may be closely related to the 
Meiaphidippus mannii group, Nagairia and/or Eris. 
Nagaina incunda G. & E. Peckham is described and 
illustrated; Dendryphantes vegettts G. & E. Peck- 
ham, Meiaphidippus flavolineatus F. P. -Cambridge, 
and Meiaphidippus expallidatus F. P.-Cambridge are 
synonymized with N. incunda. The species of the 
mannii group (temporarily retained in Meiaphidip- 
pus) that occur in the United States are also described 
and illustrated; two new combinations, Meiaphidip- 
pus chera (Chamberlin) and Meiaphidippus car- 
meriensis (Chamberlin), are established; one species, 
Meiaphidippus emmilius, is described as new; Den- 
dryphanies versicolor G. & E. Peckham is synony- 
mized with Meiaphidippus mannii (G. & E. Peck- 
ham), and Meiaphidippus franciscanus Schenkel with 
Meiaphidippus diplacis (Chamberlin). 



INTRODUCTION 

For about 50 years after the Peckham's 
(1909) revision of the jumping spider spe- 
cies north of Mexico, taxonomic work on 
North American representatives of this 
large family consisted mostly of scattered 
species descriptions by Chamberlin, 
Gertsch, Ivie, and others. Some generic re- 
visions consolidating and clarifying the 
previous work began appearing in the 
1950s (Gertsch and Ivie, 1955; Barnes, 
1955, 1958), but most genera remained 
untouched, including the three largest 
genera, Habronattus* Phidippiis, and 
Meiaphidippus, which together include 



Authors of scientific names are given in the index. 



about half of the nearly 300 species of sal- 
ticids occurring north of Mexico (accord- 
ing to the count of Richman and Cutler, 
1978). In the last three decades, increased 
interest in the family has resulted in re- 
visions of Habronattus (Griswold, 1987), 
Phidippus (Edwards, in preparation), and 
other genera (Proszynski, 1968, 1971a, 
1973a, 1980; Cutler, 1981a, 1987; Rich- 
man, 1981, 1989). However, except for 
works by Kaston (1973) on some eastern 
species and by Cutler and Jennings (1985) 
on the arizonensis group, Metaphidippus 
has remained unre vised, perhaps because 
its poorly defined limits have made the 
scope of any revision potentially trouble- 
some. When 1 first began to revise Meta- 
phidippus, I knew that I would have to 
restrict the revision to only some of the 
disparate groups placed there. The largest 
group placed in Metaphidippus, including 
the species most commonly collected in 
northern and eastern North America, was 
chosen for revision and is here moved to 
the genus Pelegrina Franganillo. 

The jumping spiders placed in Pelegri- 
na are medium-sized dendryphantines 
distributed throughout North America, 
with some species extending as far south 
as Panama. The 38 species include the well- 
known P. galathea, P. proterva, P. fla- 
vipedes, and P. aeneola. Males of Pele- 
grina are generally brown with white 
stripes (Fig. 1), and most can be distin- 
guished from other dendryphantines by 
the wide embolus with two rami retrola- 
teral to the opening (Fig. 3). The spotted 
females (Fig. 2) have large thickened flaps 
over the epigynal openings (Fig. 4). Al- 
though the eastern species were well stud- 
ied by Kaston (1973), most of the species 
occur in the western United States, Mex- 
ico, and Central America, and they re- 
ceived their last comprehensive treatments 
by G. & E. Peckham (1909) and F. O. 
Pickard-Cambridge (1901). Many of the 
western species have been inadequately 
described and illustrated, often from only 
one sex, making identification almost im- 
possible by anyone other than an araneol- 



218 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



ogist familiar with the group. Many spe- 
cies in the southwest were undescribed, 
and for most species there is Httle pub- 
Hshed information on natural history. The 
present revision has as its main goal to 
make the species known, by describing and 
illustrating them, their courtship displays, 
and their habitats. Although much prog- 
ress has been made in distinguishing spe- 
cies and matching males to females, many 
problems of geographical variation and 
uncertain male-female matching remain 
for future work, especially among Mexi- 
can Pelegrina. 

In addition to the Pelegrina species, Na- 
gaina incunda and the U.S. species of the 
Metaphidippus mannii group are de- 
scribed because they could very well be 
confused with species of Pelegrina and be- 
cause their taxonomy is in need of revision. 

This work addresses the phylogeny of 
Pelegrina and the subfamily Dendry- 
phantinae, but it has no pretensions of be- 
ing a comprehensive or modern phyloge- 
netic treatment. My phylogenetic goals are 
to propose some characters that might pro- 
vide an outline of dendryphantine rela- 
tionships, focusing on the question of the 
monophyly of Pelegrina and a few other 
groups formerly placed in Metaphidippus. 
I hope that this and the basic exploratory, 
species-level taxonomic work will provide 
the groundwork for future phylogenetic 
treatments. 

ACKNOWLEDGMENTS 

This paper formed part of a Ph.D. thesis 
for the Department of Organismic and 
Evolutionary Biology, Harvard Universi- 
ty. The work described in it has roots many 
years deep, and during these years I re- 
ceived the help of many friends and col- 
leagues. As always, David Maddison has 
been foremost among my collaborators in 
formulating ideas, on collecting expedi- 
tions, on behavioral observations, and in 
many other activities. My thesis advisor, 
Herbert W. Levi, provided a well-orga- 
nized laboratory in which to work, a model 
of dedication to arachnology, and a helpful 



encouragement that allowed me to grow 
on my own. My other friends and col- 
leagues at Harvard and in our spider lab- 
oratory — Leticia Aviles, Anna Weitzman, 
Mark Moffett, Jonathan Coddington, Mark 
Stowe, Jackie Palmer, Cecile Villars, John 
Hunter, Caty Sibble, Ardis Johnston, Laura 
Leibensperger, Dee Woessner, and Ann 
Blum — made a very supportive commu- 
nity in which to work. Leticia Aviles and 
Christopher Maddison gave me much aid 
and inspiration during the most difficult 
stages of the project. 

H. W. Levi, L. Aviles, D. Maddison, G. 
B. Edwards, P. F. Stevens, and E. O. Wil- 
son read and commented on this paper. 
Two anonymous reviewers gave useful 
suggestions. For discussion of jumping spi- 
der issues, I thank B. Cutler, G. B. Ed- 
wards, D. B. Richman, J. Proszynski, F. 
Wanless, and P. Wijesinghe. B. Cutler, G. 
B. Edwards, A. Moldenke, and D. B. Rich- 
man have sent live specimens to me for 
courtship observations. Brent Opell helped 
with trypsin clearing. H. D. Cameron gave 
me helpful advice on the formation of 
names, though he is not to be held re- 
sponsible for any ill formed, especially since 
I did not always follow his no doubt proper 
advice. Curators at various institutions lent 
me specimens; a list can be seen under 
Materials and Methods. I thank these cu- 
rators for their help. I would especially like 
to thank Luis F. de Armas of the Instituto 
de Ecologia y Sistematica, Havana, Cuba, 
for sending the Franganillo collection of 
salticids. 

This work was supported in part by a 
NSERC (Canada) Postgraduate Fellow- 
ship. For allowing me to participate in a 
major collecting trip to Mexico, I thank S. 
B. Peck and R. S. Anderson. The trip was 
funded by NSERC and had assistance from 
the Universidad Nacional Autonoma de 
Mexico and Harvard University. The De- 
partment of Organismic and Evolutionary 
Biology of Harvard supported trips to Cal- 
ifornia, Arizona, and Spain. Publication 
costs of this study were covered in part by 
the Wetmore Colles fund. 



Pelegrina Jumping Spiders • Maddison 219 



MATERIALS AND METHODS 

Collections Examined. The taxonomic 
revision is based on specimens in the fol- 
lowing collections. The abbreviation for 
the collection is followed by the name of 
the collection and the curator and others 
responsible for aiding in loaning the ma- 
terial, to whom many thanks are due: 

AMNH American Museum of Natural 
History, New York (N. Platnick, 
L. Sorkin) 

BMNH The Natural History Museum, 
London (P. Hillyard) 

CAS California Academy of Sci- 

ences, San Francisco (W. Pu- 
lawski, D. Ubick) 

DU Darrel Ubick personal collec- 

tion 

lESC Instituto de Ecologia y Siste- 

matica, Havana (Luis F. de Ar- 
mas) 

MCZ Museum of Comparative Zool- 

ogy, Cambridge (H. Levi) 

MSUW Midwestern State University, 
Wichita Falls, Texas (N. Hor- 
ner) 

TXAM Texas A&M University, College 
Station, Texas (A. Dean) 

UCB University of California, Berke- 

ley (E. Schlinger, C. Griswold) 

UWBM Burke Museum, University of 
Washington, Seattle (R. Craw- 
ford) 

WPM W. Maddison personal collec- 
tion 

ZMB Zoologishes Museum Berlin (M. 

Moritz, S. Fischer) 

Note that Canadian specimens are, in gen- 
eral, underrepresented in this revision be- 
cause two major collections of Canadian 
spiders, the Canadian National Collection 
at the Biosystematics Research Centre, Ot- 
tawa, and the Royal Ontario Museum col- 
lection, were not examined due to time 
limitations. 

Routine Examination and Illustra- 
tions. Specimens were examined in a glass 
dish with a bottom layer of half black, half 



white silicone rubber (bathtub caulking). 
The silicone rubber is superior to paraffin 
for most purposes, for it can hold even 
minuten pins firmly and later heal, and it 
offers the advantage over sand of allowing 
appendages to be pinned open. Palpi were 
mounted on Vaseline in an alcohol-filled 
depression slide with a coverslip and drawn 
at 100 X and 200 x under an Olympus 
BH-2 compound microscope using inci- 
dent fiber-optics illumination and a cam- 
era lucida. Not only did the use of a com- 
pound microscope allow higher resolution, 
but also the axial light path prevented the 
drawing difficulties caused by the side-to- 
side shifting of the image that occurs when 
focusing on a stereo dissecting microscope. 
For the external (ventral) view, epigyna 
were examined using the same technique, 
without clearing. Epigyna were dissected 
off of the specimen to allow for the small 
working distance of the compound micro- 
scope. The Vaseline on which they were 
mounted was made opaque by mixing with 
chalk dust, in order to simulate the cream- 
colored muscles and glands that would un- 
derly the epigynum on an intact specimen. 
After examination of the specimen. Vas- 
eline was removed by a xylene rinse. The 
oblique drawings of the male carapace and 
chelicerae were made mostly under the 
compound microscope, at 40 x . Most 
drawings of the female abdomen were 
made under a Zeiss stereo dissecting mi- 
croscope with a camera lucida. The draw- 
ings of the male face and female abdomen 
show the appearance in alcohol. Most 
drawings were done on coquille board with 
ink, a Conte drawing pencil, and white 
paint. Small labels with my initials (WPM) 
and the year drawn (e.g., 84) were placed 
in vials of specimens illustrated. Photo- 
graphs of living specimens were made with 
a standard 55-mm lens reversed on exten- 
sion tubes to yield approximately 2.5 x 
magnification on Kodak Technical Pan or 
Kodachrome film, using illumination by 
flash. Measurements of carapace length, 
carapace width, and body length (Galiano, 
1963; Wanless, 1978) were made from the 



220 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



dorsal view using an eyepiece reticle on a 
Leitz stereo dissecting microscope. Gen- 
erally only about five specimens of each 
sex were measured, because little reliance 
on these measurements was made in this 
study. They are intended to give only a 
general idea as to the size and proportions 
of the species. The results are presented as 
follows: minimum (median) maximum. 
Thus, if five measurements for carapace 
length for the females of one species are 
2.0, 2.1, 2.2, 2.3, and 2.3, these would be 
reported as follows: 2.0(2.2)2.3, n = 5. 

Descriptions. A species description was 
originally written from a sample of five 
males and five females or more (if avail- 
able). During subsequent identifications, 
the description was periodically checked 
to ensure that it covered the range of vari- 
ation within the species. Two characters 
described that were less thoroughly sam- 
pled are the exact region of contact of the 
forehead band with the anterior median 
eyes, which was examined in only four to 
eight males, and the details of the internal 
epigynal ducts of the female, which in 
many species was examined in only one to 
three females. 

Clearing. Clearing was used for detailed 
examination of the integument, especially 
to observe external and internal structures 
of the genitalia and mouthparts. The var- 
ious body parts were cleared by placing 
them in warm trypsin solution for 1-2 days 
to digest internal tissues. When the tryp- 
sin-clearing procedure is successful, it re- 
veals palp morphology to a level of detail 
not previously published (Figs. 3, 16-27). 
Trypsin was used instead of potassium or 
sodium hydroxide because it damages the 
cuticle very little and has shown no ten- 
dency to expand the palp except perhaps 
when hematodochae are tightly coiled as 
in Ashtabula and Bagheera. If the tissues 
of the specimen are well fixed and firm, 
then digestion will be very slow. Hence, 
for best results the specimen should be 
fresh-killed (not fixed) or "fixed" in a poor 
fixative with a low concentration of alco- 
hol. Fresh-killed, dilute ethanol-triton 



fixed, 80% ethanol fixed, and Kahle's fixed 
specimens were all used. Specimens not 
fresh-killed were first rinsed in water for 
several hours before digestion. The body 
parts were separated to allow penetration 
of the trypsin. The trypsin solution was 
made from about 1 ml purified trypsin 
(Fisher #T-360) in 10 ml water, filtering 
after mixing. The trypsin was warmed un- 
der a light bulb during digestion. After 
digestion, the parts, especially dark, heavi- 
ly sclerotized palps or epigyna, were 
bleached. They were first rinsed in water 
and then 80% ethanol, then moved to a 
solution of 1 part ethanol : 1 part 10% Aero- 
sol for a day, and then for bleaching moved 
into a solution of 1 part 30% hydrogen 
peroxide : 1 part ethanol : 2 parts 10% Aero- 
sol. Aerosol (Fisher) was added to the 
bleaching solution to inhibit the formation 
and buildup of bubbles, which otherwise 
can fill the body part and make it unusable. 
After about 1 day of bleaching, the body 
part was moved to 80% ethanol and then 
transferred to 95-100% ethanol. The body 
part was then mounted temporarily in 
clove oil or permanently in Euparal. For 
most specimens, the transferral to Euparal 
was accomplished by placing the body 
parts in Euparal thinned with either eth- 
anol or Euparal Essence and letting the 
ethanol or essence evaporate off, thus grad- 
ually taking the specimen through a series 
of stronger Euparal solutions. This was most 
easily done directly on the microscope slide, 
which was then let to dry partially in a 
dust-free area. Drying thickened the Eu- 
paral, allowing final positioning of the parts 
before adding the coverslip. After the cov- 
erslip is added, the body parts, especially 
the palpus, may move. Thus, the palp was 
placed near the edge of the coverslip to 
allow repositioning using a microneedle 
slipped under the coverslip. 

Expansion of Palps. Palps were best ex- 
panded permanently by boiling the spec- 
imen alive and then fixing it in dilute 
Kahle's fluid to harden it in an expanded 
position followed by gradual dehydraton 
to 80% alcohol. This technique is much 



Pelecrima JiMi'iNG SPIDERS • MaddisoTi 221 



like tl.-it described by Sadana (1971). Palps 
so prepared are resistant to contraction and 
can be critical-point-dried successfully 
(Figs. 7, 9). A few palpi that were already 
preserved in alcohol were expanded in a 
few minutes b\ placing them in a hot mix- 
ture of 15% hydrogen peroxide, 40% wa- 
ter, and 45% ethanol. 

Chromosomes. Chromosomes were ex- 
amined using the Feulgen technique as 
described by Maddison (1982). The results 
are given under the description of each 
species examined. 

Courtship. Courtship observations were 
obtained for Pelegrina species and nu- 
merous other dendryphantines. Specimens 
were examined within a few days after 
collecting. A male and a female were 
placed on a cotton beating sheet, usually 
not in full sunlight, and manipulated until 
the male faced the female and began dis- 
play. Behavior of the male was observed 
by eye, and notes and still photographs 
were taken. Female behavior was not re- 
corded. For most species, no filming or 
videotaping was done. As salticid behavior 
can be fast, it is difficult to take notes ac- 
curately, and there are likely errors in ob- 
servation, especially of subtle differences 
in timing and positions. Still, consistency 
of observations and videotape confirma- 
tion of my own previously taken notes for 
Pelegrina dithalea, Metaphidippus man- 
nii, Metaphidippus chera, and Phanias 
watonus all indicate that the descriptions 
should be generally accurate. There are a 
number of observations, such as the alter- 
nate palp waving in the Pelegrina fla- 
vipedes group and the triangular crouch 
pose of Pelegrina aeneola, that have been 
repeated a number of times and in which 
I have strong confidence. In the descrip- 
tions of the displays, the sample sizes for 
the observations are indicated by listing 
the number of observations for each fea- 
ture of the display. For instance, in the 
description of the courtship of P. galathea, 
the following sentence occurs: "First legs 
flicker (n = 12, 63) on each series (n = 4, 
23) up and down (n = 4, 23) and alternately 



back and forth at tips (n = 1), vigorously 
(ca. 5 c/s) (n = 1) but at low amplitude (n 
= 5, 33)." The parenthetical comments in- 
dicate the number of observations (n) and 
the number of males in which the feature 
was noted. The number of observations 
was counted as follows: a male was ob- 
served doing a bout of courtship display; 
any features of position or motion were 
considered to have thus been observed once 
during this bout. If the male stopped dis- 
playing because the female left or rejected 
him, and if he began again later (perhaps 
after I had moved them back together), 
then the next display was counted as a 
separate observation. While the more ob- 
vious features of the display may have been 
observed many times (for instance, that the 
legs were flickered was observed 12 times 
in six males in the preceding example), 
some of the more subtle details of the dis- 
play were not noticed in most displays and, 
thus, would have been observed only a few 
times (for instance, that the legs flickered 
"alternately back and forth at tips" was 
observed only once). Where there is vari- 
ation, the description lists each of the al- 
ternatives with an indication of sample 
sizes. For instance, if the legs were usually 
flickered at low amplitude, but occasion- 
ally at high amplitude or not at all, the 
description might read like this: "On each 
series legs flickered (n = 9, 13) noticeably 
(n = 1) or with fairly low amplitude (n = 
7, 33) or perhaps not at all (n = 3, 13)." 
Sample sizes are in general small. It was 
felt that it is presently more important to 
obtain a broad survey, including many 
species, than a deep analysis of a few spe- 
cies. Explanations of terms used to describe 
courtship are given in the section Expla- 
nation of Morphological and Behavioral 
Terms and in the description of behavioral 
characters (item 7) supporting the mono- 
phyly of Pelegrina. 

Phylogenetic Analysis. Though this 
work is primarily concerned with the ge- 
nus Pelegrina, an attempt was made to 
outline the broader structure of the family 
and the relationships of dendryphantines, 



222 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



partly for their own sake and partly to set 
the context for the genus Pelegrina. The 
general discussion of phylogeny within the 
family is presented separately (Maddison, 
1988, unpublished manuscript). My phy- 
logenetic proposals are presented in a nar- 
rative discussion of groups and characters; 
no numerical phylogenetic analysis was 
done. Table 2 shows the distribution of 
some of the more important characters, 
but it will be noted that some of these do 
not perfectly support the groups proposed. 
Problems with some of the characters are 
noted in the phylogenetic discussion. Fur- 
thermore, the presumption of ancestral 
state for a given character is usually not 
accompanied by rigorous outgroup anal- 
ysis (e.g., Maddison et al., 1984). We are 
still making only preliminary sketches of 
the phylogenetic structure of this large and 
poorly known family, and it would not be 
productive to delay phylogenetic hypoth- 
eses until they can be rigorously defended. 
The suggestions made should have at least 
a glimmer of truth and will, I hope, stim- 
ulate future work. 

Species Distinctions. Populations were 
considered distinct species if several con- 
sistent and discrete morphological differ- 
ences could be found among them, but 
when there were few differences, apparent 
intergradation, or little material, the de- 
cision as to whether one or more species 
are present was sometimes difficult. In sev- 
eral cases, only a single species was rec- 
ognized despite geographical variation, 
because the geographical variation was too 
confusing at present (furcata-mimus) , or 
because the differences were slight and 
possibly not consistent {aeneola / uteanus, 
northern and Floridian tillandsiae, north- 
ern and southern carmenensis, mannii/ 
versicolor) . In other cases, allopatric pop- 
ulations that are similar but that differ con- 
sistently in a number of features were rec- 
ognized as distinct (sabinema/pervaga, bi- 
cuspidata /volcana, neoleonis / tristis, 
proxima /galathea/ dithalea,chera/ tricol- 
or / diplacis) , though in each of these cases 
the decision was difficult. Pelegrina fla- 



vipedes, flaviceps, and exigua were main- 
tained distinct despite apparent hybrid- 
ization because they differ in numerous 
features. The two sympatric forms of ex- 
igua were left under one specific name 
until they can be better studied. 

Male /Female Matching. Care had to be 
taken in proposing which males and fe- 
males belonged together in the same spe- 
cies, as in other spiders (Levi, 1985). The 
problem was especially bad in Pelegrina 
species from the southwestern United 
States, Mexico, and Central America, 
where collecting has been limited. Despite 
the strong sexual dimorphism, similarity 
in body form and markings could often be 
used as evidence. Other criteria used were 
expected correlations in genitalia (e.g., 
wide, robust embolus with strong epigynal 
flaps), co-collecting in same geographical 
region, locality, or microhabitat, and sim- 
ilarity of the male and female each to those 
of another well-matched species. Com- 
ments regarding evidence used to match 
males and females are given in those spe- 
cies descriptions where it seems needed. 
Among the less certain matchings are those 
for Pelegrina sandaracina, pallidata, chai- 
mona, huachuca, and morelos. 

EXPLANATION OF MORPHOLOGICAL 
AND BEHAVIORAL TERMS 

Markings in General. Color patterns are 
generated by integument coloring and by 
covering of setae, though pale setae usually 
overlie pale integument and dark overlie 
dark. The terms hair and scale are used to 
refer to a thin, more or less cylindrical seta 
and a broad, flattened seta, respectively 
(cf. Hill, 1979). 

Markings of the Carapace. Male den- 
dryphantines are commonly dark brown 
with bands of white to yellow scales, usu- 
ally including a major longitudinal band 
on either side of the carapace and abdo- 
men. The following names are used to re- 
fer to the bands of pale scales on the car- 
apace (Fig. 1): 

side bands: Longitudinal bands on either 



Pelegrina Jumping Spiders • Maddison 223 



side of carapace, beginning beside the 
anterior lateral eyes (ALEs) and pro- 
ceeding just beneath the small eyes and 
posterior eyes and beyond, onto the tho- 
rax. 

cheek band: Oblique band on the side of 
the face, starting beneath the ALEs and 
proceeding down and posteriorly to the 
carapace margin, in Pelegrina and the 
mannii group. 

forehead band: A V-shaped marking on 
the dorsal cephalic area just behind the 
anterior median eyes (AMEs). 

marginal band: On the lower margin of 
the sides of the carapace, often extended 
from the cheek band in Pelegrina. Fe- 
males of most species show none of these 
bands distinctly; the carapace is instead 
often covered more or less uniformly 
with pale scales including a dense white 
covering on the clypeus. 

Setae Surrounding Anterior Median 
Eyes. These are of various colors, from 
white to black. In the descriptions of Pe- 
legrina males, the colors of setae around 
the circumference of the left anterior me- 
dian eye (AME) are indicated using a no- 
tation derived from hours on a clock's face. 
Usually, the colors on only the dorsal part 
of the AMEs are described, for those ven- 
trally are more variable. Thus, "white 
forehead band contacts the AME dorsally 
10:30-12:30" means that as one looks from 
the front at the left AME there are white 
setae from 10:30 o'clock to 12:30 o'clock 
that are continuous with the forehead band, 
and dark setae on either side of this. 

Markings of the Legs. The legs are yel- 
low to dark brown, but there are often 
annulate markings (Fig. 1), especially in 
males, so that each leg has pale portions 
covered with white scales alternating with 
darker portions lacking white. 

Markings of the Abdomen. The abdo- 
men is usually brown above in males, 
ringed by white side bands (Fig. 1). The 
dorsum of the abdomen often shows traces 
of the paired pale spots seen in females 
(Fig. 2). These pale spots are central and 



paired: the first pair just anterior to the 
muscle attachment of the second dorso- 
ventral muscle (number 86viii -(- ix of 
Whitehead and Rempel, 1959), the second 
pair anterior to the attachments of the third 
dorsoventral muscle, and the third, fourth, 
fifth, and sixth pairs of spots behind this. 
The fifth and sixth pairs are very small. 
Often the fourth through sixth pairs are 
each connected medially and thus form 
small chevrons. Between these pale spots 
may be spots of dark brown, sometimes 
very dark (e.g.. Fig. 353). 

Male Palpus (Figs. 3, 6-9). The descrip- 
tions generally assume that the left palp is 
being viewed from the ventral. The ad- 
jectives basal and apical when unqualified 
refer to the appearance in a contracted 
palp (i.e., basal = toward the tibia and 
apical = toward the tip of the cymbium). 
In contrast, "anatomically basal" and "api- 
cal" refer to the cymbium-embolus axis 
of connections of the palp's bulb (ie., an- 
atomically basal = toward the basal he- 
matodocha and anatomically distal = to- 
ward the tip of the embolus). 

In the subfamily Dendryphantinae, the 
shoe-shaped cymbium holds a bulb con- 
sisting of (from anatomically basal to api- 
cal) a fully expandable basal hematadocha, 
a small subtegulum, a much reduced me- 
dian hematodocha, the large tegulum, a 
fully expandable distal (embolic) hema- 
todocha, and the embolus. The basal he- 
matodocha expands so as to give a clock- 
wise rotation to the subtegulum and te- 
gulum in ventral view of the left palp, as 
in other salticids. This has been observed 
by artificial expansion on many dendry- 
phantines and during copulation of Den- 
dryphantes nigromaculatus. The subte- 
gulum is small and contains little more 
than the basal portion of the sperm duct 
reservoir (Figs. 3, 8). 

From the subtegulum, the sperm duct 
loops up, down, and around the tegulum 
(Fig. 3) in a clockwise direction to the em- 
bolus. The distal retrolateral portion of the 
tegulum, where the sperm duct enters the 
tegulum from the subtegulum, is generally 



224 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



extended distally and contains a loop of 
the sperm duct pressed against its wall. 
This portion I call the shoulder of the te- 
gulum (Fig. 3). Just proximal to the em- 
bolus and shoulder is a fold extending across 
the surface of the tegulum, the tegular 
ledge (Figs. 3, 6, 7), which serves as a pock- 
et to hold the proximal part of the embolic 
base. Wanless (1984) suggested that the 
tegular ledge (his M3) is absent from sal- 
ticids other than spartaeines, in which he 
described it. However, the fold that I am 
here calling the tegular ledge may be ho- 
mologous with his M3, for it appears as an 
extension of the embolic hematodocha cut- 
ting across the face of the tegulum and 
occurs in many groups of salticids. The 
tegulum is filled with tegular glands, which 
empty into the sperm duct via a series of 
pores in the sperm duct (see Osterloh, 1922; 
Bhatnagar and Rempel, 1962). As in other 
spiders, these pores are aligned into a band 
(Schult, 1980), which, when narrow, ap- 
pears as if it were a seam along the length 
of the sperm duct. Along the narrower part 
of the sperm duct toward the embolus, the 
glands are connected to the sperm duct via 
long ducts (Fig. 3; also known from other 
spiders; Osterloh, 1922: figs. 20, 26). These 
pores, ducts, and glands have been largely 
ignored in systematics but appear to be a 
potential source of good systematic char- 
acters (for instance, in some salticids the 
pores are arranged in a broad ribbon, 
whereas in others the band is very narrow, 
and in Sitticus the pores are arranged into 
prominent craters). 

The embolic hematodocha arises on the 
back side of the tegulum (Fig. 8), prolater- 
al to the subtegulum. Its wrinkles sweep 
apically up toward the embolus. The exact 
arrangement of the folds of the hemato- 
docha is probably of systematic value but 
is very difficult to untangle, for especially 
near the base of the embolus the folds are 
twisted and confusing even in a well- 
cleared palpus. During expansion, the em- 
bolic hematodocha expands fully to move 
the embolus counterclockwise (i.e., pro- 
laterally) and back (i.e., toward the cym- 



bium), as indicated by artificial expansions 
(Figs. 7, 9). The counterclockwise move- 
ment of the embolus itself probably ex- 
plains why the counterclockwise-coiled 
embolus can still engage the epigynal 
opening despite the clockwise movement 
of the tegulum (in salticids with the em- 
bolus fixed to the tegulum, the embolus 
has a clockwise curve, which thus coin- 
cides with the clockwise thrust of the te- 
gulum). The embolus of dendryphantines 
usually consists of a basal portion, which 
is transversely directed, and an apical por- 
tion, which is usually thin and erect and 
has the opening of the sperm duct at its 
tip (Figs. 40-47; also Figs. 64b, d). The 
embolic base consists of a more or less 
sclerotized portion of the embolic hema- 
todocha that is exposed to the ventral sur- 
face and that rests betwee the tegular ledge 
and the embolus (Figs. 3, 6, 7). Its wrinkles 
suggest that it should be considered part 
of the hematodocha. As noted, the embolus 
is coiled counterclockwise, a feature much 
more readily apparent in the euophryines 
and other subfamilies than it is in the den- 
dryphantines. In dendryphantines, the spi- 
ral is hidden and best seen in expansions 
or dissections (e.g., Fig. 35). The dendry- 
phantine embolus arises prolaterally and 
moves across toward the retrolateral side 
(the transverse basal portion of the em- 
bolus) and then folds back toward the pro- 
lateral and abruptly rises as the erect apical 
portion (Figs. 20-23). In many genera, the 
erect apical portion is fused against the 
transverse portion so that there is no open, 
freely coiling spiral. A suture on the back 
side of the embolus (the embolic suture), 
between the transverse portion and the 
erect portion, is often present and indicates 
where the folded-back spiral has not com- 
pletely fused (Figs. 3, 8, 20-23, 31-35). In 
cleared palpi, a slight bend in the sperm 
duct can be seen at this point. The coiling 
of the embolus and the embolic suture sug- 
gesting it are clearly visible in Dendry- 
phantes, Eris, Pelegrina, Phidippus, Tu- 
telina, Phanias, and many other genera. 
In Metaphidippus chera, the coiling is eas- 



Pklegrina Jumping Spidehs • Maddison 225 



ily seen (Fig. 35); in Pelegrina proterva, 
the only trace is the small suture (Fig. 34); 
in Pelegrina flavipedes, the suture is some- 
times visible and sometimes not. No trace 
of the suture or past coiling can be found 
in Terralonus cf. uniciis and Poultonella 
alboimmaculata, but Terralonus myloth- 
nis and Tiitelina elegans, which are, re- 
spectively, their close relatives, show them 
well. The loss of a trace of coiling may 
have evolved several times in the dendry- 
phantines. In some genera such as Zygo- 
hallus, Hentzia and Mabellina, the coil is 
not so compact and, instead, is more open 
(e.g.. Figs. 24-27, 37, 38, 50, 51, 58-63, 
64c, f). Figure 64 summarizes some of the 
coiling patterns seen in dendryphantines. 
The erect portion of the embolus is some- 
times a simple spike with the opening ter- 
minal, but in many dendryphantines there 
are prolongations that will be referred to 
as rami, and often small denticles occur 
on the surface of the embolus. In Pelegri- 
na, in particular, there are two rami re- 
trolateral to the opening of the sperm duct, 
the prolateral ramus very near the opening 
and the retrolateral ramus some distance 
away (Fig. 3). 

Epigynum. In most dendryphantines, 
the openings are well separated and 
S-shaped (Figs. 65-70), with entry toward 
the lateral in the anterior half and toward 
the medial in the posterior half (Fig. 5). 
The lateral rim of the opening is often 
thickened to yield a more or less convex 
teardrop-shaped area, which will be called 
the epigynal flap (Fig. 4). The inner mar- 
gins of the left and right flaps may be 
parallel to each other (e.g.. Figs. 262, 274, 
322) or be divergent from anterior to pos- 
terior (e.g., Figs. 346, 352, 398) or be con- 
vergent (e.g., Figs. 280, 297, 332, 363). In 
some Pelegrina, the flaps converge to such 
an extent that their posterior halves are 
rotated 90° and become transverse (e.g., P. 
kastoni, Fig. 317). Even more extreme ro- 
tations are seen in P. arizonensis (180°, 
Fig. 424) and P. helenae (270°, Fig. 430). 
The surface of the epigynum between and 
behind the openings varies in topography 



among the species of Pelegrina (Figs. 236- 
255). In some, the entire surface behind 
the openings is raised into a mound (e.g., 
P. clemata. Fig. 246); in others, it is much 
more concave (e.g., P. ftircata. Figs. 249, 
250). At the posterior margin of the epi- 
gynum is the notch, or guide (Figs. 4, 5), 
into which fits the male tibial apophysis, 
although in some dendryphantine groups 
(Poultonella, Hentzia) the guide has 
moved anteriorly as in pellenines (Pel- 
lenes, Hahronattus) and Bianor. 

The different parts of the copulatory 
ducts of Pelegrina species are named as 
follows (Fig. 5). From the anterior half of 
the openings, the ducts proceed first lat- 
erally and posteriorly (the first curve), then 
medially (the second curve), and then pos- 
teriorly (the third curve), and then twist 
a number of times and proceed dorsally to 
the fertilization ducts. The inner surface 
of the duct is relatively smooth in the first 
and second curves, smooth or rough in the 
third curve depending on the species, and 
rough with projections in the twisted area 
posteriorly. The flower-shaped openings 
of the accessory glands occur on the second 
curve, usually near the junction with the 
first curve (Fig. 5). The pathway from the 
copulatory opening through the copula- 
tory ducts toward the fetilization ducts is 
almost straight, lacking the separate sper- 
mathecal reservoir seen in, for instance, 
some euophryines. 

Courtship. Some introduction to the for- 
mat of descriptions of courtship has al- 
ready been given under the Materials and 
Methods section. Males, especially early in 
display, often walk not straight toward the 
female but instead sidle so that they ap- 
proach obliquely or not at all. The sidling 
may be first to the left and then to the 
right, and so on, to form a zigzag dance 
(Jackson, 1978). The male usually walks 
sporadically, taking a series of steps then 
pausing, another series of steps then paus- 
ing, and so on. Because the male's pose 
and motions of the first legs, palp, and 
abdomen often differ depending on 
whether he is walking or paused, a dis- 



226 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



tinction is made between the walking 
phase, which is called the series (of steps), 
and the pauses. While the male is walking, 
the first legs are generally raised, spread, 
and/or extended forward. The first legs 
may be waved or flickered up and down 
or forward and backward, one or repeat- 
edly. The distinction between a wave and 
a flicker is not precise; in general, "wave" 
is used when the motion is of low speed 
or frequency, "flicker" for high-frequency 
repeated motions. The abdomen may be 
twitched (Jackson, 1978) down then up, 
which in many species produces a sound 
that may function in courtship (Maddison 
and Stratton, 1988). Explanations of 
"crouch" and "raisedspread" stages of 
courtship are given in the discussion of 
monophyly (item 7) in the description of 
the genus Pelegrina. 

THE SUBFAMILY DENDRYPHANTINAE 

Pelegrina and Metaphidippus are den- 
dryphantines, which are salticids. Maddi- 
son (1988) reviewed the phylogeny of the 
family Salticidae. The majority of salticid 
species are considered to form a mono- 
phyletic group, called the Salticine divi- 
sion (Maddison, 1988), which are distin- 
guished from the remaining groups (Lys- 
somaninae, Spartaeinae, and the Cocal- 
odes group) by eye structure (Wanless, 
1984; Eakin and Brandenburger, 1971; 
Blest, 1983; Blest and Sigmund, 1984), ab- 
sence of a tarsal claw on the female palp, 
medial displacement of the gnathocoxal 
glands (Figs. 14, 15), asymmetrical tarsal 
claws, a mound of slit sense organs with 
an associated seta on the medial edge of 
the chelicera (Figs. 12, 13), and a small 
intercheliceral sclerite (Figs. 12, 13), each 
of which may be considered derived with- 
in the family. Within the Salticine divi- 
sion, there are some prominent subfamilies 
that have the embolus fixed immovably to 
the tegulum, including the Heliophaninae 
(delimited by an apparent stridulatory ap- 
paratus and a bump on the tegulum just 
clockwise [left palp] of the embolus; Mad- 
dison, 1987), the Plexippinae (including 



Plexippus, Hyllus, Evarcha, Thyene, Te- 
lamonia, Harmochirus, and part of Bia- 
nor, delimited by a modified serrula on 
the male endite and a bump on the te- 
gulum just counterclockwise [left palp] of 
the embolus), and many other familiar 
groups such as Pellenes, Salticus, Sitticus, 
Phiale, Myrmarachne, Amycus, and their 
respective relatives. However, the Eu- 
ophryinae, Dendryphantinae, and several 
smaller groups are united by an embolus 
that is free to move, being separated from 
the tegulum by a fully expandable he- 
matodocha (Figs. 29-31). In these groups, 
the embolus is also coiled counterclockwise 
(left palp). The subfamily Dendryphantin- 
ae itself is delimited by the derived con- 
ditions of a carina on the underside of the 
male chelicera, by the coil of the embolus 
folded back so as to be hidden behind the 
base of the embolus, and by S-shaped epi- 
gynal openings. 

The Dendryphantinae is a subfamily of 
several hundred species, most of these in 
the New World. Males are usually striped, 
with longitudinal bands of pale scales on 
either side of the carapace and abdomen, 
while females usually have paired spots on 
the abdomen. The chelicerae of males are 
often enlarged or elongate as is the first 
pair of legs. The posterior eyes are smaller 
than in most euophryines. 

Table 1 lists the generic names (includ- 
ing those now considered synonyms) that 
I refer at least tentatively to the subfamily. 
Some of these genera are included for the 
first time. Some genera are included with 
hesitation, either because they are obscure 
and their type species were not examined 
(e.g., Anamosa, Homalattus) or because 
they exhibited none of the supposed den- 
dryphantine synapomorphies listed later 
but have the general body form and mark- 
ings much like those of other dendry- 
phantines (e.g., Mabellina). Other genera 
are excluded with hesitation — for in- 
stance, those genera related to Ballus (Co- 
laxes, Marengo, PadiUa, Pachyballus, and 
perhaps Admestina) and to Synageles 
(Consingis, Descanso, Peckhaniia, and 



Pelegrina Jumping Spiders • Maddison 227 



Table 1. Generic 



names ok jumi'inc; 
Dendryphantinak 



SPIDERS lEN lA Il\ K1,V REEERRED 

Synonymies are not indicated. 



TO THE SUBEAMIEY 



Admirala G. & E. Peckhani, 1901 

Agassa Simon, 1901 

Amerotritte Mello-Leitao, 1944 

Anamosa G. & E. Peckham, 1895 

Atiicins Chamberlin, 1925 

Anoka G. & E. Peckham, 1893 

Ashtalntia G & E. Peckham, 1894 

Avitus G. & E, Peckham, 1896 

Bagheera G & E. Peckham, 1896 

Beata G & E. Peckham, 1895 

BeUota G. & E. Peckham, 1892 

Bryantella Chickering, 1946 

Cerionesta Simon, 1901 (n. nov. for Cydonia G & 

E Peckham, 1893, preoccupied) 
Chirothecia Taczanowski, 1878 
Dendryphantes C. L. Koch, 1837 
Donaldius Chickering, 1946 
Dryphias Simon, 1901 
Eris C. L. Koch, 1846 
Gastromicans Mello-Leitao, 1917 
Ghelna new genus 
Hentzia Marx, 1883 
Homalattoides F. P. -Cambridge, 1901 
Homalaitus White, 1841 
Lurio Simon, 1901 
Mabellina Chickering, 1946 
Maevioheata di Caporiacco, 1947 
Megatimus Thorell, 1891 
Messita G. & E. Peckham, 1896 
Metaphidipptis E. P. -Cambridge, 1901 



Nagaina G. & E. Peckham, 1896 

Oscricta Simon, 1901 

Paradamoetas G. & E. Peckham, 1885 

Parahentzia Bryant 

Paraphidippus F. P.-Cambridge, 1901 

Parnaenus G. & E. Peckham, 1896 

Partona Simon, 1902 

Pelegrina Franganillo, 1930 

Phanias F. P.-Cambridge, 1901 

Phidippus C. L. Koch, 1846 

Poultonella G. & E. Peckham, 1909 

Ramboia Mello-Leitao, 1944 

Rhene Thorell, 1869 (n. nov for Rhanis C. L. Koch, 

1848, preocc.) 
Rhetenor Simon, 1902 
Rtidra G. & E. Peckham, 1885 
Sassactis G. & E. Peckham, 1895 
Sebastira Simon, 1901 
Selimus G. & E. Peckham, 1901 
Semora G. & E. Peckham, 1892 
Tacuna G. & E. Peckham, 1901 
Terralonus new genus 
Thammaca Simon, 1902 
Tulpius G. & E. Peckham, 1896 
Tutelina Simon, 1901 
Uluella Chickering, 1946 
Wala Keyserling, 1884 
Zeuxippus Thorell, 1891 
Zygoballus G. & E. Peckham, 1885 



perhaps Cheliferoides and Leptorchestes) 
may very well be derived dendryphan- 
tines (see Maddison, 1988). 

Simon (1901, 1903) placed many of these 
dendryphantine genera in his group Den- 
dryphanteae or Rheneae, though a num- 
ber of them were scattered among other 
groups: Beata in the Simaetheae, BeUota 
in the Synageleae, Nagaina in the Belli- 
eneae, Rudra in the Rudreae, Tutelina in 
the Chrysilleae, and Zygoballus in the Zyg- 
oballeae. G. & E. Peckham (1901b) in- 
cluded in their Phidippus group of genera 
Phidippus, Paraenus, Dendryphantes, Se- 
limus, and Admirala, here considered 
dendryphantines, but as well many other 
genera now considered to belong to the 
Euophryinae and other subfamilies. Pro- 
szynski (1976: 15, 148-150) listed the gen- 
era Cheliferoides, Dendryphantes, Eris, 



Metaphidippus, Paradamoetas, Phidip- 
pus, Rhetenor, Sassacus, Thiodina, and 
Wala, in the subfamily Dendryphantinae; 
all except Cheliferoides and Thiodina are 
here considered dendryphantines. 

Specifying distinct apomorphies to jus- 
tify my concept of the subfamily is diffi- 
cult, for no characters both universal 
throughout and unique to the group are 
known; homoplasy must therefore be in- 
voked if the subfamily is to be accepted 
as proposed. Three characters derived 
within the family are proposed as syna- 
pomorphies of the subfamily: 

1. Carina on underside of the male che- 
licerae (Table 2, character 1). On the 
ventrolateral edge of the basal segment 
of the chelicera there is a fold or carina 
(Fig. 10). This carina occurs in almost 



228 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



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230 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



all genera considered dendryphantines. 
It is absent in some groups in which the 
chelicerae are much elongate (Messua, 
Hentzia, Rudra, Paradamoetas), 
though this may be related to the elon- 
gation, as Hentzia mitrata (with short 
chelicerae) does have the carina. More 
troublesome are those dendryphantine 
groups (some Phanias, Tutelina, "Eris' 
nidicolens, "Beata" octopunctata) in 
which the chelicerae are not elongate 
and yet the carina is lacking. In at least 
one of these, Phanias, there are some 
species with the carina, and its absence 
in others is presumably a secondary loss. 
The carina is lacking in all other salti- 
cids I have seen, except for Simaetha 
paetula (see Prosynski, 1984: 132) and 
Simaetha tenuior, in which a similar 
carina is present. These species have a 
Sitticus-\ike palpus, with a round te- 
gulum and immovable embolus, and 
appear to be distantly related to den- 
dryphantines. 
2. Coil of embolus compressed, so that the 
embolus folds back sharply on itself and 
superficially appears not to be coiled 
(Figs. 20-25, 31-35, 37-39, 64b-d, g; 
Table 2, character 2). Typically, the 
embolus arises prolaterally and moves 
across toward the retrolateral side (the 
transverse basal portion of the embolus) 
and then folds back toward the prolat- 
eral and abruptly rises as the erect api- 
cal portion. A suture on the back side 
of the embolus, between the transverse 
portion and the erect portion, is often 
present and indicates where the folded- 
back spiral has not completely fused. 
The coils of the embolus are not folded 
but rather open and exposed (Fig. 19), 
in the other groups, such as the eu- 
ophryines, that have a counterclock- 
wise-coiled embolus. Two main prob- 
lems with the use of this character are 
posed by various dendryphantines: in 
some, the embolus appears not even 
coiled, and in others the embolus ap- 
pears coiled but the coils are exposed 
(Fig. 64 summarizes some of the vari- 



ation seen among the dendryphan- 
tines). Despite the first problem, coiling 
is considered primitive for the subfam- 
ily (see earlier discussion under the sec- 
tion Explanation of Morphological and 
Behavioral Terms). More troublesome 
is the second problem: the occurrence 
of dendryphantine groups such as Par- 
adamoetas, the South American "Sas- 
sacus," Mahellina, Dendryphantes tro- 
picus, and Bryantella in which the coils 
of the embolus are exposed (Figs. 26, 
27, 58-63, 64e, f). However, three lines 
of evidence suggest that the exposure 
of the coils is not homologous to that of 
the euophryines and, instead, is sec- 
ondarily derived from the hidden con- 
dition. First, the exposed spiral of Den- 
dryphantines is always of a different 
form from that of the euophryines, be- 
ing placed more retrolaterally and not 
so small and tightly coiled. Second, oth- 
er characters suggest that these trou- 
blesome dendryphantines do indeed 
belong with other dendryphantines, 
such as the lack of a concave retrolateral 
loop on the sperm duct in the tegulum, 
which may be apomorphic as noted lat- 
er with respect to Pha7iias, the more 
basally placed tegular ledge, which may 
be apomorphic or plesiomorphic, and 
the occurrence of the cheliceral carina 
in at least some species (e.g., Metaphi- 
dippus vitis). Third, there is an appar- 
ent morphocline between the hidden 
spiral of Eris aurantia (Fig. 64b), 
through the more marginally open spi- 
ral of Eris militaris and a species near 
Zygoballns incertus (Figs. 64d, 53, 57), 
to the more open spiral of Zygoballns 
incertus and Paradamoetas (Figs. 64e, 
58), to the open and well-coiled spiral 
of Mahellina and Dendryphantes tro- 
picus (Figs. 62, 63, 64f). The transition 
would require merely a retrolateral shift 
of the erect portion of the embolus and 
a loss of sclerotization of the basal (lat- 
erally directed) portion of the embolus 
to leave the erect part of the embolus 
free, at which point it could coil as in 



Pelegrina Jumping Spiders • Maddison 231 



Dendryphantes tropicus and Mabelli- 
na prescotti (Figs. 62, 63). 
3. Epigynal openings S-shaped, with en- 
try toward the lateral in the anterior 
half and toward the medial in the pos- 
terior half (Figs. 5, 65, 67-70; Table 2, 
character 3). To my knowledge, this 
sinuate opening is unique among sal- 
ticids. Though most dendryphantines 
have this character, it has similar prob- 
lems to the preceding one and is cor- 
related with it. A number of dendry- 
phantines (e.g., Eris militaris, Tutelina 
elegans, Phidippiis clarus, Hentzia, 
Paradamoetas, Anicius, Zygoballus, 
Messua, Bagheera) have C-shaped or 
simple cavernous openings. In some of 
these, however, ridges descending into 
the opening are presumably remnants 
of the teardrop-shaped flaps associated 
with the sinuate openings (Eris mili- 
taris. Fig. 66; Tutelina elegans). For 
some of these, there are related species 
that have the sinuate openings {Eris 
floridana, Tutelina hartii, Phidippus 
spp., Anicius sp., Zygoballus tibialis), 
but there remain other genera with no 
remnant of the flaps or obvious close 
relatives with sinuate openings. In gen- 
eral, the species lacking a compact em- 
bolic spiral also lack the sinuate open- 
ings, perhaps because a retrolateral shift 
of the erect portion of the embolus is 
correlated with an expansion of the epi- 
gynal openings and weakening of the 
teardrop-shaped flaps. Such a correlat- 
ed change seems to mark each of Eris 
militaris, Pelegrina kastoni, Dendry- 
phantes nigromaculatus, and Phidip- 
pus octopunctatus from its close rela- 
tives. 

RELATIONSHIPS WITHIN THE 
DENDRYPHANTINAE 

The limits and interrelationships of gen- 
era within the subfamily Dendryphantin- 
ae (see Table 1 for a list) are at present 
poorly understood. The following discus- 
sion will make an attempt to resolve only 
a small part of the confusion, for I will 



focus on those phylogenetic questions that 
are most important to resolve the generic 
placement of the galathea group of spe- 
cies, which is the subject of the species 
revision that makes up the bulk of this 
paper. The galathea group has resided in 
the genus Metaphidippus, but, as ex- 
plained next, this genus is polyphyletic. 
Here I will begin (but not complete) the 
task of dismantling Metaphidippus. The 
galathea group will be moved to the genus 
Pelegrina, the harfordii group to Phanias, 
the mylothrus group to Terralonus, the 
castaneus group to Ghelna, some other 
species to Messua, but some species groups 
(mannii group, vitis group, various neo- 
tropical species) will remain temporarily 
in Metaphidippus for want of a better al- 
ternative. Table 3 lists the proposed re- 
classification of Metaphidippus and some 
other dendryphantine species. 

Metaphidippus was described in 1901 
by F. O. Pickard-Cambridge, who gave no 
clear justification for its limits. Many of 
the North American species that had been 
placed in Dendryphantes were subse- 
quently transferred to Metaphidippus, in 
part because of Bryant's (1941) conclusion 
that these species did not belong with the 
Old World Dendryphantes. In fact, her 
cited evidence was mistaken: the Euro- 
pean specimens she compared were ac- 
tually "Eris" nidicolens misidentified as 
D. hastatus (Maddison, 1988). Regardless, 
the more or less wholesale transfer of North 
American species from Dendryphantes re- 
sulted in a Metaphidippus that has been 
desperately polyphyletic, being nothing 
more than a catch-all genus of unremark- 
able North and Central American dendry- 
phantines spanning much of the diversity 
of the subfamily. Because the type species 
of Metaphidippus (M. mandibulatus F. O. 
Pickard-Cambridge) is not closely related 
to most species placed in the genus, in- 
cluding the galathea group, many species 
of Metaphidippus should be placed else- 
where. The relationships of the true Me- 
taphidippus will first be considered, after 
which the limits of Dendryphantes and 



232 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



Table 3. Summahy of placements of various New World dendryphantine species apart from 
Pelegrima species, discussed in text. ? indicates placement considered possible but unconfirmed. 

Authors of names are given in index. 



Bagheera 
Bagheera kiplingi 
B. prosper 

?Metaphidippus nigropictus 
?M. bicavatus 

Beat a 
Beata hispida 
B. inconcinna 
B. longipes 
B. maccunii 
B. magna 
B. rust tea 

Dendryphantes (partial) 
Dendryphantes nigrotnaculatus 
D. chiildensis 
D. fusconotatus 
D. hastatus 
D. rudis 

Gastromicans 
Gastromicans albopilosa 
G. hondurensis 
G. levispina 
G. noxiosa 
G. vigens 
?Hasarius lisei 

Messua 
Messua desidiosa 
M. centralis 
M. dentiger 
M. donalda 
M. lata 
M. laxa 
M. limbata 
M. niunia 
M. octonotata 
M. pura 
M. tridentata 
?M etaphidippus cupreus 
?Metaphidippus ovatus 
?M etaphidippus iridescens 
?M etaphidippus inflatus 
?M etaphidippus quadrinotatus 
^Dendryphantes felix 



Phanias 
Phanias albeolus 
P. concoloratus 
P. doniinatus 
P. flavostriatus 
P. furcifer 
P. furcillatus 
P. harfordii 
P. monticola 
P. neomexicanus 
P. watonus 
?P. salvadorensis 

Terralonus 
Terralonus californicus 
T. niylothrus 
T. unicus 
T. shaferi 
T. versicolor 
T. vittatus 
T. fraternus 

Ghelna 
Ghelna castanea 
G. barrowsi 
G. canadensis 
G. sexmaculata 

Metaphidippus niannii group 
Metaphidippus mannii 
M. carmenensis 
M. chera 
M. diplacis 
M. emniiltus 
M. tricolor 

Metaphidippus vitis group 
Metaphidippus vitis 
M. texanus 
M. mathetes 
Dendryphantes melanomerus 



Beata will be discussed. Then, the correct 
placement of various groups placed in Me- 
taphidippus will be considered. 

The Bagheera Group of Genera 

It is within the Bagheera group, com- 
mon in the Neotropics, that the true Me- 



taphidippus appear to fall. Members of 
this group, which includes such common 
species known as ^^Eris' limbata, ^'Meta- 
phidippus" prosper, "Beata" albopilosa, 
have a distinctive embolus, which appears, 
at first glance, to be coiled or curved clock- 
wise in the left palpus, opposite the coun- 



Pelegrina Jumping Spiders • Maddison 233 



terclockwise coiling I have said character- 
izes the dendry phantines and related sub- 
families (Figs. 99-101). This clockwise 
coiling is apparently superimposed upon 
the normal coiling by a twisting of the 
embolus. The main axis of the embolus has 
twisted counterclockwise (as viewed from 
tip of embolus in left palpus), thus winding 
the hematodocha and sperm duct around 
it (Figs. 64h, 99-101). The tip of the em- 
bolus, though, seems to have been left be- 
hind by the twisting, so that the apical part 
of the embolus takes on a clockwise curl- 
ing. In some species, this clockwise coiling 
is visible in the uncleared palpus (Figs. 81, 
84, 87, 93), but in others it is not (Fig. 91), 
and the palpus is little modified from the 
typical compressed counterclockwise spi- 
ral of dendryphantines. The most extreme 
clockwise coiling is seen in "Metaphidip- 
pus" prosper (Fig. 99) and ''Beata' al- 
bopilosa (see Fig. 101). 

Species in the group can be sorted pro- 
visionally into three subgroups, which may 
or may not be monophyletic, for each of 
which there exists an available generic 
name that has been mostly ignored in the 
literature: Bagheera G. & E. Peckham, 
Messua G. & E. Peckham, and Gastromi- 
cans Mello-Leitao. A fourth genus sharing 
a twisted embolus is Ashtabula G. & E. 
Peckham, though whether or not it belongs 
with the group is unclear. The twisting of 
the embolus in Ashtabula (Fig. 102) is hid- 
den beneath the tegulum and is more ex- 
treme, though similar to, that in Bagheera, 
Messua, and Gastromicans. However, 
there are several features that cast doubt 
on the placement of Ashtabula with these 
genera. Ashtabula has an extra concave 
loop on the sperm duct in the palpus, pos- 
sibly placing it near the base of the sub- 
family as noted later in connection with 
Phanias. The body in Ashtabula is not 
nearly so large and robust as in the Bag- 
heera group, instead resembling that of 
Hentzia or Anicius. More work is needed 
before the place of Ashtabula can be set- 
tled. Species of Ghelna and the arizonensis 
group of Pelegrina (discussed later) both 



have a twisted embolus, but in each it takes 
a very different form from the twisting in 
the Bagheera group of genera. Sebastira, 
Thammaca, Lurio, and Parnaenus may 
also belong with the Bagheera group of 
genera, though their emboli do not so ob- 
viously possess the twisting. A brief ac- 
count of Bagheera, Messua, Gastromicans 
and Metaphidippus is here given. 

Bagheera {Figs. 80-85). Bagheera males 
have elongate, horizontal, parallel chelic- 
erae (Fig. 80); the retromarginal teeth are 
near the base of the chelicera; in all but 
one species there is distally, near the fang, 
what appears to be a large retromarginal 
tooth but actually is not (it does not have 
the terminal canal through the cuticle that 
seems to characterize all true teeth), and 
most species have tubercles bearing setae 
on the inner margin of the basal cheliceral 
segment. Included in Bagheera are the type 
species, B. hiplingi G. & E. Peckham, 1901 
(type species by monotypy; holotype ex- 
amined; Figs. 80-83), and Bagheera pros- 
per (G. & E. Peckham) (NEW COMBI- 
NATION; Figs. 84, 85, 99) and at least two 
undescribed species. Metaphidippus ni- 
gropictus F. P. -Cambridge and M. bica- 
vatus F. P. -Cambridge may also belong in 
Bagheera. 

Messua (Figs. 86-92). Males of this ge- 
nus have elongate divergent chelicerae 
(Fig. 86) with a long and sickle-shaped 
retromarginal tooth near the fang; the pro- 
marginal teeth are near the base, well sep- 
arated from the retromarginal tooth; on 
the anterior distal margin of the basal seg- 
ment of chelicera near the fang is a flange. 
Included in Messua are the type species 
M. desidiosa G. & E. Peckham, 1896 (type 
species by monotypy; holotype and collec- 
tions of males and females from San Jose, 
Costa Rica examined; Figs. 86-89), Mes- 
sua centralis (G. & E. Peckham) (lectotype 
here designated, a male from Chiriqui), 
Messua dentiger (F. P. -Cambridge) (see 
Fig. 91), Messua donalda (Kraus), Messua 
lata (Chickering), Messua laxa (Chicker- 
ing), Messua limbata (Banks) (Figs. 90, 
100, 117), Messua moma (F. P. -Cam- 



234 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



bridge), Messua octonotata (F. P. -Cam- 
bridge), Messua pura (Bryant), and Mes- 
sua tridentata (F. P. -Cambridge). All these 
except M. desidiosa are NEW COMBI- 
NATIONS. Metaphidippus cupreus F. P.- 
Cambridge, M. ovatus F. P. -Cambridge, 
M. iridescens F. P. -Cambridge, M. injla- 
tus F. P. -Cambridge, and M. quadrino- 
tatus F. P. -Cambridge may also belong in 
Messua. Dendryphantes felix G. & E. 
Peckham might be considered either a 
Bagheera or Messua depending on any 
future lectotype designation: the body (G. 
& E. Peckham, 1901b: fig. 6a) in the type 
vial and its attached palpus are of a Bag- 
heera species, probably B. prosper, while 
the separate palpus in a microvial (G. & 
E. Peckham, 1901b: fig. 6) is of a Messua 
species. 

Gastromicans (Figs. 93-95). This genus 
is distinguished from Bagheera and Mes- 
sua in having short and vertical but very 
robust male chelicerae. Included are Gas- 
tromicans albopilosa (G. & E. Peckham), 
Gastromicans hondurensis (G. & E. Peck- 
ham), Gastromicans levispina (F. P. -Cam- 
bridge) (Figs. 93-95, 101), Gastromicans 
noxiosa (Simon), and Gastromicans vigens 
(G. & E. Peckham). All these are NEW 
COMBINATIONS. Hasarius lisei Bauab- 
Vianna & Soares probably also belongs in 
Gastromicans. Galiano (1980) synony- 
mized Gastromicans Mello-Leitao with 
Beata G. & E. Peckham because its type 
species Gastromicans squamulata Mello- 
Leitao (type species by monotypy) is syn- 
onymous with "Beata" albopilosa. But in- 
sofar as Beata albopilosa does not belong 
in the genus Beata, Gastromicans is avail- 
able as a generic name for albopilosa and 
its relatives. 

Metaphidippus (Figs. 96-98). Though 
the placement of the true Metaphidippus 
with these genera of the Bagheera group 
is to some extent problematical, such a 
placement is the best supported at present. 
Before discussing the uniting characters, it 
would be valuable to give the following 
brief description of the type species of Me- 
taphidippus, M. mandibulatus F. P. -Cam- 



bridge (type species by original designa- 
tion), whose single known male (Costa Rica, 
BMNH, examined), is strikingly unlike 
most other jumping spiders that have been 
placed in Metaphidippus. Palpus (Figs. 
97, 98): Embolus reminiscent of that of 
Eris species but with the longitudinally 
directed apical portion not fully erect, in- 
stead reclined to the prolateral (Fig. 98). 
The embolic base bears a flange covering 
the basal part of the embolus. Chelicerae: 
Long and cylindrical, horizontal and di- 
verging (Fig. 96), with two promarginal 
teeth near the base and one retromarginal 
tooth near the fang. The fang is forked 
near its base (Fig. 96). Markings (Fig. 96): 
Carapace brown, lacking side bands ex- 
cept one patch of white scales on either 
side of fovea. Wide white band along mar- 
gin. At least some metallic green-blue scales 
on cephalic area. Abdomen with thin white 
side bands broken basally; just anterior to 
each of the main dorsoventral muscle at- 
tachments is a small white patch of scales; 
in the posterior half of the dorsum are two 
pairs of small lateral white bars. The dor- 
sum has some metallic green-blue scales. 
Measurements: Body length 5.4 mm; car- 
apace length 2.4 mm, carapace width/ 
length 1.93/2.37. 

Two features that can be proposed as 
synapomorphies for the group of Bag- 
heera, Messua, Gastromicans, and Me- 
taphidippus are the following: 

1. The tibial apophysis is erect and at its 
base parallel-sided, shaped like a knife 
blade (Figs. 71-74; Table 2, character 
4). Almost all other dendryphantines 
have an apophysis tapering throughout 
its length (Figs. 75-79), including 
Fhanias and other genera that appear 
to be near the base of the subfamily 
(see later), and usually the apophysis 
points at least somewhat ventrally. The 
only other dendryphantines known to 
me with a similar knife-shaped apoph- 
ysis are Ashtabula and Poultonella. 
Poultonella does not belong with these 
genera; rather, its peculiar chelicerae 



I 



Pelegrina Jumping Spiders • Maddison 235 



assure a relationship with Tutelina. A 
few species of Messua, inchiding M. 
lata, have a more tapering apophysis. 
2. The tegular ledge runs longitudinally 
(Figs. 87, 90-93, 98; Table 2, character 
5), instead of obliquely at 0-60° from 
the transverse as seen in other dendry- 
phantines and other salticids with a teg- 
ular ledge (Figs. 40-46, 50, 52, 53). 
While this is unusual among dendry- 
phantines, it is not unique: it also occurs 
in Phanias, Anicius, and Zygoballus in- 
certus. 

Additional features that suggest a rela- 
tionship between Metaphidippus and 
Messua in particular are the long, tubular 
divergent chelicerae with a near-terminal 
retromarginal tooth and a distal anterior 
flange beside the fang base. Other den- 
dryphantines have long divergent chelic- 
erae, but in all that I have seen except some 
South American "Sassacus," the tooth ar- 
rangement is different than in Metaphi- 
dippus and Messua, with the retromar- 
ginal tooth remaining near the base or the 
promarginal teeth near the apex. This dif- 
ferent tooth placement may indicate that 
the elongation occurred in different por- 
tions of the chelicerae in these other den- 
dryphantines, and thus the elongation is 
not homologous. As well, the general body 
form and occurrence of greenish reflective 
scales are also suggestive of a close rela- 
tionship between Metaphidippus and 
Messua, though these characters are loose- 
ly defined and not necessarily unique. The 
only feature that would exclude M. man- 
dibulatus from Messua is the apparent lack 
of the reverse twisted embolus in M. man- 
dibulatus. However, one undescribed spe- 
cies from Costa Rica represented by a sin- 
gle male specimen appears very closely 
related to M. mandibulatus in having sim- 
ilar body form and markings and in having 
a slightly forked fang, and yet it has a 
slightly twisted embolus (Fig. 92). If these 
two species form a monophyletic group, 
then the lack of twisting in mandibulatus 
may be a secondary loss, which is not un- 



reasonable given that other species such as 
Messua octonotata have little trace of a 
twisted embolus, and the embolus of M. 
mandibulatus shows unusual folds and does 
recline to the prolateral as in Messua. Per- 
haps more detailed study of its peculiar 
embolus, when more specimens become 
available, will allow a more definitive an- 
swer to the question of twisting in M. man- 
dibulatus. If the genus Metaphidippus is 
only an offshoot of Messua, then Meta- 
phidippus would fall as a synonym of the 
older name Messua. However, I am re- 
luctant to effect such a synonymy at pres- 
ent given the number of Metaphidippus 
species that would be left homeless, and 
so Metaphidippus will be left standing for 
the moment. Regardless, the best-support- 
ed conclusion at present is that the name 
"Metaphidippus" properly applies to a 
small group of neotropical dendryphan- 
tines related to Messua, Bagheera, and 
Gastromicans. 

Two Genera That Have Exchanged 
Species with Metaphidippus: 
Dendryphantes and Beata 

Because most species that have been 
placed in Metaphidippus do not belong to 
the Bagheera group of genera, they cannot 
follow M. mandibulatus and, thus, need 
to be placed elsewhere. The first place we 
might look for a possible home for Me- 
taphidippus species are two genera, Den- 
dryphantes and Beata, which have in the 
past exchanged many species with Meta- 
phidippus. Many "Metaphidippus" were 
formerly placed in Dendryphantes and 
several Beata were formerly placed in Me- 
taphidippus. To discuss the proper place- 
ment of species now in Metaphidippus 
more clearly, it would be valuable to re- 
consider the limits of these two genera. 

Dendryphantes (Figs. 65, 103-108, 120). 
The genus Dendryphantes, described last 
century, has over the years accumulated 
many species, mostly on the basis of their 
being unremarkable dendryphantines. 
Many species were since moved to genera 
such as Metaphidippus, while others re- 



236 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



maining in the genus will probably even- 
tually be placed elsewhere (see the com- 
ments of Edwards, 1977). Among the New 
World species, there is only one species for 
which there is presently good evidence for 
a placement in Dendryphantes: D. nigro- 
maculatus (Keyserling), most recently 
placed in Eris (Kaston, 1973). Several Old 
World species placed in Dendryphantes, 
including D. fusconotatiis and D. chuld- 
ensis, appear very closely related to D. 
nigromaculatus (see figures of Proszyhski, 
1971b, 1982). Like the Old World D. has- 
tatus (the type species) and D. rudis, D. 
nigromaculatus has a slightly elongate 
body dully marked. Perhaps the best char- 
acter that strictly delimits Dendryphantes 
is the presence of a fold of embolic he- 
matodocha that lies across the basal part 
of the embolic base, covering the wrinkles 
there (Figs. 103-108). If this character is 
used to delimit the genus, then it would 
be a small genus of mostly Palearctic spe- 
cies. The placement of nigromaculatus in 
Dendryphantes is further supported by its 
sharing with D. rudis, D. fusconotatus, 
and D. chuldensis a much elongated prong 
coming off of the base of the embolus and 
curving toward the cymbium (Figs. 106, 
108). The embolus therefore appears to 
have two rami, much as in Pelegrina fla- 
vipedes, though not homologous according 
to reasoning given below. Species of the 
other two major groups of Old World den- 
dry phantines, the "Eris" nidicolens group 
(Fig. 61) and the genus Rhene (Fig. 52), 
lack the fold across the embolic base seen 
in Dendryphantes. Though Rhene species 
often have a prong arising from the base 
of the embolus, in Rhene it is not curled 
toward the cymbium and, instead, is erect 
as in Beata and the mannii group. Rhene 
has been considered a close relative and 
possibly a synonym of Dendryphantes 
(Proszynski, 1973b), but a number of other 
features of Rhene such as the presence of 
epiandrous fusules and the concave retro- 
marginal loop of the sperm duct of the 
palpus also cast some doubt on this place- 
ment. 



The option of returning a number of 
groups from Metaphidippus to Dendry- 
phantes has little merit at present. Moving 
these back to Dendryphantes would be 
useful only if they are likely to stay there, 
that is, if they are closely related to the 
type species of Dendryphantes. Other- 
wise, we would merely be worsening Den- 
dryphantes' status as a catch-all genus and 
adding to it the confusion of changing the 
generic placement of many common spe- 
cies. As noted, only D. nigromaculatus 
among New World species is a strong can- 
didate to stay in Dendryphantes. 

Beata (Figs. 77, 109-112). The limits of 
the genus Beata have been greatly over- 
estimated (Simon, 1903; Chickering, 1946). 
Because the type species of Beata is fissi- 
dent (it has a bifid retromarginal cheliceral 
tooth), it has not only been removed from 
the dendryphantines (Simon, 1903) but has 
also been burdened with diverse dendry- 
phantines that happen to have a similarly 
bifid tooth (Simon, 1903; Chickering, 
1946). Note that the tooth is better consid- 
ered a single bifid tooth rather than two 
fused teeth because the inner boundary of 
the cuticle does not extend to the tip of 
the second cusp. The second cusp shows 
all gradations of development in the den- 
dryphantines, with most lacking it, some 
showing a slightly swollen margin (e.g., 
Pelegrina proterva, Fig. 10), and others 
having a well-developed cusp. It is there- 
fore best to place far less emphasis on this 
character. Beata magna G. & E. Peckham 
(Fig. 109), the type species of Beata (by 
monotypy), bears few resemblances to most 
of the other fissident dendryphantines, in- 
stead having many more resemblances with 
the other robust-bodied dendryphantines 
previously placed in Dryphias, Homalat- 
toides, and Anamosa. The following char- 
acters, which appear derived within the 
subfamily, delimit this group containing 
Beata magna: 

1. Tibial apophysis narrow and bent to- 
ward the ventral, almost paralleling a 
ridge on the tibia below it (Fig. 77). 



Pelegrina Jumping Spiders • Maddison 237 



This tibial ridge is similar to that in 
Pelegrina (Fig. 78), but it is longer and 
sharper in Beata. 

2. First leg tibia dark and enlarged at least 
slightly compared to patella, even in 
females. 

3. Carapace distinctively wide and high, 
higher than in Sassacus, Agassa, and 
Rhene, wider than in Sassacus and 
Agassa. Unlike Zygoballus, but like 
Sassacus and Agassa, the carapace is 
wide well past the posterior eyes before 
it abruptly drops and narrows. 

4. Carapace scales erect (in at least some 
but perhaps not all species). 

5. Retromargin of base of embolus with 
prong rising parallel to apical erect por- 
tion of embolus (Fig. 110). Such a prong 
is also seen in Rhene (Fig. 52) and the 
Metaphidippus mannii group (Fig. 
499). 



The following species are placed in Bea- 
ta and NEW COMBINATIONS therefore 
established: Beata hispida (G. & E. Peck- 
ham) (Figs. 77, 110-112), Beata inconcin- 
na (G. & E. Peckham), Beata maccunii 
(G. & E. Peckham), and Beata rustica (G. 
& E. Peckham). Also included is Beata 
longipes F. P. -Cambridge, which may be 
the male of B. magna. Dryphias (type spe- 
cies maccuni by original designation) is a 
NEW SYNONYM of Beata. The genus 
Beata as here delimited excludes B. digi- 
tata (= Pelegrina galathea) and B. var- 
iegata (= Pelegrina variegata), B. albopi- 
losa (= Gastromicans albopilosa), B. fla- 
volineata (= Nagaina incunda), B. ce- 
phalica F. P. -Cambridge, B. jubata (C. L. 
Koch), B. munda Chickering, B. pernix 
(G. & E. Peckham), B. venusta Chicker- 
ing, B. wickhami (G. & E. Peckham), and 
B. zeteki Chickering. No new placements 
are suggested for the last-mentioned seven 
species. Other species placed in the genus 
but probably not belonging there are B. 
cinereonitida Simon, B. germaini Simon, 
B. lineata (Vinson), and B. striata Pe- 
trunkevitch. 



The Proper Placements of 
Metaphidippus Species 

Now that the relationships of the true 
Metaphidippus and the limits of Dendry- 
phantes and Beata have been reconsid- 
ered, we are in position to discuss how the 
various groups within Metaphidippus 
might be dispersed. Some of these conclu- 
sions are summarized in Table 3. 

Four groups are here removed from 
Metaphidippus, as discussed in subsequent 
sections. The galathea group is transferred 
to the genus Pelegrina (and its species re- 
vised), the harfordii group to the genus 
Phanias, the mylothrus group to the new 
genus Terralonus, and the castaneus group 
to the new genus Ghelna. 

Two species groups occurring in the 
United States are retained temporarily in 
Metaphidippus pending further study: the 
mannii group and the vitis group. The 
mannii group is discussed later in connec- 
tion with the revision of its U.S. species. 
The status of the vitis group (Figs. 27, 59), 
including Metaphidippus vitis (Cocker- 
ell), M. texanus (Banks), M. mathetes 
(Chamberlin), and Dendryphantes me- 
lanomerus Chamberlin, is not clear. These 
species have a characteristic hooked em- 
bolus (Figs. 27, 59) and are small, some- 
what elongate, and brown to black and 
shiny. Metaphidippus vitis was placed in 
Sassacus by Hill (1979) on the basis of scale 
morphology and courtship, but scale mor- 
phology is known in only few dendry- 
phantines, and a similar courtship is also 
seen in many other dendryphantines 
(raisedforward, Table 2, character 9). Fur- 
thermore, the genitalia of M. vitis are very 
different from those of the true Sassacus 
and, instead, are similar to those of the 
neotropical species placed in Sassacus such 
as S. arcuatus, which may better be placed 
in the genus Ramboia (see Bauab-Vianna 
and Soares, 1982). The vitis group, here 
retained in Metaphidippus, may eventu- 
ally find its place in a primarily neotropical 
genus. 

Some species placed in Metaphidippus 



238 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



belong in Bagheera or Messua, as already 
noted. Metaphidippus paiutus Gertsch is 
a Sassacus (Sassacus paiutus (Gertsch), 
NEW COMBINATION), possibly a syn- 
onym of S. papenhoei. Other species of 
Metaphidippus require further study be- 
fore their placement can be settled. Among 
the neotropical species listed under Me- 
taphidippus by Bonnet (1957), M. longi- 
palpus F. P. -Cambridge, M. nitidus (G. & 
E. Peckham), and perhaps M. taylori (G. 
& E. Peckham) seem to belong to Parna- 
enus, M. pallens F. P. -Cambridge in Eris, 
M. perfectus (G. & E. Peckham) (Fig. 60) 
in Selimus, and M. tropicus (G. & E. Peck- 
ham) (Fig. 62) in Bryantella. For the re- 
maining neotropical species (see Proszyn- 
ski, 1990), I have no placement to suggest. 

Phanias F. P.-Cambridge, 1901 

Type species, Phanias flavostriat us F. P.-Cambridge, 
1901, by monotypy. 

The unusual, elongate dendryphantines 
of the harfordii group (Figs. 20, 36, 47, 
70) include several species in the western 
United States but many more in the high- 
lands of Mexico. A generic name is avail- 
able for this group, Phanias F. P.-Cam- 
bridge, 1901, based on Phatiias flavostria- 
tus, described from two females from Om- 
ilteme, Mexico (BMNH, examined), and 
previously considered to belong in the 
Marpissinae. The species of the Metaphi- 
dippus harfordii group are therefore 
transferred to Phanias. The members of 
the genus Phanias share these characters, 
which may be apomorphies within the 
subfamily: 

1. Tegular ledge expanded so as to cover 
the tegular shoulder (Fig. 47). The teg- 
ular ledge of other dendryphantines and 
other salticids is not so expanded. 

2. Embolic hematodocha reduced and 
sclerotized prolaterally and basally. In 
at least some species, much of the ex- 
pansion occurs from out of the tegular 
ledge instead of from the prolateral 
dorsal surface (back side) of the tegul- 
um (Fig. 36), but this feature may be 



primitive as it is also found in Synageles 
and Admestina (Figs. 29, 30). 

3. Courtship with first legs raised, for- 
ward, and parallel (Fig. 118) and waved 
asymmetrically so that the leading leg 
on sidles is waved exclusively or more 
strongly (seen by me otherwise only in 
Anicius). 

4. Small blunt teeth on the embolus (Fig. 
20). Many other dendryphantines (e.g., 
Pelegrina montana. Fig. 204) have 
teeth on the embolus, but their teeth 
are sharp. The blunt teeth are lacking, 
however, in some species of Phanias 
(e.g., P. watonus). 

5. Longitudinal bands of white scales, in- 
stead of passing below and beside the 
posterior eyes as in other dendryphan- 
tines, pass around and directly poste- 
riorly from the posterior eyes. The dis- 
tribution of this character is not well 
known. 

The following NEW COMBINATIONS 
are established: Phanias albeolus (Cham- 
berlin & Ivie) (Figs. 20, 70), Phanias con- 
coloratus (Chamberlin & Gertsch), Phan- 
ias dominatus (Chamberlin & Ivie), Phan- 
ias furcifer (Gertsch), Phanias furcillatus 
(F. P.-Cambridge), Phanias harfordii (G. 
& E. Peckham) (Fig. 47), Phanias mon- 
ticola (Banks), Phanias neomexicanus 
(Banks), and Phanias watonus (Chamber- 
lin & Ivie). Phanias marginalis Banks (type 
specimen examined) is a Menemerus, not 
a Phanias, while Phanias salvadorensis 
Kraus may be either a Phanias or an An- 
icius. Also included in Phanias are at least 
15 undescribed species from Mexico and 
the southwestern United States. Phatiias 
may be placed near the base of the sub- 
family, for it has two features that are ar- 
guably ancestral for the subfamily, name- 
ly, the presence of epiandrous gland fu- 
sules (Machado, 1951; Table 2, character 
6; see also Maddison, 1988), which it shares 
with Hentzia, "Beata" wickhami, ^^Eris" 
nidicolens, Rhene, and groups apparently 
related to dendryphantines (euophryines, 
synagelines, ballines, Mopsus, Itata, Phle- 



Pelegrina Jumping Spiders • Maddison 239 



gra, though not Neon), and a concave 
sperm duct loop along the retromargin of 
the tegulum (Fig. 20), which it shares with 
some Hentzia, "Eris" nidicolens (Fig. 61), 
Rhene (Fig. 52), some Tutelina, Anicius, 
euophryines (Fig. 19), and Admestina. 

Terralonus new genus 

Type species, Dendryphantes mylothrus Chamber- 
lin, 1925, here designated. Name treated as mas- 
culine. 

Description and Diagnosis (Figs. 22, 44, 
68). These western North American spe- 
cies are unusual among dendryphantines 
in being ground-dwelling, usually on 
ground more or less barren of vegetation, 
often under rocks. The body shape and 
markings are distinctive and uniform 
throughout the group. They are somewhat 
elongate and have relatively low-contrast 
mottled markings of coarse brown or gray 
pubescence. The embolus is long and its 
base is more longitudinally directed than 
is usual in dendryphantines (Fig. 22), ex- 
cept for T. calif ornicus, which has a more 
typical embolus (Fig. 44). 

Included Species. The following species 
are moved to Terralonus: Maevia calif or- 
nicus G. & E. Peckham, Dendryphantes 
mylothrus Chamberlin, Dendryphantes 
unicus Chamberlin & Gertsch, Metaphi- 
dippus shaferi Gertsch & Riechert, Icius 
versicolor G. & E. Peckham, Menemerus 
vittatus Banks, and Menemerus fraternus 
Banks (type specimens of last-mentioned 
three species examined). This establishes 
the NEW COMBINATIONS: Terralonus 
calif ornicus (G. & E. Peckham), Terra- 
lonus mylothrus (Chamberlin), Terralon- 
us unicus (Chamberlin & Gertsch), Ter- 
ralonus shaferi (Gertsch & Riechert), Ter- 
ralonus versicolor (G. & E. Peckham), Ter- 
ralonus vittatus (Banks), and Terralonus 
fraternus (Banks). 

Discussion. By appearance, these are not 
typical dendryphantines — two of the spe- 
cies were described in the genus Mene- 
merus and have remained there to this 
day. Their relatives are unclear, but almost 



certainly they do not belong in the genus 
Metaphidippus, because they lack the 
characters of the Bagheera group of gen- 
era. I have chosen to describe a new genus 
for the group to remove it from its uneasy 
placement in Metaphidippus. 1 do this with 
some hesitation, given the overabundance 
of obscure genera in salticids, but the my- 
lothrus group apparently does not reach 
the Neotropics where it might have found 
an available generic name, and so it is un- 
likely to be synonymized soon. Describing 
a genus allows easier discussion of this dis- 
tinctive group. The three species associ- 
ated with the group for the first time {fra- 
ternus, vittatus, and versicolor) can there- 
fore be moved directly into Terralonus 
without being temporarily sentenced to 
Metaphidippus. 

Ghetna new genus 

Type species, Atttis castaneiis Hentz, 1846, here des- 
ignated. Name treated as feminine. 

Description and Diagnosis. These east- 
ern North American species, like the spe- 
cies of Terralonus, are ground-dwelling, 
though in more mesic habitats. They share 
a dark granulate carapace with fine golden 
scales, posterior lateral spines on the first 
tibia displaced anteriorly, reduced spines 
on the first femora, first coxae nearly 
touching, and the female palpus slightly 
swollen. The embolus, at least in the first 
two species mentioned and perhaps in all, 
is twisted so as to wind the embolic he- 
matodocha around the embolus much as 
in the Bagheera group of genera, though 
the twisting takes a very different form. 

Included species. The species Attus cas- 
taneus Hentz, Metaphidippus barrowsi 
Kaston, Icius sexmaculatus Banks, and 
Icius canadensis Banks are here moved to 
Ghelna to establish the following NEW 
COMBINATIONS: Ghelna castanea 
(Hentz), Ghelna barrowsi Kaston, Ghelna 
sexmaculata (Banks), and Ghelna cana- 
densis (Banks). 

Discussion. As with Terralonus, the rel- 
atives of Ghelna are unclear but, likewise, 



240 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



are not near the genus Metaphidippus. The 
justification for a new generic name is sim- 
ilar to that for Terralonus. 

THE GENUS PELEGRINA 
FRANGANILLO, 1930 

Pelegrina Franganillo, 1930: 44. Type species by 
original designation and monotypy Pelegrina gen- 
iculata Franganillo (= P. proxima). 

Notes on Synonymy. The problem of 
the generic name to be given to the gal- 
athea group was not an easy one to solve. 
In my thesis (Maddison, 1988), I concluded 
that a new generic name was needed for 
the group, because (1) it clearly does not 
belong with Metaphidippus and (2) the 
group is arguably monophyletic and of re- 
spectable size for a genus, and (3) it ap- 
peared that there was no genus whose type 
species fell within the group. However, 
subsequent investigation has indicated that 
an obscure Franganillo name, Pelegrina, 
should be applied to the galathea group. 
Although the revival of obscure names is 
often undesirable, in Pelegrina's case there 
is little harm because no other published 
name is available for the group. In 1930, 
Franganillo described Pelegrina and based 
it on Pelegrina geniculata Franganillo, 
which he placed in the section Unidentati, 
subfamily Heliophaninae. As discussed un- 
der the description of P. proxima, P. gen- 
iculata is here considered a junior syn- 
onym of Dendryphantes proximus G. & 
E. Peckham; thus, the name Pelegrina is 
applicable to the galathea group. 

Description and Diagnosis. Small to 
medium-sized dendryphantines distribut- 
ed throughout North and Central Ameri- 
ca. Males (Fig. 1) typically brown with 
longitudinal bands of white scales on either 
side of the carapace and abdomen. The 
inverted white V-shaped marking on the 
forehead that contacts the AMEs distin- 
guishes Pelegrina from most other den- 
dryphantines, though it is not present in 
all Pelegrina. Legs often with annulate 
markings. The relatively wide embolus 
with the tip expanded retrolateral to its 



opening and bearing two rami (Figs. 3, 
190-216, 220-224) is generally a good di- 
agnostic feature for the genus, but it is 
absent in a number of species. Tibial 
apophysis stout; just ventral to apophysis 
is usually a ridge (Fig. 78), developed into 
a second apophysis in some species (Jur- 
cata group) or a wide flange in other spe- 
cies (arizonensis group). Females gray, 
yellow, or brown with mottled markings 
of four prominent pairs of pale spots on 
the abdominal dorsum (see, e.g., Figs. 2, 
263, 269, 358, 382). Epigynal openings rel- 
atively long. Among small dendryphan- 
tines, the species of Pelegrina have per- 
haps the best-developed epigynal flaps 
(Figs. 236-255), which are the teardrop- 
shaped lateral rims of the openings. The 
flaps are usually convex and overlap the 
medial rim of opening. All species of Pe- 
legrina examined have the same chro- 
mosome complement, 2n6 = 26-1- XXO, 
as is prevalent throughout the family. 

Monophyly. Thirty-eight species are in- 
cluded in Pelegrina. Most of these species 
can be easily recognized as belonging to 
the genus by an experienced identifier on 
the basis of body form, size, and markings, 
but to articulate precisely characters that 
could serve as evidence for monophyly is 
more difficult. The following characters 
support the monophyly of the genus, 
though none provides a simple, strict de- 
limitation. Some of the characters delimit 
a group slightly smaller than the genus, 
others a group slightly larger. Thus, each 
character provides only indirect and par- 
tial evidence for monophyly. 

1. Embolus with two terminal rami retro- 
lateral to opening (Figs. 3, 190-216, 
220-224). The sperm duct opening lies 
on the prolateral side of the embolus, 
often below the tip. Retrolateral to the 
opening are two rami, one just distal to 
the opening; the other often elongate 
(see especially P. tristis. Fig. 197) and 
forming the retrolateral tip of the em- 
bolus. While other dendryphantines 
such as Tulpius, Phanias, and Tutelina 



I 



Pelegrina Jlimping Spiders • Maddison 241 



have accessory rami emerging from the 
embohis, none have such rami in the 
position or form seen in Pelegrina. This 
is perhaps the clearest character dehm- 
iting the genus, but some Pelegrina ap- 
pear to lack it (vereciinda, tillandsiae, 
btinites, orestes, arizonensis, helenae; 
Figs. 217-219, 225-227), while others 
have the rami but in a much modified 
form (flavipedes and related species; 
Figs. 201-203). These problems are dis- 
cussed later. 

2. Hematodocha of embolus bulges as far 
distally as the base of the erect portion 
of the embolus (Fig. 3; Table 2, char- 
acter 7). This feature is present 
throughout Pelegrina, including P. or- 
estes and P. bunites. In other dendry- 
phantines examined, including the 
mannii group and Eris, the hemato- 
docha joins the retrolateral edge of the 
embolic base more basally so that the 
hematodocha fails to bulge as far dis- 
tally (Figs. 22, 23). 

3. Epigynal flaps well developed, long, and 
wide and not descending into the open- 
ing posteriorly. This character is diffi- 
cult to assess, for there are other den- 
dryphantines with well-developed flaps, 
though in these the flaps do not exactly 
match those of Pelegrina, being either 
much shorter (Phidippus, Fig. 67; Bel- 
lota), narrower {Beata; Figs. 109, 112), 
or less convex (e.g., "Pseudicius'^ siti- 
culosus). The flaps of most Pelegrina 
differ from those of the mannii group 
and Eris, which have weak flaps that 
descend into the opening at posterior 
end (Figs. 66, 256, 257); the flaps in the 
two most problematical Pelegrina spe- 
cies, P. bunites and P. orestes, are 
somewhat like those of the mannii 
group. 

4. Wrinkles present on anterior margin of 
male cheliceral fang (Fig. 11; Table 2, 
character 8). Running parallel to the 
serrate edge of the fang (Hill, 1977b), 
just anterior to it, is a line of transverse 
wrinkles, which appears like an irreg- 
ular secondary serrate edge. In Pele- 



grina, except P. orestes, it reaches dis- 
tally to the fang opening. This contrasts 
with Dendryphantes, the mannii group, 
and most other dendryphantines, which 
lack such wrinkles. The only other sal- 
ticids seen with such a "secondary ser- 
rula" are (a) Phanias (four species ex- 
amined), in which the wrinkles are re- 
stricted to a depression that does not 
reach the opening; (b) Selimus sp. and 
'^Beata" octopunctata, in which the 
wrinkles are long and regular; and (c) 
Eris militaris, in which the wrinkles do 
not quite reach the opening. Except for 
E. militaris, the wrinkles are of differ- 
ent form, suggesting they are not ho- 
mologous. 

5. Distinct cheek bands on the male face 
(Figs. 1, 258, 264, and so on). Though 
other dendryphantines may have pale 
scales on the side of the face under the 
ALEs, in most these pale scales do not 
form a distinct band separated from the 
side bands by a dark area. Such a sep- 
aration of the cheek and side bands is 
also seen in some species of the mannii 
group (Figs. 493, 514, 534). It is lacking 
in other dendryphantines except one 
species from Venezuela examined, pos- 
sibly an Admirala sp., though the char- 
acter has not been surveyed intensively. 
Some species of Pelegrina (P. varie- 
gata. Fig. 447), however, do not have 
a distinct cheek band. 

6. An inverted V-shaped mark of pale 
scales on the forehead, contacting the 
AMEs (Figs. 1, 258, 264, and so on). 
Most Pelegrina males show this fore- 
head band, though some lack it (e.g., 
P. aeneola). While the character has 
not been surveyed thoroughly, it is lack- 
ing in other dendryphantines except for 
a few species (e.g., Tutelina hartii). 

7. Male courtship with prolonged 
"crouch" display. In the courtship of 
Pelegrina, Eris militaris, the Metaphi- 
dippus mannii group, and Nagaina in- 
cunda, there is a crouch display, in 
which the first legs are held low and 
forward, usually horizontal and below 



242 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



the level of the body (Figs. 125-128, 
134, 140, 162 for Pelegrina; Figs. 124, 
178, 184 for Eris and mannii group; 
Peckham and Peckham, 1889; Rich- 
man, 1982: fig. 3; Table 2, character 9). 
The more distal segments may be raised 
(e.g., Pelegrina kastoni, Fig. 140), but 
at least the femur is low. The male pro- 
ceeds toward the female in this pose, 
waving the first legs at low amplitude 
if at all. The body is held horizontal, in 
many species close to the substrate 
(hence the name "crouch"). This pose 
contrasts with that seen in most other 
dendryphantines (Table 2; Peckham 
and Peckham, 1889, 1890; Crane, 
1949a; Richman, 1982; Jackson, 1978; 
Hill, 1977a; and my own unpublished 
observations on about 50 species in 
which the first legs are generally raised 
and spread (the raisedspread display. 
Figs. 113-115, 121; e.g., Phidippus, 
Paradamoetas, Zygoballus) or raised 
and held forward (the raisedforward 
display. Figs. 118-120; e.g., Phanias, 
Sassacus papenhoei, Dendryphantes 
nigromaculatus) , or held horizontal and 
spread very wide (the lowspread dis- 
play. Figs. 116, 117; e.g., Messua lim- 
bata, Tulpius). The distinctions among 
these poses are, of course, vague. It 
should be noted that Pelegrina does 
have a raisedspread display, generally 
performed with the carapace held high 
and the abdomen low, but this is per- 
formed usually when the male is far 
from the female. A Pelegrina male thus 
often begins with a raisedspread display 
and then proceeds to a crouch display 
as he approaches the female. Insofar as 
other dendryphantines often reach with 
the legs parallel, forward, and low just 
before mounting and copulation (stage 
II courtship. Crane, 1949b), as in the 
crouch display, it may be claimed that 
the crouch display is merely stage II 
courtship and, thus, not restricted to Pe- 
legrina and relatives. Indeed, the crouch 
display may represent a prolonged stage 
II. If so, however, it is still distinctive 



from the brief stage II seen in other 
salticids, in being much more pro- 
longed, performed farther from the fe- 
male, and having a more rigid appear- 
ance. The crouch display has been ob- 
served in all Pelegrina and mannii 
group species examined, with the fol- 
lowing exceptions: in P. furcata a strik- 
ingly different display (the semaphore 
display) is seen, whereas in Pelegrina 
arizonensis, P. chalceola, and Meta- 
phidippus diplacis only a raisedspread 
display was seen. The lack of observed 
crouch display in these latter species 
calls attention to the difficulty of using 
the character. When seen, the crouch 
display is distinctive and can be con- 
sidered present. However, when not 
seen, it may still be characteristic of the 
species but not observed because the 
male simply failed to perform it, re- 
maining longer than usual in the raised- 
spread display. Nevertheless, the crouch 
display was scored as absent in these 
Pelegrina species because the males 
were observed for a reasonable sample 
of displays. Another problem with the 
character is the occurrence of a similar 
first leg pose in Tutelina and Hentzia 
(Figs. 122, 123). However, the exact leg 
poses and motions in these other genera 
differ in a number of respects from those 
of Pelegrina. 
8. Ridge under tibial apophysis (Figs. 78, 
389, 421, 427; Table 2, character 10). 
Under the tibial apophysis is a ridge 
that in extreme cases can make the 
apophysis appear bifid (e.g., Pelegrina 
bicuspidata). The ridge is present 
throughout Pelegrina (except fla- 
vipedes and similar species) but is also 
present in some species of the mannii 
group (mannii, diplacis), Beata (Fig. 
77), and in Tulpius hilarus. It is poorly 
developed in Eris militaris (Fig. 79), 
Bellota wheeleri, and Metaphidippus 
vitis. It is lacking in other dendryphan- 
tines including Dendryphantes (Figs. 
75, 76). 



Pelegrina Jumping Spiders • Maddison 243 



The preceding characters give reasonable 
support to the monophyly of the genus 
Pelegrina, though none provides a strict 
dehmitation. Even if the genus as consti- 
tuted here is not monophyletic, the char- 
acters provide good evidence that it is at 
least mostly monophyletic, to the extent 
that monophyly could probably be 
achieved by including or excluding only a 
few troublesome species. The following 
discussion regards the Pelegrina species 
that fail to show some of the characters 
supposedly delimiting the genus and why 
I have chosen to include these species in 
the genus. Whether or not species in the 
mannii group might be included in Pe- 
legrina is discussed in the introduction to 
the revision of the species of the mannii 
group. 

Pelegrina arizonensis and P. helenae. 
These species lack the two rami on the 
embolus. Instead, the embolus is blade- 
shaped, shifted retrolaterally, and concave 
on the exposed (ventral) surface (Figs. 217, 
218, 422, 428). However, there are a num- 
ber of other characters that would other- 
wise place the two species within Pelegri- 
na. These species have distinct cheek and 
forehead bands, a ridge just ventral to the 
tibial apophysis, and the male fang with 
wrinkles as in other Pelegrina. The pe- 
culiar embolus might be derived from that 
of a typical Pelegrina embolus by twisting 
about its longitudinal axis so as to reverse 
pro- and retrolateral edges and to present 
the embolus's concave surface, normally 
facing inward to the cymbium, outward 
toward the front. This is indicated by the 
presence of a ridge cutting across the face 
of the embolus joining the prolateral sur- 
face of the base with the retrolateral edge 
of the embolus, the position of the opening 
on the retrolateral side, the concave ex- 
posed face of the embolus, and a pro- 
nounced furrow on the embolis hemato- 
docha as if folded inward. Cutler and Jen- 
nings (1985) noted that "internal epigynal 
structure of his [Proszynski, 1982:1] 
D[endryphantes]. czekanowskii bears a 
close resemblance to the internal epigynal 



structures seen in the M. arizonensis 
group." Perhaps the similarity they no- 
ticed was the looping of the duct just inside 
the opening. The epigynum of D. czeka- 
nowskii is much like that of D. nigroma- 
culatus and D. fusconotatus. This looping 
in both Dendryphantes and the arizonen- 
sis group is related to the rotation of the 
epigynal flaps medially at the posterior end. 
In Dendryphantes, the rotation reaches 90°; 
in the arizonensis group, it is much more 
extreme, 180-270°. This rotation must not 
be considered a critical character; similar 
rotations are seen in the Pelegrina pervaga 
group (P. kastoni), in Phidippus octo- 
punctatus, and in Agassa (as compared to 
Sassacus). Indeed, in other respects the 
epigyna of the arizonensis group and Den- 
dryphantes are rather different, with the 
flaps being on the surface in the former 
(as in other Pelegrina, Fig. 251), whereas 
they descend beneath the opening as a sim- 
ple ridge in the latter (Fig. 65). 

Pelegrina flavipedes, P. flaviceps, and 
P. exigua. The biramous embolus of this 
group (Figs. 201-203) might be interpret- 
ed either as arising from an embolus like 
that of Pelegrina sabinema (Fig. 198) by 
more deeply splitting the two terminal rami 
of the embolus or as arising from an em- 
bolus like that of Dendryphantes rudis and 
D. nigromaculatus (Figs. 106, 108) by pro- 
longation of the retrolateral projection on 
the shoulder of the embolus. The fla- 
vipedes group lacks a ridge under the tibial 
apophysis, which otherwise seems char- 
acteristic of Pelegrina, thus supporting the 
interpretation that the flavipedes group 
does not belong with Pelegrina. However, 
there is more compelling evidence that the 
flavipedes group is derived from within 
Pelegrina. Like the pervaga group, the fla- 
vipedes group members are conifer dwell- 
ers with yellow chelicerae. Like Pelegrina, 
they have wrinkles anterior to the fang 
serrula and an embolic hematodocha bulg- 
ing distally, cheek bands on the male face, 
and a crouch display in male courtship 
(Fig. 127). Like Pelegrina and a few other 
dendryphantines, the embolic base is well 



244 BtiUetin Museum of Comparative Zoology, Vol. 154, No. 4 



sclerotized, with few wrinkles over most 
of its exposed surface. Finally, if the fla- 
vipedes group were derived from those 
Dendryphantes with long retrolateral pro- 
longations on the embolus shoulder (rudis, 
nigromaculatus) , the group should have 
the synapomorphy for Dendryphantes, the 
fold across the embolic base. The fla- 
vipedes group lacks this fold. 

Pelegrina furcata. This species has two 
terminal rami on the embolus, robust epi- 
gynal flaps, and cheek bands, but its court- 
ship display is unlike that of any others in 
the genus. The first legs are held wide and 
high, unlike Pelegrina, Eris militaris, and 
the mannii group, but like most other den- 
dryphantines, and waved in a distinctive 
semaphore-like fashion (Fig. 121). 

Pelegrina verecunda. Arizonan speci- 
mens lack the two distinct terminal rami 
on the embolus (Fig. 219), but Chihuahuan 
specimens identified as this species have 
the rami. 

Pelegrina tillandsiae. This species lacks 
the two terminal rami on the embolus (Fig. 
225) and is in many respects atypical. It is 
tentatively included in Pelegrina because 
its epigynum shows strong flaps that, as in 
Pelegrina, do not descend into the open- 
ings (Fig. 254). 

Pelegrina orestes and P. hunites. These 
species present the greatest problems with 
inclusion in Pelegrina, and I might have 
treated them as belonging to the mannii 
group or elsewhere. They lack the two ter- 
minal rami on the embolus characteristic 
of Pelegrina (Figs. 226, 227), though at 
least in P. orestes the embolus is obliquely 
truncated distal to the opening and has one 
ramus well separated from the opening. 
On the other hand, the epigynal flaps of 
hunites are flatter and narrower than in 
other Pelegrina, more as in the mannii 
group (Figs. 225, 481). Pelegrina hunites, 
though, lacks the bulge above the tibial 
apophysis characteristic of the mannii 
group and has three characters that might 
argue for the placement of hunites in Pe- 
legrina: the occurrence of wrinkles on the 
cheliceral fang, the bulging of the embolic 



hematodocha to the shoulder of the em- 
bolus, and the forehead band contacting 
the AMEs. On balance, then, a case can 
be made for tentatively describing hunites 
in Pelegrina. The situation with orestes is 
more difficult. Pelegrina orestes lacks the 
cheliceral fang wrinkles of Pelegrina, and 
one character, the presence of a ridge on 
the chelicera (Fig. 483), gives positive ev- 
idence to place orestes in the mannii group, 
though the ridge is especially weak and 
not present in all males. However, orestes 
lacks the bulge just dorsal to the tibial 
apophysis characteristic of the mannii 
group and does have the bulging embolic 
hematodocha characteristic of Pelegrina. 
Because of this, orestes will be described 
in Pelegrina, though it may eventually 
have to be moved. 

Natural History. Species of Pelegrina 
are found in various habitats from the Arc- 
tic to the tropical lowlands of Central 
America. While most species in Mexico 
and Central America appear to occur in 
the highlands (cloud forest and oak-pine 
zones), there are some lowland tropical 
species (P. sandaracina and P. yucate- 
cana). All species in the genus are pri- 
marily dwellers on foliage, being only oc- 
casionally found on the ground. Most other 
dendryphantines are also foliage dwellers, 
though some dendryphantines, in partic- 
ular those with more elongate and dully 
marked gray or brown bodies, are ground 
or bark dwellers (Terralonus, Ghelna, some 
Phanias species). A number of species of 
Pelegrina appear to be most common on 
or restricted to particular sorts of plants: 
the flavipedes group to various confiers, 
the pervaga group and P. halia to junipers, 
P. clemata and P. helenae to sagebrush 
(Artemisia tridentata), and P. tillandsiae 
to Spanish moss (Tillandsia usneoides). 
Other species do not appear so specialized 
to particular plants, yet in my collecting 
they do seem to prefer certain habitats: P. 
proterva occurs in forest understory, at least 
in the south of its range; P. galathea, in 
fields; P. variegata, in desert scrub; P. 
montana, in streamside vegetation; and P. 



Pelegrina Jumping Spiders • Maddison 245 



insignis, on low plants in fields and bogs. 
A number of southwestern species are most 
commonly collected from oaks. One gen- 
eralist species is P. aeneola, which is found 
on trees and herbs of various sorts in the 
Pacific Northwest, though not usually in 
the arid regions. The silken retreats and 
egg sacs are constructed among the foliage 
on which the adults are collected. 

PHYLOGENY WITHIN PELEGRINA 

While insufficient evidence was found 
to indicate the basal divisions of Pelegrina, 
the delimitation of a number of smaller 
groups can be made (see cladogram given 
in Fig. 129). One clearly delineated group 
is the flavipedes group, whose three mem- 
bers (flavipedes, flaviceps, and exigua) 
share the following characters derived 
within the genus: 

1. Embolus deeply bifid (Figs. 201-203). 
All other species of Pelegrina have the 
division between the terminal rami of 
the embolus not nearly so deep as in 
the flavipedes group. Other dendry- 
phantines have either a simple embolus 
or one with accessory rami different 
from those in Pelegrina and the fla- 
vipedes group. 

2. Chelicerae of male yellow with dark 
spot in medial concavity (Figs. 319, 324, 
329, 334). The dark concavity is unique 
to this group. 

3. Asymmetrical circling of palps in male 
courtship. During the crouch display, 
the palps are waved fairly slowly at high 
amplitude in circles such that as one is 
rising the other is falling (Fig. 127). 
Though other Pelegrina may wave their 
palps out of phase, in none examined 
is this asymmetrical waving made so 
obvious by the large slow waves. No 
other dendryphantines examined have 
such waving, except some Phidippus. 

Another group similarly well defined is 
the pervaga group, consisting of pervaga, 
sabinema, and kastoni, which share the 
following: 



1. Distinct markings on yellowish male 
palpus, consisting of prominent patches 
of white scales on the femur, tibia, and 
cymbium, interrupted by dark hairs on 
the patella. Other Pelegrina have no 
white scales on the tibia, or fewer on 
the tibia than on the patella. 

2. Cheek band extended posteriorly par- 
allel to the side band, separated from 
the side band by dense band of dark 
hairs (Figs. 304, 309, 314). This yields 
the appearance of white-black-white 
carapace side bands. Other Pelegrina 
have dark setae between side and cheek 
bands, but in none is the cheek band so 
horizontal and the dark hairs so dense. 
One species in the mannii group, Me- 
taphidippus emmiltus, has a superfi- 
cially similar pattern. 

These two groups, the flavipedes and 
pervaga groups, may together form a 
monophyletic group, as delimited by the 
following: 

1. Chelicerae yellow in males. Other Pe- 
legrina have dark brown chelicerae ex- 
cept southern males of P. tillandsiae. 
Brown chelicerae are present in other 
similar dendryphantines such as Eris, 
Nagaina, Beata, and the mannii group 
except Metaphidippus emmiltus, which 
also has yellow chelicerae. 

2. Conifer dwelling. All members of the 
flavipedes group, and P. pervaga and 
P. kastoni, are known to dwell more or 
less exclusively on conifers. The habitat 
of P. sabinema is unknown. This con- 
trasts with the habitat of most other 
Pelegrina, which inhabit broadleaf 
trees, shrubs, and herbs. However, P. 
proterva, P. aeneola, and P. furcata are 
known to frequent conifers, whereas P. 
halia appears restricted to conifers. Also, 
the polarity of this character is unclear. 
Outside the genus, Metaphidippus em- 
miltus is a juniper dweller, whereas Eris 
species are often collected from coni- 
fers. 

The neoleonis group, including tristis 



246 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



from Arizona and neoleonis from Mexico, 
is distinguished by the broad, dark rotated 
epigynal flaps. Though their appearance 
is much more Hke that of typical brown- 
legged Pelegrina, they share the following 
characters with the preceding two groups 
of yellow-legged species: 

1. Retrolateral ramus of embolus much 
elongate and curled to the prolateral 
(Figs. 196-203). The only other Pele- 
grina with such a long ramus is the 
furcata group, though in that group all 
but one species have it curling to the 
retrolateral. The exception is morelos, 
which has a prolateral curl (Fig. 215), 
but because it seems very close to fur- 
cata the prolateral curl is probably con- 
vergent. 

2. Embolus very broad. Such a broad em- 
bolus is not found in other Pelegrina 
except peckhamorum. 

3. First curve of duct of epigynum broad. 
Though unusual, a duct as broad is 
found in P. huachuca and P. morelos. 

The relationships of this proposed fla- 
vipedes-pervaga-neoleonis clade are not 
altogether clear, though the sickle-shaped 
retrolateral ramus of the embolus of pro- 
terva, galathea, edrilana, and pallidata 
may be viewed as a preliminary version 
of the very long ramus of this clade. Most 
of these species together with dithalea, 
proxima, and peckhamorum have an angle 
on the embolus just basal to the tip of the 
retrolateral ramus (Figs. 259, 265, 271, 277, 
283, 289, 310) which may be a synapo- 
morphy for a large clade (Fig. 129), but 
whether or not the angle is absent from all 
other Pelegrina is unclear (e.g., see Fig. 
215). 

Another clearly delineated group is the 
arizonensis group (Cutler and Jennings, 
1985), including the two species P. ari- 
zonensis and P. helenae. The apomorphies 
supporting the group are as follows: 

1. Epigynal flaps far rotated, at least 180° 
(Figs. 424, 430). No other dendryphan- 
tines known to me have a similar ro- 



tation; flap rotations in Terralonus and 
Ghelna, for example, are in the oppo- 
site direction. 

2. Erect portion of embolus displaced to 
retrolateral side and tegulum rotated 
somewhat clockwise (Figs. 422, 428). 
The embolus displacement is seen in 
some other dendryphantines, but the 
tegulum rotation is perhaps unique. 

3. Embolus twisted about longitudinal axis 
so as to reverse pro- and retrolateral 
edges and to present the back side for- 
ward. 

4. Ridge under tibial apophysis uniquely 
developed into flange (Figs. 421, 427). 

5. Markings of female abdomen some- 
what lineate (Figs. 425, 431). Lineate 
markings are also seen in P. tillandsiae 
and the Metaphidippus mannii group. 

The furcata group of Central America 
and southwestern North America includes 
the widespread and common P. furcata as 
well as a number of rare species (huachu- 
ca, morelos, bicuspidata, volcana, ochra- 
cea). Apomorphies supporting the group 
are as follows: 

1. Ridge under tibial apophysis unusually 
strongly developed, so as to form a sec- 
ond apophysis (Fig. 389; even stronger 
in the other species in the group). This 
is found in all species, although it is 
lacking in some specimens of furcata. 

2. Wrinkles on the retrolateral basal edge 
of the embolic base either transverse or 
ascending apically toward the retrola- 
teral (Figs. 390-394, 404, 406, 411, 416). 
This contrasts with the wrinkles of other 
Pelegrina and similar dendryphan- 
tines, which have wrinkles descending 
basally toward the retrolateral. Pele- 
grina insignis may have wrinkles sim- 
ilar to those in the furcata group. 

3. Epigynal flaps very convex, unlike oth- 
er dendryphantines except Pelegrina 
proxima and peckhamorum. 

4. Epigynum concave behind flaps, unlike 
the case in Eris, the mannii group, and 
Nagaina, though also seen in Pelegrina 
proxima, peckhamorum, and balia. 



Pelegrina Jumping Spiders • Maddison 241 



Pelegrina furcata itself has a very pe- 
culiar courtship display. Whether or not 
this feature is shared by other members of 
this group awaits their examination. 

The montana group includes three spe- 
cies: montana, msigriis, and chaimona. 
These species share the following: 

1. Concavity on back of embolus restrict- 
ed to distal half of erect portion. In 
other Pelegrina species, the concavity 
on the back of the embolus (Figs. 7, 34, 
35) extends from the base of the erect 
portion to its tip. Some Pelegrina, how- 
ever, have no clear concavity at all (e.g., 
P. aeneola, P. chalceola). 

2. Small, sharp denticles on front surface 
of embolus (Figs. 204-206). Other den- 
dryphantines have denticles on the em- 
bolus, but they are usually on the re- 
trolateral surface or are of different 
form. This character has not been well 
surveyed, however. 

Among the remaining species of Pele- 
grina are many that have a narrow em- 
bolus with small rami {aeneola, clemata, 
chalceola, halia, variegata, verecunda, 
sandaracina, and others). However, this 
may be the primitive condition for the 
genus. No clear subgroups were found 
among these species. 

IDENTIFYING SPECIES OF PELEGRINA 
AND THE METAPHIDIPPUS MANNII 
GROUP 

In general, the genitalia provide the best 
means of identifying species, but facility 
in recognizing the distinguishing features 
may require some experience. Males are 
much more easily identified than females, 
as the palpus and face markings provide 
more readily described and interpretable 
differences than the differences in epi- 
gyna. Take note especially of the width of 
the erect portion of the embolus and the 
size and orientation of the two terminal 
rami. Indeed, it is usually possible to iden- 
tify males simply by referring to the two 
pages of Figures 190-235. A single key for 
all species is given for males. 



Females, however, pose many more 
problems for identification. Though the 
abdominal markings and epigyna of fe- 
males vary in a number of features, the 
differences can be subtle and difficult to 
describe. One might think of the abdom- 
inal markings as falling in two major cat- 
egories: those in which the paired white 
spots dominate the dorsum (Figs. 263, 275, 
293, 382, 441, 451) and those in which the 
dark patches between and beside the white 
spots dominate the dorsum (Figs. 281, 287, 
347, 358, 364, 377, 387). Epigynal features 
to note are the topography of its surface 
and the size, convexity, color, and place- 
ment of the teardrop-shaped flaps cover- 
ing the openings. Even once experienced 
with these characters, an identifier can still 
have difficulties with some specimens. The 
problems are lessened within a given geo- 
graphical region. Because of this, separate 
female keys are given for five regions: the 
eastern United States and Canada, the 
Great Plains, Pacific Coast Untied States 
and western Canada, Arizona and Mexico, 
and Central America. Parts of the south- 
western United States are therefore with- 
out a key to females of Pelegrina, namely, 
Texas and the Rocky Mountain states. For 
Montana and Wyoming, the Pacific Coast 
key can be used (except possibly for prairie 
species). For Colorado, Utah, and New 
Mexico, most identifications can be accom- 
plished using the Pacific Coast and Ari- 
zona keys, though the Great Plains key will 
be needed occasionally. For Texas, the Ar- 
izona and Great Plains keys will usually 
suffice, but the eastern United States key 
will be needed on occasion. In general, 
mannii group females are not included in 
the keys. Metaphidippus mannii is in- 
cluded in the Pacific Coast key, but five 
other species in the group that occur in 
southern California are not included; man- 
nii and chera are in the Arizona key, but 
carmenensis is excluded. 

The keys are written for adult speci- 
mens, but the keys for females will be of 
some aid to identifying immatures based 
on markings. A key for immature Pele- 



248 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



grina from Minnesota has been given by 
Cutler (1981b). 

Figures 258-538 provide the most com- 
prehensive set of illustrations of the spe- 
cies, but important aid can be obtained 
from Figures 130-189, which show living 
specimens. Figures 190-235, which sum- 
marize the male emboli, and Figures 236- 
257, which show the surface topography 
of the epigyna. 

Key to the Males of All Species of 

Pelegrina and Those 

Metaphidippus mannii Group Species 

Occurring in the United States* 

1. Erect portion of embolus extremely nar- 
row or tapers to point, lacking two ter- 
minal rami (Figs. 226-235); lateral 
margin of chelicera usually with ridge 
near base of fang (Figs. 483, 493, 503, 
509, 514, 529); at least small patch of 
white or orange scales on chelicerae; 
western United States and southwest- 
ern Canada, south into Central Amer- 
ica {Metaphidippus mannii group, in 
part, and some Pelegrina) 2 

Erect portion of embolus in most species 
wide at tip and with two terminal rami 
(Figs. 190-225); chelicera lacking ridge 
near base of fang (e.g.. Figs. 258, 264); 
scales on chelicerae varied; widely dis- 
tributed 10 

2(1). Long patch of white scales on chelicerae, 
longer than V2 length of chelicerae 
(Figs. 478, 493, 514, 529) 3 

White or orange patch small (Figs. 483, 

503, 509) 7 

3(2). Embolus wide at base of erect portion 

(Figs. 226-231) 4 

Embolus very narrow (Figs. 516-523, 

535) '. 6 

4(3). Forehead white band lacking so that se- 
tae above AMEs are dark (Figs. 493, 
503); retrolateral edge of base of em- 
bolus with prolongation (Figs. 494, 499, 
505) 5 

White forehead band present and con- 
tacting AMEs above (Fig. 478); retro- 
lateral edge of base of embolus lacking 



* The mannii group species in Me.xico and Central 
America are not included; they can be distinguished 
from Pelegrina by their narrower embolus tip, which 
lacks the two rami. Nagaina incunda, described later, 
is brown and yellow striped with the first legs brown 
and the posterior legs yellow (Fig. 174). 



prolongation (Fig, 479); central Ari- 
zona south to Oaxaca .. 37. biinites (part) 

5(4). Carapace and abdomen shiny dark or 
coppery brown contrasting strongly 
with dense white cheek band and chel- 
iceral patches, with small or no white 
side bands (Figs. 178, 180, 493); pro- 
longation on retrolateral edge of base 
of embolus blunt or small (Fig. 494) 
where sympatric with diplacis; British 
Columbia to California east to Idaho 

and central Arizona 40. mannii 

Carapace and abdomen with more ex- 
tensive white side bands (Figs. 182, 
503); cheek band not distinct from side 
band (Fig. 503); prolongation of retro- 
lateral edge of base of embolus distinct 
and long (Figs. 504, 505); along Pacific 
Coast of southern California and Baja 
California 41. diplacis (part) 

6(3). Erect portion of embolus straight (Fig. 
233); face dark under eyes (Fig. 514) 

43. chera 

Erect portion of embolus curves ven- 
trally (Fig. 234); face extensively cov- 
ered with white scales (Fig. 529) ex- 
cept in Baja California Sur 

44. carmenensis 

7(2). Erect portion of embolus obliquely trun- 
cated, broad at base (Figs. 227, 484); 
carapace and abdomen dusted with 
beige to light brown scales (Figs. 172, 

483); Arizona and Mexico 

38. Orestes (part) 

Erect portion of embolus not so broad or 
truncated (Figs. 505, 510); markings 
dark brown with white (Figs. 182, 184, 
503, 509) or mostly yellow (Fig. 176) 
8 

8(7). Legs yellow, first legs fringed with white 
(Fig. 176); anterior median eyes ringed 
with red; chelicerae vertical and rel- 
atively weak (Fig. 534); erect portion 
of embolus very thin (Fig. 535); south- 
ern California to New Mexico 

45. emmiltus (part) 

Legs with dark brown markings (Figs. 
182, 184, 503, 509); anterior median 
eyes ringed with dark setae; chelicerae 
at least slightly divergent; embolus 
thick or thin; Pacific Coast of Califor- 
nia and Baja California 9 

9(8). Embolus wide at base of erect portion 
(Figs. 231, 504, 505); scales on chelic- 
erae white; body with bronze reflec- 
tions (Fig. 182); prolongation on re- 
trolateral edge of base of embolus large 
and distinct (Figs. 504, 505); southern 

California and Mexico 

41. diplacis (part) 

Embolus thin (Figs. 232, 510); scales on 



Pelegrina Jumping Spiders • Maddison 249 




Maps 1-7. Distributions of Pelegrina species. 1 . Pelegrina galathea in North and Central America. 2. P. proxima in the Caribbean. 
3. P. dithalea in Arizona. 4. P. edrilana in Mexico. 5. P. proterva in North America. 6. P. neoleonis in Mexico. 7. P. fnsf/s in 
Arizona. 



250 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 




Maps 8-15. Distributions of Pelegrina species. 8. P. peckhamorum in the eastern United States. 9. P. sabinema in the 
southwestern United States. 10. P. pervaga in the central United States. 11. P. kastonitn the southwestern United States. 12. 
P flavipedes in North America. 13. P. flaviceps in eastern North America. 14. P. exigua in the eastern United States. 15. P. 
clavator in Mexico. 



Pelecrina Jumping Spiders • Maddison 251 



chelicerae orange; body dark and dull 
(Fig. 184); prolongation on retrolateral 
edge of base of embolus small or absent 
(Fig. 510); central and northern Cali- 
fornia 42. tricolor 

10(1). Clypeus with patch of white or yellow 
scales between AMEs (not merely hairs 
overhanging chelicerae or circling an- 
terior median eyes) 11 

- Clypeus without white or yellow setae 

between AMEs, except perhaps for 
hairs surrounding anterior median eyes 

and overhanging chelicerae 17 

11(10). Chelicerae with large white or yellow 
patch of scales medially, at least for Vz 
length of chelicerae (Figs. 159, 437, 
442, 457); Mexico and Central Amer- 
ica _ _ 12 



i2(ii; 



13(12) 



14(11) 



15(14) 



16(14) 



17(10). 



18(17) 



Chelicerae lacking pale scales (Figs. 282, 
288, 319, 334); United States east of 
Rocky Mountains and Canada 14 

Retrolateral ramus of embolus much lon- 
ger than prolateral (Fig. 221) 

32. pallidata 

Retrolateral ramus of embolus small, 
subequal to prolateral (Figs. 220, 224) 13 

Pale markings usually yellowish; embo- 
lus small and tapers to tip (Fig. 224); 
lowland 35. sandaracina 

Pale markings white; embolus broadly 

truncate (Fig. 220); montane 

31. elevator (part) 

Chelicerae yellow (Figs. 319, 334); em- 
bolus deeply divided (Figs. 320, 330); 
cymbium lacks white scales; dwellers 
on conifers 15 

Chelicerae brown (Figs. 282, 288); em- 
bolus not deeply divided (Figs. 260, 
266); cymbium with dorsal patch of 
white scales; habitat varies 16 

First tibia yellow or with thin black stripe; 
retrolateral ramus of embolus thick, 
only slightly thinner than prolateral 
(Figs. 201, 320) 12. flavipedes 

First tibia dark; retrolateral ramus much 
thinner than prolateral (Figs. 203, 330) 
14, exigua (part) 

Embolus tapers to tip (Fig. 284), with 
long hooked retrolateral ramus (Fig. 
283) 5. proterva 

Embolus very broad at tip (Fig. 290), 
retrolateral ramus short (Fig. 289) 
6. peckha moru m 

Chelicerae yellow (Figs. 304, 309, 314, 
324, 329, 534); dwellers on conifer and 
Spanish moss 



Chelicerae brown (e.g.. Figs. 258, 264) 24 
Medial black spot on chelicerae (Figs. 
324, 329); embolus deeply divided 
(Figs. 325, 330); eastern and central 
United States and Canada _ 19 



- No medial black spot on chelicerae (Figs. 

304, 309, 314, 534); southern United 

States and northern Mexico 20 

19(18). Forehead flat (Fig. 329); forehead dark 
brow n in alcohol; bod\ and legs brown 
(Fig. 146); chelicerae yellow laterally 
(Fig. 329); southeastern United States 
north to Massachusetts and New York 

14. exigua (part) 

Forehead bulbous (Fig. 324); forehead 
yellow in alcohol; body and legs pale 
(Fig. 144); chelicerae with dark spot 
laterally (Fig. 324); northeastern Unit- 
ed States and southeastern Canada 

13. flavieeps 

20(18). Erect portion of embolus very thin (Fig. 
535); AMEs ringed with red; first legs 

fringed with white (Fig. 176) 

45. emmiltus (part) 

- Erect portion of embolus thicker (Figs. 

305, 310, 315, 474); AMEs ringed with 

white or brown; legs not fringed 21 

21(20). Cymbium yellow; band of dark setae un- 
der carapace side band (Figs. 304, 309, 
324); embolus wide at tip (Figs. 305, 
310, 315); dwelling on conifer; Kansas 

west to Arizona 22 

Cymbium dark distally; no band of dark 
setae under carapace side band (Fig. 
472); embolus tapers to narrow tip (Fig. 
474); dwelhng on Spanish moss, Flor- 
ida and North Carolina west to Texas 

36. tillandsiae (part) 

22(21). Clypeus brown (Figs. 304, 309); retro- 
lateral ramus of embolus long (Figs. 
198, 199, 305, 310); embolus broad at 

base of erect portion 23 

Clypeus with white band except cen- 
trally (Fig. 314); retrolateral ramus of 
embolus short (Fig. 200); embolus 
rectangular, narrow, and displaced re- 

trolaterally (Fig. 315) 11. kastoni 

23(22). Abdomen brown above; embolus (Fig. 

305) wider than in pervaga 9. sabinema 
Abdomen with central longitudinal pale 
stripe as in females (Fig. 313); embolus 
(Fig. 310) narrower than sabinema 

10. pervaga 

24(17). Ridge under tibial apophysis usually de- 
veloped into acute second apophysis 
(Fig. 389); wrinkles on embolic base 
transverse or ascending apically to- 
ward the retrolateral edge (Figs. 390, 
404, 406, 411, 416); southwestern 
United States to Panama (Jurcata 

group) 25 

At most small ridge or broad flange un- 
der tibial apophysis (Figs. 78, 421, 427); 
wrinkles on embolic base descending 
basally toward the retrolateral edge 



252 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 




Pelegrina Jumping Spiders • Maddison 253 



29 



26 



27 



(e.g.. Figs. 315, 438); widely distrib- 
uted _ 

25(24). Rami of embolus small and subequal 

(Figs. 213, 214) 

Retrolateral ramus of embolus long, much 
longer than prolateral (Figs. 212, 215, 
216) 

26(25). White patches on chelicerae extend to at 
least 1/2 their length (Fig. 403); embolus 
more or less straight (Figs. 213, 404); 
Panama 23. volcano 

- White patches on chelicerae small (Fig. 

405); embolus bent to retrolateral (Figs. 
214, 406); Guatemala and Mexico 
24. bicuspidata 

27(25). Retrolateral ramus curls to prolateral 

(Figs. 215, 411) 26. morelos 

Retrolateral ramus points retrolaterally 
or distally (Figs. 212, 216, 390-394, 
416) 28 

28(27). Retrolateral ramus points more distally, 
about twice as long as prolateral ramus 
(Figs. 212, 390-394); widely distrib- 
uted 22. fiircata (part) 

- Retrolateral ramus points more retrolat- 

erally, more than four times longer 
than prolateral ramus (Figs. 216, 416); 

southern Arizona 27. huachuca 

29(24). Large white patches on chelicerae at least 
to V2 their length (Figs. 437, 447, 452, 
478); embolus twists to tip; Arizona, 
Mexico, and Central America 30 

- Chelicerae with at most a small medial- 

basal patch of scales (e.g.. Figs. 258, 
348); embolus varied; widely distrib- 
uted geographically 33 

30(29). Embolus broad and truncated (Figs. 220, 
438); dorsum of abdomen mostly 
brown between side bands (Fig. 164); 
montane 31. clavator (part) 

- Embolus narrower, tapers to tip; dorsum 

of abdomen may have large white spots 

(Fig. 166); varied habitats 31 

31(30). Embolus tapers to narrow tip with ter- 
minal opening (Fig. 226); abdominal 
dorsum brown between white side 

bands (Fig. 170); montane habitats 

37. bunites (part) 

- Embolus tip wider, opening subterminal 

(Figs. 223, 224); abdominal dorsum 
with mixed pale and dark spots (Fig. 
166) as in female; deserts and tropical 

lowlands „ _.... 32 

32(31). Side and cheek bands fused (Fig. 447); 



chelicerae robust; embolus appears to 
taper in ventral view but in an oblique 
view the two small subecjual rami are 
easily seen (Fig. 222); abdomen with 
strong white spots (Fig. 166); arid 
regions of Mexico and Central Amer- 
ica _ 33. variegata (part) 

Side and cheek bands separate (Fig. 452); 
chelicerae narrower; embolus weaker 
with two rami not easily visible; ab- 
domen with unusual transverse bands 
(Fig. 169); seasonal tropical forests of 
Yucatan Peninsula 34. yucatecana 

33(29). Embolus with long retrolateral ramus 
(e.g., Figs. 196, 197, 209); western 

United States and Mexico _ 34 

Embolus with short retrolateral ramus 
(e.g.. Figs. 190, 206, 210), or rami not 
distinct (e.g.. Figs. 219, 225); widely 
distributed 38 

34(33). Retrolateral ramus curled prolaterally 
(Figs. 196, 197); embolus very broad 

at base 35 

Retrolateral ramus erect or pointing pro- 
laterally (Figs. 193, 209, 212); embolus 
narrower at base _. 36 

35(34). Prolateral ramus of embolus obtuse (Figs. 
196, 259); retrolateral ramus blunt and 
with bump (Fig. 295); embolus nar- 
rower at base than in iritis (Fig. 196); 

Mexico 8. neoleonis 

Prolateral ramus acute (Figs. 197, 300); 
retrolateral ramus sharp; embolus 

broader at base (Fig. 197); Arizona 

- 7. tristis 

36(34). Side and forehead bands on carapace re- 
duced or absent (Figs. 156, 365); west- 
ern United States and Canada 

19. aeneola (part) 

- Side and forehead bands on carapace well 

developed (Figs. 158, 276, 388); south- 
western United States, Mexico, and 

Central America 37 

37(36). Retrolateral ramus of embolus longer and 
diverging from prolateral (Fig. 212); 
widely distributed 22. jurcata (part) 

- Retrolateral ramus of embolus vertical 

(Fig. 193); embolus narrows abruptly 

near tip; central Mexico _ 

__ 4. edrilana (part) 

38(33). Erect portion of embolus arises on retro- 
lateral side (Figs. 422, 428); flange un- 
der tibial apophysis (Figs. 421, 427; 
arizonensis group) 39 



Maps 16-20. Distributions of Pelegrina species. 16. P. montana in North America. 17. P. insignis in Nortti America. 18. P. 
chaimona in Mexico and Arizona. 19. P. ciemata in western North Amehca. 20. P. balls and P. chalceola in western North 
America. 



254 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 




Map 25 



y Map 27 



Pelegrina Jumping Spiders • Maddison 255 



Erect portion of embolus arises centrally; 
ridge under tibial apophysis not de- 
veloped into flange (Fig. 78) 40 

39(38). Embolus tip sharp (Fig. 422); tibial, 
apophysis flange broad and short (Fig. 

421) 28. arizonensis 

Embolus tip blunt (Fig. 428); tibial 
apoph) sis flange narrow and elongate 
(Fig. 427) 29. helenae 

40(38). Chelicerae lacking pale scales 41 

Chelicerae with small patch of white or 
yellow scales 46 

41(40). Embolus narrows abruptly just basal to 
opening (Fig. 190); retrolateral ramus 
is small hook (Fig. 259); southern On- 
tario, eastern United States south to 

Central America 1. galathea 

Embolus parallel-sided or widens near 
tip; retrolateral ramus is small angle 
or apparently absent; Canada and 
mountainous areas of United States 42 

42(41). Retrolateral ramus of embolus longer 
than prolateral and leaning retrolater- 
ally (Figs. 208, 209); forehead band 

absent 19. aeneola (part) 

Both rami of embolus small; forehead 
band present or absent 43 

43(42). Embolus swollen near tip (Figs. 204, 344) 

15. montana 

Embolus with sides parallel or slightly 
tapering near tip (Figs. 207, 210, 211) 
_ 44 

44(43). Forehead band well developed and con- 
tacting AMEs (Figs. 152, 359); tegul- 
um with prominent prolateral bump 

(Fig. 361) 18. clemata (part) 

- Forehead band absent or if present then 
not contacting AMEs (Figs. 378, 383); 
tegulum with at most small prolateral 
bump (Figs. 379, 384) 45 

45(44). Cheliceral fang with flange (Fig. 378); 
carapace side bands broad (Fig. 378), 
embolus bends slightly (Figs. 210, 379); 
California and northern Arizona north 

to Washington 20. balia 

Cheliceral fang lacking flange (Fig. 383); 
side bands narrower (Fig. 383); em- 
bolus straight (Figs. 207, 211); south- 
ern Arizona to southern Illinois 

2 1 . chalceola 

46(40). Abdomen with striking lineate markings 
as in female (Fig. 477); embolus ta- 
pering to sharp tip in ventral view; 



living on Spanish moss 

36. tillandsiae (part) 

- Abdomen brown or spotted above; em- 

bolus usually broad at tip though var- 
ies; habitat varied 47 

47(46). Erect portion of embolus very thin (Fig. 
535); anterior median eyes ringed with 
red; first legs yellow fringed vvitli white 

(Fig. 176) 45. emmiltus (part) 

Erect portion of embolus thicker; eyes 
ringed with white or brown; legs not 
fringed 48 

48(47). Abdomen with paired black spots on 
brown dorsum; pale markings yellow- 
ish; embolus long and rectangular, 
leaning slightly retrolaterally (Figs. 
205, 349, 350) truncate at tip and with 
retrolateral ramus apparently absent 
(Fig. 205); Canada and northeastern 

United States 16. insignis 

Abdomen lacking distinct black spots; 
pale markings white or yellowish; em- 
bolus shorter (Fig. 219) or if long, then 
straight and with more prominent rami 
(e.g.. Figs. 192, 206, 207, 222) 49 

49(48). Abdomen with large and distinct paired 
white spots as in female (Fig. 166); side 
and cheek bands fused (Fig. 447); che- 
licerae robust (Fig. 447); erect portion 
of embolus parallel-sided and with two 
subequal rami (Fig. 222); deserts of 

Mexico and Central America 

33. variegata (part) 

- Abdomen with only small (Figs. 130, 132, 

152) or indistinct (Figs. 162, 172) 
paired pale spots, side and cheek bands 
separate; chelicerae not so robust; em- 
bolus varied 50 

50(49). Side bands of carapace and abdomen 
weak or absent and abdomen mottled 
(Figs. 162, 172); embolus small, lack- 
ing two distinct rami (Figs. 219, 227) 
51 

- Side bands well developed and abdomen 

not mottled (Figs. 130, 132, 152) 52 

51(50). Pale markings white or gray; erect por- 
tion of embolus widens gradually on 
prolateral Iside as it contacts basal por- 
tion (Fig. 433); tip of embolus rounded 
(Fig. 219) 30. vereciinda 

- Pale markings orange or tan; erect por- 

tion of embolus widens abruptly on 
prolateral side as it contacts basal por- 



Maps 21-27. Distributions of Pelegrina species. 21. P. aeneola in western North America. 22. P. furcata in Mexico and the 
southwestern United States. 23. P. furcata group members in Mexico and Central America. 24. P. huachuca in Arizona. 25. P. 
arizonensis and P. helenae in western North America (see Cutler and Jennings, 1985, for additional records). 26. P. verecunda 
in western North America. 27. P. tillandsiae in southeastern North America. 



256 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 




Pelegrina Jumfinc; Spidehs • Maddison 257 



tion so as to make a distinct corner 
(Fig. 484); tip of embolus pointed ret- 

rolateral to opening (Fig. 227) 

_ 38. orestes (part) 

52(50). Retrolateral ramus longer than prolater- 

al (Figs. 190, 193) 53 

Rami subequal (Figs. 191, 192, 206, 207) 54 

53(52). Embolus sides parallel below opening 
(Fig. 190); retrolateral ramus narrow 

(Fig. 259); widespread 

1. galathea (part) 

- Embolus inflated below opening (Fig. 
193); retrolateral ramus wide (Fig. 
277); central Mexico 4. edrilana (part) 

54(52). Abdomen with two central broken lon- 
gitudinal pale stripes in addition to side 
bands as in female (Fig. 269); Carib- 
bean 2. proxirna 

Abdomen lacking central longitudinal 
bands; western North America 55 

55(54). Embolus parallel-sided or tapers slightly 
to tip (Figs. 207, 360, 361); basal to 
opening the prolateral side is straight 
(Fig. 207); sagebrush of western Un- 
tied States and Canada 

_ 18. clemata (part) 

Embolus widens slightly near tip (Figs. 
192, 206); just basal to opening on pro- 
lateral side is angle (Figs. 192, 206); 
Arizona and Mexico 56 

56(55). Rami of embolus well separated (Fig. 
192); side bands have extensions join- 
ing between posterior eyes (Fig. 132); 
no denticles on exposed surface of em- 
bolus (Fig. 192) 3. dithalea 

Rami close together (Fig. 206); side bands 
without extensions; surface of embolus 
with denticles (Fig. 206) 17. chaimona 



Key to the Female Pelegrina of the 

Eastern United States and Canada 

(East of the Mississippi River and 

Manitoba) 

1. Posterior margin of epigynal flap truncated 
so as to be transverse, and standing high 
over surface behind it (Figs. 236, 262); 
epigynal flaps convex, parallel; legs dis- 
tinctly annulate (Fig. 131); abdomen 
marked with four pairs of prominent 
white spots with small black spots behind 
them (Figs. 131, 263) _ 1- galathea 



- Posterior margin of epigynal flap rounded, 

not transverse, or if transverse then flaps 
flat and flush with surface Ix'hind them 
(Figs. 238, 239, 241-245, 254); legs not 
distinctly annulate; abdomen w ith white 
spots smaller, often thinner and elongate 

(e.g.. Figs. 287, 323, 353) 2 

2(1). Abdomen with prominent paired black 
spots on orange-brown background (Figs. 
161, 353); epigynal flaps divergent (Fig. 
352); epigynal surface rises dramatically 
from low area around flaps to high pos- 
terior margin (Fig. 245); legs and face 
yellowish; mostly northern 16. insignis 

- Abdomen lacking prominent paired black 

spots though may have brow n or reddish 
patches; epigynal flaps parallel, conver- 
gent, or divergent; epigynal surface in 
most species with little relief (e.g., Figs. 
238, 241); legs and face varied; locality 
varied 3 

3(2). Epigynum with ridge just behind each flap 
(Figs. 244, 346); posterior notch often 
rectangular; body large and dark with 
very small paired white spots on dark 
abdominal dorsum (Fig. 347); Canada 

and mountains of United States _ 

_ _ 15. montana 

Epigynum lacking ridges behind flaps; pos- 
terior notch triangular; body smaller; ab- 
domen varied 4 

4(3). Abdomen strongly striped longitudinally 
yellow and brown (Fig. 477); epigynal 
flaps pale; living on Spanish moss of the 

southeastern United States 36. tillandsiae 

Abdomen not striped yellow and brown 
longitudinally, usually spotted; epigynal 
flaps varied _ - 5 

5(4). Epigynal surface and flaps very flat (Figs. 
24i-243); flaps not much darker than rest 
of epigynum except for narrow rim (Figs. 
322, 327, 332); carapace often with shiny 
scales and pale spot above and between 
anterior median eyes (e.g., Fig. 143); co- 
nifer dwellers {flavipedes group) 6 

Epigynal surface and flaps with more relief 
(Figs. 237, 238); flaps usually distinctly 
darker than rest of epigynum (Figs. 286, 
292); carapace lacking shiny scales .._ 8 

6(5). Forehead dark above and between .\MEs; 
head often bulbous; legs pale yellow, usu- 
ally with thin longitudinal dark lines on 



Maps 28-37 Distributions of species of Pelegrina. the Metaphidippus mannll species group, and Nagainaincunda^ 28. P. 
variegata in Mexico and Central America. 29. P. pallidata, P. yucetecana. and P. sandaracina in Mexico and Central America. 
30 Pbunites and P. orestes in Arizona and western Mexico. 31 . M. mannim western North Amenca. 32. M. tricolor n Califom a. 
33' M. diplacis in California and Baja California. 34. M. carmenensis in Mexico and the southwestern United States. 35^ M. 
ctiera in Mexico and the southwestern United States. 36. P. emmlltus in California and New Mexico. 37. Nagaina incunda in 
Mexico and Central America. 



258 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



femora; epigynum with second curve of 
spermathecal duct wider than in fla- 
vipedes but no so wide as in exigua (Figs. 
340, 341); northeastern United States 
bordering Canada and southeastern Can- 
ada 1 3. flaviceps 

- Forehead with pale spot above and be- 

tween anterior median eyes; head not 
bulbous; legs generally lacking longitu- 
dinal lines or if persent then wide and 
mostly on anterior legs; epigynum oth- 
erwise; distribution generally farther 

north or farther south than flaviceps 7 

7(6). Epigynal flaps parallel (Figs. 241, 322); sec- 
ond curve of spermathecal duct narrow 
and oblique (Figs. 321, 338, 339); cara- 
pace narrow; mostly northern (Canada 
and northern United States) though found 

occasionally on southern mountains 

12. flavipedes 

- Epigynal flaps convergent (Figs. 243, 332); 

second curve of spermathecal duct very 
broad and transverse (Figs. 331, 336, 337) 
carapace broader; mostly southern Unit- 
ed States north to Massachusetts and New 

York 14. exigua 

8(5). Abdomen marked with large square brown 
spots on or between paired pale spots 
(Figs. 2, 135, 287); epigynal flaps con- 
vergent and fairly flat, short (Figs. 238, 
286); surface rises quickly behind flaps 

to broad mound (Fig. 238) 5. proterva 

Abdomen uniformly light brown with small 
white spots (Figs. 137, 293); epigynal flaps 
long and fairly convex (Figs. 239, 292); 
surface rises gradually behind flaps to 

mound at posterior (Fig. 239) 

6. peckhamorum 

Key to the Female Pelegrina of the 

Great Plains (between the 

Rocky Mountains and the 

Mississippi River)* 

1. Epigynal flaps rotated 180° (Fig. 424); ab- 
domen with strongly lineate markings 
(Fig. 425) 28. arizonensis 

- Epigynal flaps rotated at most 45°; abdom- 

inal markings not so clearly lineate 2 

2(1). Legs distinctly annulate, and abdomen 
marked with four pairs of prominent 
white spots with small black spots behind 
them (Figs. 131, 263); epigynal flaps con- 
vex, parallel, posterior margin truncated 



so as to be transverse and standing high 
over surface (Figs. 236, 262) 1. galathea 

- Legs not distinctly annulate; abdomen with 

more prominent dark areas on either side 
of smaller pale spots (though pervaga with 
pale spots coalesced into single large spot); 
epigynal flaps varied, but posterior mar- 
gin not truncated 3 

3(2). Area behind epigynal flaps raised into high 
mound (Figs. 245, 246); carapace densely 
covered with white or yellow scales (Figs. 
153, 161 ) 4 

- Area behind epigynal flaps more or less flat 

(Figs. 238, 240); carapace thinly covered 
with pale scales (e.g.. Fig. 135) 5 

4(3). Scales on carapace white; legs beige and 
brown; abdomen with large dark patches 
on either side of central paired spots but 
lacking strong black spots (Fig. 364), an- 
teriormost pale spots fused into short lon- 
gitudinal bands; epigynal surface behind 
flaps raised into broad dark shiny round 
mound (Fig. 246); flaps convergent (Figs. 
246, 363); usually collected from sage- 
brush 18. clemata 

Scales on carapace yellowish; legs yellow; 
abdomen with paired black spots (Fig. 
353); epigynal surface behind flaps raised 
gradually but steeply into high mound 
along posterior of epigynum (Fig. 245); 
flaps divergent (Figs. 245, 352); low herbs 
in fields and bogs 16. insignis 

5(3). Abdomen marked with large square brown 
spots between small paired pale bands 
(Figs. 135, 287); epigynal flaps conver- 
gent and fairly flat, short and dark (Figs. 
238, 286); surface rises quickly behind 
flaps to broad mound (Fig. 238); cara- 
pace narrow; widespread (5) proterva 

- Abdomen with large central pale spot (Fig. 

313); epigynal flaps large and flat, fairly 
pale (Figs. 240, 312); carapace very wide; 
Kansas to Texas 10. pervaga 



Key to the Female Pelegrina of the 

Pacific Coast of the United States and 

Western Canada* 

Epigynal flaps rotated 270° so that flaps are 
transverse (Fig. 430); abdomen with li- 
neate markings (Figs. 155, 431); com- 
monly found on sagebrush 29. helenae 

Epigynal flaps rotated less than 45°; abdo- 



* Not included are some tree-dwelling species whose 
ranges reach into the Great Plains: Pelegrina fla- 
vipedes and exigua, which occur on conifers in the 
north and east; P. peckhamorum, on oaks in the south- 
east; and chalceola, in Texas to extreme southern 
Illinois. 



* Includes California, Nevada, Oregon, Washing- 
ton, Idaho, British Columbia, and Alberta. Included 
is Metaphidippus mannii, as well, but not the other 
mannii group species, which are restricted to the 
southern part of the area of the key. Not included is 
P. verecunda (see Arizona key). 



Pelegrina Jumping Spidkrs • Maddison 259 



men with markings not lineate except 

occasinally in cleniata, habitat varied 2 

2(1). Epigynal surface and flaps very flat (Figs. 
241, 256); body with shiny bronze or cop- 
per scales (Figs. 143, 179) 3 

- Epigynal flaps more convex and epigynal 

surface with more relief (Figs. 238, 244- 
248); flaps usually distinctly darker than 
rest of epigynum; body usually without 

metallic sheen; habitat varied 4 

3(2). Orange scales bet\\een and beside AMEs 
just above clypeus; body fairK smooth 
with shiny coppery scales (Fig. 179); usu- 
ally on oaks, holly, Arctostaphijlos, and 
other shrubs and trees with leathery leaves 
40. mann ii 

- White or dark scales around eyes; carapace 

often with pale spot above and between 
AMEs (e.g., Fig. 143); body with rougher 
appearance; on conifers 12. flavipedes 

4(2). Epigynal flap angled where flap bends down 
toward opening (Figs. 247, 374, 376); 
surface rises immediately behind flap to 
broad plateau covering posterior of epi- 
gynum (Fig. 247); carapace thinly cov- 
ered with white scales that often form 
an inverted T behind the AMEs (Fig. 
157); abdomen often with anterior me- 
dial paired spots coalesced into one large 
white spot (Figs. 157, 377); common on 

various plants including conifers 

19. aeneola 

Epigynal flaps not angled; surface of epi- 
gynum varied; carapace lacking T-shaped 
marking on head; abdomen with anterior 
medial paired spots separate (Figs. 347, 
364, 382); habitat varied 5 

5(4). Scales on carapace yellowish; legs yellow; 
abdomen with paired black spots (Fig. 
353); epigynal surface behind flaps raised 
gradually but steeply into high mound 
along posterior of epigynum (Fig. 245); 
flaps divergent (Figs. 245, 352); low herbs 

in fields and bogs 16. insignis 

Scales on carapace white, beige, or tan; epi- 
gynal surface behind flaps either more 
or less flat (Figs. 244, 248) or raised 
quickly behind flaps into mound (Figs. 
238, 246); habitat varied 6 

6(5). Area behind epig>nal flaps raised into a 
broad mound (Figs. 238, 246); flaps con- 
vergent; carapace with whitish scales 7 
Area behind flaps more nearly flat or con- 
cave (Figs. 244, 248); flaps divergent or 
convergent; carapace dark or covered 
with yellowish scales ^ 

7(6). Area behind epigynal flaps strongly raised 
into round dark shiny mound (Figs. 246, 
363); flaps convergent; carapace densely 
covered with white scales (Fig. 153); ab- 
domen with large dark patches on either 



side of central paired spots (Fig. 364), 
anteriormost pale spots fused into short 
longitudinal bands; commonly found on 
sagebrush 18. clemata 

- Area behind flaps only moderately raised 
into broad mound (Fig. 238); carapace 
not densely covered with white; abdo- 
men marked with large square brown 
spots between paired pale spots (Figs. 2, 
135, 287); found on various shrubs and 
trees 5. proterva 

8(6). Body dark, with very small pale spots on 
abdomen (Fig. 347); white scales be- 
tween AMEs; epigynum dark; surface 
rising immediately behind flap to ridge 
(Fig. 244); collected from waterside 

shrubs and trees 15. montana 

Body mottled beige and tan, with large 
yellowish spots on abdomen (Fig. 382), 
orange scales between AMEs; epigynum 
pale; surface rising gradually behind flaps 
(Fig. 248); juniper-dwelling 20. balia 

Key to the Female Pelegrina and mannii 
GROUP OF Arizona* 

1. Epigynal flaps thin and rotated 90°, lying 

in cavity (Fig. 317); markings gold and 
beige; junipers of southern mountains 

1 1 . kastoni 

Epigynal flaps rotated less than 60° (or, 
if rotated 90°, rarely in tristis, then 
flaps very broad); markings varied 2 

2(1). Epigynal flaps broad and flat (Figs. 302, 
307); epigynal surface more or less flat 

3 

Epigynal flaps narrower; epigynal sur- 
face varied 4 

3(2). Anterior end of epigynal opening deep, 
with the surface there pale and de- 
scending deeply under flap; flaps dark 

brown; southern Arizona 7. tristis 

Anterior end of epigynal opening shal- 
low, with the surface not so shallow 
nor descending so deeply as in tristis; 
flaps usually light brown; northern Ar- 
izona and New Mexico 9. sabinema 

4(2). Epigynal surface flat or convex behind 
flaps (Figs. 247, 252, 255-257); flaps 

often flat 5 

Epigvnal surface concave behind flaps 
(Figs. 248-250); flaps convex 14 

5(4). Epigynal flaps narrow and flat (Fig. 257), 
often transparent and difficult to see 



* Not included in the key are northern species that 
may occur in Arizona but have been at most rarely 
collected there; Pelegrina montana, flavipedes, in- 
signis, and clemata. Metaphidippus carmenensis is 
a species similar to chera with one known specimen 
from Arizona. It is not included in the key. 



260 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



(Figs. 525, 526); flaps divergent; com- 
mon in desert vegetation though oc- 
curs at higher elevations 43. chera 

- Epigynal flaps wider and more pig- 

mented, usually more robust than in 
chera, flaps parallel, convergent or di- 
vergent; generally found in oak-coni- 
fer habitats above 1,200 m elevation 6 
6(5). Epigynal flaps narrow, flat and mostly 
parallel e.xcept for sharp bend inward 
near posterior end (Figs. 255, 481); 
body vellow, sometimes with paired 
dark spots on abdomen almost as in 
insignis (Fig. 171) 37. bunites 

- Epigynal flaps without sharp posterior 

bend; if abdomen yellow then lacking 

large paired dark spots 7 

7(6). Abdomen very pale, yellowish, with 
markings consisting of little more than 
small dark speckles (Figs. 436, 487, 502) 8 

- Abdomen more darkly marked with 

brown or gray (Figs. 275, 358, 377, 
387 ) _.._ _ 1 

8(7). Epigynal flaps divergent and narrow (Fig. 

501) _ 40. mannii (part) 

Epigvnal flaps parallel or convergent 
(Figs. 435, 486) _ _ _ .... 9 

9(8). Epigynal flaps pale (Fig. 486) 38. orestes 

Epigynal flaps small and dark (Fig. 435) 

_ 30. verecunda 

10(7). Epigynal flap angled about midway along 
its length where flap bends down to- 
ward opening (Fig. 247); surface rises 
immediately behind flap to broad pla- 
teau covering posterior of epigynum 
(Fig. 247); carapace thinly covered 
with white scales that often form an 
inverted T behind the AMEs (Fig. 157); 
abdomen often with anterior medial 
paired spots coalesced into one large 
white spot (Fig. 157) 19. aeneola 

- Epigynal flaps not angled so abruptly in 

middle; epigynal surface varied; car- 
apace lacking T-shaped marking on 
head; abdomen with paired spots sep- 
arate 11 

11(10). Epigynal flaps divergent and narrow (Fig. 

501) _ 40. mannii (part) 

Epigynal flaps parallel or convergent, not 
so narrow (Figs. 274, 357, 386) 12 

12(11). Abdomen marked much as in galathea, 
with four pairs of prominent white 
spots with small black spots behind 
them (Figs. 133, 275); epigynal flaps 
short, fairly flat, and parallel (Fig. 274) 

3. dithalea 

Abdomen dark areas more prominent 
than paired white spots (Figs. 358, 387); 
epigynal flaps varied 13 

13(12). Carapace covered with reflective scales; 
abdomen brown with large paired 



darker brown spots (Fig. 387), setae 

around AMEs darkest dorsally 

21. chalceola (part) 

- Carapace covered with white scales; ab- 

domen with pale longitudinal side 
bands enclosing brown dorsum with 
paired white spots (Fig. 358), setae 

around AMEs all white . ._ _ 

_ 17. chaimona (part) 

14(4). Epigynal flaps dark, long, narrow, and 
convex (Fig. 418); epigynal surface 
with strong relief consisting of raised 
bumps just medial to each flap, a con- 
cavity behind flaps rising to posterior 
edge (Fig. 418); first curve of epigynal 

ducts broad and long (Fig. 417) 

_ 27. huachuca 

Epigynal flaps not so long, nor is epig\ nal 
surface so strongly sculptured; first 
curve of ducts narrower 15 

15(14). Epigynal flaps strongly convex (Figs. 249, 
250); posterior end rounded and stand- 
ing high abo\e surface (Figs. 249, 250); 
second curve of duct broad (Figs. 397, 
400) _ 22. fiircata 

- Epigynal flaps less convex (Figs. 248, 357, 

381, 386); posterior end not standing 
high above surface (Fig. 248); second 
curve of duct narrower (Figs. 356, 380, 

385 ) _ 1 6 

16(15). Abdomen marked with large round white 
spots (Fig. 382); carapace wide; epi- 
gynal flaps narrow and pale (Fig. 381) 
_ 20. balia 

- Abdomen with small white spots if any 

(Figs. 358, 387); carapace varied; epi- 
gynal flaps broader and shorter (Figs. 

357, 386 ) 1 7 

17(16). Carapace covered with brown reflective 
scales; abdomen brown with large 
paired darker brown spots (Fig. 387); 
setae around anterior median eyes 
darkest dorsally ..„ 21. chalceola (part) 

- Carapace covered with white scales, ab- 

domen with pale longitudinal side 
bands enclosing brown dorsimi with 
paired white spots (Fig. 358); setae 
around anterior median eyes all white 
17. chaimona (part) 

Key to the Pelegrina and Nag.mna 

FEMALES OF ME.XICO AND 

Central America* 

1. Body and legs mostly yellow (Fig. 163, 

173, 175, 436, 461, 487, 492), with small 



* Females of the Metaphidippus mannii groups 
are not included. These can usually be distinguished 
from Pelegrina females by their weaker epigynal 
flaps, which descend into the openings posteriorly. 



Pelegrina Jumping Spiders • Maddison 261 



dark markings if anv; epigvnum and 
flaps mostly flat (Figs, 252,' 253, 255) 2 

- Bod\ and legs well marked with brown 

and gray (e.g.. Figs. 298, 318, 409, 414, 
425); epigynal surface flat or more or 
less concave (e.g.. Fig. 250) 9 

2(1). First femur, patella, and/or tibia with 
small subterminal dark transverse bar 
(e.g., Peckham and Peckham, 1896: fig. 
10); clypeus covered with yellow scales 
except for barren patch beneath AMEs, 
beneath which on chelicera is dark line; 
epigynal flaps weak (Fig. 491); dis- 
turbed lowland habitats 

39. Nagaina incunda 

Legs uniform in color or if annulate, with 
dark annulae more extensive; clypeus 
densely covered with pale scales even 
below AMEs; habitat varied 3 

3(2). Epigynal surface more or less flat except 
for longitudinal ridge between flaps 
(Fig. 253); flaps convergent, narrow, 
only slightly convex (Figs. 253, 450) .. 
33. variegata (part) 

- Epigynal surface lacking central ridge 

(Figs. 252, 255); flaps varied 4 

4(3). Body and legs uniformly orange-yellow 
except sometimes for discrete small 
dark spots on abdomen; epigynum 
transparent so that spermathecae eas- 
ily visible without dissection (Figs. 460, 
463); flaps convergent; southern Mex- 
ico and Central America 

35. sandaracina 

Body and legs pale yellowish beige, not 
so orange; epigynum varied 5 

5(4). Abdomen uniformly yellowish, with 
small discrete dark speckles only (Figs. 

436, 487 ) ' 6 

Abdomen mostly yellow but any dark 
markings are larger spots and patches 
(e.g.. Figs. 171, 409, 446) 7 

6(5). Epigynal flaps pale, transparent (Fig. 

486), convergent _ _.... 38. orestes 

Epigynal flaps dark (Fig. 435), conver- 
gent to divergent 30. verecunda 

7(5). Epigynal flaps strongly convex (Fig. 408); 

epigynum concave behind flaps 

25. ochracea 

Epigynal flaps flat (Figs. 255, 482, 445); 
epigynal surface more or less flat 8 

8(7). Epigynal flaps narrow, with abrupt bend 
near posterior end (Figs. 255, 481); Ar- 
izona to Oaxaca 37. bunites (part) 

- Epigynal flaps wider, convergent, but 

without abrupt bend (Fig. 445); Chia- 
pas to Nicaragua 32. pallidata (part) 

9(1). Epigynal flaps rotated 180° (Fig. 424); 
abdomen with strong lineate markings 
(Fig. 425) 28. arizonensis 

- Epigynal flaps rotated at most 90°; ab- 



dominal markings not so clearly li- 
neate 10 

10(9). Epigynal flaps dark, wide, flat, and 
strongly convergent (Fig. 297); mon- 
tane 8. neoleonis 

Epigynal flaps not so dark and wide; hab- 
itat varied 1 1 

11(10). Epigynal flaps rotated 90° and in pits 
(Fig. 317); body yellowish (Fig. 141); 

northern Mexico, on junipers 

1 1 . kastoni 

Epigynal flaps rotated less than 60°; body 
varied; distribution varied 12 

12(11). Abdomen with peculiar transverse 
markings (Figs. 169, 456); fourth pair 
of spots in particular a transverse stripe; 
legs strongly annulate (Fig. 169); face 
thinly covered with pale scales; epi- 
gynal flaps pale and convergent (Fig. 
455); Yucatan Peninsula . 34. ijucatecana 

- Abdomen without such transverse mark- 

ings; fourth pair of spots not a trans- 
verse stripe; legs, face and epigynal 

flaps varied 13 

13(12). Epigynal flaps with abrupt bend near 
posterior end (Figs. 255, 481); epigyn- 
al flaps and surface more or less flat .. 
37. bunites (part) 

- Epigynal flaps without abrupt bend near 

posterior end; epigynal surface varied 14 
14(13). Epigynal surface more or less flat except 
for longitudinal ridge between flaps 
(Fig. 253); flaps convergent, narrow, 
only slightly convex; abdomen marked 
with large white spots (Figs. 167, 451) 

33. variegata (part) 

Epigynal surface usually rises to poste- 
rior edge; if flat then lacking longi- 
tudinal ridge; flaps varied; markings 

varied 15 

15(14). Epigynal flaps strongly convex (e.g.. Figs. 
236, 249, 250), parallel or slightly con- 
vergent 16 

- Epigynal flaps flat or only slightly convex 

(similar to those in Figs. 247, 252); may 

be strongly convergent _ 20 

16(15). Epigynal surface concave behind flaps 
(Figs. 249, 250), rising gradually to 

posterior margin (Jurcata group) 17 

Epigynal surface rises quickly behind 
flaps to mound covering most of pos- 
terior (Fig. 236) 19 

17(16). Epigynal flaps fairly short, pale (Fig. 408); 

southern Mexico and Guatemala 

25. ochracea 

Epigynal flaps generally longer, dark 
(Figs. 298, 413) 18 

18(17). First curve of duct narrow, second curve 
very broad (Figs. 397, 400); abdominal 
markings shiny, pale spots generally 
small (Figs. 396, 398, 402) 22. jurcata 



262 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



- First curve of duct wide, second curve 

narrow (Fig. 412); strong pale spots on 

dark abdominal dorsum (Fig. 414) 

26. morelos 

19(16). Epigynal flaps long and convergent, not 
truncate posteriorly (Figs. 4, 280); lon- 
gitudinal dark bands on abdomen 
prominent (Fig. 281) 4. edrilana 

- Epigynal flaps shorter, parallel, truncate 

posteriorly (Figs. 236, 262); abdomen 
marked with prominent white spots, 
without prominent dark bands (Figs. 

131, 263) 1. galathea (part) 

20(15). Epigynal flaps convergent, higher me- 
dially and tilted down laterally (Fig. 
440); epigynal surface high between 
and behind teardrops, lower lateral to 
this (Fig. 440) 31. clavator 

- Epigynal flaps not so tilted; epigynal sur- 

face flat or only slightly higher me- 
dially than laterally 21 

21(20). Epigynal flaps truncate posteriorly (Fig. 
236), high above surface at posterior 

end 1. galathea (part) 

Epigynal flaps not truncate posteriorly 
nor so high above surface 22 

22(21). Abdomen marked much as in galathea, 
with four pairs of prominent white 
spots with small black spots behind 
them (Figs. 133, 275); epigynal flaps 

fairly flat and parallel (Fig. 274) 

3. dithalea 

Abdomen with smaller white spots (Figs. 
358, 446); epigynal flaps usually con- 
vergent (Figs. 357, 445) 23 

23(22). Epigynum very flat (as in verecunda. Fig. 
252); dark band along inner margin of 
epigynal opening very wide (Fig. 444) 

32. pallidata (part) 

Epigynum with some relief; dark band 
along inner margin of epigynal open- 
ing narrow (Fig. 356) 17. chaimona* 

DESCRIPTIONS OF THE SPECIES OF 
PELEGRINA 

The Pelegrina species of Canada and 
the northern and eastern United States can 
be considered reasonably well known, but 
the same cannot be said for the species of 
Arizona, Mexico and Central America. In 
Arizona are many species, some poorly col- 
lected such as P. huachuca, P. chaimona, 
P. tristis, P. chalceola, and P. dithalea. 



* A number of unmatched females from Mexico 
may be P. chaimona or a species easily confused with 
it. 



Even if no additional species are discov- 
ered in Arizona, there is the danger that 
males and females of some of the known 
species have been mismatched. In Mexico 
and Central America, the situation is worse, 
where there are probably several species 
that will remain undescribed for some time 
to come. Already there are known some 
female Pelegrina from southern and cen- 
tral Mexico that apparently represent spe- 
cies not described here. I shall not give 
names to them here so as to avoid making 
more species names based on difficult to 
determine females and because with ad- 
equate collecting we may discover that 
they are females of already-described 
males. I do, however, give figures of some 
of them (Figs. 464-471). Figures 464-466 
show a single female from Neriaco, Mexico 
(state unknown), which may represent an 
extreme southern form of P. chalceola. 
Figures 467 and 468 show a form from 
Guerrero, Jalisco, and Michoacan that may 
be a southern form of P. dithalea. Figures 
469-471 show a form occurring in collec- 
tions from Durango. 

The descriptions follow a more or less 
consistent format except that occasionally 
a feature is noted in a few species that is 
not noted in any others: for instance, 
strongly annulate legs are noted under P. 
yucatecana, but leg annulation is usually 
not even mentioned, and in P. balia the 
flange on the cheliceral fang is noted but 
the fang is ignored in most other descrip- 
tions. In the case of leg annulation and 
male abdominal markings, the species 
should be assumed to be characterized by 
the usual Pelegrina condition (legs annu- 
late, but fairly indistinctly, and male ab- 
domen brown above, with at most small 
white spots, and ringed by white side 
bands) unless otherwise mentioned. In the 
case of the other characters, such as the 
flange in balia, the distribution of the fea- 
ture in all species is not fully known. Such 
a character is described to aid in separating 
the species from similar species that are 
known to lack it (in this example, chalceola 
lacks the flange). 



Pelecrina Jumping Spiders • Maddison 263 



Information on the labels of type ma- 
terial is cited, and, where possible, the au- 
thor of handwritten labels is identified. 
Banks, Chamberlin, Kaston, and Levi types 
still have with them the author's original 
labels, handwritten except those of Kaston, 
whose typewriter was distinctive. F. Pick- 
ard-Cambridge's and some of the Peck- 
hams' types no longer have their original 
labels. F. P. -Cambridge's labels have been 
replaced by labels handwritten in pencil, 
perhaps by Pocock or Browning (Levi, 
personal communication). Some of the 
Peckhams' labels were rewritten by Bry- 
ant, but most labels of Pelegrina types are 
apparently original. Some are in George 
Beckham's handwriting, but most are in a 
handwriting that is probably that of Eliz- 
abeth Peckham, for it occurs in other orig- 
inal labels in the Peckham Collection and 
in some of George Peckham's correspon- 
dence to Henshaw. 

1 . Pelegrina galathea 

(Walckenaer, 1837) 

new combination 
Figures 5, 10, 11, 13, 35, 78, 125, 
130, 131, 190, 236, 258-263; Map 1 

Attus galathea Walckenaer, 1805; 23 (cites Bosc's MS 
figure, pi. 1, fig. 4, 9) (nomen nudum). 

Attus galathea Walckenaer, 1837: 456, sp. 100. Type 
material lost or destroyed. Walckenaer (1837) cited 
Bosc's MS pi. 1, fig. 4, and also Abbott's fig. 405 
(9), but as Walckenaer (1805) refered only to Bosc's 
figure, this is to be taken as figure of type. Insofar 
as A. galathea is such a common and well-known 
species, and Bosc's ambiguous figure could be in- 
terpreted as another species, a NEOTYPE is here 
designated, 1<? in MCZ with label "NORTH CAR- 
OLINA: Raleigh, garden, 24-31 May 1943, Brim- 
ley." 

Attus nuhilis Hentz, 1846: 358, pi. 21, fig. 15, 9. Type 
material lost or destroyed. 

Euophrys leucophaea C. L. Koch, 1846: 216, fig. 1261, 
(3. Holotvpe 13 from Pennsylvania in ZMB with 
labels "E. Leucophaea ZMB'i794," "1794," "Hol- 
otypus," examined. Was dried; now rehydrated. 
NEW SYNONYMY. 

Icius crassiventer Keyserling, 1884: 503, fig. 11, 2. 
Holotype in MCZ 19 with labels "18 Icius crassi- 
venter Keys., 2 Massachusetts." and "18.", exam- 
ined. NEW SYNONYMY. 

Dendryphantes ornatus Banks, 1892: 75, pi. 4, fig. 
29a, pi. 5, fig. 29, 2. Holotype in MCZ 12 with labels 



"Dendryphantes ornatus Bks type," "Ithaca, N. Y." 
and "Nathan Banks Coll.," examined. 

Dendryphantes hondurensis: — G. & E. Peckham, 
1896, in part: 48, pi. 4, fig. 4a, 9. Type material in 
MCZ 22 from Belize labeled "449 Dendryphantes 
hondurensis Peck., Type, British Honduras 2 1423, 
G. W. & E. G. Peckham Coll." (in Bryant's han3^- 
writing) which both belong to the genus Gastromi- 
cans, and 15 29 labeled "461 Dendryphantes hon- 
durensis Peck., Guatemala, G. W. & E. G. Peckham 
Coll." (in Bryant's handwriting), of which IS 19 
are P. galathea and 19 is in the genus Messua, 
e.xamined. One Gastromicans 9 from Belize is here 
designated LECTOTYPE of D. hondurensis, and 
thus D. hondurensis is not properly a synonym of 
P. galathea. 

Metaphidipptis capitatus: — F. P. -Cambridge, 1901: 
272; Bonnet, 1957: 2810, in part. 

Metaphidippus digitatus F. P. -Cambridge, 1901: 269, 
pi. 24, figs. 12, 12a-c, S. Type material in BMNH 
Is and fragments of two other S labeled "Dendry- 
phantes digitatus, sp. n. Type S, Guatemala (Sarg.)" 
and 2S labeled "Dendryphantes digitatus, sp. n. S's, 
Mexico (Teapa) H. S", examined. NEW SYN- 
ONYMY. 

Beata digitata: —Simon, 1903: 841. Roewer, 1954: 
1007. Bonnet, 1955: 873. 

Dendryphantes capitatus: — G. & E. Peckham, 1909: 
469, pi. 38, fig. 5, possibly also pi. 36, figs. 4, 4a, 9. 

Metaphidippus galathea: — Chamberlin and Ivie, 
1944: 203. Kaston, 1973: 117, figs. 47-50, <59. 

Dendryphantes galathea: — Roewer, 1954: 1203. 

Notes on Synonymy. (1) I interpret 
Bosc's ambiguous figure (photograph of 
plates in MNHN Paris seen) as the species 
we call P. galathea, following recent us- 
age. Abbott's figure 405 probably shows a 
9 proterva. (2) Walckenaer's Attus atten- 
tus and Attus furtivus might also refer to 
this species. (3) The epigynum figured by 
the Peckhams for D. hondurensis was that 
of a P. galathea female, but despite this 
the female Gastromicans from Belize was 
chosen as lectotype because of the name 
hondurensis (suggesting British Honduras 
was intended as type locality), the label 
"Type," and because their figure 5 is not 
of P. galathea. Their description appears 
to apply to the mixture of species in the 
vials. (4) Kaston used the name nubilis for 
his numerous identifications of material 
around 1940. 

Diagnosis. A widespread species, for- 
merly confused with others in eastern 
North America, from which it is distin- 



264 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



guished by the embolus shape of males and 
speckled abdominal dorsum, annulate legs, 
and convex epigynal flaps of females. Car- 
apace wider than in most eastern species. 
Similar especially to the Caribbean prox- 
ima and southwestern dithalea. Can be 
separated from proterva by (males) the 
narrower embolus with smaller hook, 
darker face, and broader carapace and by 
(females) the abdominal markings and 
convex epigynal flaps. 

Male. Palpus (Figs. 190, 259, 260): Em- 
bolus rectangular, narrowing abruptly just 
basal to opening, with small pointed, 
curving hook at retrolateral tip (Figs. 190, 
259). Markings (Figs. 130, 258): Forehead 
band often well developed, with each 
branch forked and extending back to pos- 
terior eyes (Fig. 258). Cheek band weak. 
Clypeus brown, lacking central white spot 
on clypeus, with hairs overhanging chelic- 
erae dark except sometimes a few white 
hairs medially. White forehead band con- 
tacts AMEs dorsally 10:30-12:30. Chelic- 
erae lacking pale scales except in some 
southern males. Femur of palpus only 
slightly paler than more distal segments, 
cymbium dark brown and lacking white 
scales except in some southern males. Fem- 
ora of second, third, and fourth legs often 
more uniformly dark than in proterva, light 
brown base graduating to dark brown apex, 
though in some 63, especially in south, base 
abruptly pale. Abdomen shows trace of 
white spot pattern of 92. Measurements: 
Body length 3.0(3.4-3.8)4.0 mm; carapace 
length 1.4(1.6-1.8)2.0 mm, width/length 
0.77(0.79)8.1; n = 6<5 from Michigan and 
Georgia. 

Female. Epigynum (Figs. 261, 262): 
Flaps convex (Fig. 236), inner edges often 
parallel and close together, back edge of- 
ten perpendicular to body axis and stand- 
ing much higher than surface immediately 
behind it (Fig. 236). Surface rises fairly 
quickly behind flaps so that posterior sur- 
face is mostly raised, unlike the more con- 
cave surface of proxima and peckhamo- 
rum though not so uniformly high as in 
proterva. First curve of duct broad, but 



not so much as in proterva; second curve 
proceeds medially. Markings (Figs. 131, 
263): Carapace covered above with white 
to gray scales. Clypeus relatively thinly 
covered with white setae. Abdominal 
markings dominated by central pale spots 
each of which is shadowed by dark behind. 
Measurements: Body length 4.0(4.6)5.7 
mm; carapace length 1.7(1.8)1.9 mm, 
width/length 0.78(0.82)0.82; n = 59 from 
Georgia, Alabama, and Michigan. 

Chromosomes. 2nS = 26 acrocentrics + 
XXO {16 with full count plus 13 with only 
XXO observed, Toronto, Ontario). 

Courtship (73 observed from seven lo- 
cations: Rowan Co., Kentucky; San Jacin- 
to, Gonzales, and Hidalgo Co., Texas; San 
Luis Potosi: 99°42'W, 22°28'N; near Tux- 
pan, Veracruz; and north of Ciudad Ca- 
margo. Chihuahua). Raisedspread (n = 9, 
53). Crouch (Fig. 125; n - 14, 63): Body 
low (n = 5, 33) and horizontal (n = 14, 63). 
First legs held forward and horizontal (n 
= 12, 53), or slightly raised (n = 1), or raised 
about 45° (n = 1); bowed and touching or 
almost touching at tips (n = 9, 43), or 
straight forward (n = 4, 13), or slightly 
spread, though more parallel and lower as 
3 gets closer (n = 1). First legs flicker (n = 
12, 63) on each series (n = 4, 23) up and 
down (n = 4, 23) and alternately back and 
forth at tips (n = 1), vigorously (ca. 5 c/s) 
(n = 1) but at low amplitude (n = 5, 33). 
Palpi held down (n = 9, 53), either resting 
on first leg femora (n = 1), tucked beside 
chelicerae (n = 3, 13) or over chelicerae 
(n = 2, 13) and pointing inward (n = 4, 
23). Palpi waved (n = 10, 43) up and down 
(n = 3, 13) on each series (n = 6, 23) vig- 
orously (ca. 5 c/s) but at low amplitude (n 
= 1). Repertoires: 13 raisedspread only; 23 
crouch only; 43 raisedspread and crouch. 

Distribution (Map 1). Eastern North 
American north to southern Ontario, west 
to the Rocky Mountains, south to Florida 
and Costa Rica. 

Records. Many specimens, especially in MCZ and 
AMNH, from; CANADA: ONTARIO: Burlington, 
Hamilton, Port Credit, Windsor. UNITED STATES 
(county records): NEW HAMPSHIRE: Cheshire, 



Pelegrina Jumping Spiders • Maddison 265 



Hillsborough. Strafford: VERMONT: Windham; 
MASSACHUSETTS: Barnstable, Dukes, Essex, Mid- 
dlesex, Nantucket, Norfolk, Suffolk; RHODE IS- 
LAND: Newport; CONNECTICUT: Fairfield, Hart- 
ford, Litchfield, Middlesex, New Haven, Tolland; 
NEW YORK: Dutchess, Nassau, Suffolk, Tompkins, 
Wyoming; NEW JERSEY: Bergen, Cape May, 
Gloucester, Hunterdon, Middlesex, Morris; PENN- 
SYLVANIA: Adams, Berks, Bucks, Erie, Montgom- 
ery; OHIO: Ashtabula, Champaign; DELAWARE: 
Sussex; MARYLAND: Baltimore, Montgomery, 
Washington; DISTRICT OF COLUMBIA: Washing- 
ton; WEST VIRGINIA: Mercer; VIRGINIA: Alle- 
gheny, Botetourt, Fairfax, Suffolk, Surry, Portsmouth, 
Richmond, Washington; KENTUCKY: Rowan; 
TENNESSEE: Benton, Unicoi; NORTH CAROLL 
NA: Avery, Buncombe, Camden, Craven, Durham, 
Johnston, Macon, Mecklenburg, Nash, New Hanover, 
Pender, Transylvania, Wake, Washington, Yancey; 
SOUTH CAROLINA: Oconee, Orangeburg; GEOR- 
GIA: Chattahoochee, Clarke, Cobb, Glynn, Thomas, 
Ware; FLORIDA: Alachua, Escambia, Hillsborough, 
Indian River, Jefferson, Leon, Madison, Orange, Palm 
Beach, Pinellas, Polk, Putnam; ALABAMA: Baldwin, 
Colb, Coosa, Dallas, Mobile, Tallapoosa; MISSISSIP- 
PI: Harrison, Rankin; LOUISIANA: Baton Rouge, 
Caddo, Jefferson, St. Charles; MICHIGAN: Calhoun, 
Gratiot, Hillsdale, Jackson, Livingston, Micosta, Mid- 
land, Montcalm, Muskegon, Newaygo, Oakland, 
Washtenaw, Wayne; INDIANA: Clay, Howard, Mar- 
ion, Starke; ILLINOIS: Adams, Champaign, Peoria; 
MISSOURI: Berry, Boone, Jackson, Nevada, St. 
Charles, St. Louis, Vernon; ARKANSAS: Carroll, 
Conway, Hempstead, Lincoln, Washington; KAN- 
SAS: Bourbon, Cherokee, Jefferson, Riley; OKLA- 
HOMA: Cleveland, Kiowa, Payne; TEXAS: Aransas, 
Bexar, Brazos, Cameron, Comanche, Dallas, Denton, 
Galveston, Grayson, Harris, Hidalgo, Jim Wells, 
Karnes, Kleberg, Leon, Llano, McLennan, Nueces, 
Taylor, San Jacinto, San Patricio, Wichita; COLO- 
RADO: Boulder, Denver, Sedgwick; NEW MEXICO: 
Dona Anna, Rio Arriba, MEXICO: TAMAULIPAS: 
Santa Gracia, Reynosa; SAN LUIS POTOSL near Ciu- 
dad del Maiz; NUEVO LEON: Villa de Santiago; 
COAHUILA: Gloria; CHIHUAHUA: 21 km N of 
Ciudad Camargo, Delicias; VERACRUZ: just S of 
Tuxpan, Fortin; CHIAPAS: Tuxtla Gutierrez. GUA- 
TEMALA: Amatitlan, Capetillo. COSTA RICA: Chi- 
ral Paraiso, Cartago. BERMUDA: Grasmere. 

Natural History. In eastern North 
America, this species is generally found in 
sunlit places such as oldfields, in contrast 
to P. proterva, which is generally more of 
a forest dweller. In Chihuahua, P. galathea 
lives in riparian vegetation. Horner (1972) 
has investigated the bionomics and im- 
portance of P. galathea in biological con- 
trol in sorghum. Steiner and Greenstone 



(1991) examined segregation of isozyme 
markers in P. galathe^ 



lea. 



2. Pelegrina proxima 
(G. & E. Peckham, 1901) 
new combination 
Figures 191, 237, 264-269; Map 2 

Dendryphantes proxima G. & E. Peckham, 1901b 
(January; see G. & E. Peckham, 1909: 457): 327, 
pi. 28, figs. 3, 3a, 3$. Types in MCZ IS 19 2imm. 
"Dendryphantes proxima Pkm, 1901. Cuba Type. 
S 9." and "G. W. Peckham Coll." (label is original; 
handwritten, probably by Elizabeth Peckham), ex- 
amined. The type vial also contains one palpus of 
another species, perhaps Metaphidippus mannii, 
which is probably misplaced. 

Dendryphantes prudens G. & E. Peckham, 1901a 
(May): 15, pi. 4, figs. 13, 13a, 13b, $. Types in MCZ 
2S 19 with labels "1131 Dendryphantes prudens 
Peckhams, B.0155, Jamaica, Kingston 31423, 94123" 
(in George Beckham's handwriting) and "B.0155," 
examined. Roewer, 1954: 1199. 

Dendryphantes (Metaphidippus) proximus: — Pe- 
trunkevitch, 1911: 640. 

Pelegrina geniculata Franganillo, 1930: 45, fig. 17, 
9. Types from Sierra Maestra, Cuba, in lESC, orig- 
inally labeled only by a numerical code but 19 here 
designated as lectotype with labels "PF 548, " "Pe- 
legrina geniculata Franganillo, Lectotype, desig. 
W. Maddison 1990" (see comments regarding the 
generic name Pelegrina, earlier). 49 here desig- 
nated as paralectotypes, 3 deposited in lESC, and 
1 deposited in MCZ. Franganillo, 1936: 138,fig. 76. 
NEW SYNONYMY. 

Metaphidippus proximus: — Bryant, 1940: 501 (= 
prudens). Bonnet, 1957: 2817. 

Metaphidippus prudens: — Bryant, 1943: 496, figs. 56, 
57, 63, .59. Bryant, 1950: 189. Bonnet, 1957: 2817. 

Dendryphantes proximus: — Roewer, 1954: 1199. 

Notes on Synonymy. Bryant synony- 
mized prudens with proxima in 1940 but 
then, in 1943 and 1950, used the name 
prudens without explanation. The synon- 
ymy of Pelegrina geniculata is based on 
Franganillo's description and an exami- 
nation of all surviving specimens of the 
Franganillo collection, kindly sent to me 
from the lESC by Luis F. de Armas via 
Herbert Levi and Charles Dondale. The 
collection consists of 26 numbered vials 
containing at least 17 species (Table 4). 
The number and diversity of species rep- 
resented is approximately what might be 
expected from Franganillo's papers; thus. 



266 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



Table 4. Franganillo's collections of salticids. The identifications are by me (with vial number; 

E.G., PF 548, IN parentheses). 



Agobardus cubensis (Franganillo) sensu Bryant; 1 penultimale 3 22 (PF 546) 

Agobardus sp.: 12 (PF 551) 

Corythalia cf. arcuata sensu Bryant: 12 (PF 539) 

Corythalia cf. squamata Bryant: 2S (PF 540) 

Corythalia sp. (not C. arcuata sensu Bryant): 26 42, 1 imm. (PF 539), 26 12 (PF 540) 

Hentzia palmarum. \S (PF 543) 

Hentzia cf. tibialis: 16 

Hentzia sp.: 12 (PF 544), 12 (PF 544-2), 12 (PF 562), 1 imm. (PF 564) 

Lijssomanes antillanus. 22, 2 imm. (PF 535), 32 (PF 536), 56 12 (PF 542), 12, 4 imm. (PF 543) 

Lyssomanes sp., 1 imm. (PF 532) 

Menemerus bivittatus: 32, 1 imm. (PF 541), 1 penultimate 2 (PF 567), 1 penultimate 6, 12 (PF 568), 1<5 

(PF 569) 
Metacyrba taeniola. 22 (PF 575) 
Metacyrba sp., 12 (PF 575) 
Pelegrina proxima, 52 (PF 548), 1.? (PF 569) 
Nilakantha or Thiodina sp.: 1 penultimate 6 (PF 538) 
Nilakantha sp.: 12 (PF 544-2) 
Phidippus audax: 16, 1 penultimate 6 (PF 571) 
Platycryptus sp., 16 (PF 560) 

Plexippus paykulli: 26 (PF 534), 32 (PF 535), 12 (PF 566) 
Synemosyna smithii. 12 (PF 550). 



the collection may remain more or less 
complete. The collection lacks labels in- 
dicating locality or species (Alayon, 1982); 
thus, it is possible that we will never iden- 
tify the type specimens of Pelegrina gen- 
iculata with complete certainty. However, 
I will argue that Pelegrina geniculata is a 
junior synonym of Dendryphantes proxi- 
mus and that, in particular, the types are 
the females in vial PF 548. Franganillo's 
description is rather detailed in some re- 
spects, and a figure of the epigynum was 
provided. The described size, shape of the 
carapace, nature of the clypeus, chelicer- 
ae, sternum, eyes, and legs all fig proxima. 
The placement of Pelegrina in the Uni- 
dentati implies a single simple tooth on the 
retromargin of the cheliceral fang furrow, 
consistent with proxima and inconsistent 
with some genera such as Hentzia. The 
description of Pelegrina geniculata con- 
tains nothing that would rule out proxima, 
and several features that in particular point 
to this species. These are as follows: 

1. Leg spination: Franganillo's descrip- 
tion of the spination of the first and 
fourth legs can apply only to P. proxima 



2. 



among the species in Franganillo's col- 
lection. Table 5 lists species in his col- 
lection and their spination. The spina- 
tion of 3-3 on the tibia and 2-2 on the 
metatarsus described for Pelegrina 
geniculata narrows down the species to 
a dendryphantine, and the fourth tibia 
spination matches P. proxima exactly. 
Table 5 also lists a few other Cuban 
salticids not in Franganillo's collection. 
Among species known from Cuba but 
not listed in the table, Bryant's (1940) 
descriptions indicate that none have the 
leg spination described for Pelegrina, 
except Sidusa turquinensis, Icius ivick- 
hami, Phidippus spp., and Neon nigri- 
ceps, which can be ruled out as Pele- 
grina geniculata on other grounds. 
Though spination can sometimes be un- 
reliable (Maddison, 1987), spination dif- 
ferences such as those seen between 
dendryphantines and the other subfam- 
ilies listed in Table 5 are reasonably 
reliable. 

Epigynum: Franganillo's (1930) epi- 
gynal figure shows two dark teardrop- 
shaped objects and posterior notch. No 



Pelegrina Jumping Spiders • Maddison 267 



Table 5. Leg spination of Cuban salticids. Specimens marked by asterisk (*) are from Franganiro's 

COLLECTION. ALL SPECIMENS IN FRANGANILLO's COLLECTION ARE INCLUDED EXCEPT LYSSOMANES AND 

Synemosyna, whk:h are c:learly not Pelegrina by the description. The spination pattern of 
Pelegrina genicvlata is taken from Franganillo's (1939) description.^ 























— — - 






Meta- 




























tarsus 






First Pai 


r Tibia 






Metatarsus 






"ourth Pa 


ir Tibia 


Non- 




























termi- 




av 


pv 


al 


pi 


av 


pv 


al 


pi 


av 


pv 


al 


pi 


nal 


Pelegrina geniculata. 




























(Franganillo's description) 


3 


3 








2 


2 








2 


1 





2 


1 


Dendryphantinae 




























Pelegrina proxima* 


3 


3 








2 


2 








2 


1 





2 





Hentzia sp., 29* 


3 


3 








2 


2 








1 


1 











Phidippiis audax, 16 IpS* 


3 


3 








2 


2 








2 


1 


1 


2 


3-4 


Eris flava 


3 


3 








2 


2 








1 


1 


1 


1 


2 


Zygohallus sp. 


3 


3 








2 


2 








2 


1 


1 


1 


? 


Euophryinae 




























Agobardtts cubensis, 29* 


4 


4 








2 


2 


2 


2 


2 


1 


3 


3 


4 


Agobardus sp., 19* 


4 


4 


2 





3 


3 


1 


1 


2 


1 


3 


3 


4 


Corythalia sp. A, 29* 


2 


3 


2 





2 


2 


2 


2 


2 


1 


2 


2 


5 


Others 




























Habronattus spp. 


3 


2 








2 


2 








2 


1 


2 


3 


■? 


Marpissa pihei 


4 


4 








p 


? 


? 


? 


1 














Menemerus bivittatus, 39* 


4 


2 








2 


2 








1 


1 











Metactjrba taeniola, 29* 


2 











2 


2 























Metacyrba sp., 19* 


2 











2 


2 























Nilahantha sp.* 


2 


1 








2 


2 














1 


1 





Platycryptus sp* 


4 


3 








2 


2 








1 


1 





1 





Plexippns payhnlli, 39* 


4 


3 








2 


2 








2 


1 


3 


3 


7 



^ Abbreviations: av, pv, al, pi, anterior and posterior of ventral and lateral; p3, penultimate male. 



epigynum from Cuba known to me 
would have this appearance except that 
of P. proxima, whose long epigynal flaps 
and posterior notch are unique on Cuba. 
Interpreting the teardrop-shaped ob- 
jects as spermathecae or other struc- 
tures cannot help the figure to be ap- 
plied to any other Cuban species known 
to me. Certainly no euophryines have 
epigyna like this, nor any of the other 
dendryphantines known from Cuba. 
The only difficulty with considering the 
epigynal figure conclusive is that in 1936 
the same figure was published inverted 
as figure 76, 2. If the 1936 orientation 
were correct, then the epigynum would 
not apply to P. proxima, but it would 
also not apply to any other salticid 
known to me. 
3. Abdominal markings: Franganillo de- 
scribes a series of longitudinal bands. 



Centrally, there is one band described 
as "leonada." L. Aviles (personal com- 
munication) suggests that this term may 
mean "the color of a lion," namely, light 
brown. According to the description, 
this central band is flanked by pale 
bands, which are flanked by more light 
brown bands. The pale bands are 
toothed and have one particularly large, 
oblique, curved tooth near the spinner- 
ets. This describes P. proxima females 
exactly (Fig. 269), the large teeth being 
the fourth pair of spots (see Fig. 2). 
Other Cuban salticids that have some- 
what similar markings are Platycryptus 
sp., Menemerus bivittatus, and some 
of the Agobardus species, but their 
markings are not exact matches and 
these species can be ruled out on other 
grounds such as size and spines. 
4. Specimens cited: Vial PF 548 contains 



268 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



several adults, all females of P. proxi- 
ma. The description notes the material 
being six females; the vial contains five 
females. No other vial in the collection 
contains so many females without ac- 
companying males or immatures. 

This evidence taken together indicates that 
the description of Pelegrina geniculata 
applies to P. proxima and in particular to 
the females in vial PF 548. Accordingly, 
one of these females has been designated 
as a lectotype of Pelegrina geniculata. 

Diagnosis. The only known Caribbean 
Pelegrina, differing from the similar gal- 
athea in having more lineate abdominal 
markings and in details of genitalia. 

Male. Palpus (Figs. 191, 265, 266): 
Embolus rectangular, not narrowing so 
abruptly near the opening as in galathea; 
retrolateral ramus an angle not prolonged 
into a hook (Figs. 191, 265). Markings (Fig. 
264): Cheek band very weak. Clypeus 
brown, with hairs overhanging chelicerae 
dark with a few white medially. White 
forehead band contacts AMEs dorsally 
10:30 to 12:30. Chelicerae with small me- 
dial patch of pale scales. Cymbium usually 
lacking white scales. Abdomen often show- 
ing a trace of the longitudinal white bands 
of females. Measurements: Body length 
2.8(3.0)4.2 mm; carapace length 1.5(1.6)2.1 
mm, width/length 0.78(0.79)0.80; n = 56 
from Havana, Cuba. 

Female. Epigynum (Figs. 237, 267, 268): 
Flaps long, fairly convex, dark, not trun- 
cated behind as in galathea. Surface be- 
hind flaps more or less concave, rising 
gradually so that posterior mound restrict- 
ed to guide area or absent (Fig. 237). First 
curve of duct narrow; second curve pro- 
ceeds medially. Markings (Fig. 269): 
Carapace covered above thinly with white 
scales. Clypeus densely covered with white 
scales. Abdominal white spots arranged into 
two median longitudinal white bands on 
brown background. Measurements: Body 
length 3.5(3.7)4.8 mm; carapace length 
1.5(1.6)1.9 mm, width/length 0.76(0.76) 
0.79; n = 59 from Havana, Cuba. 



Distribution (Map 2). Known from the 
larger Caribbean islands. 

Records. BAHAMAS: Grand Bahama Island, Free- 
port (19, AMNH); Rum Cay, near Port Nelson (25, 
AMNH). CUBA: Havana (many 59, MCZ); Havana: 
Santiago de las Vegas (63 59, MCZ); Marianao Habana 
(29, AMNH); Soledad, Cienfuegos (5<5 49, MCZ, 
AMNH); Oriente: Santiago de Cuba (19, MCZ); Trin- 
idad Mtns., Hanabanillo Falls (15, MCZ); Holquin 
(59, MCZ); Banes (19, MCZ); 7 km N of Vinales (25 
19, AMNH); Vega Alta, Santa Clara (45, AMNH); San 
Vicente, Pinar del Rio (45 29, AMNH). JAMAICA: 
Christiana (19, AMNH); Claremont (15, AMNH); 
Spanish Town (29, MCZ); St. Andrew: Mona (55 19, 
MCZ); St. Andrew: Liquanea (19, MCZ); St. Ann, 1.6 
km E of Moneague (19, MCZ). DOMINICAN RE- 
PUBLIC: S. O. de las Matas (15, MCZ); La Vega (19, 
MCZ); Ciudad Trujillo (19, MCZ). HAITI: Diquini 
(15, MCZ); Enerv, Bata (15, AMNH); hills nr. Port- 
au-Prince (15, MCZ); Quest (15 19, MCZ). 

3. Pelegrina dithalea new species 

Figures 132, 133, 192, 270-275; Map 3 

Holotype male and paratype female in MCZ, with 
label "ARIZONA: Santa Cruz Co., Sycamore Can- 
yon, ca. 9 mi [14 km] W of Peiia Blanca Lake, W 
of Nogales, ca. 4000 ft. el. [1,220 m], 19 Jun 1985 
W. Maddison 85-060, sweeping in canyon where 
stream flowing." 

Etymology. An arbitrary combination 
of letters, to be treated as a noun in ap- 
position. 

Diagnosis. Similar in markings to gal- 
athea, from which it differs by the em- 
bolus that lacks the hooklike retrolateral 
ramus and that widens toward the tip. Em- 
bolus resembles that of chaimona, but the 
rami are farther apart (Figs. 192, 206). 

Male. Palpus (Figs. 192, 271, 272): Em- 
bolus widens slightly from near base to 
near tip. Rami subequal, though retrolat- 
eral is more prominent. Markings (Figs. 
132, 270): On carapace, white bars from 
side bands to fovea usually strong and fused 
into inverted V mark. Cheek band weak. 
Clypeus brown, hairs overhanging chelic- 
erae white centrally, dark laterally. White 
forehead band contacts AMEs dorsally 
10:30-12:30. Chelicerae with small medial 
patch of white scales. Cymbium with few 
white scales. Legs fairly distinctly annu- 
late. Abdomen shows traces of white spots 



Pelegrina Jumping Spiders • Maddison 269 



of female. Measurements: Body length 
3.6(4.0)4.2 mm; carapace length 1.9(2.0)2.1 
mm, width/length 0.77(0.78)0.81; n = 53 
from Sycamore Canyon, Arizona. 

Female. Epigynum (Figs. 273, 274): 
Flaps slightly convex; posterior edge not 
standing so high above surface behind them 
as in galathea. Surface rises immediately 
into gentle mound covering all of poste- 
rior. Second curve of duct proceeds me- 
dially. Markings (Figs. 133, 275): Cara- 
pace covered by scales mostly gray-white, 
and some brown scales around fovea, and 
just medial to posterior eyes. Clypeus 
densely covered with white scales. Abdom- 
inal markings gray-brown with large white 
spots and small dark spots, much like gal- 
athea (Fig. 275). Measurements: Body 
length 3.9, 4.3, 5.3 mm; carapace length 
2.0, 2.1, 2.1 mm, width/length 0.75, 0.77, 
0.79; n = 32 from Santa Cruz and Pima 
Counties, Arizona. 

Male /Female Matching. The two sexes 
were co-collected in Sycamore Canyon and 
Kitt Peak, have similar markings on the 
abdomen, and are similar in markings and 
form to galathea. 

Courtship (2<? observed from Sycamore 
Canyon, Arizona). Crouch (n = 8, 26): Body 
low and horizontal (n = 2, IS). First legs 
fairly wide to bowed and parallel (n = 6, 
2(5), low (n = 8, 2(5) to raised a bit (n = 4, 
1(5), waved on series (n = 8, 26) at low 
amplitude (n = 4, 26). Palpi held down (n 
= 2, 13), waved up and down on series so 
as to drum on substrate (n = 2, 13), still on 
pause (n = 2, 13). 

Distribution (Map 3). Southern Arizona. 

Records. UNITED STATES: ARIZONA: Pima Co.: 
Quinlan Mtns., picnic area nr. Kitt Peak Observatory, 
1,900-2,000 m el, 20 June 1985 (33 12, MCZ); Santa 
Cruz Co.: Sycamore Canyon, ca. 14 km W of Pena 
Blanca Lake, ca. 1,200 m el., 19 June 1985 (123 19, 
MCZ); Santa Rita Mtns., Madera Canyon, nr. Bog 
Springs Cmpgd., ca. 1,500 m el., 17 June 1985 (12, 
MCZ), 

Natural History. In oak woodland at all 
three Arizona localities. At Sycamore Can- 
yon, beating vegetation, especially shaded, 
deep in canyon where stream still flowing 



in June. At Kitt Peak, beating oaks and 
other shrubs and trees. 

4. Pelegrina edrilana new species 
Figures 4, 193, 276-281 ; Map 4 

Holotype male with one immature in AMNH with 
label "MEXICO: Tlalpam, D.F. [Distrito Federal], 
Apr. 17, 1946, J, C. Pallister." 

Etymology. An arbitrary combination 
of letters, to be treated as an adjective. 

Diagnosis. An enigmatic species from 
southcentral Mexico with a palpus in some 
ways resembling each of galathea, proter- 
va, and peckhamorum. The swollen base 
of the erect portion of the embolus is wider 
than in galathea, thought not so extreme 
as in proterva. The retrolateral ramus is 
wider than in galathea, though not so long 
and hooked as in proterva. 

Male. Palpus (Figs. 193, 277, 278): Em- 
bolus swollen at the base of the erect por- 
tion; narrowing distally near opening. 
Retrolateral ramus extended into short stout 
hook (Figs. 193, 277). Markings (Fig. 276): 
Cheek band fairly dense but not so dense 
as proterva. Clypeus brown; hairs over- 
hanging chelicerae dark except for few 
white medially. White forehead band con- 
tacts AMEs dorsally 10:30-12:30. Chelic- 
erae with narrow medial patch of pale 
scales from base to V2 length. Cymbium 
lacking white scales. Measurements: Body 
length 3.7(3.7-3.8)3.9 mm; carapace length 
1.7(1.8)1.9 mm, width/length 0.72(0.76) 
0.78; n = 53 from Distrito Federal and 
Durango, Mexico. 

Female. Epigynum (Figs. 4, 279, 280): 
Flaps long and convex, turning slightly in- 
ward, shiny and generally pale. Surface 
rises to mound quickly behind flaps; in 
many specimens the mound has two dis- 
tinct front corners (Fig. 280). Females 
from Oaxaca (Figs. 4, 279), which may 
represent a distinct species, have some- 
what longer epigynal flaps and a gentler 
mound on the posterior surface. First curve 
of duct wide; second curve proceeds 
obliquely anteriorly. Markings (Fig. 281): 
Carapace covered with white scales. Clyp- 



270 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



eus densely covered with white scales. Ab- 
domen marked somewhat as proterva, with 
brown background having white side bands 
and central spots. Measurements: Body 
length 3.4(4.8-5.2)5.4 mm; carapace length 
1.7(1.9)2.0 mm, width/length 0.75(0.77) 
0.80; n = 79 from Distrito Federal, Mexico. 

Male/Female Matching. Males and fe- 
males were co-collected and have a com- 
mon distribution; they have similar size 
and markings. 

Distribution (Map 4). Mexico. Most 
specimens from the Distrito Federal; also 
known from Durango, Oaxaca, and San 
Luis Potosi. 

Records. MEXICO: SAN LUIS POTOSI: Guana- 
juato border on Hwy 57, ca. lOOMS'W, 2r35'N (19, 
MCZ); DURANGO. Pales Colorados, 5 August 1947 
(1<3 19, AMNH); DISTRITO FEDERAL: Contreras, 
2,500 m, 23 July 1947 (29, AMNH); Mexico City, 
January 1941 (1<J, AMNH); Pedregal, 8 August 1947 
(is, AMNH); San Jeronimo, 11 June, 21 June, and 1 
July 1946 (2<5 89, AMNH); Tlalpam, 2,300 m, 21 July 
1947 (19, AMNH); 17 April 1946 {IS, AMNH); OA- 
XACA: 2 km S of El Tule, 1,500 m el, ca. 96°40"W, 
17°02'N (39, MCZ). 

Natural History. Collected from Acacia 
savannah in Oaxaca and from mesquite in 
San Luis Potosi. 

5. Pelegrina proterva (Walckenaer, 1837) 

new combination 
Figues2, 3, 6-9, 15,34, 134, 
135, 194, 238, 282-287; Map 5 

Attus protervus Walckenaer, 1837: 443. Type 15, lost, 
shown in figure 402 of Abbot (1792), whose caption 
reads "Taken 8th June, in a Dirt daubers Nest the 
only one I have seen." 

?Attus capitatus Hentz, 1845: 200, pi. 17, fig. 15, S. 

?Attus octavus Hentz, 1846: 365, pi. 22, fig 15, 9. 

Attus aestivalis G. & E. Peckham, 1883: 2, figs. 2, 
2a-c, 59. Types lost or destroyed (Bryant, 1941). 

Dendryphantes capitatus: — G. & E. Peckham, 1909: 
469, pi. 36, figs. 4b, c, pi. 38, fig. 5a, possibly also 
pi. .36, figs. 4, 4a, <39. Roewer, 1954: 1202. 

Dendryphantes atopodon Chamberlin, 1925a: 234. 
Holotype in MCZ 16 with label "Dendryphantes 
atopodon Chamb., S holotype, Va.: Scott's Runn, 
July, R. V. Chamberlin, Coll.", examined. Roewer, 
1954: 1206. Bonnet, 1956: 1392. 

Metaphidippus protervus: — Chamberlin and Ivie, 
1944: 204 (not fig. 23). Kaston, 1973: 117, figs. 43- 
46, (59. 



Metaphidippus capitatus: — Bonnet, 1957: 2810, in 
part. 

Notes on Synonymy. (1) Walckenaer's 
first description of Attus protervus (p. 443, 
after Abbot's fig. 402) probably refers to 
this Pelegrina; his second (p. 465, after 
Abbot's fig. 463) probably to Maevia in- 
clemens (see Walckenaer, 1837: 425; 
Chamberlin and Ivie, 1941: fig. 23). Attus 
attentus Walckenaer (species number 61, 
Abbot's fig. 157) may also be P. proterva. 
Attus capitatus Hentz, though considered 
a synonym of proterva by Chamberlin and 
Ivie (1941), might equally well be P. gal- 
athea, Eris militaris, or another species. 
(2) Euophrys concolor Banks was synon- 
ymized with P. proterva by Edwards 
(1980), apparently because one of the two 
specimens in the type vial is a 2 F. pro- 
terva. Banks' description clearly refers to 
the other specimen (by color, length of 
fourth leg, and epigynum with single 
opening and two posterior circles), which 
is therefore the holotype. This holotype is 
of the species now called Sitticus cursor 
Barrows, which should henceforth be called 
Sitticus concolor (NEW COMBINA- 
TION), with the name cursor relegated to 
synonymy (NEW SYNONYMY). Since 
Banks refers to only a single female, the 
proterva 9 was probably added to the vial 
subsequently. (3) Kaston used the name 
octavus for his numerous identifications of 
material around 1940. 

Diagnosis. The white face markings of 
males, abdominal markings of females, and 
the genitalia distinguish this species, which 
is abundant in woodlands throughout much 
of Canada and found south to Florida. 
Long confused with galathea under the 
name capitatus, the two species are similar 
in embolus but distinct in numerous ways. 
Pelegrina proterva can be separated from 
galathea by (males) the broader embolus 
with larger hook, strongly marked face, 
and narrower carapace and by (females) 
the abdominal markings and flat epigynal 
flaps. Male markings much like those of 
peckhamorum, from which proterva dif- 



Pelegrina Jumping Spiders • Maddison 271 



fers in having a narrower embolus and 
flatter epigynum. Palpus much like that of 
the central Mexican edrilana, differing in 
details. 

Male. Palpus (Figs. 3, 6-9, 283, 284): 
Erect portion of embolus inflated basally, 
wide and transparent centrally. Terminal 
portion near opening much narrower than 
basal portion. Retrolateral ramus a long 
curved hook (Fig. 283). Markings (Figs. 
134, 282): Carapace with strong white 
markings, including forehead and side 
bands. Cheek band dense and distinct from 
side bands. Clypeus with prominent dia- 
mond of white scales between AMEs and 
overhanging chelicerae; lateral to this the 
clypeus and hairs overhanging chelicerae 
are dark. White forehead band contacts 
AMEs dorsally; setae ringing AMEs white 
7:00-12:00 and 2:00-4:00. Chelicerae 
lacking pale scales. Femur of palpus dis- 
tinctly paler than more distal segments. 
Cymbium with generally dense patch of 
white scales centrally. Second, third, and 
fourth legs with femur bases abruptly pale. 
Abdominal dorsum usually light brown 
with dark spots between side bands. Mea- 
surements: Body length 3.3(3.6)4.2 mm; 
carapace length 1.6(1.6)1.9 mm, width/ 
length 0.74(0.78)0.81; n = 73 from Ontar- 
io, Iowa, and Saskatchewan. 

Female. Epigynum (Figs. 238, 285, 286): 
Flaps fairly flat, dark, and slightly con- 
vergent. Surface smooth, rises fairly 
abruptly behind flaps into wide though 
gentle mound covering most of posterior. 
First curve of duct broad; second curve 
proceeds obliquely anteriorly. Markings 
(Figs. 2, 135, 287): Carapace covered with 
white and some brown scales. Face some- 
what darker than galathea, relatively thin- 
ly covered with white scales, especially in 
southern females. Legs only slightly an- 
nulate, beige to light brown with darker 
markings reddish brown markings. Ab- 
domen pale on sides, with two reddish 
brown longitudinal bands above broken by 
oblique to transverse white stripes. Mea- 
surements: Body length 4.4(5.1-5.3)5.6 
mm; carapace length 1.6(1.9)2.0 mm. 



width/length 0.75(0.76)0.79; n = 6$ from 
Iowa and Ontario. 

Geographical Variation. Southern fe- 
males have a darker carapace with distinct 
side bands, a face more sparsely covered 
with pale scales, and an abdomen with the 
central pale spots coalesced medially into 
a chevron stripe, which is also seen in some 
southern males. Some males from Florida, 
Georgia, and South Carolina have a nar- 
rower embolus and lack the white scales 
between the AMEs on the clypeus. 

Chromosomes. 2n(5 = 26 acrocentrics + 
XXO (13, Mollis, New Hampshire). 

Courtship (103 observed from Shenan- 
doah Co., Virginia; Middlesex and Barn- 
stable Counties, Massachusetts; Dorchester 
and Caroline Counties, Maryland; Thun- 
der Bay, Ontario; Binscarth, Manitoba; see 
also Peckham and Peckham, 1889: 45, fig. 
18). Has the crouch display with body of- 
ten high and first legs low. Raisedspread 
(n = 13, 53). Crouch (Fig. 134; n = 15, 73): 
Body horizontal, held low (n = 1), at about 
normal height (n = 6, 33) or high (n = 8, 
3 3). First legs forward and low (n = 15, 
73), either horizontal (n = 3, 23) or even 
lower than body (n = 10, 43), bowed and 
parallel (n = 4, 23), or slightly spread (n 
= 3, 13). On one observation the legs were 
slightly raised at first but just before he 
touched 2 they were lower than body. First 
legs waving little if a all (n = 2, 13) or not 
at all (n = 4, 13). Palpi held down (n = 7, 
43) or forward (n == 7, 33); moved forward 
(n = 10, 53) and waved up and down on 
each series (n = 15, 73). Abdomen de- 
pressed on each series (n = 1), twitches at 
least occasionally (n = 4, 13). Repertoires: 
33 raisedspread only; 53 crouch only; 23 
raisedspread and crouch. Several times the 
display proceeded directly from the 
raisedspread stage to touching the female 
without a distinct crouch display (n = 8, 
43). 

Distribution (Map 5). Across much of 
Canada and northeastern United States, 
south in the east to Florida and Texas. 
Although the following records include 
only few from Canada, this is due to my 



272 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



examining only collections at museums in 
the United States; proterva is actually very 
common throughout much of Canada. 

Records. Manv specimens, especially in MCZ and 
AMNH, from: CANADA: NOVA SCOTIA: KentviUe; 
QUEBEC: St. Louis de France, Quebec City; ON- 
TARIO: Martin River 58 km N of North Bay, Ottawa, 
Belleville, Barrie, 36 km E of Thunder Bay, Sudbury 
Dist.: Espanola, 15 km E of Espanola, nr. Bruce Mines 
nr. Sault St, Marie, 14 km S of Pte. Au Baril (Parry 
Sound Dist.), 20 km E of Manitoba border on Hwy 
17; Lake Temagami, Port Credit; SASKATCHE- 
WAN: Waskana Creek, North Battleford; MANI- 
TOBA: Sandilands Provincial Forest, Binscarth, Gyp- 
sumville; BRITISH COLUMBIA: Salmon Arm. 
UNITED STATES (county records): MAINE: Han- 
cock, Lincoln; NEW HAMPSHIRE: Belknap, Carroll, 
Cheshire, Grafton, Hillsborough, Sullivan; VER- 
MONT: Addison, Caledonia, Chittenden, Rutland, 
Windham, Windsor; MASSACHUSETTS: Barnsta- 
ble, Berkshire, Dukes, Esse.x, Franklin, Hampden, 
Middlesex, Nantucket, Norfolk, Suffolk, Worcester; 
RHODE ISLAND: Washington; CONNECTICUT: 
Fairfield, Middlesex, New Haven, New London, Tol- 
land; NEW YORK: Hamilton, Nassau, New York City 
Thompkins, Westchester; NEW JERSEY: Bergen 
Cape May, Hunterdon, Passaic; PENNSYLVANIA 
Adams, Bucks, Carbon, Centre, Forest, Monroe 
Montgomery, Schuylkill, Warren, York; OHIO 
Champaign; MARYLAND: Caroline, Charles 
Dorchester; WEST VIRGINIA: Mercer; VIRGINIA 
Fairfax, Rockingham, Shenandoah, Suffolk, Wash- 
ington; KENTUCKY: Hardin, Rowan; TENNESSEE: 
Grundv, Sevier, Unicoi; NORTH CAROLINA: Bun- 
combe^ Durham, Macon; SOUTH CAROLINA: Hor- 
ry; GEORGIA: Cobb, Polk, Thomas; FLORIDA: Ala- 
chua; ALABAMA: Clarke, Dekalb, Jackson; MICH- 
IGAN: Berrien, Calhoun, Charlevoix, Cheboygan, 
Crawford, Eaton, Emmet, Genesee, Hillsdale, Isa- 
bella, Jackson, Kent, Lake, Livingston, Mackinac, 
Marquette, Midland, Muskegon, Osceola, Sanilac, 
Washtenaw, Wayne; INDIANA: Jasper, Marion, 
Starke; WISCONSIN: Ashland, Chippewa, Crawford, 
Dane, Door, Douglas, Grant, Green Lake, Iowa, Jef- 
ferson, Lincoln, Manitowac, Marathon, Price, Rich- 
land, Rusk, Sauk, Shawano, Taylor, Waushara; IL- 
LINOIS: Champaign, Cook, Mason; MINNESOTA: 
Blue Earth, Freeborn, Marshall, Olmsted, Steele, Wi- 
nona; IOWA: Boone, Cla> ton, Hancock, Winnebago, 
Woodbury; MISSOURI: Boone, Cole, Jackson, John- 
son, St. Louis; NEBRASKA: Lancaster, Loup, Saline; 
KANSAS: Decateur, Rile\; TEXAS: Anderson, Den- 
ton, Hardin, Sabine, San Jacinto; MONTANA: Ra- 
valli, Stillwater; COLORADO: Fremont, Larimer. 

Natural History. Found on various trees 
and shrubs, usually in or near forests; less 
often found in fields and on herbs than is 



P. galathea. Dondale (1961) describes the 
life history of P. proterva in Nova Scotia. 

6. Pelegrina peckhamorum 
(Kaston, 1973) 
new combination 

Figures 126, 136, 137, 195,239, 

288-293; Map 8 

Metaphidippus peckhamorum Kaston, 197.3: 115, figs. 
39-42, (52. Holotype 6 and paratype $ in AMNH 
with labels "Holotype S + allotype 9, Metaphidip- 
pus peckhamorum n. sp., det by B. J. Kaston (1949)" 
and "col. by B. Malkin, Lakehurst, N. J. 25 May 
1941," e.xamined. BrignoU, 1983: 643. 

Diagnosis. A relatively rare eastern spe- 
cies with male body form and markings 
very much like proterva but outstanding 
for its very broad embolus. The female is 
best distinguished from other eastern spe- 
cies by the indistinct markings and large, 
slightly concave epigynum, with flaps that 
are more convex than in proterva and that 
have the posterior edge not truncate as in 
galathea. 

Male. Palpus (Figs. 195, 289, 290): Em- 
bolus very broad, tapering but still broad 
at tip. Rami well separated; retrolateral 
ramus not elongate as in proterva. Mark- 
ings (Figs. 136, 288): Cheek band dense 
and discrete. Clypeus with prominent di- 
amond of white scales between AMEs and 
overhanging chelicerae; lateral to this the 
clypeus and hairs overhanging chelicerae 
are dark. White forehead band contacts 
AMEs dorsally; setae ringing AMEs white 
7:00-12:30 and 2:00-4:00. Chelicerae 
lacking pale scales. Femur of palpus dis- 
tinctly paler than more distal segments. 
Cymbium with white scales centrally. Leg 
femora distinctly paler basally. Measure- 
ments: Body length 3.0(3.6)3.7 mm; car- 
apace length 1.4(1.7)1.8 mm, width/length 
0.74(0.76)0.78; n = 53 from Barnstable 
County, Massachusetts. 

Female. Epigynum (Figs. 239, 291, 292): 
Large. Flaps long, fairly convex, usually 
convergent. Surface rises very gradually 
behind flaps; most of posterior area con- 
cave. First curve of duct broad; second 



Pelecrina Jumping Spiders • Maddison 273 



curve proceeds obliquely anteriorly. 
Markings (Figs. 137, 293): Carapace cov- 
ered with yellowish scales. Clypeus cov- 
ered thinly with yellowish white scales. 
Abdomen more uniform in color than pro- 
terva, light bro\\'n with pale spots. Mea- 
surements: Body length 3.6(4.0)5.4 mm; 
carapace length 1.7(1.9)2.1 mm, width/ 
length 0.73(0.77)0.78; n = 52 from Mas- 
sachusetts and Arkansas. 

Male/Female Matching. Males and fe- 
males have been co-collected in New Jer- 
sey, Arkansas, and Massachusetts; other- 
wise, they are the only unmatched $9 in 
the northeast. 

Courtship (45 observed from Cape Cod, 
Massachusetts). Has the crouch display with 
body high and first legs low as in P. pro- 
terva. Raisedspread (n = 17, 35). Crouch 
(Fig. 126; n = 12, 33): Body held normal- 
high (n = 6, 16) or high (n = 6, 26). First 
legs held horizontal (n = 12, 3<3) and lower 
than body (n = 3, 16), waved little if at all 
(n = 9, 26). Palpi held down (n = 12, 33); 
still on pause (n = 2, 16), waved on series 
(n = 4, 26) up and down (n = 5, 15), spe- 
cifically from down to forward (n = 2, 26), 
with medium-high amplitude (n = 5, 13). 
Abdomen still on series (n = 4, 23) but 
twitched on pause (n = 4, 23). Repertoires: 
13 raisedspread only; 23 raisedspread and 
crouch; 13 crouch only. 

Distribution (Map 8). Known from Mas- 
sachusetts, New York, New Jersey, Ohio 
(Kaston, 1973), Indiana (Kaston, 1973), 
Tennessee, Arkansas, and Texas. 

Records. UNITED STATES: MASSACHUSETTS: 
Barnstable Co.: Chatham (IS 3 9, MCZ), South Chat- 
ham (6<3, MCZ), nr. North Truro at junction of Hwy 
6 and Head of the Meadow Road (10^ 32, MCZ); 
Dukes Co.: Oak Bluffs (1<3, MCZ); NEW YORK: Dav- 
isville; Suffolk Co.: Riverhead {2S, AMNH), Coram 
(2.3 12, AMNH); NEW JERSEY: BurUngton Co.: 11 
km W of New Gretna [46 22; AMNH), Lebanon State 
Forest {16, AMNH); Middlesex Co.: Old Bridge (1<5, 
AMNH); Morris Co.: Chatham, Great Swamp (12, 
AMNH); Ocean Co.: Lakehurst {226 112, AMNH), 
Lake Horicon nr. Lakehurst (32, AMNH), 6 km W 
of Lakehurst (23 32, AMNH); TENNESSEE: Knox 
Co.: University of Tennessee farm 3 {16. AMNH); 
ARKANSAS: Washington Co.: 24 km S of Prairie 
Grove in Cove Creek Valley of the Boston Mtns. (28.3 



62, MCZ), 24 km W of Prairie Grove {56 12, MCZ); 
TEXAS: Leon Co.: SW of Oakwood {16, AMNH). 

Natural History. May specialize on oaks. 
On Cape Cod, Massachusetts, collected by 
beating oaks and cranberries in understory 
of pine forest (1 record), sweeping oak- 
pitch pine (2 records), and beating oaks (1 
record). 

7. Pelegrina neoleonis new species 
Figures 138, 196, 294-298; Map 6 

Holotype male and paratype female in MCZ with 
label "MEXICO: NUEVO LEON: Chipinque Mesa 
just S of Monterrey, ca. 4500 ft. [1,370 m]; ca. 
100.4°W 25.6°N, 2 Jun 1983 W. Maddison & R. S. 
Anderson 83-034, beating and sweeping forest un- 
derstory." 

Etymology. After the state from which 
most known specimens come. 

Diagnosis. A Mexican species similar to 
tristis with a distinctive long, curved retro- 
lateral ramus on the embolus. The erect 
portion of the embolus is narrower than in 
tristis. No characters have yet been found 
to distinguish the female from that of tris- 
tis, except locality. 

Male (from Nuevo Leon). Palpus (Figs. 
196, 295): Embolus distinctive; broad, with 
retrolateral ramus extended into long hook, 
much as in tristis, but ramus bears small 
bump and is blunt at tip; prolateral ramus 
obtuse or only slightly acute. Markings 
(Figs. 138, 294): Cheek band weak. Clyp- 
eus brown; hairs overhanging chelicerae 
dark. White forehead band contacts AMEs 
dorsally 10:30-12:30. Chelicerae lack pale 
scales. Femur of palpus distinctly pale than 
more distal segments. Cymbium with none 
to few white scales. Femur of third leg 
pale on basal Vs. Measurements: Body 
length 3.6, 3.7 mm; carapace length 1.7, 
1.8, 1.9 mm, width/length 0.75, 0.75, 0.76; 
n = 33 from Nuevo Leon and San Luis 
Potosi. 

Female. Epigynum (Figs. 296, 297): 
Flaps large and dark, flat and inwardly 
rotated. Surface gently convex, highest 
medially behind flaps, except for surface 
diving deeply under flaps. First curve of 
duct very broad, expanded to the side and 



274 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



posterior so that second curve begins well 
posteriad of posteriormost portion of flap 
(in contrast with the sympatric clavator); 
second curve proceeds anteriorly. Inner 
surface of third curve rough, with numer- 
ous projections. Markings (Fig. 298): Car- 
apace dark above, covered with transpar- 
ent reflective scales; sides covered thinly 
with white scales. Clypeus densely covered 
with white scales. Legs brown. Abdomen 
fairly dark, with only small pale spots. 
Measurements: Body length 4.3(5.9- 
6.1)6.2 mm; carapace length 1.9(2.2- 
2.3)2.3 mm, width/length 0.74(0.77- 
0.79)0.83; n = 4$ from Nuevo Leon, Hi- 
dalgo, and Oaxaca. 

Male/Female Matching. This associa- 
tion is indicated by co-collecting in Nuevo 
Leon; by the large epigynal flaps, which 
would be expected in a species with such 
a robust embolus; and by the similarity of 
male and female with those of tristis. 

Geographical Variation. The single male 
from San Luis Potosi differs from those of 
Nuevo Leon in having a sharper retrolater- 
al prong on the embolus, more extensive 
white scales surrounding AMEs, a small 
patch of pale scales medially on chelicerae, 
and the femur of palpus relatively dark. 

Courtship (33 observed from Chipinque 
Mesa, Nuevo Leon; Cerro Potosi, Nuevo 
Leon; and Xilitla, San Luis Potosi). Raised- 
spread (n = 3, 2(5). Crouch (n = 6, 33): 
Body held in normal to low position (n = 
1). First legs bowed and forward (n = 4, 
23), raised to ca. 30° (n = 2, 23), or hori- 
zontal (n = 1), or femora low but tips curl 
upward (n = 1). Leg tips not touching (n 
= 2, 23), apparently not waved (n = 3, 23), 
or waved only slightly (n = 2, 13) on series 
(n = 1). Palpi down (n = 6, 33), over che- 
licerae (n = 1) or curled under tips of che- 
licerae (n = 1), waved (n = 6, 33) up and 
down (n = 1) or outward (n = 1) on series 
(n = 3, 23) ca. 5-7 c/s (n = 1). Abdomen 
depressed a bit on series (n = 1), or at end 
of series (n = 1). Repertoires: 13 crouch 
only; 23 raisedspread and crouch. 

Distribution (Map 6). Northeastern 
Mexico south to Oaxaca. 



Records. MEXICO: SAN LUIS POTOSI; 21 km W 
of Xilitla on Hwy 120, 99°05'W, 21°18'N, 12 June 
1983 (15, MCZ); NUEVO LEON: Chipinque Mesa 
just S of Monterrey, 100. 4°W, 25.6°N, 2 June 1983 
{IS 12, MCZ); Cerro Potosi, ca. 100°14'W, 24°52'N, 4 
June 1983 (15 12, MCZ); HIDALGO: Pachuca (12, 
MCZ); OAXACA: 50 km NW of Oaxaca, 97°00'W, 
17°14'N, 6 August 1983 (22, MCZ). 

Natural History. Beating oaks and pines 
in oak-pine area (3 records); sweeping 
shrubs, cloud forest (1 record). Elevations 
at four locations in Nuevo Leon and San 
Luis Potosi from 1,400 to 2,900 m. 

8. Pelegrina tristis new species 
Figures 197, 299-303; Map 7 

Holotype male and paratype female in AMNH with 
labels "ARIZONA: Cochise Co., Round Park, Chir- 
icahua Mtns., June 28, 1967. 9300 ft. [2,840 m], 
Gertsch, Hastings." 

Etymology. Latin adjective for "sad," 
referring to the large size of the teardrop- 
shaped flaps over the epigynal openings. 

Diagnosis. A large, dark, plainly marked 
species known from southern Arizona, 
similar in genitalia to neoleonis and sa- 
binema. The erect portion of the embolus 
is broader than in either of those species, 
and the rami are sharper than in neoleonis. 
Females are generally not so yellow as in 
sabinema, and the epigynal openings are 
deeper, in that the surface descends more 
deeply under the anterior part of the flaps. 

Male. Palpus (Figs. 197, 300): Embolus 
extremely broad, so that retrolateral mar- 
gin joins without angle to retromargin of 
embolar base. Both rami sharply pointed; 
retrolateral ramus extended into long hook, 
lacking subterminal bump. Markings (Fig. 
299): Carapace dark, with reduced fore- 
head band. Cheek band very weak to ab- 
sent. Clypeus brown, with dark hairs over- 
hanging chelicerae. White forehead band 
absent or much reduced, fails to contact 
AMEs, which are ringed with dark above. 
Chelicerae lacking pale scales. Palpus al- 
most uniformly brown, femur not distinct- 
ly paler. Cymbium lacking white scales. 
Legs relatively uniform brown, femora en- 
tirely dark. Measurements: Body length 
3.7(4.3)4.6 mm ; carapace length 1 .8(2. 1)2.1 



Pelegrina Jumping Spiders • Maddison 275 



mm, width/length 0.75(0.76)0.82; n = 4<5 
from Chiricahua and Santa Catalina Mtns., 
Arizona. 

Female. Epigyniim (Figs. 301, 302): 
Flaps large, dark, and convergent, often 
far rotated, sometimes as far rotated as in 
neoleonis (Fig. 297). Just medial to the flap 
at the anterior end the surface is pale and 
descends deep under flap (Fig. 302, ar- 
row). Except for this concavity, the epi- 
gynal surface is gently convex, highest me- 
dially behind flaps. First curve of duct very 
broad; second curve proceeds anteriorly. 
Markings (Fig. 303): Carapace covered 
above thinly with white to dark transpar- 
ent reflective scales. Clypeus densely cov- 
ered with white scales. Abdomen light to 
medium brown with small central pale 
spots. Narrow dark brown spots beside 
these pale spots form longitudinal dark 
stripes. Measurements: Body length 
5.0(5.7)6.2 mm; carapace length 1.8(2.0)2.3 
mm, width/length 0.77(0.78)0.82; n = 59 
from Chiricahua, Huachuca, and Santa 
Rita Mtns., Arizona. 

Male/Female Matching. This match- 
ing is indicated by microsympatry in Chir- 
icahua Mtns., by robust embolus and flaps, 
by similar large size, and by similarity of 
genitalia to male and female of sabinema, 
which are reasonably surely matched. 

Distribution (Map 7). Southern Arizona. 

Records. UNITED STATES: ARIZONA: Santa Rita 
Mtns.: Madera Canyon (49, AMNH, MCZ); Huachuca 
Mtns.: Garden Canyon (19, AMNH); Santa Catalina 
Mtns.: Bear Wallow to Mt. Lemmon (19, AMNH), 
Chiricahua Mtns.: Round Park, Southwestern Re- 
search Station 8 km W of Portal, Barfoot Park, and 
Rustler's Park (3<3 49, AMNH). 

Natural History. Collected at 1,500- 
2,800 m elevation (3 records). Females have 
been collected in June (2 records), July (4 
records), and August (3 records). 

9. Pelegrina sabinema new species 
Figures 198, 304-308; Map 9 

Holotype male in AMNH with label "ARIZONA, 
Showlow, July 1967, W. J. Gertsch. " 

Etymology. An arbitrary combination 
of letters, to be treated as an adjective. 



Notes on Specific Distinctness. Pele- 
grina sabinema is much like pervaga, and 
indeed I long considered it only the west- 
ern form of pervaga, but the more strongly 
developed white-black-white carapace 
stripes and narrower embolus of pervaga 
suggest that pervaga may be the sister spe- 
cies to kastoni, with sabinema the sister to 
those two. The embolus and markings of 
P. sabinema are slightly more like those 
of tristis and neoleonis, which may be con- 
sidered outgroups. 

Diagnosis. Differs from pervaga in hav- 
ing less swollen carapace sides, an abdo- 
men lacking the pale central stripe on the 
abdomen, wider embolus, weaker male 
cheek band, less dense band of dark hairs 
beneath male carapace side bands, darker 
and more robust epigynal flaps, and yellow 
female legs. Differs from tristis in having 
yellow legs, yellow male chelicerae, nar- 
rower embolus, dense covering of pale 
scales on female carapace, and shallower 
epigynal openings. 

Male. Palpus {Figs. 198, 305): Embolus 
very wide at base of erect portion, thought 
still with a distinct angle between retro- 
margins of erect portion and base; retro- 
lateral ramus long and blunt. Carapace 
often broad though sides not swollen as in 
pervaga. Markings (Fig. 304): Cheek band 
weak, runs horizontally and posteriorly be- 
neath band of dark hairs beneath white 
side bands. Clypeus brown, hairs over- 
hanging chelicerae dark. White forehead 
band contacts AMEs rather far medially, 
from 9:00 to 12:00. Chelicerae yellow, 
lacking pale scales. Palpus yellow with 
white scales on femur, tibia and cymbium 
interrupted by dark hairs on patella and 
base of cymbium. Legs uniformly yellow. 
Abdomen brown centrally with white side 
bands, showing trace of paired dark spots 
of female. Measurements: Body length 
3.3(3.6)3.8 mm; carapace length 1.7(1.7)1.9 
mm, width/length 0.78(0.80)0.83; n = 53 
from New Mexico and Arizona. 

Female. Epigynum (Figs. 306, 307): 
Flaps flat and large, often far rotated, as 
in tristis and neoleonis, though usually not 



276 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



so dark as in those species. Openings shal- 
lower than in tristis; that is, just medial to 
the flap at the anterior end the surface is 
not so pale and does not dive deep under 
flap. Epigynal surface flat. First curve of 
duct very broad; second curve proceeds 
less anteriorly than in tristis. Markings 
(Fig. 308): Carapace well covered with 
white to yellowish scales. Clypeus very 
densely covered with white scales. Legs 
yellow. Abdomen yellowish with brown 
centrally, paired white spots. Usually 
paired dark brown spots in posterior half 
are beside white spots. Measurements: 
Body length 4.0(4.2)5.5 mm; carapace 
length 1.6(1.7-1.8)1.9 mm, width/length 
0.78(0.80-0.81)0.86; n = 6$ from New 
Mexico and Arizona. 

Male/Female Matching. Male and fe- 
males were matched by similar yellow col- 
or, by robust embolus and flaps, and by 
co-collecting and common distribution in 
New Mexico and northern Arizona, where 
no other unmatched females and males are 
known. 

Distribution (Map 9). New Mexico, 
northern Arizona, southern Colorado, and 
western Texas. 

Records. UNITED STATES: TEXAS: Jeff Davis 
Co.: 24 km NW of Fort Davis (19, AMNH); COL- 
ORADO: Montezuma Co.: Mesa Verde National Park 
(1<5, AMNH); NEW MEXICO: Bernalillo Co. (29, 
AMNH); Lincoln Co.; nr. Ruidoso, Ruidoso Cmpgd. 
(22, AMNH); Los Alamos Co.: nr. Los Alamos (19, 
AMNH); Sandoval Co.: (19, AMNH), Sandia Mtns., 
Juan Tabo area (29, AMNH); Santa Fe Co. (2<? 19, 
AMNH), Glorieta Mesa nr. Rowe (1<5 19, AMNH), 5 
km N of Galiseo (13, AMNH), Route 66 just E of 
Edgewood (19, AMNH), 19 km S of Lamy (19, 
AMNH); Taos Co.: 27 km S of Taos (13 19, AMNH); 
ARIZONA: Coconino Co.: Flagstaff {IS 19, UCB), 
Navajo Co.: Showlow (13, AMNH), 

Natural History. Collected at 7,000 ft 
elevation (2 records), from piny on pine- 
juniper (1 record). 

1 0. Pelegrina pervaga 
(G. & E. Peckham, 1909) 
new combination 
Figures 199, 240, 309-313; Map 10 

Dendryphantes pervagiis G. & E. Peckham, 1909: 
474, pi. 37, figs. 9, 9a, 2. Holotype in MCZ 12 with 



labels "Dendryphantes pervagus P., 9 Wallace, 
Kansas type" (label is original; handwritten, prob- 
ably by Elizabeth Peckham) and "G. W. Peckham 
Coll.", examined. Roewer, 1954: 1214. 

Dendryphantes (Metaphidippus) prevagus [sic]: — 
Petrunkevitch, 1911: 640. 

Metaphidippus pervagus: — Bonnet, 1957: 2817. 

Diagnosis. A striking species with swol- 
len carapace sides and central pale stripe 
on the abdomen in both sexes. Very similar 
to sabinema, from which it is distinguished 
by the features discussed under that spe- 
cies. 

Male. Palpus (Figs. 199, 310); Embolus 
wide basally; retrolateral ramus long, 
pointing distally, having subterminal 
bump. Carapace sides swollen. Markings 
(Fig. 309): White carapace side band bor- 
dered below by narrow band of black hairs. 
Below this, the dense white cheek bands 
do not reach clypeus, which is dark. Hairs 
overhanging chelicerae dark. White fore- 
head band contacts AMEs far medially, 
from 9:00 to 11:00. Chelicerae yellow, 
lacking pale scales. Palpus yellow, with 
white scales on end of femur, on tibia and 
cymbium, interrupted by dark hairs on 
patella. Legs light yellowish brown with 
some darker annulations. Abdomen with 
central longitudinal pale stripe as in fe- 
male. Measurements: Body length 
3.7(3.9)4.4 mm; carapace length 1.8(1.8)2.0 
mm, width/length 0.83(0.85)0.88; n = 5<5 
from Erath Co., Texas. 

Female. Epigynum (Figs. 240, 311, 312): 
Flaps fairly large and flat, generally pale. 
Surface flat. First curve of duct wide; sec- 
ond curve proceeds obliquely anteriorly. 
Markings (Fig. 313): Carapace wide and 
covered with whitish scales. Clypeus cov- 
ered densely with white scales. Abdomen 
with distinctive pale patch on middle of 
dorsum. Measurements: Body length 
4.3(4.8)5.9 mm; carapace length 1.8(2.0)2.2 
mm, width/length 0.80(0.82)0.85; n = 59 
from Erath Co., Texas. 

Distribution (Map 10). Texas, Oklaho- 
ma, and Kansas. 

Records. UNITED STATES: KANSAS: Wallace 
Co.: Wallace (19, MCZ); OKLAHOMA: Commanche 
Co.: Visitor's Center, Wichita Mtns. (13, WPM); TEX- 



Pelecrina Jumping Spiders • Maddison 211 



AS: Erath Co.: 11 km NE of Stephenville (63 72, 
TXAM). 

Natural History. Collected from juni- 
pers in Texas (three records). 

1 1 . Pelegrina kastoni new species 
Figures 140, 141, 200, 314-318; Map 11 

Metaphidippus n. sp. nr. aeneolus: — Jung and Roth, 
1974: 33 (specimens identified by W. J. Gertsch, 
examined). 

Holotvpe male and paratype female in MCZ with 
label "ARIZONA: Santa Cruz Co., Santa Rita Mtns., 
gate at 26 km of Whipple Obs[ervatory]. Rd. on 
Mt. Hopkins 7100 ft el. [2,170 m], 17 June 1985 
W. Maddison 85-059, beating Cercocarpus and ju- 
niper." 

Etymology. This beautiful species is 
named after the late B. J. Kaston, whose 
excellent work on the eastern species of 
Pelegrina added much to our understand- 
ing of the genus. 

Diagnosis. The yellowish appendages 
and carapace stripes of males are similar 
to those of pervaga and sabinenia, but the 
embolus and white bands on clypeus fail- 
ing to meet at center are distinctive. The 
golden and beige females have distinctive 
epigynal flaps rotated 90° inward. 

Male. Palpus. (Figs. 200, 315): Embolus 
rectangular but twisted at tip, relatively 
narrow, arising from retrolateral side of 
base. Markings (Figs. 140, 314): White 
carapace side band is bordered below by 
narrow band of black hairs, as in pervaga. 
Below this are thin, dense white cheek 
bands that extend across clypeus like Clark 
Gable moustache, broken in center. White 
forehead band either fails to contact AMEs, 
or at most contacts AMEs locally at 10:30- 
12:30. Chelicerae yellow, lacking pale 
scales. Palpus yellow, with markings much 
as pervaga and sabinema, with white scales 
on end of femur, on tibia and cymbium 
alternating with dark hairs on patella and 
base of cymbium. Legs yellow to light 
brown except first metatarsus distinctly 
darker, brown to black. Measurements: 
Body length 4.1(4.2)4.5 mm; carapace 
length 2.1(2.1)2.2 mm, width/length 
0.79(0.80)0.83; n = 53 from Santa Rita 
Mtns., Arizona. 



Female. Epigynum (Figs. 316, 317): 
Flaps rotated a full 90°, with deep openings 
just anterior to them, resting in pits, almost 
as in Dendryphantes nigromaculatus 
though flaps maintain their prominence as 
in other Pelegrina. Surface flat or slightly 
convex except for pits containing flaps. 
First curve of duct narrow and proceeding 
medially. Markings (Figs. 141, 318): Car- 
apace wide, covered with yellowish scales. 
Clypeus densely covered with white scales. 
Legs light orange-brown. Abdomen brassy 
with beige markings. Measurements: Body 
length 4.9(5.2)6.5 mm; carapace length 
2.1(2.3)2.4 mm, width/length 0.80(0.81) 
0.83; n = 59 from Santa Rita Mtns., Ari- 
zona. 

Male /Female Matching. The matching 
is indicated by extensive co-collecting on 
junipers in Arizona, by the wide carapace 
and yellowish color, and by the correlated 
retrolateral shift of the embolus and ro- 
tation of flaps. 

Chromosomes. 2n6 = 26? acrocentrics 
+ XXO (16, Madera Canyon, Arizona). 

Courtship (33 observed from Santa Rita 
Mountains, Arizona). The side-to-side 
waving of legs and palpi during crouch is 
distinctive. There is no clear distinction 
between raisedspread and crouch displays, 
but what may be raisedspread stage oc- 
curred as follows (n = 2, 23): First legs 
bowed and forward, tips apart (n = 1) to 
nearly touching (n = 3, 23), not moving (n 
= 3, 23). Palpi down (n = 3, 23), waving 
irregularly (n = 2, 13). Crouch (n = 9, 13): 
Body low (n = 8, 13) or at normal height 
(n = 1). First legs forward and bowed with 
tips touching, femora approximately hor- 
izontal but leg raised distally (n = 8, 13). 
First legs waving at very low amplitude 
very high frequency (n = 8, 13). Later, as 
he gets closer, the first legs are more par- 
allel, and the first legs and palpi are waved 
slowly, ca. 1-2 c/s, all four first to right 
then to left (n = 1). Palpi when close (late 
crouch) held forward and waved left to 
right as noted above (n = 1). Abdomen 
twitching (n = 8, 13). Repertoires: 23 



278 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



raisedspread or early crouch only; 16 
crouch only. 

Distribution (Map 11). Southern Ari- 
zona, southwestern New Mexico, and Chi- 
huahua. 

Records. UNITED STATES: ARIZONA: Santa 
Cruz Co.: Madera Canyon (6<J 159, AMNH, MCZ), 
Mt. Hopkins (3<? 52, MCZ), Sycamore Canyon W of 
Nogales (2<5 29, MCZ), 29 km E of Nogales (19, 
AMNH); Cochise Co.: Chiricahua Mtns., Southwest- 
ern Research Station 8 km W of Portal (8<5 59, AMNH), 
5 km N or Portal (19, AMNH), Cave Creek Canyon 
{IS, AMNH), Cienega Lake (19, AMNH), Huahucha 
Mtns., Garden Canvon (19, AMNH); Gila Co.: Tonto 
Creek Camp nr. Kahb's ranch (2$ 19, AMNH). NEW 
MEXICO: Hidalgo Co.: Animas Valley (13, AMNH). 
MEXICO: Chihuahua (1<5, MCZ). 

Natural History. Primarily found on ju- 
niper. In my collections (17-19 June 1985) 
from oak-juniper-pine woodlands in south- 
central Arizona, 96 179 were beaten from 
junipers, 29 from oaks, and 16 39 from 
Cercocarpus. Collected at elevations from 
1,200 to 2,200 m. Most males collected in 
June (May — 2 records, June — 8, July — 2, 
November — 1); most females in June and 
July (June — 7 records, July — 5, August — 
2). Jung and Roth (1974) collected this spe- 
cies in their zone 2 in the Chiricahua 
Mountains (1,460-1,700 m elevation). 

12. Pelegrina flavi pedes 
(G. & E. Peckham, 1888) 
new combination 

Figures 142, 143, 201, 241, 319-323, 

338,339; Map 12 

Dendryphantes flavipedes G. & E. Peckham, 1888: 
42, pi. 3, fig. 29a, $. Holotype in MCZ \6 with labels 
"Dendryphantes flavipedes Pkm, 1888. Canada. 
Type. S" (label is original; handwritten, probably 
by Elizabeth Peckham) and "G. W. Peckham Coll.", 
examined. G. & E. Peckham, 1909: 471, pi. 38, figs. 
3, 3a-c, <59. Roewer, 1954: 1210. 

Dendryphantes (Metaphidippus) flavipedes: — Pe- 
trunkevitch, 1911: 630. 

Metaphidippus flavipedes: — Chickering, 1944: 174, 
figs. 66-69, S9. Bonnet, 1957: 2813. Kaston, 1973: 
112, figs. 21-25, (59. 

The flavipedes group at first glance ap- 
pears to consist of three easily distin- 
guished species (Kaston, 1973): a striped 
species distributed across Canada (fla- 
vipedes), a yellow species with bulbous 



head in the northeastern United States 
(flaviceps), and a dark southern species 
(exigua). The situation is not nearly so sim- 
ple as this, however, for hybridization may 
occur at the borders of their ranges (dis- 
cussed under flaviceps) and two species 
may be confused under the name exigua 
(discussed under exigua). 

Diagnosis. The handsomely striped yel- 
low and brown males of flavipedes are dis- 
tinguished from flaviceps and most exigua 
by the strong cheek band, three white spots 
above anterior eyes, narrow carapace, 
smaller medial black spot on the chelic- 
erae, and lack of bulbous head. The striped 
form of exigua might be confused for fla- 
vipedes except by the much broader retro- 
lateral ramus of the embolus of flavipedes. 
The females have a brassy sheen distin- 
guishing them from other northern Pele- 
grina. Pelegrina flavipedes females differ 
from flaviceps females in being darker and 
having a narrower, more obliquely di- 
rected second curve of the epigynal ducts; 
they differ from exigua females in having 
more parallel epigynal flaps and a much 
narrower second curve of the epigynal 
ducts. See notes under flaviceps regarding 
possible hybridization. 

Male. Palpus (Figs. 201, 320): Embolus 
divided deeply into two rami; prolateral 
ramus of embolus twisted at tip; retrola- 
teral ramus thick. Markings (Figs. 142, 
319): Carapace well marked with dense 
side band, wide, dense cheek band, and 
three patches of white scales on forehead, 
a large one between two AMEs and smaller 
ones between each AME and ALE. Be- 
cause the forehead band contacts AMEs 
medially, the setae ringing the AMEs above 
are dark from 10:30 to 1:30. Clypeus with 
patch of white scales between AMEs; hairs 
overhanging chelicerae white medially 
brown laterally. Chelicerae yellow, with a 
small black spot medially. Femur of palpus 
through tibia yellow, contrasting with 
brown cymbium, which lacks white scales. 
Legs generally yellowish with more or less 
distinct longitudinal stripe on first femur; 
in a few specimens with stripes on all fem- 
ora but not as thin as flaviceps. Measure- 



Pelegrina Jumping Spiders • Maddison 279 



merits: Body length 3.6(3.7)4.3 mm; car- 
apace length 1.7(1.9)2.0 mm, width/length 
0.74(0.76)0.79; n = 5<3 from Neepawa, 
Manitoba. 

Female. Epigynum (Figs. 241, 321, 322): 
Flaps flat, parallel or only slightly conver- 
gent. Surface flat. First curve of duct broad; 
second curve narrow, beginning from pos- 
terior end of first curve and proceeding 
anteriorly toward the midline; flowerlike 
gland openings on dorsal face of duct. 
Markings (Figs. 143, 323): Carapace cov- 
ered with transparent brown scales with a 
brassy sheen. Beige spots on forehead be- 
tween and beside AMEs recall white spots 
of male and are usually stronger than in 
exigua. Setae directly above AMEs brown. 
Clypeus generally densely covered with 
white scales. Abdomen with brassy sheen 
and fourth pair of white spots formed into 
distinct chevron. Measurements: Body 
length 4.4(4.7)4.8 mm; carapace length 
1.8(1.9)2.0 mm, width/length 0.74(0.77) 
0.77; n = 52 from Neepawa, Manitoba. 

Chromosomes. 2n<5 = 26 acrocentrics + 
XXO (16 Nipigon, Ontario; 16 Edmonton, 
Alberta). 

Courtship (43 observed from Edmon- 
ton, Alberta, and Neepawa, Manitoba). 
With unusual alternate waving of palpi 
during crouch display. Raisedspread (n = 
1). Crouch (n = 7, 4(5): Body held at normal 
height (n = 5, 23) or somewhat raised (n 
= 1). First legs forward and horizontal (n 
= 2, 23) with tips raised (n = 1) or not (n 
= 1), or whole leg raised slightly (n = 5, 
23). First legs apparently not waving (n = 
2, 23). Palpi held down (n = 1) and some- 
what forward (n = 2, 23). Palpi wave (n 
= 6, 33) up and down alternately (n = 5, 
23), fairly slowly, on series (n = 1); spe- 
cifically, palpi tips wave in small circles (n 
= 1), left clockwise and right counterclock- 
wise or vice versa (n = 1). Abdomen 
twitches with low amplitude on series (n 
= 1). Repertoires: 33 crouch only; 13 
raisedspread and crouch. 

Distribution (Map 12). Across much of 
Canada and northeastern United States, 
south along the Rocky and Appalachian 
Mountains. 



Records. Many specimens, especially in MCZ and 
AMNH, from: CANADA: NORTHWEST TERRI- 
TORIES: Mackenzie: Prelude Lake 113°55'W, 
62°33'N; Lady Evelyn Falls 117°19'W, 60°57'N; 
NEWFOUNDLAND: Humber River; PRINCE ED- 
WARD ISLAND: Tracadie; QUEBEC: Quebec City; 
NOVA SCOTIA: Weymouth, Harrington, Baddeck 
(Cape Breton), North Sydney; ONTARIO: Ottawa, 
Marten River 58 km N of North Bay, Parry Sound 
Dist.: 14 km S of Pte. Au Baril Station, Algoma Dist.: 
near Bruce Mines; Sudbury Dist.: Espanola; Thunder 
Bay Dist.: 7 km E of Nipigon; Kenora Dist.: Granite 
Lake; Lakefield, Chapleau, Sowerby, Lake Tema- 
gami, 56 km E of Hearst, Sioux Lookout, Gawas Bay, 
Kamiskotia Lake, Turkey Point, Moberly, Uxbridge, 
Spanish River, Iron Bridge, St. Williams, Batchawana, 
Nipigon, Dorset, L. Opeongo, Cababogie (Tweed), 
Haileybury, Pancake Bay nr. Batchawana, Emo, 
Fairbank Lake Province Park, Nestorville, Golden 
Lake, Minden, SouthTea Lake (Algonquin); MANI- 
TOBA: Kettle Rapids, Cedar Lake, Lyons Lake, 19 
km E of Neepawa, Sandilands Provincial Forest; AL- 
BERTA: Edmonton, Jasper, Banff, Athabasca Land- 
ing, Fitzgerald, North Lake Athabasca; BRITISH 
COLUMBIA: Wells Cray Park, Columbia Lake, 
Salmon Arm, Arrow Lakes. UNITED STATES 
(county records): MAINE: Aroostook, Penobscot, Pis- 
cataquis, Washington; NORTH CAROLINA: Avery, 
Buncombe; MICHIGAN: Allegan, Baraga, Calhoun, 
Charlevoix, Cheboygan, Chippewa, Crawford, Delta, 
Emmett, Grand Traverse, Ingham, Mackinac, Mar- 
quette, Oakland, Washtenaw; WISCONSIN: Chip- 
pewa, Lincoln; ILLINOIS: Lake; MINNESOTA: 
Clearwater, Hennepin; MONTANA: Jefferson, Ra- 
valli, Sanders; IDAHO: Fremont; WYOMING: Crook, 
Lincoln, Park, Teton; COLORADO: Archuleta, Boul- 
der, Gilpin; NEW MEXICO: Sandoval, San Miguel, 
Taos; WASHINGTON: Okanagan, Stevens. 

Natural History. A conifer dweller, col- 
lected from spruce (11 records from On- 
tario, Manitoba, and Alberta), including 
white spruce (2 records); pines (4 records 
from Ontario, British Columbia, and 
Michigan), including lodgepole pine (1 
record) and jackpine (1 record); junipers 
(3 records from Ontario); and larch (2 re- 
cords from Ontario and Illinois). 

13. Pelegrina flaviceps 
(Kaston, 1973) 
new combination 

Figures 144, 145, 202, 242, 324-328, 

340,341; Map 13 

Metaphidippus flaviceps Kaston, 1973: 110, figs. 15- 
20, (59. Holotype 6 and paratype 9 in AMNH with 
labels "Holotype S + allotype 9, Metaphidippus 
flaviceps n.sp., det by B. J. Kaston (1949)" and 



280 



Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



"Clarendon, Vt., 2 Sept. 1939, E. M. Greenspan," 
examined. Brignoli, 1983: 643. 

Notes on Specific Distinctness. Appar- 
ent hybridization between this species and 
the other two in the group makes their 
separation doubtful. Males from northern 
Massachusetts (Pepperell, Groton, East 
Templeton), southern New Hampshire 
(Hollis), and Ithaca, New York, show grades 
of intermediacy between exigua and flav- 
iceps. Of 25 males collected at one site in 
Pepperell, 9 have dark legs and a flat ce- 
phalic area (as in exigua farther south), 10 
have dark legs and a bulbous cephalic area, 
1 has yellow legs and a flat cephalic area, 
and 5 have yellow legs and a bulbous ce- 
phalic area (as in flaviceps farther north). 
At this site, even those with yellow legs 
and bulbous carapace have a wider cara- 
pace than is usual for flaviceps. Courtship 
behavior of the apparent hybrids from 
Pepperell appears like that o{ flaviceps and 
exigua (35 observed). In Michigan and 
Maine, on the northern edge of the range 
of flaviceps, males otherwise like fla- 
vipedes have been found with the bulbous 
cephalic area. 

Diagnosis. Males can be distinguished 
from those of flavipedes and exigua by the 
bulbous cephalic area, dark lateral spots 
on the chelicerae (Fig. 324), carapace dust- 
ed with pale scales (Fig. 144), and pale 
yellow legs. Females can be difficult to 
distinguish from those of flavipedes and 
exigua; they are generally paler, lack white 
spots on the forehead, often have narrow 
dark lines on the femora, and may have 
the cephalic area slightly swollen. The epi- 
gynal flaps and ducts are intermediate be- 
tween those oi flavipedes and exigua. The 
medial black spot on the chelicerae is much 
more distinct in flaviceps females than in 
flavipedes females. 

Male. Palpus (Figs. 202, 325): Embolus 
divided deeply into two rami; prolateral 
ramus of embolus not as twisted as in ex- 
igua. Carapace is swollen dorsally in ce- 
phalic area; carapace narrowest of group, 
especially at front. Markings (Figs. 144, 
324): Carapace and abdomen dusted with 
pale scales, so as to make side bands in- 



distinct. Lacks white spots on forehead. 
Cephalic area appears yellow in alcohol, 
though may be dark in life (Fig. 144) and 
is covered with small dark hairs. Cheek 
band dense and wide. Clypeus brown, hairs 
overhanging chelicerae dark. Setae sur- 
rounding AMEs brown to light brown en- 
tirely. Chelicerae yellow with long, deep, 
dark black spot medially, and brown spot 
laterally near the base. Just lateral to the 
medial black spot the surface is flat to 
slightly concave. Palpus light brown 
throughout, cymbium not noticeably 
darker. Cymbium lacks white scales. Legs 
pale yellow with thin black prolateral lon- 
gitudinal stripe on all femora except some- 
times lacking on posterior legs. First leg 
with fringe of white hairs. Measurements: 
Body length 3.7(3.8)3.8 mm; carapace 
length 1.8(1.8)1.9 mm, width/length 
0.72(0.73)0.74; n = 53 from Sagadahoc Co., 
Maine. 

Female. Epigynum (Figs. 326, 327, 340, 
341): Flaps parallel to convergent. Epi- 
gynal surface flat. First curve of duct broad, 
not so long as in flavipedes; second curve 
broader and more medially directed than 
in flavipedes, though not nearly so broad 
as in exigua; flowerlike gland openings on 
dorsal face of duct. Carapace: Sometimes 
with swollen cephalic area. Markings (Fig. 
328): Carapace uniformly dark on fore- 
head, lacking patches of pale scales behind 
anterior eyes. Clypeus densely covered 
with white scales. Chelicerae with prom- 
inent medial black spot reminiscent of 
males. Some females with thin longitudi- 
nal lines on leg femora. Abdomen usually 
indistinctly marked, sometimes with dark 
areas formed into longitudinal bands on 
either side of middle paler area. Measure- 
ments: Body length 3.7(4.0)4.6 mm; car- 
apace length 1.7(1.8)1.9 mm, width/length 
0.74(0.74)0.75; n = 59 from Sagadahoc Co., 
Maine, and Durham, New Hampshire. 

Courtship (43 observed from Reid State 
Park, Maine). With unusual alternate cir- 
cling of palpi during crouch stage. Raised- 
spread (n = 2, 2(5): continued until 2 or 3 
body lengths from female when male went 
into crouch stage (n = 2, 23). Crouch (n = 



Pelegrina Jumping Spiders • Maddison 281 



10, 46): Body held high (n = 6, 23) or low 
(n = 1). First legs forward and bowed (n 
= 5, 3(3), tips close (n = 2, 13); legs move 
more parallel as he gets close (n = 1). Fe- 
mur low but raised distally (n = 5, 33), or 
legs may be horizontal and lower than body 
(n = 4, 13). Palpi down and forward (n = 
3, 13). Palpi flicker (n = 9, 43) for few 
seconds (n = 4, 33), then pause for few 
seconds (n = 3, 23). Flicker-pause cycle 
repeats (n = 5, 33), 3-4 (n = 1) or more 
than 8 times (n = 1). Palpus flicker is of 
low amplitude and high frequency (n = 
2, 23) and is superimposed on larger up 
and down slightly circular wave; right pal- 
pus moving up as left down and vice versa 
(n = 2, 23). Abdomen twitched occasion- 
ally (n = 9, 43) when palpi flickered (n = 
5, 33). For much of display, male stood 
without walking, thought when he did 
palpi and abdomen were flickered during 
series (n = 1) and during pause (n = 3, 23). 
Once, male proceeded to mount after about 
3-4 palpus flicker cycles of crouch (n = 
1). Repertoires: 23 crouch only; 23 raised- 
spread and crouch. 

Distribution (Map 13). Northeastern 
United States and southeastern Canada. 

Records. Over 1003 1009 in AMNH and MCZ from; 
CANADA: QUEBEC: Lake Champlain; ONTARIO: 
Toronto, Newmarket, Ottawa. UNITED STATES 
(county records): MAINE: Cumberland, Hancock, 
Penobscot, Sagadahoc, Waldo; NEW HAMPSHIRE: 
Carroll, Cheshire, Coos, Grafton, Strafford; VER- 
MONT: Caledonia, Rudand, Windham; NEW YORK: 
Albany, Cortland, Essex, Franklin, Greene, Hamil- 
ton, Lewis, Nassau, Oneida, Onondaga, Schoharie, 
Steuben, Tompkins, Yates; PENNSYLVANIA: Pike; 
WEST VIRGINIA: Preston; MICHIGAN: Charle- 
voix, Kalkaska. 

Natural History. A conifer dweller, col- 
lected from spruce (2 records), junipers 
and pine (1 record), spruces, firs, and pine 
(1 record), and hemlocks (1 record). 

14. Pelegrina exigua (Banks, 1892) 
new combination 

Figures 127, 146-149, 203, 243, 

329-337,342; Map 14 

Dendryphantes exiguus Banks, 1892: 75, pi. 5, fig. 
30, 9. Holotype in MCZ 19 with labels "Dendry- 
phantes exiguus Bks type," "Ithaca, N.Y., ' and 



"Nathan Banks Coll.", e.xamined. Roewer, 1954: 
1209. 

Dendryphantes virginis Chamberlin, 1925a: 23.3. 
Type material said to be in MCZ by Chamberlin 
but no holotype found therein. Paratypes from 
Woodridge, District of Columbia (33 19), Bladen- 
burg, Maryland (23), examined. Bonnet, 1956: 1402. 

Metaphidippus virginis: — Muma, 1944; 11. 

Metaphidippus exiguus: — Kaston, 1945; 10. Kaston, 
1973; 112, figs. 26-29, 39. 

Metaphidippus flavipedes: — Bonnet, 1957: 2813. 



Notes on Synonymy. The holotype fe- 
male of D. exiguus is peculiar in having 
the palpus tarsi swollen, as in antepenul- 
timate instar males, yet has a well-devel- 
oped epigynum. There is doubt as to the 
placement of this specimen, for the epi- 
gynum is much more like that of flaviceps 
than of southern species that has been called 
exiguus, with the flowerlike glands on the 
face of the duct, not hidden by a fold, and 
the duct not nearly so broad (Fig. 342) as 
is usual in the southern species. However, 
the markings of the type are like those of 
the southern species (white scales between 
anterior eyes on forehead, no stripes on 
legs, wide carapace, abdominal markings 
more uniformly dark). Males collected 
from Ithaca, New York (including a male 
in the MCZ with a Bryant label indicating 
that it was collected with Banks's female 
type of exiguus), are in most respects like 
exigua but have a head bump like flavi- 
ceps. It therefore appears that the type 
locality of exigua is along the hybrid zone 
with flaviceps (see notes under flaviceps). 
Specimens of pure flaviceps occur in towns 
near Ithaca. Until variation at the type 
locality is better understood, Kaston's ap- 
plication of the name exigua to the south- 
ern species will be maintained. 

To further complicate the application 
of the name exigua, two distinct color forms 
are found in the south that may very well 
represent distinct species. The typical (dull) 
form (Figs. 146-147, 329, 333) is more 
uniformly dark in carapace and append- 
ages; the striped form (Figs. 148, 149, 334, 
335) has much more extensive white mark- 
ings and distinct markings on the legs. The 
two forms appear perfectly discrete: all 
undamaged specimens are easily sorted to 



282 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



one or the other. However, no clear dif- 
ferences in genitaha have been found, both 
forms having the embolus and spermathe- 
cae as considered diagnostic for exigua. 
The spermathecal ducts may be more con- 
voluted in the dull form, but an adequate 
and convincing characterization of any dif- 
ference has proved elusive. The only con- 
vincing structural difference so far noted 
is in the width of the dark depression on 
the inner margin of the male's chelicera. 
The one striped male observed had a 
courtship display as in dull males, to the 
level of detail observed. Only the dull form 
has been seen from some coastal states 
(New York, New Jersey, North Carolina), 
while only the striped form has been seen 
from Missouri and Illinois. However, the 
two forms occur sympatrically in Virginia, 
Maryland, and Arkansas. In Virginia and 
Maryland, they have even been caught 
from conifer trees in the same field. In 
Massachusetts, there is variation in the 
amount of white markings but the varia- 
tion does not appear to sort itself into two 
discrete forms. Until stronger evidence is 
available to separate the forms, they will 
be kept under the name P. exigua. The 
name D. exiguus and apparently also D. 
virginis apply to the dull form. In the fol- 
lowing description the markings of the two 
forms will be described separately. 

Diagnosis. Males of exigua can be dis- 
tinguished from flavipedes and flaviceps 
by the mostly dark brown legs, the thin 
retrolateral ramus (Chamberlin, 1925a), 
and the more twisted prolateral ramus. The 
black spot on the chelicerae is much broad- 
er than in flavipedes. Females of exigua 
have more strongly convergent epigynal 
flaps and an extremely broad second curve 
of the internal ducts. 

Male. Palpus (Figs. 203, 330): Embolus 
divided deeply into two rami; retrolateral 
ramus thinner than prolateral; prolateral 
ramus strongly twisted. Markings: Dull 
form (Figs. 146, 329): Carapace relatively 
dark, area above first eye row brown. 
Cheek band much reduced, to streak be- 
side ALEs. Clypeus brown, hairs over- 



hanging chelicerae dark. Setae surround- 
ing AMEs white laterally 2:00-4:00, often 
medially 9:00-10:00; brown otherwise. 
Chelicerae yellow with front surface 
slightly concave and with black patch on 
inner margin shorter than in flaviceps but 
nonetheless deep, distinct, and wide; che- 
licera lacks prominent basal lateral brown 
spot. Palpus brown, basal segments not dis- 
tinctly paler. Patella, tibia, and cymbium 
lack white scales. Legs more or less uni- 
formly dark, without distinct longitudinal 
lines. Abdomen dorsum brown, with weak 
side bands. Striped form (Figs. 148, 334): 
Carapace with white spots between eyes 
of anterior eye row as in flavipedes. Cheek 
band thin but long. Clypeus with patch of 
white scales between AMEs, hairs over- 
hanging chelicerae white centrally and 
dark laterally. Setae surrounding AMEs 
white laterally 2:00-4:00, medially 7:00- 
11:00; brown otherwise. Chelicerae yellow 
with black depression on inner margin nar- 
rower than in dull form. Basal segments 
of palpus paler than cymbium. Palpus fe- 
mur, patella, and tibia with white scales; 
cymbium lacking white scales. First leg 
femur and patella dark ventrally but with 
white scales dorsally; tibia mostly dark es- 
pecially on anterior surface. Other legs pale 
with dark markings. Abdomen dorsum 
dark brown, with strong white side bands. 
Measurements: Body length 4.4(4.5)4.9 
mm; carapace length 1.9(2.0)2.2 mm, 
width/length 0.81(0.84)0.84; n = 53 from 
Massachusetts, Maryland, Virginia, Ken- 
tucky, and North Carolina. 

Female. Epigynum (Figs. 243, 331, 332, 
336, 337, 342): Flaps convergent, poste- 
riorly at about 45° rotation. Epigynal sur- 
face flat. Ducts most notable for the very 
broad second curve such that the anterior 
edge of this curve begins near anterior end 
of flap; flowerlike gland openings along 
anterior edge of curve. Markings: Dull 
form (Figs. 147, 333): Carapace brown, 
covered with transparent brown scales ex- 
cept for some beige to white scales. Like 
flavipedes, there are patches of pale scales 
behind and between eyes of anterior row; 



Pelegrina Jumping Spiders • Maddison 283 



scales surrounding AMEs dark dorsally. 
Clypeus densely covered with white scales. 
Abdomen almost uniform brown, with 
chevrons not so distinct as in flavipedes. 
Striped form (Figs. 149, 335): Carapace 
often darker than in dull form, covered 
with transparent brown scales except for 
some beige to white scales. Behind and 
between anterior eyes are patches of pale 
scales; scales surrounding AMEs dark dor- 
sally. Clypeus densely covered with white 
scales. Legs pale with some dark markings; 
first tibia dark as in males. Abdomen dark 
centrally, with contrasting pale side bands. 
Measurements: Body length 5.3(5.7)5.8 
mm; carapace length 2.0(2.1)2.2 mm, 
width/length 0.79(0.80)0.82; n - 59 (dull 
form) from Rowan Co., Kentucky. 

Courtship (76 observed from Shenan- 
doah Co., Virginia; Caroline and Mont- 
gomery Counties, Maryland; all dull form 
except one S striped from Montgomery 
Co.). With unusual alternate waving of 
palpi during crouch display. Raisedspread 
(n = 16, IS). Crouch (n = 10, 53): Body 
low (n = 3, 2(5) or raised (n = 1). Male 
walks in series with long pauses during 
which palpi are waved (n = 1), or walk is 
more or less continuous with constant pal- 
pus and leg waving and abdomen twitch- 
ing (n = 2, 1(5). First legs forward and 
parallel (n = 9, 4<5), slightly bowed (n = 2, 
2(5) or stretched forward (n = 7, 3(5), with 
tips almost touching (n = 3, 26) or not (n 
= 2, 1(5). First legs horizontal (n = 2, 1(5: 
apparently in intense display), or slightly 
raised (n = 6, 3(5), to ca. 30° (n = 2, 23). 
Tips of first legs moved side to side (both 
same direction) during leg waving (n = 5, 
33), in phase with palpus waving (n = 1). 
Palpi forward (n = 3, 33) and down (n = 
1) or slightly raised (n = 2, 23). Palpi waved 
alternately (n = 5, 33) up and down (n = 
5, 33) several times at about 2 left and 2 
right palpus waves per second for 2-5 sec 
(n = 1). Abdomen twitched (n = 3, 23) 
while palpi waved (n = 2, 23). Repertoires: 
23 raisedspread only; 53 raisedspread and 
crouch. 



Distribution (Map 14). Eastern United 
States except for far north. 

Records. Many specimens, especially in MCZ and 
AMNH, from: UNITED STATES (county records): 
Dull form: MASSACHUSETTS: Barnstable, Essex; 
CONNECTICUT: FairBeld, New Haven; NEW 
YORK: Nassau, Suffolk, Tompkins; NEW JERSEY: 
Burlington, Hunterdon, Ocean; WEST VIRGINIA: 
Jefferson; VIRGINIA: Augusta, Bedford, Brunswick, 
Essex, Fairfax, Norfolk, Page, Shenandoah, Wash- 
ington; MARYLAND: Caroline, Montgomery, Prince 
Georges; DISTRICT OF COLUMBIA: Woodridge; 
KENTUCKY: Rowan; TENNESSEE: Unicoi; NORTH 
CAROLINA: Burke, Craven, Durham, Rockingham 
SOUTH CAROLINA: Oconee; GEORGIA: Thomas 
ALABAMA: DeKalb, Tuscaloosa; MISSISSIPPI: Scott 
ARKANSAS: Bradley, Calhoun; TEXAS: San Augus- 
tine. Striped form: VIRGINIA: Augusta, Essex, 
Washington; MARYLAND: Georges, Montgomery; 
ALABAMA: Madison; ILLINOIS: Hardin; MISSOU- 
RI: Boone, Cole; ARKANSAS: Washington. Form not 
distinguished: MASSACHUSETTS: Barnstable, Es- 
sex, Middlesex, Norfolk, Suffolk. 

Natural History. Usually found on co- 
nifers, known from pines (8 records), ju- 
nipers (8 records), occasionally on other 
plants such as oak (1 record) and walnut 
(2 records). 

1 5 . Pelegrina montana 
(Emerton, 1891) new combination 

Figures 1, 154, 204, 244, 343-347; 

Map 16 

Dendryphantes montanus Emerton, 1891: 11. Types 
in MCZ 2$ 29 with labels "Dendryphantes mon- 
tanus [underlined in red], Mt. Washington, N. H." 
and "J H. Emerton Coll. , examined. G. & E. 
Peckham, 1909: 459, pi. 37, figs. 4, 4a-c, $9. Roew- 
er, 1954: 1213. Bonnet, 1956: 1396. 

Dendryphantes {Metaphidippus) montanus: — Pe- 
trunkevitch, 1911: 636. 

Metaphidippus montanus: — Chickering and Bacorn, 
1933: 526. Kaston, 1973: 115, figs. 37, 38, <39. 

Diagnosis. A large, dark northern and 
montane species. The embolus that ex- 
pands near tip and the rough epigynum 
with ridges behind the flaps are diagnostic. 

Male. Palpus (Figs. 204, 344): Embolus 
narrow near base and flares at tip, almost 
spoon-shaped, concave dorsally. Retrolat- 
eral ramus reduced. Small denticles cover 
the embolus surface basally. Markings 
(Figs. 1, 154, 343): Carapace with diffuse 



284 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



white side band and white forehead band. 
Cheek band very weak to absent. Clypeus 
brown; hairs overhanging cheUcerae dark. 
White forehead band contacts AMEs dor- 
sally 10:30-1:00. Chelicerae lacking pale 
scales. Cymbium lacking white scales. Ab- 
domen more or less uniformly dark dor- 
sally, much darker than insignis; anterior 
white spots absent and posterior reduced 
to thin lateral bars. Measurements: Body 
length 4.1(4.6)5.3 mm; carapace length 
2.0(2.1)2.5 mm, width/length 0.74(0.80) 
0.82; n = 5(5 from Colorado, Vermont, New 
Hampshire, and Quebec. 

Female. Epigynum (Figs. 244, 345, 346): 
Flaps short, convex, parallel or divergent. 
Most distinctive are the small but conspic- 
uous ridges (seen as dark streaks in Fig. 
346) immediately behind and lateral to 
flaps. Surface not raised into a bulge pos- 
teriorly. Notch usually deep, sometimes 
longer than flaps, often shaped like a 
rounded rectangle. First curve of duct very 
narrow; second curve proceeds obliquely 
posteriorly unlike most other members of 
genus. Markings (Fig. 347): Carapace dark, 
thinly to sparsely covered with white scales. 
Clypeus densely covered with white scales. 
Abdominal dorsum largely brown, medi- 
ally slightly paler, with only small paired 
white spots, without black spots; white lat- 
erally. Measurements: Body length 
5.5(5.9-6.9)7.1 mm; carapace length 
2.4(2.5)2.7 mm, width/length 0.75(0.78) 
0.79; n = 59 from Colorado, Vermont, and 
New Hampshire. 

Courtship (IS observed from Butte, 
Montana). Crouch (n = 6): Body low (n = 
4). First legs horizontal (n = 4) and parallel 
(n = 4); not waving (n = 6). Palpi tucked 
in to side (n = 4), down (n = 2); waved 
slightly (n = 6). Abdomen twitching oc- 
casionally (n = 6). 

Distrihution (Map 16). Newfoundland 
west to British Columbia, Yukon Territory 
south to Colorado in the west and New 
Hampshire and New York in the east. Kas- 
ton's (1973) report of a record from Illinois 
is probably based on a pair of insignis from 
Urbana in the AMNH identified by him 



in 1949 as montana; the male has the tip 
of both emboli broken (see Cutler, 1979) 
and thus resembles montana. 

Records. Most in MCZ, from: CANADA: YUKON: 
Tklo-Klut, 10 km E Old Crow (1<5 49), King Edward 
range N of Old Crow (1(5); NEWFOUNDLAND: In- 
dian River (23 19), Terra Nova National Park (13); 
QUEBEC: Anticosti, Fox Bay (23 19), Lake Mistassini, 
Ayikwapit Peninsula (13); MANITOBA: Cedar Lake 
(13 19), H. B. Railway 214 (13); ALBERTA: Edmon- 
ton (19), Waterton Lakes National Park, nr. Waterton 
Lake (13, WPM), Jasper (19); BRITISH COLUMBIA: 
Pink Mountain (39), Muskeg nr. Little Prairie (29). 
UNITED STATES (county records): NEW HAMP- 
SHIRE: Chesire (13), Coos (23 39); VERMONT: Chit- 
tenden (13 29); NEW YORK: Cattaraugus (23, 
AMNH), Essex (13 19, AMNH), Greene (23, AMNH); 
VIRGINIA: Giles (19); MONTANA: Beaverhead (39), 
Glacier (19), Jefferson (19), Park (19); IDAHO: Custer 
(13 49), Franklin (19); WYOMING: Albany (19), Te- 
ton (13 39); COLORADO: Boulder (69), Custer (13), 
Grand (13), Gunnison (13), Larimer (13); UTAH: Uin- 
ta (19), Wasatch (19); NEW MEXICO: Bernalillo (19); 
ALASKA: Shaw Creek, mile 289 Richardson Hwy 

as). 

Natural History. From birch, willow, 
poplar, and other deciduous bushes and 
trees beside streams and rivers and in bogs 
in British Columbia, Yukon, Montana (4 
records). In aspen-lodgepole pine meadow 
in Colorado (1 records). In Yukon below 
450 m elevation (1 record), Montana and 
Idaho 1,800-2,000 m (2 records), Colora- 
do, Utah, and Wyoming 2,400-2,900 m (9 
records). 

16. Pelegrina insignis 
(Banks, 1892) new combination 

Figures 128, 150, 151, 205, 245, 

348-353; Map 17 

Dendryphantes insignis Banks, 1892:74, pi. 5, figs. 
28, 28a, 9. Holotype in MCZ 19 with labels "Den- 
dryphantes insignis Bks type," "Ithaca, N. Y, ' and 
"Nathan Banks Coll.", examined. 

Metaphidippus niontanus: — Chickering, 1944: 176, 
figs. 70-73, 39, 

Metaphidippus octavus: — Kaston, 1945: 10. 

Metaphidippus insignis: — Kaston, 1948: 476, figs. 
1750-1752, 39. Kaston, 1973: 114, figs. 34-36, 39. 
Cutler, 1979: 279-274. 

Dendryphantes capitatus: — Roewer, 1954: 1202. 

Metaphidippus clematus: — Levi and Levi, 1951, in 
part: 232, possibly figure 39 (paratype series in- 
cludes insignis females). 



Pelegrina Jumping Spiders • Maddison 285 



Metapl dippus capitatus: — Bonnet, 1957; 2810, in 
part. 

Notes on Synonymy. The specimen cit- 
ed above may be labeled incorrectly as 
holotype, for Banks's description and fig- 
ures seem to depict females of proterva 
because of the white pubescence (not yel- 
low as in what we now call "insignis"), 
the red-ringed leg segments (not uniform- 
ly yellowish), the large reddish spots on 
the abdomen (not smaller and black), and 
the long, parallel epigynal flaps (not short 
and divergent). Nonetheless, since the 
specimen marked holotype is clearly of the 
species we now call "insignis," it seems 
best to leave the matter as it stands and 
presume that Banks was somewhat erro- 
neous in this description. 

Diagnosis. Notable for the yellowish 
markings with strong black spots on the 
abdomen. Pelegrina montana and cle- 
mata are similar but the long spatulate 
embolus and epigynal topography of in- 
signis are distinctive. 

Male. Palpus (Figs. 205, 349, 350): Em- 
bolus flares slightly in distal half; usually 
bent slightly toward the retrolateral. Ret- 
rolateral ramus reduced. The tip of the 
embolus is very thin and sometimes par- 
tially or entirely broken off (see Cutler, 
1979). Small denticles cover the embolus 
surface basal to opening. Markings (Figs. 
150, 348): Carapace often suffused with 
brassy scales dorsally, with patches of white 
between the fovea and posterior eyes in 
addition to ample white side bands and V 
mark behind AMEs. Cheek band weak. 
Clypeus brown; hairs overhanging chelic- 
erae dark. White forehead band contacts 
AMEs dorsally 10:30-12:30. Chelicerae 
with small patch of yellowish scales me- 
dially near base. Palpus femur covered with 
white scales; more distal segments darker 
and usually without white scales. Cym- 
bium with none to a few white scales. Ab- 
domen shows paired black spots of female; 
anterior paired medial white spots distinct; 
brassy sheen medially. Measurements: 
Body length 3.4(3.6)4.1 mm; carapace 
length 1.7(1.8)2.1 mm, width/length 



0.77(0.77)0.79; n = 53 from New Bruns- 
wick, Massachusetts, Minnesota, and Sas- 
katchewan. 

Female. Epigynum (Figs. 245, 351 , 352): 
Flaps short and divergent or parallel. Sur- 
face behind flaps raised into bulge only 
medially and posteriorly, so that surface 
rises gradually behind flaps, or if surface 
rises abruptly it does so at some distance 
behind flaps. Posterior bulge often consid- 
erably higher than level of flaps. Notch 
triangular and sharp. First curve of duct 
narrow; second curve proceeds obliquely 
posteriorly or medially. Markings (Figs. 
151, 353): Carapace covered densely with 
yellowish white scales. Clypeus densely 
covered with yellow scales. Legs more or 
less uniformly yellow. Abdomen dorsum 
with prominent paired black spots, oth- 
erwise yellow-brown to red-brown with 
paired spots and lateral markings of yellow 
and white scales. Measurements: Body 
length 3.8(4.1)5.3 mm; carapace length 
1.7(1.9)2.1 mm, width/length 0.74(0.78) 
0.79; n = 59 from Saskatchewan and Min- 
nesota. 

Chromosomes. 2nS = 26 acrocentrics + 
XXO {IS from Taber, Alberta). 

Courtship (26 observed from Taber, Al- 
berta, and North Battleford, Saskatche- 
wan). With triangular leg position during 
crouch display. Raisedspread (n = 1). 
Crouch (Fig. 128; n = 8, 23): Body at low 
(n = 5, 15) to normal (n = 3, \$) height. 
First legs horizontal and forward (n = 8, 
23), spread slightly (n = 3, 13) or bowed 
with tips close (n = 5, 13). First legs ex- 
tended forward on series and waved slight- 
ly; on pauses, first legs held back so as to 
form a roughly triangular shape as in ver- 
ecunda and aeneola (n = 5, 13), though 
the leading leg is sometimes held in during 
series (n = 3, 13). Palpi down (n = 8, 23) 
and tucked in (n = 5, 13); waved up and 
down (n = 8, 23), on series (n = 5, 13). 
Repertoires: 13 crouch only; 13 raised- 
spread and crouch. 

Distribution (Map 17). New Brunswick 
west to Alberta, south to New York and 
Colorado. 



286 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



Records. In AMNH, MCZ, and WPM from: CAN- 
ADA: NEW BRUNSWICK: nr, Chipman; ONTAR- 
IO: Barrie; 5 km S of Richmond near Ottawa; Cam- 
bridge; SASKATCHEWAN: Wells Lake nr. Alberta 
border; North Battleford; Estevan; ALBERTA: Islay; 
8 km W of Writing-on-Stone Provincial Park; Med- 
icine Hat; betvv. Cereal and Oven; Taber. UNITED 
STATES (county records): MAINE: Hancock, Lin- 
coln; VERMONT: Windham; MASSACHUSETTS: 
Middlesex, Norfolk, Worcester; CONNECTICUT: 
Fairfield, Hartford, New Haven, New London, Tol- 
land; NEW YORK: Seneca, Tompkins; NEW JER- 
SEY: Bergen, Ocean; MICHIGAN: Hillsdale, In- 
gham, Lenawee, Midland, Oakland; WISCONSIN: 
Dane; ILLINOIS: Champaign; MINNESOTA: Hen- 
nepin, Olmsted, Polk, Ramsey; IOWA: Boone; 
NORTH DAKOTA: Burleigh, Nelson, Ransom; 
COLORADO: Fremont. 

Natural History. On Chamaedaphne, 
Betula, and other vegetation in bogs in 
New Brunswick and Ontario; in oldfields 
and prairies in Ontario, Alberta, Minne- 
sota, and Wisconsin. In an open habitat in 
North Battleford, Saskatchewan, P. pro- 
terva was common and restricted to the 
taller shrubs (taller than 1 m), whereas 
P.insignis was common and restricted to 
the short shrubs (shorter than 0.5 m) among 
the grasses. In habitats in Alberta and Mas- 
sachusetts, P. insignis has also been found 
on short shrubs and herbs. In late June near 
Templeton, Massachusetts, numerous fe- 
males were found with egg sacs in nests in 
living but curled leaves of goldenrod and 
other herbs. 

17. Pelegrina chaimona new species 
Figures 206, 354-358; Map 18 

Metaphidippus n. sp. nr. verecundus: — Jung and Roth, 
1974: 33 (specimens identified by W. J. Gertsch, 
examined). 

Holotype male in AMNH with label "ARIZONA: 5 
mi [8 km] W of Portal, Cochise County, SWRS 
[Southwestern Research Station], June 9, 1968, V. 
Roth." 

Etymology. An arbitrary combination 
of letters, to be treated as an adjective. 

Diagnosis. Male closely resembles P. 
montana, but erect portion of embolus is 
more or less parallel-sided. The embolus is 
shorter and stouter than that of insignis 
and has a distinct retrolateral ramus. The 
embolus bears some resemblance to that 



of dithalea, but the rami are closer to- 
gether (Figs. 206, 192). The female of 
chaimona is poorly known. 

Male. Palpus (Figs. 206, 355): Embolus 
rectangular, widening gradually to base on 
retromargin so that there is no distinct an- 
gle between erect portion and base. Rami 
small and not much separated. Erect por- 
tion of embolus arises more retrolaterally 
than in dithalea. Embolus surface with 
small denticles basally. Markings (Fig. 
354): Cheek band generally weak. Clypeus 
brown; hairs overhanging chelicerae dark 
except white medially. White forehead 
band contacts AMEs dorsally 10:30-12:30. 
Chelicerae with small medial patch of 
white scales. Cymbium with none to a few 
white scales. Measurements: Body length 
3.9(4.1-4.2)4.3 mm; carapace length 
1.6(2.0)2.1 mm, width/length 0.77(0.78) 
0.80; n = 55 from Chiricahua Mtns., Ari- 
zona. 

Female. Epigynum (Figs. 356, 357): 
Flaps fairly short, convergent or parallel. 
Surface raised just inside flaps midway 
along their length; behind flaps surface low, 
rises into prominent mound posteriorly. 
First curve of duct narrow; second curve 
proceeds medially. Markings (Fig. 358): 
Carapace with white scales dorsally. Clyp- 
eus densely covered with white scales. Ab- 
domen pale laterally and dark centrally 
with paired white spots. Measurements: 
Body length 4.6, 5.0, 54. mm; carapace 
length 1.8, 1.8, 2.1 mm, width/length 0.78, 
0.78, 0.79; n = 32 from Arizona and Chi- 
huahua. 

Male/Female Matching. With the fe- 
males of chalceola, dithalea, and kastoni 
well associated with males, there remain 
two males (chaimona and huachuca) and 
two females unmatched in southern Ari- 
zona. The matching of these is in doubt, 
but the following evidence supports the 
matching of the females described above 
with the male of chaimona: the flaps of 
the female are of typical robustness, as 
would be expected to match the embolus 
of chaimona; the first curve of the epi- 
gynal duct is narrow like the related in- 



Pelecrina Jumping Spiders • Maddison 287 



signis and montana; the cephalic plate is 
somewhat rugose as in males; these females 
have only been found in extreme eastern 
Arizona and south, as have the males. 

Distribution (Map 18). Southeastern Ar- 
izona, Chihuahua, and Nuevo Leon. 

Records. UNITED STATES: ARIZONA: Cochise 
Co.: Chiricahua Mtns.: Southwestern Research Sta- 
tion 8 km W of Portal (76 12, AMNH), Cottonwood 
Creek, Rucker Canyon (15, AMNH), Cave Creek 
Canyon (I.?, AMNH). MEXICO: CHIHUAHUA: Pri- 
mavera (1<?, AMNH), San Jose Babicora (26, AMNH), 
summit NE of San Jose Babicora (19, AMNH), Ma- 
dera (19, AMNH); NUEVO LEON: Cerro Potosi (1<5, 
MCZ). 

Natural History. Collected from Chrys- 
othamnus (IS) and sweeping herbs (13) at 
elevations from 1,650 to 3,200 m. Males 
collected in May (13), June (63), July (33); 
females collected in July (39). Jung and 
Roth (1974) collected this species in their 
zone 2 in the Chiricahua Mountains (1,460- 
1,700 m elevation). 

1 8. Pelegrina clemata 

(Levi & Levi, 1951) new combination 

Figures 152, 153, 207, 246, 

359-364; Map 19 

Metaphidippus clematus Levi and Levi, 1951: 232, 
figs. 37, 39, 40, 42 (fig. 39 may be insignis), 69. 
Holotype 6 and paratype 9 in AMNH with label 
"Metaphidippus clematus Levi 9 6, 6 holotype 9 
allotype. Medicine Hat, Alta., Aug. Carr," exam- 
ined. 

Dendryphantes clematus: — Roewer, 1954: 1209. 

Diagnosis. Markings whitish instead of 
yellow distinguish it from insignis. Em- 
bolus narrower at tip than in montana, 
insignis, chaimona, and dithalea, much as 
in baila and chalceola, though these have 
different markings and a wider carapace 
than clemata. The tegulum's prominent 
bulge is unusual. The dark epigynum with 
a prominent shiny mound behind the flaps 
is distinctive. 

Male. Palpus (Figs. 207, 360, 361): Erect 
portion of embolus straight, tapering or of 
equal width from base to tip, with small 
rami at tip; prolateral ramus is more prom- 
inent than retrolateral ramus, which is 
small and not projecting retrolaterally. 



Embolus lacks the spoonlike dorsal con- 
cavity present in the montana group. Te- 
gulum swollen prolaterally into prominent 
bump. Markings (Figs. 152, 359): Cara- 
pace with white side bands and white V 
mark behind AMEs; sometimes with a 
weak marginal white band. Cheek band 
weak. Clypeus brown; hairs overhanging 
chelicerae dark. White forehead band con- 
tacts AMEs dorsally 10:30-12:30. Chelic- 
erae occasionally with a few white scales 
medially. Palpus femur and distal seg- 
ments including cymbium all with at least 
some white scales. Abdomen without cen- 
tral paired white spots; some males show 
paired black spots. Measurements: Body 
length 3.8(4.0)4.2 mm; carapace length 
1.9(2.0)2.2 mm, width/length 0.76(0.77) 
0.79; n = 53 from Outlook, Saskatchewan. 

Female. Epigynum (Figs. 246, 362, 363): 
Dark and shiny. Flaps convergent. Entire 
area behind flaps raised into a bulge, so 
that surface rises abruptly behind flaps. 
Notch triangular or rounded and short, of- 
ten only half the length of flaps. First curve 
of duct narrow; second curve goes medi- 
ally. Markings (Figs. 153, 364): Carapace 
covered with white scales. Clypeus densely 
covered with white scales. Abdomen dor- 
sum brown with white paired spots and 
lateral markings; usually with paired black 
spots, though not so distinct as insignis. 
The first three pairs of white spots may be 
joined to form two longitudinal bands. 
Measurements: Body length 4.7(5.8)5.8 
mm; carapace length 1.9(2.1)2.1 mm, 
width/length 0.73(0.76)0.79; n = 52 from 
Outlook, Saskatchewan. 

Chromosomes. 2n3 = 26 acrocentrics + 
XXO (13 from Richfield, Utah). 

Courtship (143 observed from Outlook, 
Saskatchewan; Morrin, Alberta; and Piute 
Co., Utah). Raise dspre ad (n = 21, 113). 
Crouch (n = 29, 93): Body at normal height 
(n = 8, 33) to low (n = 2, 13). First legs 
forward and horizontal (n = 15, 73) or 
slightly raised (n = 8, 33), or raised to 45° 
with femur low (n = 3, 23); nearly parallel 
(n = 5, 23), or spread fairly wide (n = 4, 
23). First legs not waved (n = 11, 43). Palpi 



288 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



down (n = 23, 8S) and beside chelicerae 
(n = 3, 16) or curled under (n = 5, 15); 
flickered during series (n = 23, 83), vig- 
orously (n = 5, 1$) up and down (n = 4, 
2(5), still on pause (n = 4, 2(5). Abdomen 
twitched (n = 15, 5(5) during series (n = 9, 
36), or perhaps at end of series (n = 3, 16), 
still on pause (n = 4, 26); twitched contin- 
uously when walking to mount (n = 1). 
Repertoires: 36 raisedspread only; 26 
crouch only; 86 raisedspread and crouch. 
Distribution (Map 19). Western Canada 
and United States. 

Records. In MCZ, AMNH, UWBM, MSU, and WPM 
from: CANADA: SASKATCHEWAN: Outlook; AL- 
BERTA: Medicine Hat; Morrin Recreation Area on 
Red Deer River; BRITISH COLUMBIA: Bull River, 
ca. 115.5° W, 49.5° N; Cranbrook. UNITED STATES 
(county records): MONTANA: Sanders; WYOMING: 
Carbon, Lincoln, Park, Sheridan, Teton; COLORA- 
DO: Dolores, Gunnison, Huerfano, Lake, Larimer, 
Saguache; UTAH: Box Elder, Cache, Millard, Piute, 
Rich, Sevier, Summit, Utah, Weber; IDAHO: Bear 
Lake, Blaine, Canyon, Cassia, Fremont, Gem, Onei- 
da, Payette; NEVADA: Elko; NEW MEXICO: Taos; 
WASHINGTON: Grant, Kittitas, Yakima; ORE- 
GON: Baker, Deschutes, Grant, Harney, Malheur; 
CALIFORNIA: Contra Costa, Eldorado, Los Angeles, 
Mono, Placer, Sierra, Siskiyou. 

Natural History. From sagebrush (Ar- 
temisia) in Washington, Oregon, Colora- 
do, Utah, and Wyoming (10 records). Also 
taken from field vegetation (Washington), 
Purshia (Oregon), Chrysothamnus (Cali- 
fornia), Haplopappus, and Sarcobatus 
(Utah), 1 record each. 

19. Pelegrina aeneola 
(Curtis, 1892) new combination 

Figures 156, 157, 208, 209, 247, 

365-377; Map 21 

Dendryphantes aeneoliis Curtis, 1892: 332. Types in 
MCZ 3(5, 62 with labels "Dendryphantes aeneolus 
Curtis 1892. California. Co-type. S 9" (label is orig- 
inal; handwritten, probably by Elizabeth Peckham) 
and "G. W. Peckham Coll.", examined. Curtis (1892: 
335) implies that the type locality is the San Fran- 
cisco Bay area. G. & E. Peckham, 1909: 468, pi. 
36, figs. 1-lb, and pi. 38, 6, 6a, <?9. Roewer, 1954: 
1205. 

Dendrijphantes bifida Banks, 1895: 96. Types in MCZ 
22, 3 pS with labels "Dendryphantes bifida Bks. 
type", "Olympia, Wash" and "Nathan Banks Coll.", 
examined. This type series is incomplete, for Banks 
described the adult male. 



Dendryphantes (Metaphidippus) aeneolus: — Pe- 
trunkevitch, 1911: 622. 

Dendryphantes uteanus Chamberlin and Gertsch, 
1929: 110, figs. 50, 51, S. Holotype in AMNH 1<3 
with labels "Dendryphantes uteanus, S holotype," 
"Metaphidippus aeneolus (Curtis), 6 wlll.n40, 
HOLOTYPE Dendryphantes uteanus Chamb. & 
Gert.", "Utah: Lamb's Can 6-10-28 W. J. G.", and 
"28Ff. N40:W111," examined. Roewer, 1954: 1216. 
Bonnet, 1956: 1402. NEW SYNONYMY. 

Metaphidippus aeneolus: — Chamberlin and Ivie, 
1941: 26. Bonnet, 1957: 2809, 

Diagnosis. A common species of the 
western United States and Canada. Males 
are notable for their dark markings, with 
reduced white scales on the carapace. The 
rami of the embolus are more widely sep- 
arated than in clemata, balia, or chalceola. 
Females can be identified by carapace and 
abdominal markings, the epigynal topog- 
raphy and angled flaps. The epigynal sur- 
face rises more abruptly behind the flaps 
than in balia, but not so raised in a bulge 
as in insignis or clemata. The epigynal 
flaps are much more robust than in the 
sympatric Metaphidippus mannii, from 
which aeneola also differs in markings, in- 
cluding having white scales between AMEs. 

Male. Palpus (Figs. 208, 209, 366-372): 
Embolus leaning toward the retrolateral, 
broadest at tip; two rami widely separated. 
Markings (Figs. 156, 365): Carapace dark 
and shiny, with few or no white scales 
except beneath ALEs and small eyes (oc- 
casional SS have white side bands, though 
usually sparse). Clypeus brown; hairs over- 
hanging chelicerae dark brown. Forehead 
band absent; setae ringing AMEs white at 
least laterally 2:00-3:00; otherwise, thin 
scales brown to white. Chelicerae lack pale 
scales. Cymbium lacking white scales. Ab- 
domen dark; white side bands often faint 
or absent posteriorly. Measurements: Body 
length 4.2(4.3)4.7 mm; carapace length 
2.0(2.1)2.2 mm, width/length 0.75(0.79) 
0.81; n = 76 from Oregon, Nevada, and 
Utah. 

Female. Epigynum (Figs. 247, 373-376): 
Flap with inner edge angled where the 
posterior half of flap bends down into de- 
pression. Surface rises abruptly, almost im- 
mediately posterior to the flaps, but pos- 



Pelegrina Jumping Spiders • Maddison 289 



terior portion of epigynum is fairly flat. 
Notch often triangular with a sharp an- 
terior point, but many 99 have rounded 
notch. First curve of duct narrow; second 
curve proceeds medially. Markings (Figs. 
157, 377): Carapace only thinly covered 
with white scales; bronze scales also es- 
pecially on cephalic plate. Most females 
with inverted T marking on cephalic plate 
consisting of white band of scales starting 
between AMEs, proceeding posteriorly, 
then spreading laterally to behind small 
eyes. Otherwise, bronze behind AMEs and 
ALEs, and bronze between posterior eyes. 
Clypeus densely covered with white scales, 
paler between AMEs than in balia and 
mannii, which have orange scales. Abdo- 
men white markings usually small except 
often large central pale spot. Background 
dark, often bronze or gray, occasionally 
with paired dark spots on either side of 
the midline. Measurements: Body length 
4.5(4.8)6.1 mm; carapace length 2.0(2.1)2.3 
mm, width/length 0.77(0.79)0.85; n = 79 
from Oregon and Arizona. 

Geographical Variation. In western 66 
(aeneola proper, British Columbia, Wash- 
ington, Oregon, California), embolus nar- 
row with prolateral face gently curved or 
slightly bent (Fig. 208); in eastern 66 (form 
uteanus. South Dakota, Montana, Wyo- 
ming, Colorado, Utah, New Mexico, Ari- 
zona), embolus is wider with prominent 
angle on prolateral face just basal to the 
opening, and rami more divergent (Fig. 
209). The few specimens available from 
the intermediate area (Idaho, Nevada, and 
southeastern California) and occasional 
specimens from Oregon show some inter- 
gradation. No other differences in 66 and 
none in 99 have been found between west- 
ern and eastern populations; hence, I con- 
sider them conspecific. 66 from the Colum- 
bia basin of Washington, some from south- 
ern California, and occasional males from 
Oregon and northern California have ex- 
tensive but not very dense white side bands 
on the carapace. 

Chromosomes. 2n6 = 26 acrocentrics + 
XXO (23 from Apple Canyon, Riverside 
Co., California). 



Courtship (93 observed from Yakima 
and Kittitas Counties, Washington, and 
Riverside Co., California). First legs make 
triangular shape during crouch display, as 
in P. verecunda. Raisedspread (n = 21, 
53). Crouch (n = 22, 83): Body low (n = 
6, 33). First legs horizontal (n = 17, 73) but 
femora held back and to sides and distal 
segments pointed forward and with tarsi 
nearly touching so as to make a triangle 
shape (n ^ 12, 53), though occasionally 
femora held forward and tips not touching 
(n = 3, 13). First legs flickered on series (n 
= 9, 43), but only slightly (n = 3, 13) or 
not waved (n = 7, 33); also flickered when 
very close (n = 6, 43) at which time legs 
extended (n = 4, 33). Palpi tucked in (n = 
6, 23), flickering on series (n = 6, 23) or 
when male very close (n = 2, 23). Reper- 
toires: 13 raisedspread only; 43 crouch only; 
43 raisedspread and crouch. 

Distribution (Map 21). Western United 
States, extending into Canada and Mexico. 

Records. Many specimens, especially in MCZ, 
AMNH, UWBM,'and UCB, from: CANADA: BRIT- 
ISH COLUMBIA: Alice Lake Province Park; Creston; 
Furry Creek; Massett; Victoria; Wellington, Quali- 
cum, Nanaimo; ALBERTA: Waterton Lakes National 
Park. UNITED STATES (county records): SOUTH 
DAKOTA: Custer, Horsethief, Jackson, Pennington; 
MONTANA: Flathead, Lewis and Clark, Park; IDA- 
HO: Bear Lake, Boise, Bonner, Franklin, Latah, 
Oneida, Payette, Washington; WYOMING: Sheri- 
dan; COLORADO: Alamosa, Custer, Douglas, Fre- 
mont, Hinsdale, Juab, La Plata, Larimer, Logan, 
Montezuma, Utah; UTAH: Box Elder, Cache, Davis, 
Grand, Juab, Piute, Salt Lake, Uinta, Weber; NE- 
VADA: Washoe; NEW MEXICO: BernaHllo, Lin- 
coln, Otero, Rio Arriba, Sandoval, San Miguel, Santa 
Fe, Taos, Torrance, Valencia; ARIZONA: Apache, 
Cochise, Coconino, Graham, Mohave, Yavapai; 
WASHINGTON: Asotin, Chelan, Clallam, Columbia, 
Douglas, Grant, Grays Harbor, Jefferson, King, Kit- 
titas, Pierce, San Juan, Skagit, Skamania, Snohomish, 
Stevens, Thurston, Walla Walla, Yakima; OREGON; 
Benton, Deschutes, Douglas, Grant, Harney, Jackson, 
Jefferson, Josephine, Klamath, Lake, Lane, Marion, 
Multinomah, Umatilla, Union, Wallowa; CALIFOR- 
NIA: Alameda, Contra Costa, El Dorado, Kern, Las- 
sen, Los Angeles, Marin, Mendocino, Modoc, Mon- 
terey, Nevada, Orange, Plumas, Riverside, San Ber- 
nardino, San Diego, San Mateo, Santa Barbara, Santa 
Clara, Santa Cruz, Santa Cruz Island, Shasta, Sierra, 
Siskivou, Trinitv, Tulare, Ventura, Yuba. MEXICO: 
BAJA CALIFORNIA DEL NORTE: Parque Nacion- 
al Sierra San Pedro Martin. 



290 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



Natural History. Habitat: In Washing- 
ton, collected from pines including Pinus 
ponder osa, understory of riparian poplar 
woodland, understory ferns, Ceanothus, 
alders in bog, lakeside marsh, houses; at 
elevations of less than 50 m to more than 
1,700 m. More than 15 aduhs males and 
females were recovered by D. H. Mann 
and others on snow at 1,700-3,000 m, ap- 
parently having ballooned (Pierce Co., 
Washington). In Oregon, commonly col- 
lected from Abies grandis and Pinus pon- 
derosa; also from Pseudotsuga, Picea, Pi- 
nus contorta, hemlock, alder, Salix, brack- 
en fern, Calocedrus, Taxus, oak, Ceano- 
thus, and Larix occidentalis. Curtis (1892) 
reported it common in San Francisco area 
gardens on honeysuckle, rose bushes, live 
oaks, and Laurestina. In Nevada, beating 
pinyon pine; in Arizona on pine. Life cy- 
cle: In Oregon's Malheur National Forest, 
B. Fichter and A. Moldenke collected 44<5 
779 from 12 to 17 June 1982, 66 259 from 
18 to 23 July 1982, and 819 from 20 to 29 
September 1982. In the San Francisco area, 
"males and females appear as adults as 
early as April, but the former become rare 
after the first of June and the latter after 
the first of September. The females begin 
laying eggs in May" (Curtis, 1892: 335). 
Curtis reported one or two egg sacs per 
females with about 50 eggs that hatched 
on average in 25 days. In Los Angeles 
County, California, the 20<55 examined by 
me were collected from November through 
June; the 2199 from February through June 
plus two in September. Behavior: Curtis 
(1892) described the spider's entrance to 
and defense of its retreat, its ballooning, 
and its reaction to a sluggish pet lizard. 
Land (1969a, b, 1971) investigated visual 
behavior and eye structure and function 
of P. aeneola. 

20. Pelegrina balia new species 
Figures 210, 248, 378-382; Map 20 

Holotype male and paratype female in UCB with 
label "CA[lifornia]: S[an]ta. Barbara Co., Ballinger 
Cyn., 17 mi, [27 km] SE. New Cuyama, el. 3000' 
[915 m], V-9-1980, C. E. Griswold." 



Etymology. After the Greek adjective 
balios, meaning "dappled" (Woods, 1966). 

Diagnosis. A large western species with 
light brown and beige markings. The male 
can be identified by the broad side bands 
of the carapace and the flange on the fang. 
The embolus is narrow, very similar to that 
of chalceola, but it leans slightly and the 
erect portion broadens gradually into base. 
The tibial apophysis of balia usually points 
more distally than in chalceola. The fe- 
male can be identified by the spotted ab- 
domen and the epigynum, which differs 
from that of the sympatric aeneola in hav- 
ing the surface rise gradually behind the 
flaps. 

Male. Palpus (Figs. 210, 379): Embolus 
narrow and tall, leaning slightly to retro- 
lateral; erect portion broadens gradually 
into base. Retrolateral ramus points retro- 
laterally. Retrolateral edge of embolic base 
with membranous fold, unlike chalceola. 
Chelicerae robust and divergent; fang with 
pronounced flange on cutting edge (Fig. 
378, arrow). Markings (Fig. 378): unlike 
most other Pelegrina males in having 
markings more spotted than striped. Car- 
apace with distinctive, broad creamy white 
side bands. Cheek band weak and broad. 
Clypeus brown; hairs overhanging chelic- 
erae dark with some white medially. White 
forehead band fails to contact AMEs dor- 
sally, which are ringed by dark or at most 
thin white setae. Chelicerae lacking pale 
scales. Cymbium lacking white scales. Ab- 
domen either mottled as in female, with 
conspicuous basal band and second lateral 
bar, or with side spots fused into very wide 
cream side bands. Measurements: Body 
length 3.8(4.3)5.1 mm; carapace length 
1.8(2.1-2.3)2.3 mm, width/length 0.79 
(0.81-0.82)0.85; n = 66 from California, 
Oregon, and Washington. 

Female. Epigynum (Figs. 248, 380, 381): 
Flaps usually light brown, convergent, 
though sometimes dark. Epigynal surface 
mostly concave except or bump just medial 
to flaps; surface rises behind flaps gradu- 
ally to a medial and posterior bulge. First 
curve of duct pale; second curve goes me- 



Pelegrina Jumping Spiders • Maddison 291 



dially; third curve smooth on inner sur- 
face. Markings (Fig. 382): Carapace cov- 
ered densely with yellow-white scales, of- 
ten with dark streak on thorax side. Clyp- 
eus densely covered with white, between 
AMEs orange or tan (white in some Ore- 
gon 29). Abdomen mottled with large pale 
spots: with pale basal band fused to first 
lateral bar; second oblique bar swollen; 
posterior lateral bars inconspicuous; cen- 
tral pale spots round, edge or connected 
with dark brown. Measurements: Body 
length 4.7(5.2)6.1 mm; carapace length 
2.1(2.2)2.3 mm, width/length 0.79(0.80) 
0.81; n = 52 from California. 

Male/Female Matching. This match- 
ing is indicated by the similarity in robust 
carapace and mottled markings and by co- 
collecting. 

Distribution (Map 20). California, north 
to Washington and east to Arizona and 
Colorado. 

Records. About 70 specimens in AMNH, UCB, and 
MCZ from; UNITED STATES (county records): 
WASHINGTON: Spokane; OREGON: Baker, Des- 
chutes, Harney, Jackson, Klamath, Lane, Wheeler; 
CALIFORNIA: El Dorado, Fresno, Inyo, Kern, Las- 
sen, Los Angeles, Mariposa, Mendocino, Modoc, Mono, 
Plumas, Riverside, San Bernardino, Santa Barbara, 
Shasta, Siskiyou, Sonoma, Stanislaus, Trinity, Ven- 
tura; COLORADO: Mesa; ARIZONA: Yavapai. 

Natural History. On Juniperus occi- 
dentalis (3 records), Pseudotsuga (2 re- 
cords), Cupressus macnabiana (1 record), 
Abies (1 record), and Calocedrus (1 rec- 
ord) in Oregon and northern California. 
From juniper woodland in California (5 
records). Over the entire range, S6 were 
collected in April, 10 in May, 3 in June 
and 1 in September. 

21 . Pelegrina chalceola new species 
Figures 139, 211, 383-387; iVIap 20 

Metaphidippus n. sp. nr. montaniis: — Jung and Roth, 
1974: 33 (specimens identified by W. J. Gertsch, 
examined). 

Holotype male and several immatures in MCZ with 
label "ARIZONA: Santa Cruz Co., upper Madera 
Canyon, Santa Rita Mtns., ca. 5500 ft [1,680 m]. 13 
Aug' 1983. W. Maddison 83-158 oak woodland, 
beating oaks." 



Etymology. An arbitrary combination 
of letters designed to resemble the name 
of the similar species P. aeneola both in 
structure and in referring to the bronze 
color (Greek, chalceos). 

Diagnosis. A dark, shiny southwestern 
species resembling aeneola and balia. The 
narrow, tall embolus is much like that of 
balia, from which chalceola differs by the 
lack of the pronounced flange on the male 
fang, and the much darker body with a 
bronze sheen in both males and females. 

Male. Palpus (Figs. 211,384): Erect por- 
tion of embolus narrow and tall, straight, 
usually broadens abruptly into embolic 
base so as to leave angle between erect 
portion and base. Retrolateral ramus points 
retrolaterally. Retrolateral edge of embolic 
base is simple and schlerotized, lacking the 
fold seen in balia. Markings (Figs. 139, 
383): Carapace dark, with narrow white 
side bands often reduced behind posterior 
eyes. Cephalic area with transparent 
bronze scales. Forehead lacks white band, 
though in some males there is small patch 
of pale scales between and behind AMEs. 
Cheek band weak to almost absent. Clyp- 
eus brown; hairs overhanging chelicerae 
white centrally and tan laterally, to all 
brown. Setae ringing AMEs brown dor- 
sally. Chelicerae robust but vertical, with 
none to a few pale scales medially. Cym- 
bium lacking pale scales. Third leg dark 
on distal % to entirely dark. Abdomen dor- 
sally brown with paired black spots and 
thin white side bands; third and fourth 
pairs of white spots when present are lat- 
erally directed bars. Measurements: Body 
length 3.9(4.0)4.9 mm; carapace length 
1.8(2.1)2.3 mm, width/length 0.80(0.82) 
0.83; n = 5(5 from Arizona. 

Female. Epigynum (Figs. 385, 386): 
Flaps short, slightly convergent, posteri- 
orly in concavity. Epigynal surface gently 
convex behind flaps. Second curve of duct 
goes medially; third curve with rough in- 
ner surface. Markings (Fig. 387): Brown 
with bronze sheen. Carapace covered with 
reflective white to tan scales. Spots of pale 
scales between anterior eyes similar to fla- 



292 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



vipedes. Clypeus densely covered with 
white scales. AME eye ring darkest dor- 
sally, tan to brown, as in male. Abdomen 
with large paired dark spots. Measure- 
ments: Body length 4.4, 4.4, 4.7, 4.7 mm; 
carapace length 2.1, 2.2, 2.3, 2.3 mm, 
width/length 0.78, 0.78, 0.81; n = 42 from 
Arizona. 

Male/Female Matching. Among Ari- 
zonan species, the female and male share 
a distinctive wide box-shaped carapace, 
dark bronzed appearance, and markings 
on abdomen. The AME ring dark above 
and light elsewhere also unites male and 
female. The underside of the abdomen is 
also fairly pale, distinguishing it from the 
female of hiiachuca. 

Geographical Variation. Two male Pe- 
legrina from Durango tentatively identi- 
fied as chalceola differ from specimens 
from the United States in being large (body 
length 5.2,5.2 mm; carapace length 2.5,2.5 
mm, width/length 0.82,0.83), with em- 
bolus shorter and leaning more to the ret- 
rolateral in the distal half, and with a bump 
on the side of the chelicera almost as in 
the mannii group but not so well devel- 
oped. 

Courtship (15 observed from Madera 
Canyon, Arizona). Raisedspread (n = 6): 
Carapace high (n = 6); abdomen down (n 
= 6) and trailing (n = 2). First legs raised, 
forward, spread (n = 6), but legs moved 
to more parallel as he got closer (n = 4), 
waved little if at all (n = 6). Palpi down, 
waving little if at all (n = 6). Male pro- 
ceeded directly from raisedspread to 
reaching to touch the female, without go- 
ing through a crouch display (n = 4). 

Distribution (Map 20). Southern Ari- 
zona east to southern Illinois. 



Records. UNITED STATES: ARIZONA: Cochise 
Co.: Chiricahua Mtns., Southwestern Research Sta- 
tion (8.5, AMNH); Chiricahua Mtns. (19, AMNH); Hu- 
achuca Mtns., Montezuma Pass (13, AMNH); Santa 
Cruz Co.: Santa Rita Mtns., Madera Canyon (1<5, MCZ; 
16, AMNH), Santa Rita Mtns. (29, AMNH); TEXAS 
Denton Co.: Lake Dallas opposite Hatchery (13, MCZ) 
Erath Co.: Stephenville (33, TXAM); ARKANSAS 
Washington Co.: Boston Mtns., Cove Creek Valley, 



24 km S of Prairie Grove (IS, MCZ); ILLINOIS: Har- 
din (9), 

Natural History. Collected beating oaks 
in oak woodland (1<3, Arizona) and from 
juniper (13, Texas) at altitudes from 300 
m (Arkansas) to 1,650 m (Arizona). Jung 
and Roth (1974) collected this species in 
their zones 1 and 2 in the Chiricahua 
Mountains (1,200-1,700 m). 

22. Pelegrina furcata 
(F. P.-Cambridge, 1901) 
new combination 

Figures 158, 159, 212, 249, 250, 

388-402; Map 22 

Metaphidippus furcatus F. P.-Cambridge, 1901: 267, 
pi. 24, figs. 8, 8a, S. Type material in BMNH 23 
with label "Dendryphantes furcatus, sp. n. m's, Ori- 
zaba, Mexico (H. S. )" and 23 with label "Dendry- 
phantes furcatus, sp. n. Type 3, S>ntype 3., Gua- 
temala (Sarg)," examined. Despite the type label 
on the latter specimens, the holotype may be better 
considered to be among the former, given Cam- 
bridge's indication of the distribution as Orizaba. 
Bonnet, 1957: 2813. 

Dendryphantes furcatus: — G. & E. Peckham, 1909: 
473. Roewer, 1954: 1203. 

Dendryphantes mimus Chamberlin, 1925b: 135, figs. 
53, 54, 3. Holotype in MCZ 13 with label "Den- 
dryphantes mimus Chamb., 3 holotype, N. M.: Pe- 
cos, R. V. Chamberlin Coll. 1047, " examined. 
Roewer, 1954: 1212. Bonnet, 1956: 1396. NEW 
SYNONYMY. 

Diagnosis. A species common in the 
Mexican highlands, with a striking court- 
ship display and distinctive embolus hav- 
ing two blunt rami. The epigynum, with 
convex flaps, concave surface, and wide 
second curve of the ducts, is distinctive. 

Male. Palpus (Figs. 212, 389-394): Em- 
bolus heavy and slanting, with retrolateral 
ramus extended and truncate. Markings 
(Figs. 158, 88): dark brown with distinct 
sheen and contrasting white side bands. 
Carapace side bands usually connect to 
white scales over anterior eye row to make 
a continuous band of white encircling the 
front of the carapace (though not seen in 
male drawn. Fig. 388). Cheek band mod- 
erately weak. Clypeus brown; hairs over- 
hanging chelicerae brown to white me- 
dially, brown laterally. White forehead 



Pelecrina Jumping Spiders • Maddison 293 



band contacts AMEs dorsally 10:30-12:30 
or 1:00. Chelicerae with some pale scales 
medially. Cymbium lacking pale scales. 
Measureinents: Southern Arizona: Body 
length 3.5(4.2)4.5 mm; carapace length 
1.7(2.1)2.2 mm, width/length 0.74(0.77) 
0.77; n = 53 from Santa Rita Mtns., Ari- 
zona. Northern Arizona: body length 3.5, 
3.6, 3.9 mm; carapace length 1.7, 1.8, 1.8 
mm, width/length 0.75, 0.78, 0.79; n = 33 
from Yavapai Co., Arizona. 

Female. Epigijnum (Figs. 249, 250, 395, 
397, 398, 400, 401): Flaps strongly convex 
and often dark. Surface concave behind 
flaps, without mound, rising gradually to 
lip at back edge. First curve of duct nar- 
row; second curve broad initially but nar- 
rows as it proceeds medially. Markings 
(Figs. 159, 396, 399, 402): Body often with 
slight bronze sheen; variable in markings. 
Carapace covered with brassy reflective 
scales, sometimes dark, sometimes mixed 
with white. Clypeus only thinly covered 
with white scales except in northernmost 
populations (form mimus). Measure- 
ments: Southern Arizona: Body length 
4.5(4.7-4.9)5.4 mm; carapace length 
2.0(2.1)2.2 mm, width/length 0.75(0.78- 
0.79)0.79; n = 6$ from Santa Rita Mtns., 
Arizona. Northern Arizona: Body length 
4.0(4.6)5.7 mm; carapace length 1.8(1.8)2.0 
mm, width/length 0.72(0.75)0.77; n = 59 
from Yavapai Co., Arizona. 

Geographical Variation. Four geo- 
graphical forms might be recognized. (1) 
The most widespread form occurs from 
Guatemala north to northern Mexico, with 
narrower embolus and thinner epigynal 
flaps that are divergent or parallel (furcata 
s.s.,; Fig. 393). The retrolateral ramus of 
the embolus is truncated obliquely. Most 
females through this range are well marked 
with pale spots, as in form mimus (Fig. 
396). (2) A second form, very similar to 
the widespread form, occurs in the Santa 
Rita, Santa Catalina, and Chiricahua 
Mountains of southern Arizona and prob- 
ably in northern Mexico (Figs. 392, 397- 
399). The embolus is also narrow and the 
flaps divergent or parallel, but the retro- 



lateral ramus of the embolus is truncated 
transversely, so that its distal tip makes a 
line perpendicular to the axis of the palpus. 
Females are dark. (3) A third form occurs 
in northern Arizona (Yavapai Co.), Colo- 
rado, and New Mexico, having a wider 
embolus and convergent flaps (mimus: 
Figs. 390, 391, 395, 396). The retrolateral 
ramus of the embolus is truncated trans- 
versely, as in form (2). The difference be- 
tween the northern (mimus) and southern 
(furcata) Arizona specimens is rather strik- 
ing, for the females are also smaller and 
paler in the north. Though mimus might 
be considered a distinct species, specimens 
in New Mexico present a confusing mix- 
ture of characteristics of forms (1), (2), and 
(3). (4) A fourth form is found in western 
Oaxaca (43 159, 31 km N or Guelatao de 
Juarez, ca. 96.5°W, 17.5°N, 2,600 m el., 3 
August 1983, W. Maddison & R. S. An- 
derson, MCZ), with very wide embolus, 
dark females, and extremely robust flaps 
(Figs. 388, 400-402). This form occurs 
within 50 km of the widespread form. In 
total, the variation among these popula- 
tions is confusing, and though several spe- 
cies may be present, only one will be rec- 
ognized until better studied. 

Chromosomes. 2n3 = 26 acrocentrics + 
XXO (23 from Madera Canyon, Arizona). 

Courtship (103 observed from Nuevo 
Leon, Hidalgo, Queretaro, Puebla, Oaxa- 
ca, Chiapas, and the Santa Rita Mountains 
of Arizona). Very unusual for the genus, 
with vigorous leg waving and body jerking 
in a stage I will call the semaphore stage. 
Semaphore (n = 24, 103): Body high to 
very high (n = 24, 103). Male walked si- 
dling (n = 19, 83) in series (n = 14, 53). 
First legs wide, nearly 180° apart, approx- 
imately horizontal (n = 21, 83) or below 
horizontal (n = 1), though occasionally not 
much more than 90° apart and raised to 
60° (n = 1), waved vigorously up and down 
almost to vertical (n = 21, 93), though 
sometimes only to ca. 40° (n = 1), at ca. 
3-4 c/s (n = 2, 23) or 5 c/s (n = 1) on each 
sidle (n = 9, 43). The leg wave is vertical 
and slightly posterior to bring the legs up 



294 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



and back (n =7, 43); the left and right sides 
wave in unison (n = 4, 33), or occasionally 
asynchronously (n = 1). Palpi hanging 
down (n = 18, SS) and parallel (n = 9, 46) 
and a bit forward (n = 7, 33). Palpi wave 
with low-medium amplitude (n = 5, 33) 
on each sidle (n = 4, 23), up and down like 
pushing and pulling motion (n = 1), or 
largely still (n = 1, 23). Abdomen trails a 
bit on sidles (n = 5, 23), ca. 10-30° (n = 3, 
13), but more or less horizontal (n = 16, 
53). Occasionally 3 pauses from vigorous 
leg waving and jerks whole body (n = 11, 
63) approximately 4-5 times (n = 3, 33) or 
3 times (n = 1) while the first legs are 
spread wide and horizontal (n = 11, 63). 
These jerks came after a few sidles (n = 
1). The body may be lowered for the jerks 
(n = 1). Reach (n = 3, 23): The male pro- 
ceeded directly from this semaphore stage 
into the reach to touch the female (n = 3, 
23). During reach, body jerked a few times 
(n = 1). First legs held parallel and forward 
(n = 2, 23). Palpi held parallel and forward 
(n = 2, 23). 

This description is from the widespread 
form (1) with the following exceptions. 
Displays of 43 from the distinctive popu- 
lation from Guelatao de Juarez, Oaxaca 
(form (4)), showed the same form of sem- 
aphore display with legs spread wide, vig- 
orously waved up during series (n = 11, 
43). No whole-body jerks were noted, how- 
ever. One male from southern Arizona 
(form (2)) showed the same semaphore dis- 
play (n = 4), though no whole-body jerks 
were noted. 

Distribution (Map 22). Throughout the 
highlands of Mexico, extending north to 
Colorado and south to Guatemala. 



Records. Mostly in MCZ and AMNH: UNITED 
STATES (county records): COLORADO: Boulder 
(4<?), El Paso (IS), Rio Grande (13); NEW MEXICO: 
Bernalillo {U 32), Catron (1<5), Colfax (2<5 19), Grant 
(1(5), Lincoln (33 49), Los Alamos {IS), Mora (29), 
Otero {1$ 29), Sandoval (IS 29), San Miguel (IS), Santa 
Fe (1.5), Socorro (2<3), Valencia {\$ 39); ARIZONA: 
Cochise (more than 15 S 99), Santa Cruz (183 179), 
Graham (19), Pima (33 29), Yavapai (53 89). M£A7- 
CO: NUEVO LEON: Cerro Potosi (23 49); Monter- 



rery (13); CHIHUAHUA: Canon Prieta, near Pri- 
mavera (19); HIDALGO: 4 km NE of Tlanchinol (23 
69); 3.4 km SW of Cuesta Colorada (43 29); 10 km 
SW of Santa Maria, 99°00'W, 2r06'N (13); 8 km N 
of Encarnacion, 99.12°W, 20.55°N (29); Huachinango 
(19); Apulco (13 29); Champuhuacan (19); Maguey 
Verde, 99.12°W, 20.49°N; PUEBLA: 8 km N of Te- 
ziutlan (13 39); near Xicotepec de Juarez, 97°59'W, 
20"'17'N (13) 5 km N of Hvw 130 on road to Naupan 
(13); QUERETARO: ca. 99°10'W, 21°15'N (53 39); 
GUERRERO: Chilapa (19); VERACRUZ: 3 km N of 
Fortin de las Flores (13); 6 km NE of Coscomatepec 
(23); 7 km N of Huatusco (13 39); Orizaba (3399); 
DISTRITO FEDERAL: Santa Rose (13), Contreras 
(13); OAXACA: 23 km SW of Valle Nacional on Hvvy 
175 (43 59); 27 km SW of Valle Nacional on Hwy 
175 (13 29); 31 km N of Guelatao de Juarez (43 159); 
CHIAPAS: 5 km W of San Cristobal de las Casas (33 
39); Grutas de San Cristobal (19); San Cristobal (43 
49). GUATEMALA: locality unknown (23). 

Natural History. Collected from oak (7 
records), grasses, herbs, and shrubs in 
clearings (5 records), pine (3 records), ju- 
niper (1 record), Ceanothus (1 record), 
Cercocarpus (1 record), in oak-pine cloud 
forest zones. Collected at elevations of 
1,000-1,400 m (8 records, 1,500-2,000 m 
(7 records), and 2,100-3,000 m (7 records). 

23. Pelegrina votcana new species 
Figures 213, 403, 404; Map 23 

Holotype male in MCZ with labels: "PANAMA: El 
Volcan, A. M Chickering" and "R. P. El Volcan, 
Aug. 9-14, 1950." 

Etymology. An arbitrary combination 
of letters referring to the type locality, to 
be treated as an adjective. 

Diagnosis. Known from only two males 
from Panama; much like bicuspidata but 
the long embolus is not so bent as in that 
species. 

Male. Palpus (Figs. 213, 404): Erect por- 
tion of embolus broadens gradually into 
base; rami small and subequal. Tibial 
apophysis appears double because ridge 
prolonged into second apophysis, more ex- 
treme than in furcata (Fig. 389). Markings 
(Fig. 403): Typical for genus with cara- 
pace side bands, forehead band. Cheek 
band dense. Clypeus brown; hairs over- 
hanging chelicerae white medially and 
brown laterally. White forehead band con- 



Pelegrina Jumping Spiders • Maddison 295 



tacts AMEs dorsally 10:30-12:30. Chelic- 
erae with long medial patch running al- 
most at least % length. Palpus femur dis- 
tinctly paler than more distal segments. 
Cymbium with none or few white scales. 
Legs fairly distinctly annulate. Anterior 
three pairs of abdominal spots longitudi- 
nally directed, 4 through 6 transverse. 
Strong abdominal side bands. Measure- 
ments: Body length 3.5, 3.7 mm; carapace 
length 1.7, 1.8 mm, width/length 0.76, 
0.77; n = 2S from Panama. 



Chiapas S.) Measurements: Body length 
2.7, 3.2 mm; carapace length 1.3, 1.5 mm, 
width/length 0.77, 0.83; n = 23 from Chia- 
pas and Guatemala. 

Female. None matched to male. Cam- 
bridge's P. ochracea may represent the fe- 
male of P. bicuspidata. 

Records (Map 23). In addition to the holotype, one 
other male is known, from Mexico; Chiapas: pine 
forest, 24 km NW of Arriaga 94.01°W, 16.25°N, 27 
August 1966 (AMNH). 



24. Pelegrina bicuspidata 
(F. P.-Cambridge, 1901) 
new combination 
Figures 214, 405, 406; Map 23 

Metaphidippus bicuspidatits F. P.-Cambridge, 1901: 
269, pi. 24, 6gs. 13, 13a, b, 6. Holotype in BMNH 
1(5 with labels "Dendryphantes bicuspidatus, sp. 
Type S, Guatemala (Sarg)" and "1905, 268.", ex- 
amined. Bonnet, 1957: 2810. 

Dendryphantes bicuspidatus: — Roewer, 1954: 1191. 

Diagnosis. A rarely collected species 
from southern Mexico and Guatemala. The 
distinctive embolus is long and bent, unlike 
that of volcana. 

Male. Palpus (Figs. 214, 406): Embolus 
heavy, abruptly bends basal to opening. 
Rami small and subequal. Tibial apophysis 
appears double because ridge prolonged 
into second apophysis, more extreme than 
in furcata (Fig. 389). Markings (Fig. 405): 
Typical for the genus, with white cheek, 
forehead, and side bands. Cheek band 
moderately dense. Clypeus brown; hairs 
overhanging chelicerae white medially, 
brown laterally. White forehead band con- 
tacts AMEs dorsally 10:00-12:30. Chelic- 
erae with thin patch of pale white scales 
medially extending to % length. Palpus fe- 
mur distinctly paler than more distal seg- 
ments. Cymbium lacking white scales. Legs 
with fairly distinct annulation; back of tib- 
ia 2 uniformly pale; femur 3 dark in distal 
^3. Abdomen dorsum more or less solid 
brown, without trace of dark spots, sur- 
rounded by discrete white side bands on 
abdomen. (Description based mostly on 



25. Pelegrina ochracea 
(F. P.-Cambridge, 1901) 
new combination 
Figures 407-409; Map 23 

Metaphidippus ochraceus F. P.-Cambridge, 1901: 
272, pi. 25, figs. 6, 6a, 9, Holotype in BMNH 19 
with label "Dendryphantes ochraceus, sp. n. Type 
9, Guat. (Sarg)," examined. Bonnet, 1957: 2816. 
Chickering's M. ochraceus (1946: 312) is not the 
same as this species, nor is it a Pelegrina. 

Dendryphantes ochraceus: — Roewer, 1954: 1198. 

Diagnosis. The epigynum is similar to 
that of P. furcata, but the flaps are not 
quite so convex and are shorter. As already 
noted, this could be the female of P. bi- 
cuspidata. 

Female. Epigynum (Figs. 407, 408): 
Flaps convex, parallel, and fairly short; 
light brown; behind them the surface is 
gently concave with mound restricted to 
near posterior margin, and median ridge 
extending from flaps to posterior mound. 
First curve of duct narrow; second curve 
proceeds medially. Markings (Fig. 409): 
Carapace red-brown, thinly covered with 
white scales. Clypeus covered with white 
scales, scales surrounding AMEs white. 
Legs lacking strong annulae, tan to light 
brown. Abdomen hght brown with pale 
markings (Fig. 409). Measurements: Ho- 
lotype body length 4.0 mm; carapace 
length 1.6 mm, width/length 0.75. 

Records (Map 23). MEXICO: OAXACA: Oaxaca, 
Base San Felipe Mtn., 16-17 September 1947 (19, 
AMNH); CHIAPAS: San Crist6bal, 13 September 1947 
(19, AMNH). GUATEMALA (19, BMNH). 



296 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



26. Pelegrina morelos new species 
Figures 215, 410-414; Map 23 

Holotype male in AMNH with label "7 mi [11 km] 
N Cuernavaca, Morelos, Mexico, July 3, 1941, A M 
and L I Davis. " 

Etymology. A noun in apposition, after 
the type locality. 

Diagnosis. Much like furcata, but the 
retrolateral ramus of the embolus is pro- 
longed and curves to the prolateral. The 
female matched to the holotype has much 
more contrasting markings than in furcata 
and epigynal ducts with a much narrower 
second curve. 

Male. Palpus (Figs. 215, 411): Embolus 
with two blunt rami much like those of 
furcata, but the retrolateral is long and 
curves to the prolateral. Tibial apophysis 
appears double because ridge prolonged 
into second apophysis. Markings (Fig. 410): 
Carapace side bands dense and discrete. 
Cheek bands weak. Clypeus brown; setae 
overhanging chelicerae dark except a few 
white hairs medially. White forehead band 
contacts AMEs dorsally 10:30-12:30. Che- 
licerae with a few white scales medially. 
Palpus cymbium brown, lacks white scales. 
Abdomen with white side bands and paired 
white spots reminiscent of female. Mea- 
surements: Body length 38 mm; carapace 
length 1.9 mm, width/length 0.78. 

Female. Epigynum (Figs. 412, 413): 
Flaps long and convex, dark. Epigynal sur- 
face slightly concave, rises gradually be- 
hind flaps to higher posterior margin. First 
curve of duct broad; second curve narrow, 
proceeds medially. Markings (Fig. 414): 
Carapace brown, thinly covered with white 
scales. Clypeus covered with white scales 
though scales surrounding AMEs are or- 
ange laterally and medially. Legs annu- 
late. Abdomen strongly marked with four 
distinct pairs of white spots on dark back- 
ground; fourth pair large and transverse. 
Measurements: Body length 4.7 mm; car- 
apace length 1.9 mm, width/length 0.80. 

Male/Female Matching. The female 
described is matched tentatively with the 
male, for both are similar to furcata in 
genitalia, they have similar abdominal 



markings with strong lateral fourth pair of 
spots, and they occur sympatrically. 

Records (Map 23). In addition to the male holotype, 
the single female known is from: Mexico: Morelos: 
Cuernavaca, July 1953 (AMNH). 

27. Pelegrina huachuca new species 
Figures 216, 415-419; Map 24 

Holotype male in AMNH with label "ARIZONA: 
8000 ft. [2,440 m], Carr Canyon, Huachuca Mts., 
June 3, 1952." 

Etymology. A noun in apposition, after 
the type locality. 

Diagnosis. Most distinctive for its large 
branched embolus bearing some resem- 
blance to that of P. furcata. Females 
matched with the male have long flaps in 
a distinctively sculptured epigynum. 

Male. Palpus (Figs. 216, 416): Embolus 
large and unusual, with retrolateral ramus 
extended retrolaterally as a long blade. 
Tibial apophysis appears double because 
ridge prolonged into second apophysis. 
Markings: Typical for the genus, with 
cheek, side, and forehead bands on the 
carapace and side bands on the abdomen. 
Cheek band moderately dense. Clypeus 
brown; hairs overhanging chelicerae dark 
with some white centrally. White forehead 
band contacts AMEs dorsally. Chelicerae 
with a few pale scales medially. Legs with 
indistinct annulation. Measurements: Body 
length 3.7 mm; carapace length 1.8 mm, 
width/length 0.77; n = 13 from Huachuca 
Mtns., Arizona. 

Female. Epigynum (Figs. 417, 418): 
Flaps dark, long, and parallel. Surface 
bulges just medial to flaps about half way 
along their length; behind this the surface 
is concave, rising gradually into pro- 
nounced medial and posterior bulge. First 
curve of ducts very broad; second curve 
goesanteriomedially. Markings (Fig. 419): 
Carapace covered with white scales with 
some light brown. Clypeus densely white; 
white between AMEs. AME scales yellow- 
ish white above, white below. Abdomen 
brown; central pale spots are laterally di- 
rected bars, side bands. Measurements: 
Body length 4.5, 5.1 mm; carapace length 



Pelegrina Jumping Spiders • Maddison 297 



2.1, 2.2 mm, width/length 0.79, 0.80; n = 
29 from Santa Catalina and Santa Rita 
Mtns., Arizona. 

Male/Female Matching. As discussed 
under P. chaimona, there is doubt regard- 
ing the male/female association of chai- 
mona and huachuca. The following evi- 
dence supports the matching of the fe- 
males already described with the male of 
huachuca: the flaps of the female are long 
and convex, the surface has distinct con- 
cavities and bulges, and the first curve of 
the duct is very broad, as would be ex- 
pected to match the robust embolus of hu- 
achuca ; the cephalic plate is smoother than 
in chaimona males and females, as in the 
male huachuca; the females have been 
found in central Arizona, as was the male. 

Distribution (Map 24). Southcentral and 
southeastern Arizona. 

Records. UNITED STATES: ARIZONA: Cochise 
Co.: Huachuca Mtns., Garden Canyon Road, base of 
Sawmill Canyon, 19 March 1989 {S3, MCZ), Hu- 
achuca Mtns., Carr Canyon, 3 June 1952, 2,400 m el. 
(1(5, AMNH), Chiricahua Mtns., upper Cave Creek, 
10 May 1969, 1,800 m el. (1$, AMNH), Chiricahua 
Mtns., Onion Saddle, 2,370 m el, 20 March 1989; 
Pima Co.: Santa Catalina Mtns. (12, AMNH); Santa 
Cruz Co.: Santa Rita Mtns., upper Madera Canvon, 
1,700 m el., 13 August 1983 (19, MCZ). 

Natural History. Collected from oaks (4 
records). 



28. Pelegrina arizonensis 
(G. & E. Peckham, 1901) 
new combination 

Figures 160, 161, 217, 251, 420-425; 

Map 25 

Dendryphantes arizonensis G. & E. Peckham, 1901b: 
326, pi. 28, fig. 2, S. Holotype in MCZ IS with 1 
imm. with labels "Dendryphantes arizonensis Pkm, 
1901. Arizona. Type. 6." (label is original; hand- 
written, probably by Elizabeth Peckham) and "G. 
W. Peckham Coll.", examined. G. & E. Peckham, 
1909: 463, pi. 36, fig. 7, S. Roewer, 1954: 1206. 

Dendryphantes glacialis Schefi^er, 1905, figs. 3, 4, 8, 
(59. Type material lost (Cutler and Jennings, 1985), 
though the Peckham collection has some material 
labeled Dendryphantes glacialis from Manhattan, 
Kansas, possibly sent by Scheffer, examined. G. & 
E. Peckham, 1909: 463, pi. 37, figs. 7, 7a, b, 69. 
Roewer, 1954: 1210. 



Dendryphantes (Metaphidippns) arizonensis: — Pe- 

trunkevitch, 1911: 622. 
Dendnjphantes mimus: — Chamberlin, 1925b, in part: 

135, fig. 52 9. 
Metaphidippns arizonensis: — Bonnet, 1957: 2810. 

Cutler and Jennings, 1985: 3, figs. 3-11, (59. 
Metaphidippus glacialis: — Bonnet, 1957: 2814. 

Diagnosis. Like helenae this species has 
genitalia unusual for the genus, with the 
erect portion of the embolus arising retro- 
laterally and the epigynal flaps far rotated. 
Differs from helenae in having a sharp 
pointed embolus, short tibial apophysis, and 
flaps rotated only 180°. 

Male. Palpus (Figs. 217, 421, 422): Em- 
bolus arising toward retrolateral side, 
blade-shaped and with exposed surface 
concave, with retrolateral ridge extending 
into distal point. Tibial apophysis almost 
hidden behind wide flange beneath it. 
Markings (Figs. 160, 420): Cheek bands 
weak. Markings on face quite variable. 
Clypeus brown, sometimes with a few 
white hairs, hairs overhanging chelicerae 
white to brown. White forehead band con- 
tacts AMEs dorsally 10:30-12:00. Chelic- 
erae lacking pale scales Cymbium with 
none to a few white scales. Abdomen show- 
ing lineate markings of females, with two 
medial longitudinal stripes in addition to 
the side bands. Measurements: Body 
length 4.0(4.3)4.3 mm; carapace length 
2.0(2.0)2.0 mm, width/length 0.80(0.81) 
0.83; n = 5 (5 from Minnesota. 

Female. Epigynum (Figs. 251, 423, 424): 
Flaps rotated 180° so that ancestrally pos- 
terior end is anterior. Surface flat except 
for concavity in front of flaps. First curve 
of duct broad, on medial side of opening 
because of flap rotation; second curve pro- 
ceeds laterally. Markings (Figs. 161, 425): 
Carapace with white scales dorsally. Clyp- 
eus densely covered with white scales. Ab- 
dominal markings strikingly lineate, with 
two central pale bands flanked by two thin 
rows of dark spots, flanked by brown bands 
and pale side bands. Measurements: Body 
length 4.7(4.9)5.9 mm; carapace length 
2.0(2.0)2.2 mm, width/length 0.77(0.80) 
0.84; n = 59 from Minnesota. 

Courtship {IS observed from Anoka Co., 



298 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



J 



Minnesota): The one male observed showed 
no crouch display. Raisespread (n = 16): 
Abdomen twitched on pause (n = 3). First 
legs waved irregularly on series (n = 9); as 
he got closer legs lowered, put forward and 
more parallel but no discrete crouch dis- 
play was seen (n = 7). Palpi wave irreg- 
ularly on series (n = 4). 

Distribution (Map 25). Minnesota and 
Alberta south to Zacatecas and Tlaxcala. 

Records. CANADA: ALBERTA: Medicine Hat {IS 
19, AMNH). UNITED STATES: MINNESOTA: 
Anoka Co.: 5 km E of Bethel (15 39, MCZ; IS 39, 
AMNH); NORTH DAKOTA: Burleigh Co.: Menoken 
Indian Village (1<5, MCZ); Williams Co.: WiUiston (1<5 
39, WPM); KANSAS: Rilev Co.: Manhattan {IS 19, 
MCZ); COLORADO: Denver (1<5, MCZ); TEXAS: 
Brewster Co.: Marathon (1<3 19, AMNH); Alpine {IS, 
AMNH); NEW MEXICO: Bernalillo Co.: Sandia Mtns. 
{2S, AMNH); Hidalgo Co.: 3 km N of Rodeo {IS, 
AMNH); Lincoln Co.: T6N R6ES74 (1<5); near Nogal 
(13); Sierra Co.: 5 km E Hillsborough {IS, MCZ); 24 
km N of Ruidoso {7S 69, AMNH); Torrance Co.: 1.6 
km E Clines Corners (19, AMNH); Valencia Co.: 3 
km E of Grants {IS 19, AMNH); ARIZONA: Coconino 
Co.: Sitgreaves National Forest (1<5, AMNH); Graham 
Co.: Thatcher {IS 29, MCZ); Santa Cruz Co.: 29 km 
NW of Nogales (19, AMNH) MEXICO: ZACATE- 
CAS: 11 km SE of Salinas (San Luis Potosi) on Hwy 
45, ca. 101°39'W, 22°34'N (29, MCZ); DURANGO: 
Rodeo (59, AMNH); Santa Maria del Oro (19, AMNH); 
TLAXCALA: 13 km W of Calpulapan (29, AMNH). 
Cutler and Jennings (1985) give additional records. 

Natural History: Collected from sand 
prairie (Minnesota, 1 record), Hymeno- 
clea (New Mexico, 1 record), ponderosa 
pine (Arizona, 2 records), grass-mesquite 
(Arizona, 1 record), and beating junipers 
(1 record. New Mexico) at elevations of 
1,700-2,200 m in Arizona, Zacatecas, and 
Durango. Jennings (1973) describes egg re- 
treats of this species in Tragopogon and 
Pinus ponderosa. Cutler and Jennings 
(1985) give additional habitat information 
and characterize P. arizonensis as a species 
of grasslands. 

29. Petegrina hetenae {Banks, 1921) 
new combination 
Figures 155, 218, 426-431; Map 25 

Dendryphantes helenae Banks, 1921: 101, fig. 5, S9. 
Type material in CAS IS 19 with labels "Dendry- 
phantes helenae Bks type," "San Francisco, Cal. 



IV-7-'18," and "Coll and don by Helen Van Du- 
zee," examined. Roewer, 1954: 1211. 

Dendryphantes saiisalitanus Chamberlin, 1925b: 137, 
figs. 57, 58, S. Type in MCZ 15 with label "Den- 
dryphantes sausalitanus Ch, S holotype, Cal.: Sau- 
salito 1909, R. V. Chamberlin Coll., 1045," ex- 
amined. 

Metaphidippus helenae: — Gertsch, 1934: 18. Bonnet, 
1957: 2814. Cutler and Jennings, 1985: 5, figs. 12- 
17, 59. 

Diagnosis. Differs from arizonensis in 
having blunt embolus, tibial apophysis on 
an elongate projection, and epigynal laps 
rotated very far, to 270°. 

Male. Palpus (Figs. 218, 427, 428): Erect 
portion of embolus is blade-shaped and 
blunt and arises toward retrolateral side of 
base. Tibial apophysis and flange elevated 
on narrow projection (Fig. 427). Markings 
(Fig. 426): generally dark with weak white 
side bands on carapace and abdomen. 
Cheek band very weak. Clypeus brown; 
hairs overhanging chelicerae dark, some- 
times pale medially. White forehead band 
contacts AMEs dorsally 10:30-1:00. Che- 
licerae lacking pale scales. Cymbium with 
none to a few white scales. Longitudinal 
markings on abdominal dorsum, with cen- 
tral longitudinal lighter brown band 
flanked by black bands. Measurements: 
Body length 4.0(4.2)4.3 mm; carapace 
length 1.8(2.0)2.1 mm, width/length 
0.77(0.80)0.82; n = 53 from Nevada and 
Oregon. 

Female. Epigynum (Figs. 429, 430): 
Flaps rotated 270°, so that ancestrally pos- 
terior end is lateral. Surface flat. First curve 
of duct anterior to opening because of flap 
rotation; second curve proceeds posteri- 
orly. Markings (Figs. 155, 431): Carapace 
thinly to densely covered with white or 
gray scales. Clypeus densely covered with 
white scales. Abdominal markings some- 
what lineate, but not so strongly as in ar- 
izonensis, with the posterior dark spots 
more prominent. Measurements: Body 
length 4.2(5.0)5.5 mm; carapace length 
1.9(2.0)2.2 mm, width/length 0.76(0.80) 
0.82; n = 59 from Oregon, Nevada, and 
Washington. 

Distribution (Map 25). Wyoming to 
Washington south to Utah and California. 



Pelecrina Jumping Spiders • Maddison 



299 



Records. UNITED STATES: IDAHO; Blaine Co.. 
Carev (IS, AMNH); Custer Co.: Salmon River, 19 km 
N Challis (19, AMNH); Gem Co.: 11 km W of Horse- 
shoe Bend, 116°18'W, 43°57'N (56, AMNH); 13 km 
W of Horseshoe Bend (12, AMNH); Jerome Co.: Twin 
Falls (1(3, AMNH); Owyhee Co. : Bruneau Canyon Hot 
Creek Falls (19, AMNH); WYOMING: Bighorn Co.: 
10 km E of Shell (1<J 29, WPM); UTAH: Little Cot- 
tonwood Campground, near Salt Lake City (33, 
AMNH); Sevier Co.: Richfield (IS 29, AMNH); NE- 
VADA: Humboldt Co.: 48 km N of Winnemucca (19, 
AMNH); Washoe Co.: N of Reno (3<5 19, MCZ); Reno 
(25, MCZ); WASHINGTON: Franklin Co.: just W of 
Palouse Falls, 46.66°N, 118.23°W, (39, MCZ); Grant 
Co.: Wahluke Wildlife Recreation Area, 46.705°N, 
119.42rW (29, MCZ); OREGON; Baker Co.; Baker 
(19, AMNH); Crook Co.; 8 km W of Prineville (2S 
79, AMNH); Deschutes Co.; Redmond (49, AMNH); 
Harney Co.; Tencent Lake (IS 39, AMNH), Manns 
Lake (13, AMNH); Klamath Co.; above Algoma (13, 
AMNH), Bly Mountain (19, AMNH); Malheur Co.; 
Succor Creek Canyon (2<5 19, AMNH); E of Ontario, 
116°57'W, 44''2'N (19, AMNH); Umatilla Co.; 19 km 
SW of Echo (19, AMNH); Wasco Co.: Mosier (19, 
AMNH); CALIFORNIA; Marin Co.: Sausalito (S, 
MCZ); Mono Co.; Benton (29, AMNH); Riverside Co.; 
Lake Hemet 116°59'W, 33''43'N; San Diego Co.; 3 
km E of Pine Springs (13, AMNH); Pine Valley (19, 
AMNH); San Francisco Co.: San Francisco (13 29 
MCZ); Sierra Co.; Peavine (39, AMNH). Additional 
records are given by Cutler and Jennings (1985). 

Natural History. Collected from sage- 
brush {Artemisia tridentata; 7 records). 
Cutler and Jennings (1985) give additional 
habitat information. 

30. Pelegrina verecunda 
(Chamberlin & Gertsch, 1930) 
new combination 

Figures 162, 163, 219, 252, 432-436; 

Map 26 

Sassacus uteanus: — Chamberlin and Gertsch, 1929, 
in part: fig. 54 (this figure may have been mis- 
placed, given that it is unlikely the authors would 
have confused their species S. uteanus and D. ver- 
ecundus). 

Dendryphantes verecundus Chamberlin and Gertsch, 
1930; 144. Holotype 13 in AMNH with labels "Den- 
dryphantes verecundus 3 / Utah; Dry Canyon Ho- 
lotype/ 6-14-29 Gertsch" and "29Bb. N40 ; Will," 
examined. 

Dendryphantes verecundus: — Roewer, 1954; 1216. 
Bonnet, 1956: 1402. 

M etaphidippus verecundus: — Jung and Roth, 1974; 
33. 

Diagnosis. A small indistinctly marked 
species with dark males and pale females 



from the southwest. May be confused with 
the sympatric orestes but smaller, more 
gray than orange, lacking the lateral chel- 
iceral ridge, and having the short embolus 
broadening gradually into base prolater- 
ally. 

Male. Palpus (Figs. 219, 433): Embolus 
short, obliquely truncate, broadens grad- 
ually into base prolaterally; rami indis- 
tinct. Markings (Figs. 162, 432): Carapace 
with scattered white scales; side bands 
weak. Cheek band weak. Clypeus brown; 
thin white to brown hairs overhanging 
chelicerae. White forehead band contacts 
AMEs dorsally 10:30-12:30. Chelicerae 
with some pale scales, scattered but es- 
pecially medially. Palpus entirely brown, 
with femur not distinctly paler than cym- 
bium. Cymbium lacking white scales. Ab- 
domen side bands indistinct, often with 
white markings centrally. Measurements: 
Body length 2.7(3.1)3.6 mm; carapace 
length 1.3(1.5)1.7 mm, width/length 
0.77(0.79)0.80; n = 5<5 from Yavapai Co., 
Arizona. 

Female. Epigynum (Figs. 252, 434, 435): 
Flaps dark, parallel, slightly convex, pos- 
teriorly flush with surface. Surface flat. 
First curve of duct relatively narrow, dark; 
second curve proceeds slightly anteriorly. 
Markings (Figs. 163, 436): Carapace cov- 
ered with white scales. Clypeus densely 
covered with white. Legs pale yellowish. 
Abdomen pale, with many small dark 
speckles sometimes coalescing into gala- 
thea-\ike pattern. Measurements: Body 
length 3.7(4.0)4.5 mm; carapace length 
1.5(1.6)1.8 mm, width/length 0.77(0.78) 
0.80; n = 59 from Yavapai Co., Arizona. 

Male/Female Matching. This associa- 
tion is indicated by co-collecting, distri- 
bution, similar size, and indistinctness of 
markings. 

Courtship (25 observed from Yavapai 
Co., Arizona). First legs held in a trian- 
gular bowed position during pause of 
crouch display, as in P. aeneola. Crouch 
(n = 15, 23): Body horizontal (n = 15, 2S) 
and low (n = 13, 2<5) to high (n = 1). First 
legs held low and forward, bowed, with 



300 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



tips almost touching (n = 15, 23); on series 
legs extended forward and waved vigor- 
ously (n = 15, 25) though somewhat irreg- 
ularly (n = 8, is); on pause legs retracted 
(n = 12, 26) or not (n = 3, 13). Palpi held 
down (n = 11, 23). On series palpi held in 
and under and motionless (n = 3, 13); at 
end of series/start of pause palpi hang 
down and wave a few times (n = 3, 13). 
Repertoires: 23 crouch only. 

Distribution (Map 26). Utah south into 
northern Mexico. 

Records. UNITED STATES (county records); 
UTAH: Kane (12, AMNH), Salt Lake (175 1?, AMNH), 
Sevier (33, AMNH), Utah (3<5 79, AMNH), Wasatch 
(113 39, AMNH), Washington (93 19, AMNH, MCZ), 
Weber (23, AMNH); ARIZONA: Cochise (53 129, 
AMNH, MCZ), Coconino (13, AMNH), Santa Cruz 
(19, MCZ), Yavapai (83 119, AMNH, MCZ); NEW 
MEXICO: Bernalillo (23, AMNH), Catron (19, 
AMNH), Grant (39, AMNH), Sandoval (13, AMNH); 
CALIFORNIA: Mono (13, AMNH). MEXICO: CHI- 
HUAHUA: Las Delicias (13 49, AMNH); PPrimavera 
(19, AMNH); 21 km N of Ciudad Camargo on Hwy 
45, 105°13'W, 27°52'N (29, MCZ); ?40 km SW of 
Camargo (19, AMNH); PDURANGO: Ojo de los En- 
cinos (19, AMNH). 

Natural History. Specimens collected at 
elevations from 1,100 to 1,800 m (12 re- 
cords). In Arizona, beaten from Quercus, 
Cercocarpus, Alnus, Salix, Chrysotham- 
nus, pine, and spruce (6 records). Jung and 
Roth (1974) found this species in their zone 
2 of the Chiricahua Mountains. 

31 . Pelegrina clavator new species 
Figures 164, 165, 220, 437-441 ; Map 15 

Holotype male and paratype female with label 
"MEXICO: NUEVO LEON: Chipinque Mesa just 
S of Monterrey, ca. 4500 ft. [1,370 m]; ca. 100.4°W 
25.6°N, 2 Jun 1983, W, Maddison & R. S. Anderson 
83-034, beating and sweeping forest understory." 

Etymology. A Latin noun in apposition, 
"club-bearer," referring to the large blunt 
embolus. 

Diagnosis. A Mexican species having a 
distinctive broad, truncate embolus and 
angled flaps flanking a central mound on 
the epigynal surface. 

Male. Palpus (Figs. 220, 438): Embolus 
broad, truncate, with rami small. Embolus 



twists toward tip. Markings (Figs. 164, 
437): Carapace with large forehead band. 
Cheek band dense and distinct from side 
band. Clypeus brown in two specimens, 
brown with white scales between AMEs in 
another specimen; hairs overhanging che- 
licerae brown to tan. White forehead band 
contacts AMEs dorsally; setae ringing 
AMEs white except 12:00-1:30. Chelic- 
erae with dense patch of white scales from 
base to % length, longer and wider than 
in variegata. Cymbium lacking white 
scales. Measurements: Body length 4.1, 4.2, 
4.3, 4.4 mm; carapace length 2.1, 2.1, 2.1, 
2.2 mm, width/length 0.76, 0.77, 0.77, 0.80; 
n = 43 from Nuevo Leon. 

Female. Epigynum (Figs. 439, 440): 
Flaps flat and convergent, medial edge at 
level of high central plateau and lateral 
edge in concavity so that flap slopes down 
laterally. Epigynal surface high between 
flaps as noted. First curve of duct broad 
but does not extend so far posterior as in 
the sympatric neoleonis, so that second 
curve begins at or only a bit posterior to 
posteriormost portion of flap; second curve 
proceeds obliquely medial-anterior. Flow- 
erlike gland opening on anterior face of 
second curve. Inner surface of third curve 
fairly smooth. Markings (Fig. 441): Car- 
apace covered above with white scales. 
Clypeus covered densely with white scales. 
Abdomen marked somewhat like P. gal- 
athea. Measurements: Body length 4.0, 4.2 
mm; carapace length 2.0, 2.0 mm; width/ 
length 0.76, 0.77; n = 29 from Nuevo Leon. 

Male/Female Matching. Males and fe- 
males were matched by co-collecting at 
two localities in Nuevo Leon, by common 
distribution; and by the similar robust car- 
apace and abdominal markings. 

Geographical Variation. Females from 
Tamaulipas and Veracruz have smaller 
flaps less deeply set into epigynum, with 
less flaring ducts, and may represent an- 
other species. 

Courtship (23 observed from Chipinque 
Mesa, Nuevo Leon). Raisedspread (n = 3, 
13). Crouch (n = 3, 23): First legs foward, 
slightly spread, horizontal (n = 1) or raised 



Pelegrina Jumping Spiders • Maddison 301 



fairly high to ca. 40° (n = 2, IS), lowered 
when close (n = 2, 13); not noticeably 
waved (n = 3, 2S). Palpi down (n = 3, 26), 
waved on series (n = 3, 25) and when very 
close and reaching (n = 2, 13). Abdomen 
twitched occasionally, possibly at pause (n 
= 2, 13). Repertoires: 13 crouch only; 13 
raisedspread and crouch. 

Distribution (Map 15). Nuevo Leon 
south to Veracruz. 

Records. MEXICO: TAM.AULIPAS: ca. 1.5 km E 
of Tula, 99.5°W, 22.9°N, 8 June 1983 (12, MCZ); 
Sierra de Tamaulipas, 4-7 August 1945 (16, AMNH); 
SAN LUIS POTOSI: 32 km E of Ciudad del Mais, 
23 March 1940 (12, AMNH); NUEVO LEON; Chip- 
inque Mesa, just S of Monterrey, 100.4°W, 25.6°N, 2 
June 1983 and 7 April 1946 (33 'l2, MCZ; 12 AMNH); 
Villa de Santiago, Hacienda Vista Hermosa, 19 June 
1940 (1<5, MCZ); VERACRUZ: 2 km SE of Naolinco 
on Hwy 127, 96.9°W, 19.6°N, 20 June 1983 (12, MCZ). 

Natural History. Collected from un- 
derstory shrubs of broadlead forest at ca. 
1,400 m elevation (2 records); also known 
from 600 to 800 m elevation (3 records). 

32. Pelegrina pallidata 
(F. P.-Cambridge, 1901) 
new combination 
Figures 221, 442-446; Map 29 

Metaphidippu.s pallidatus F. P.-Cambridge, 1901: 
270, pi. 24, figs. 17, 17a, 2. Holotype in BMNH 12 
with label "Dendr\ phantes pallidatus, sp. nov. Guat. 
Sarg Type 2," examined. Bonnet, 1957: 2816. 

Dendryphantes pallidatus: — Roewer, 1954: 1198. 

Notes on Synonymy. The specimens de- 
scribed here are identified with Cam- 
bridge's pallidata primarily on the basis 
of similarities in the details of the epigyn- 
um. This identification is made with some 
hesitation, for the type material of palli- 
data consists of a single poorly marked 
female whose epigynum was lost by me in 
the course of examination, but my notes 
and the figures of C. L. Scioscia (personal 
communication) regarding the epigynum 
are fairly detailed and provide evidence 
for the identification. Compared to the 
other Mexican and Central American spe- 
cies with small, convergent flaps and a fair- 
ly flat epigynal surface (variegata, san- 



daracina, yucatecana, verecunda) the fe- 
males described below and the type of pal- 
lidata are unique in having (1) an unusually 
broad dark band along the margin of the 
opening (Fig. 445), (2) first curve of duct 
wide and long, (3) the flowerlike gland 
openings placed on the anterior surface of 
the second curve and more medially (clos- 
er to junction with third curve than first 
curve), and (4) fertilization ducts arising 
anterior to the center of the lumen of the 
spermatheca. 

Male (Tentatively Associated with Fe- 
male). Palpus (Figs. 221, 443): Embolus 
twists apically. Retrolateral ramus rela- 
tively long and curved, as in P. pervaga 
and tristis, but embolus much smaller than 
in those species. Markings (Fig. 442): Car- 
apace medium to pale brown, with well- 
developed side and cheek bands. Clypeus 
brown except for patch of white scales be- 
tween AMEs that overhangs chelicerae. 
White forehead band contacts AMEs dor- 
sally 10:30-12:00. Chelicerae with white 
scales medially. Cymbium with some white 
scales. Legs beige with brown annulae. Ab- 
domen shows white spots of female. Mea- 
surements: Body length 3.4 mm; carapace 
length 1.7 mm, width/length 0.77, n = 13. 

Female. Epigynum (Figs. 444, 445): 
Flaps convergent, not very convex. Epi- 
gynal surface flat. First curve of duct fairly 
broad and long; second curve goes anter- 
iomedially. As already noted, there is an 
unusually broad band along the margin of 
the opening (Fig. 445, arrow), the flow- 
erlike gland openings are placed on the 
anterior surface of the second curve close 
to junction with third curve, and the fer- 
tilization ducts arise anterior to the center 
of the lumen of the spermatheca. Mark- 
ings (Fig. 446): Cambridge's holotype is 
now uniformly pale, though may be partly 
faded. The other available specimens have 
the carapace covered with yellowish white 
scales, though not densely. Clypeus dense- 
ly covered with white scales; AMEs en- 
tirely ringed by white scales. Legs uniform 
orange-brown except Nicaragua 9, which 
has some brown spots. Sternum distinctly 



302 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4 



darker than coxae. Abdomen with paired 
white spots on brown dorsum; each spot 
in first pair fused with spot in second pair. 
Venter dark between epigynum and spin- 
nerets. Measurements: Body length 4.0, 
4.3 mm; carapace length 1.6, 1.7 mm, 
width/length 0.76, 0.78; n = 29, female 
holotype and 19 from Nicaragua. 

Male/Female Matching. This is indi- 
cated by co-collecting in Nicaragua and 
similarity of markings. 

Distribution (Map 29). Southern Mexico 
to Nicaragua. 

Records. MEXICO: CHIAPAS: 5 km W of San 
Cristobal de Las Casas on Hvvy 190, ca. 92°41'W, 
16°44'N, 27-28 July 1983, W, Maddison & R. S. An- 
derson (19, MCZ). GUATEMALA (19, BMNH); Chi- 
chicastenango, 6-7 August 1947, C. & P. Vaurie (29, 
AMNH). NICARAGUA: Matagalpa, 4 October 1952, 
R. B. Swain (1<S 19, AMNH). 

Natural History. The female from near 
San Cristobal was collected beating oak, 
madrofio, and pine in oak-pine woodland 
at 2,100 m elevation. 

33. Petegrina variegata 
(F. P.-Cambridge, 1901) 
new combination 

Figures 166, 167, 222, 253, 447-451; 

Map 28 

Metaphidippus variegatus F. P.-Cambridge, 1901: 
268, pi. 24, figs. 10, 10a, 11, 11a, S9. Holotype in 
BMNH 1(5 with 19 with label "Philaeus variegatus 
F.Cb., Type <5, gynetype 9 Mexico. Amula [Guer- 
rero]. H. S.", examined. 

Beata variegata: — Simon, 1903: 841. Roewer, 1954: 
1008. Bonnet, 1955: 874. Chickering's Beata var- 
iegata (1946: 267, figs. 226, 227) is not this species, 
nor is it a Pelegrina. 

Diagnosis. Probably the most common- 
ly collected Mexican species, reminiscent 
of galathea. Males distinctive for their 
strong white spotting on the abdomen and 
robust chelicerae. Females can be identi- 
fied by the epigynal topography. 

Male. Palpus (Figs. 222, 448): Embolus 
relatively narrow, parallel-sided, twisted 
so that tip appears to taper in ventral view, 
but oblique view shows two small, sub- 
equal rami; embolus widens abruptly at its 



base on retrolateral side, so as to make 
distinct angle. Markings (Figs. 166, 447): 
Carapace with extensive white markings. 
Cheek band broad and dense, fused with 
side band. Clypeus brown, with setae over- 
hanging chelicerae white medially, some 
brown hairs laterally. White forehead band 
contacts AMEs dorsally; setae ringing 
AMEs white except from 12:30 to 2:00. 
Chelicerae robust, though not elongate, 
with white patch on medial surface from 
base to about V2 length. Cymbium with 
central patch of white scales. Legs dis- 
tinctly annulate. Abdomen not striped as 
in most Pelegrina males but rather with 
paired white spots almost as in 9. Mea- 
surements: Body length 3.4(3.9-4.2)4.4 
mm; carapace length 1.7(1.9-2.1)2.2 mm, 
width/length 0.79(0.80-0.81)0.84; n = 63 
from Oaxaca, Nuevo Leon, and Nayarit. 

Female. Epigynum (Figs. 253, 449, 450): 
Flaps slightly convex, usually convergent 
and somewhat rotated. Epigynal surface 
rather flat, without pronounced posterior 
mound; medial surface at about same 
height throughout. Between the flaps is a 
medial longitudinal ridge; nearer the flaps, 
the surface is lower. First curve of duct 
pale, narrow; second curve proceeds me- 
dially, bearing flowerlike gland opening 
on dorsal surface of duct. Markings (Figs. 
167, 451): Carapace covered with gray- 
white scales, sometimes mixed with light 
brown. Clypeus densely covered with yel- 
lowish white scales. Abdominal markings 
much like galathea, with white or beige 
spots on tan to gray background. Mea- 
surements: Body length 3.5(4.1-4.2)4.9 
mm; carapace length 1.6(1.8)1.8 mm, 
width/length 0.76(0.78)0.83; n = 79 from 
Oaxaca. 

Male/Female Matching. This associa- 
tion is indicated by extensive co-collecting 
and by the similarity of markings on ab- 
domen. 

Courtship (46 observed from two loca- 
tions in Tamaulipas and Oaxaca): Raised- 
spread (n = 3, 33). Crouch (n = 19, 43): 
Body horizontal (n = 19, 43), normal to 
low height (n = 2, 13). First legs held for- 



Pelegrina Jumping Spiders • Maddison 303 



ward, horizontal to 10° raised, bowed, tips 
slightly convergent, parallel or slightly 
spread, not touching (n = 19, 43). On each 
series legs flickered (n = 9, \6) noticeably 
(n = 1) or with fairly low amplitude (n = 
7, 33) or perhaps not at all (n = 3, 13). Palpi 
held down (n = 12, 43), pointing inward 
and resting over chelicerae (n = 7, 23), on 
each series flickered with fairly low am- 
plitude (n = 12, 13) outward (n = 1) or up 
and down (n = 7, 23). Abdomen twitches 
(n = 6, 13) at end of each series (n = 3, 
13) or in pause (n = 4, 23). 

Distribution {Map 28). Nuevo Leon 
south to Panama. 

Records. Most in AMNH; some in MCZ, from: 
MEXICO: TAMAULIPAS: 11 km E of Ocampo, 
99°16'W, 22°49'N (19); 35 km SSW of Mante (3<5); 
Paso del Abra 99.0rW, 22.45°N (IS); Mante (IS 5$); 
23 km S of Villa Juarez (19); Hidalgo (U); 19 km SE 
of Ciudad Victoria (1$); Rio Guajolotes, 64 km S of 
Victoria {2S 29); Sisal, 24 km S of Victoria {26 29); 
Ciudad Victoria (13 39); 18 km N of Victoria (1<J); 
SAN LUIS POTOSi: Covadonga, WSW of Valles 
99.05°W, 21.57''N (19); Valles (29); Taninul, Valles (1<S 
19); El Salto (IS); 19 km E Ciudad del Maiz (19); Pujal 
(19); NUEVO LEON: Santa Rosa Canyon 29 km W 
of Linares (2S 29); Montemorelos (19); CHIHUA- 
HUA: 8 km S of Chihuahua (1<5); Catarinas (13); SIN- 
ALOA: 64 km S of Culiacan (IS); 48 km N of Mazatlan 
(13 19); 10 km E of Villa Union (1<5); Culiacancito 
107.32''W, 24.50°N (23); DISTRITO FEDERAL: 
Xochimilco (13); MORELOS: Cuernavaca (19); NAY- 
ARIT: Tepic (143 119); 43 km S of Tepic (19); 56 km 
S of Tepic (13 19); La Mesa de Nayarit (29); San Bias 
(13); Jalisco (19); Jesus Maria (23); JALISCO: Zapot- 
lanejo (13); Zapotlanejo (13); COLIMA: 32 km N of 
Cohma (13 19); GUERRERO: Iguala (13 19); Chil- 
pancingo (13); Teloloapan (79); VERACRUZ; Plan del 
Rio (23 19); Tierra Colorado (23 19); OAXACA: 2 km 
S of El Tule (23 89); 3 km W of Tapanatepec (13); 
Oaxaca (13); San Felipe, N of Oaxaca City (33 49); 
Paso Real, Rio Tonto (19); Tehuantepec (43); Monte 
Alban (13); 3 km SE of Niltepec, 94.33°W, 16.32°N 
(13); Soladad (23); CAMPECHE: Campeche (23 59); 
YUCATAN: Progresso (13); Motul (13); Chichen Itza 
(23); CHIAPAS: Arriaga, N of Arriaga Mtns. (19); 24 
km NW of Arriaga 94.01°W, 16.25''N; Cintalapa (83 
99); Ocozucuantla (.33 49); Rio de las Flores, 30 km 
NE of Cintalapa (73 89); Tuxtla Gutierrez (43 19); Las 
Cruzes (.33 19). HONDURAS: Zamorano. NICARA- 
GUA: San Marcos (13 49). COSTA RICA: San Jose 
(19). PANAMA: 8 km S of El Valle (23). 

Natural History. Collected beating Aca- 
cia, composites, and other vegetation in 



desert scrub at 1 ,500 m elevation (Oaxaca); 
beating shrubs and trees in fairly dry bot- 
tom of river valley at 600 m elevation 
(Nuevo Leon); and from a pine forest 
(Chiapas). Known from 220 to 1,700 m 
elevation throughout Mexico (8 records). 

34. Pelegrina yucatecana new species 
Figures 169, 223, 452-456; Map 29 

Holotvpe male and paratype female in MCZ with 
labels "MEXICO: YUCATAN: 3 km E of Chichen 
Itza ruins on Hwy 180, ca. 88°34'W 20°40'N, 19- 
20 July 1983 W. Maddison & R. S. Anderson, 8.3- 
115 seasonal forest, beating understory and trailside 
shrubs and small trees." 

Etymology. An adjective, formed after 
yucateco (Spanish) or yucatecan (English), 
referring to the Yucatan Peninsula. 

Diagnosis. An interesting species with 
unusual transverse abdominal markings; in 
genitalia resembling variegata and san- 
daracina but differing from both in details. 

Male. Palpus (Figs. 223, 453): Embolus 
short, with rami very small. Erect portion 
of embolus with sides parallel; widens 
abruptly at base so that a distinct angle is 
made between the erect portion and base 
along the prolateral margin. Markings (Fig. 
452): As only known 3 is teneral, its proper 
colors are not exactly known, though ap- 
pears brown with white markings. Mar- 
ginal band well developed. Carapace with 
forehead band an acute V, proceeding 
more posteriorly from AMEs than later- 
ally. Cheek band dense, and distinct from 
side band. Clypeus brown, with setae over- 
hanging chelicerae dark. White forehead 
band contacts AMEs dorsally 10:30-12:00. 
Chelicerae with dense media