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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. XCV, No. 1

REVISION OF THE AFRICAN LIZARDS OF THE
FAMILY CORDYLIDAE

By Arthur Loveridge

With Twelve Plates

CAMBRIDGE, MASS. U. S. A.
PRINTED FOR THE MUSEUM

November, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
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Of the Bulletin, Vols. I to XCIV, and Vol. XCV, No. 1, have ap-
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These publications are issued in numbers at irregular intervals.
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A price list of the publications of the Museum will be sent upon ap-
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Cambridge, Massachusetts.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCV, No. 1

^REVISION OF THE AFRICAN LIZARDS OF THE
FAMILY CORDYLIDAE

By Arthur Loveridge

With Twelve Plates

CAMBRIDGE, MASS. U. S. A.
PRINTED FOR THE MUSEUM

November, 1944

No. 1. — Revision of the African Lizards of the Family Cordylidae

By Arthur Loveridge

CONTENTS

Page

Introduction 1

Index to the Species recognized 6

Key to the Genera 8

Bibliography 110

INTRODUCTION

That the name ZONURIDAE should have been employed by
Boulenger (1884a), was most unfortunate, for Zonurus Merrem (1820)
was antedated by Cordylus Laurenti (1768), as has been pointed out by
Stejneger (1936b). As article 5 of the International Rules of Nomen-
clature requires that the family name be based on the type genus,
CORDYLIDAE must replace ZONURIDAE, however much such
major changes are to be deplored.

In 1885 Boulenger recognized the family as comprised of 14 species,
as against 48 forms enumerated in the present revision. In 1930 Power,
when preparing a key to the South African species of the genus "Zonu-
rus", listed 18 members and, though two of these {capensis and robertsi)
are here transferred to Pseudocordylus, no fewer than 28 species and
races of Cordylus are accepted in the present paper. Apart from desul-
tory comments, no revisionary work1 has been done on Pseudocordylus
or Platysaurus, but Boulenger (1899c) furnished a synopsis of the 7
species of Chamaesaura, now reduced to 5 forms by synonymy.

The present revision, like others in the series2, is an attempt to
furnish a synopsis of all additions to our knowledge of the family since
1885, and a serious endeavour to arrange its members according to
probable lines of evolution. In this connection I entertain doubts as to

1 The present revision (completed in the summer of 1942) was just going to press (1944)
when a copy of FitzSimons' (1943) "The Lizards of South Africa," was received. The two
new races of Cordylidae which he describes have been included or discussed in this revision
and a few footnotes and comments inserted, but his book must be consulted for other data,
English names for most of the species, and many fresh locality records. While we are in sub-
stantial agreement as to the entities of most forms, my phylogenetic conclusions are so widely
divergent from the arrangement devised by FitzSimons that it must be left to others to decide
which more truly represents the probable lines of descent.

2 Revision of the African Lizards of the Family Amphisbaenidae, 1941, Bull. Mus. Comp.
Zool., 87, pp. 353-451.

Revision of the African Lizards of the Family Gerrhosauridae, 1942, Bull. Mus. Comp. Zool.,
89, pp. 483-543., etc.

4 bulletin: museum of comparative zoology

whether the order in which the species of Cordylus are arranged, should
not be largely reversed; it is difficult to believe that so well protected a
species as giganteus should have given rise to less spinose forms without
osteoderms, though such species might gain in activity. However,
without adequate grounds for change I have accepted Boulenger's
order though attracted by the idea that Platysaurus, through Pseudo-
cordyhis, gave rise to some species of Cordylus like caeruleopunctatus
lacking osteoderms. Cape Province, inhabited by 19 species or races,
or the mountainous Transvaal with 17, would certainly appear to be
the centre of speciation.

This opportunity is taken of expressing my deep appreciation of the
kindness shown by Dr. V. FitzSimons of the Transvaal Museum in
checking over my redescriptions of five of the Cordylus described by
him and inserting additional information regarding them; for amending
the spelling of many place names and answering numerous questions
as will be seen from the footnotes. It is with much pleasure that one
of the two novelties:

Platysaurus guttatus fitzsimonsi subspec. nov.

is named after the author of "The Lizards of South Africa." The
other:

PSEUDOCORDYLUS LANGI Spec. nOV.

is named for Mr. Herbert Lang whose activities have enriched so many
collections and greatly contributed to our knowledge of the reptiles
of Africa.

I wish also to thank Dr. Walter Rose for generous permission to re-
produce figs. 2 and 3, on pis. vi and xii respectively, which are taken
from his book "Veld and Vlei."

loveridge: African lizards

Geographical Distribution
of the family Cordylidae

Cordylus giganteus

w. warreni
" w. barberlonensis
" w. perkoensis
" w. vandami

w. depressus

w. laevigaius

w. breyeri

caeruleopunclalus

vittifer
" c. rivae
" c. Iropidosternum

c. Jones ii
" c, angolensis

c. rhodesianus

c. lawrenci
" c. tasmani

c. minor
" c. cordylus

c. niger

peersi

ukingensis

macropholis

cataphraclus
" pusfulatus

namaquensis

campbelli
" p. polyzonus
" p. Jordani
Pseudocordylus capensis

robertsi

langi spec. nov.

m. melanonotus

m. namaquensis

m. microlepidotus

m. fasciatus
Platysaurus g. rhodesianus

g. fdzsimonsi subsp. nov.

g. guttalus

g. lorquatus

g. wilhelmi

g. minor
" g. orienlalis

capensis
Chamaesaura aenea

a. anguina

a. lenuior

miopropus

macrolepis

Total number of forms recorded

3
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6 bulletin: museum of comparative zoology

Index to the Species Recognized

Page

*Cordylus giganteus Smith 16

*Cordylus warreni warreni (Boulenger) 19

*Cordylus warreni barbertonensis (van Dam) 20

Cordylus warreni perkoensis (FitzSimons) 21

*Cordylus warreni vandami (FitzSimons) 23

*Cordylus warreni depressus (FitzSimons) 24

Cordylus warreni laevigatas (FitzSimons) 26

Cordylus warreni breyeri (van Dam) 27

*Cordylus caeruleopunctatus (Methuen & Hewitt) 28

*Cordylus vittifer (Reichenow) 30

Cordylus cordylus rivae (Boulenger) 32

*Cordylus cordylus tropidosternum (Cope) 33

*Cordylus cordylus jonesii (Boulenger) 36

Cordylus cordylus angolensis (Bocage) 38

*Cordylus cordylus rhodesianus (Hewitt) 40

Cordylus cordylus lawrenci (FitzSimons) 41

*Cordylus cordylus tasmani (Power) 42

Cordylus cordylus minor FitzSimons 43

*Cordylus cordylus cordylus (Linnaeus) 44

*Cordylus cordylus niger Cuvier 48

*Cordylus peersi (Hewitt) 50

*Cordylus ukingensis (Loveridge) 51

*Cordylus macropholis (Boulenger) 52

Cordylus cataphractus Boie 53

Cordylus pustulatus (Peters) 57

* Cordylus namaquensis (Methuen & Hewitt) .".... 58

Cordylus campbelli (FitzSimons) 59

Cordylus polyzonus polyzonus Smith 60

*Cordylus polyzonus jordani (Parker) 64

* Pseudocordylus capensis (Smith) 70

* Pseudocordylus robertsi (van Dam) 71

* Pseudocordylus langi spec, nov 73

* Pseudocordylus microlepidotus melanotus (Smith) 75

Pseudocordylus microlepidotus namaquensis Hewitt 78

* Pseudocordylus microlepidotus microlepidotus (Cuvier) 79

* Pseudocordylus microlepidotus fasciatus (Smith) 80

*Platysaurus guttatus rhodesianus FitzSimons 86

*Platysaurus guttatus fitzsimonsi subspec. nov 88

* Represented in the collections of the Museum of Comparative Zoology; examples of species
without asterisk are earnestly desired.

LOVERIDGE: AFRICAN LIZARDS 7

*Platysaurus guttatus guttatus Smith 89

*Platysaurus guttatus torquatus Peters 91

*Platysaurus guttatus wilhelmi Hewitt 92

*Platysaurus guttatus minor FitzSimons 93

Platysaurus guttatus orientalis FitzSimons 94

*Platysaurus capensis Smith 96

*Chamaesaura aenea (Fitzinger) 101

*Chamaesaura anguina anguina (Linnaeus) 102

*Chamaesaura anguina tenuior Giinther 105

*Cha?naesaura miopropus Boulenger 107

*Chamaesaura macrolepis (Cope) 108

Family CORDYLIDAE

1845. Zonuridae Gray (part), Cat. Lizards Brit. Mus., p. 45.
1845. Chamaesauridae Gray, Cat. Lizards Brit. Mus., p. 61.
1884a. Zonuridae Boulenger, Ann. Mag. Nat. Hist. (5), 14, pp. 119, 121.
1923. Zonuridae Camp, Bull. Am. Mus. Nat. Hist., 48, pp. 297, 331.
1937b. Cordylidae Mertens, Abhand. Senckenberg. Naturf. Ges., No. 435,
p. 8.

For further synonymy see Boulenger, 1885e, Cat. Lizards Brit. Mus.,
2, p. 251, from whom the following definition is adapted after incor-
poration of most of the findings of Camp (1923), who should be con-
sulted for further details.

Habit robust, moderate, or serpentif orm ; head covered with sym-
metrical shields; eyes present; eyelids well developed; tympanum dis-
tinct or deeply sunken; dentition pleurodont, teeth numerous, small,
hollow at the base, with long cylindrical shafts; palate toothless;
tongue short, villose, scarcely protractile, entire or very feebly nicked
at the end; body scales, if not granular, forming regular transverse
series; lateral fold present or absent; limbs well developed or rudi-
mentary or absent anteriorly; femoral pores present (though often in-
distinct in females) ; tail stout and spinose, or moderate, or excessively
long and fragile.

Skull with both postorbital and frontosquamosal arches present;
head with dermal bony shields in Cordylus; nasals distinct; frontal,
parietal, and premaxillary single; lower jaw composed of six bones;
palatines and pterygoids widely separated medially, both bordering
the infraorbital fossa; postorbital and frontosquamosal arches bony;

8 bulletin: museum of comparative zoology

supra temporal fossae roofed over by dermal ossifications ; zygosphenal
articulation rudimentary; clavicle slender, not dilated proximally, in-
tercla vicle cruciform with tendency to dilation ; sternum without fonta-
nels; no abdominal ribs; non-tuberculate osteoderms present in most
species of Cordylus.

Range. Africa south of Giacorsa, Ethiopia, i.e. about 5° N. (Re-
ported from Madagascar in error by Cope).

Remarks. Camp (1923) places CORDYLIDAE (as ZONURIDAE)
in a super-family Zonuroidea at the end of the section Anguimorpha,
division Autarchoglossa of the suborder Sauria, thus removing it far
from the IGUANIDAE, but placing it near the ANGUIDAE, the
two families between which it was accorded an intermediate position
by Boulenger (1885e). He would further recognize two subfamilies,
i.e. Zonurinae and Chamaesaurinae, the former would now have to be
termed Cordylinae.

Key to the Genera

1. Habit serpen tiform; limbs short, often rudimentary, or
fore limbs absent; uninjured tail at least thrice the length
of head and body; ear-opening moderate or small; no

collar fold; ventrals lanceolate like dorsals Chamaesaura

(p. 98)
Habit normal; limbs well developed, pentadactyle; tail
less than twice the length of head and body; ear-opening
large; a collar fold at least laterally distinct; ventrals
squarish or transversely enlarged 2

2. Dorsal lepidosis entirely granular; usually a color pattern
of three, light, longitudinal lines, or series of spots, on

back, though sometimes absent in old males Platysaurus

(p. 82)
Dorsal lepidosis heterogeneous or composed of large scales 3

3. Nape covered with granules, or if dorsals extend to occiput
they are much reduced; back usually covered with
granules intermixed with nodular scales or dorsals, the
latter small, soft, and devoid of underlying osteodermal

plates Pseudocordylus

(p. 66)
Nape covered with well developed scales like the dorsals,
the latter strong, usually1 with underlying osteodermal

plates Cordylus

(p. 9)

1 Absent in caeruleopunctatus.

loveridge: African lizards 9

Genus Cordylus

1763. Cordylus Gronovius, Zoophyl., p. 13 (type cordylus Linnaeus). Set
aside by action of International Commission of Zoological Nomencla-
ture (Opinion 89).

1768. Cordylus Laurenti (part), Syn. Rept., p. 51 (type verus Laurenti =
cordylus Linnaeus).

1820. Zonurus Merrem, Versuch Syst. Amphib., p. 57 (type cordylus Lin-
naeus).

For further synonymy (but omit Hemicordylus Smith), see Boul-
enger, 1885e, Cat. Lizards Brit. Mus., 2, p. 252.

Head and body depressed; limbs well developed; tail moderate.
Head shields regular; nostril pierced in the nasal; eyelids well de-
veloped; ear-opening large; 4 parietals; sides of neck covered with
scales; no collar fold; dorsals large, usually bony1, forming regular
transverse series extending to occiput; ventrals large, quadrangular or
subtriangular, juxtaposed or imbricate, forming longitudinal and
transverse series; femoral pores present in both sexes; digits slightly
keeled inferiorly; tail spinose.

Those species which I have examined appear to have the following
characteristics in common, consequently these have been omitted from
the specific descriptions.

Head longer than broad; rostral at least twice as broad as high;
postnasal present only as an aberration; a preocular; anterior supra-
ocular longest, the second broadest; frontal subpentagonal or sub-
hexagonal, slightly broader anteriorly; occipitals present (except in
caeruleopunctatus) ; mental rather large; a slight lateral fold; a pair of
enlarged preanals (though in p. polyzonus and p. jordani occasionally
subequal, and said to be so in the juvenile type of c. rivae); limbs
above with large, keeled spinose, imbricate scales.

Range. Africa, in savanna areas south of Ethiopia, i.e. about 5° N.

Remarks. Stejneger (1936b, p. 137) has set forth the reasons for
recognizing Cordylus Laurenti, in contradistinction to Cordylus
Gronovius, and regarding Merrem's monotypic Zonurus as a synonym.

Despite the fact that for half-a-century herpetologists have been
pointing out the instability in shape (pentagonal, hexagonal, etc.) of
head shields in this genus, and consequently their uselessness for
taxonomic purposes, others continue to stress these unimportant varia-
tions as if they were of value. Which upper labial is lowest and which

1 No osteoderms in young giganteus {fide Broom) , or caeruleopunctatus which, in the absence of
occipitals and reduced nuchals, shows affinities with Pseudocordylus.

10 bulletin: museum of comparative zoology

highest also appears largely fortuitous and might well be dropped from
future descriptions though I have included here all the available data
for some species. It might be worth pointing out that the subocular
descends to the lip in the warreni group, c. niger, peersi, and the two
forms of polyzonus.

Suboculars normally number 3, with 2 or 4 occurring as variants;
in the East African forms of C. cordylus, however, 2 has become normal
at the expense of the third which has been reduced and pushed to one
side, 2 also appears to be normal for a dwarf form (macropholis) known
only from Kleinzee, Little Namaqualand. Van Dam has recorded 5 for
C. barbertonensis, but this is apparently due to his inclusion of a small
scale not truly a subocular.

Upper labials are normally 5 or 6, with 7 occurring as a common
variant ; only in the dwarfed macropholis are they reduced to 4 with 5
still the normal. Lower labials again are normally 5 or 6, with 7 occur-
ring as a rare variation, here it is the dwarf ukingensis (known to me
only from the holotype) which has reduced to 4. In the hope that the
results might prove useful I counted the gulars lying between the
angles of the jaws in all specimens in the Museum of Comparative
Zoology, however as counts were not available for eight species I was
unable to take advantage of this character.

The term postfrontals is employed as preferable to frontoparietals
used by Boulenger and some others. As the head is longer than broad
in all species except in old males of cataphractus, where it is as long as
broad, this character has been dropped from the descriptions.

It will be noted that in the majority of species the keels, or spines,
on the dorsum and dorsal aspect of the tail are less developed than on
the flanks or sides of tail, hence it has not been considered necessary to
repeat this for each species. The relative smoothness of the dorsals
supposedly facilitates the lizard's entry and withdrawal from beneath
boulders, while the lateral spinosities, especially when the lizard in-
flates, tend to catch on the sides of a rock crevice and prove a supple-
mentary hindrance to the owner being withdrawn.

The state of the "lateral fold," characterized as "distinct," "weak,"
"slight," or "strong" by various authors, is largely a reflection of the
state of nourishment of the individual being described. In an emaci-
ated lizard taken after a prolonged drought it will appear "strong," in
a gorged specimen or gravid female, scarcely perceptible. For this
reason I have ignored the lateral fold for taxonomic purposes, though
undoubtedly some degrees of variation in development occur as be-
tween certain species.

loveridge: African lizards 11

Color in members of the genus Cordylus seems to be particularly
susceptible to environmental changes, as is evidenced by Peers, notes on
cataphractus (vide p. 55). Too much importance, therefore, should
not be attached to color in this genus.

This lizard — cataphractus — is the only member of the genus about
which we have anything approaching a complete life history, and it is
to Peers (1930), whose fascinating account should be consulted for
further details, that we are indebted. His paper might well serve as
a model to stimulate young South African naturalists to concentrate
on other members of the genus with a view to filling in the deplorable
gaps in our knowledge of the family.

Key to the Species

1. Lower eyelid opaque; supranasals absent 2

Lower eyelid with transparent disk; supranasals
present or absent (in one species only) 22

2. Rostral in contact with frontonasal; occipital spines

present 3

Rostral in contact with frontonasal only rarely, usually
separated from it; occipital spines absent 6

3. Median subocular not reaching lip; occipital spines
enormous; scales beneath forelimbs smooth; caudal
whorls subequal, gradually diminishing towards tip of
tail; range: Transvaal; Orange Free State; Cape
Province giganteus

(p. 16)
Median subocular reaches lip between two labials;
occipital spines short; scales beneath forelimbs keeled;
large caudal whorls separated by smaller ones 4

4. 34-42 dorsal scales between occiput and base of tail;

range: Zululand w. warreni

(p. 19)
24-32 dorsal scales between occiput and base of tail;
range : Transvaal 5

5. 14-16 ventral scales across the belly races of warreni1

10-12 ventral scales across the belly; range: Waterberg

Mountains, northwestern Transvaal w. breyeri

(p. 27)

1 See full descriptions of various races found in mountains of eastern Transvaal.

12 bulletin: museum of comparative zoology

6. No occipitals; nuchal scales of first six rows greatly-
reduced, minute; range: Cape Province caeruleopunctatus

(p. 28)
Occipitals present; nuchal scales moderate or large. . . 7

7. Nuchals comprising foremost, i.e. postoccipital, row,
twice as large as those in second row; range: Bechua-
naland; Transvaal; Zululand; Natal vittifer

(p. 30)
Nuchals comprising foremost, i.e. postoccipital, row,
subequal, certainly not twice as large as those in second
row 8

8. Interparietal enclosed between two pairs of parietals . . 9
Interparietal on a line with anterior parietals and in
contact with (rarely separated from1) the postfrontals . 18

9. Dorsals elongate, the two vertebral rows not or but

scarcely enlarged; laterals unlike dorsals 10

Dorsals squarish, the two vertebral rows much en-
larged; laterals like dorsals 15

10. Head moderately depressed; temporals moderate;

gulars small 11

Head much depressed; temporals large; gulars moder-
ate or large •. 14

11. Femoral pores 3; nostril said to be pierced in centre of
nasal; range: Ethiopia c. rivae

(p. 32)
Femoral pores 5-8; nostril in lower centre of postero-
inferior corner of nasal; range: Kenya Colony and
countries to the south of Kenya 12

12. Ventrals in 28-34 transverse rows; size larger, length
from snout to vent in adults 82-92 mm. ; range : Kenya

south to Mozambique and Southern Rhodesia c. tropidosternum

(p. 33)
Ventrals in 22-27 transverse rows; size smaller, length
from snout to vent in adults 64-74 mm 13

13. Southern Mozambique; Southern Rhodesia (south of
Bulawayo); Bechuanaland Protectorate (also intro-
duced at Kimberly, Cape Province) c. jonesii

(p. 36)
Southwestern Angola (known only from scanty
description); probably northern South West Africa;

possibly north-central Belgian Congo also c. angolensis

(p. 38)

1 Sometimes in namaquensis.

loveridge: African lizards 13

14. Posterior parietals much larger than the anterior;
dorsals in 27-28 transverse rows; ventrals in 14 longi-
tudinal rows; range: Southern Rhodesia northeast of

Bulawayo c. rhodesianus

(p. 40)
Posterior parietals subequal to the anterior; dorsals in
24 transverse rows; ventrals in 12 longitudinal rows;

range : Little Namaqualand c. lawrenci

(P- 41)

15. Laterals on flanks slightly smaller than dorsals; range:
Uitenhage Division, Cape Province c. tasmani

(p. 42)
Laterals on flanks just as large as the dorsals 16

1 6. Rostral 2 3^-4 times as broad as high ; median subocular
not, or but rarely, descending to the lip; color above
brown or olive with or without markings; range: Cape
Province (exclusive of parts of Uitenhage Division) . . c. cordylus1

(p. 44)
Rostral, 2-2 x/i times as broad as high; median sub-
ocular descending to the lip between fourth and fifth
labials; color above uniformly jet black 17

17. Head shields smooth or slightly rugose; nasal moderate,
not or but slightly swollen; temporals large, rarely
keeled; the two vertebral rows of dorsals enlarged;
femoral pores 5-9 ; range : Cape Peninsula c. niger

(p. 48)
Head shields very strongly rugose; nasal large, very
strongly swollen; temporals very large, strongly
keeled; vertebral rows of dorsals not differentiated
from adjacent dorsals; femoral pores 8-12; range:
Little Namaqualand peersi

(p. 50)

18. 11-14 gulars between angles of jaws; the two vertebral
rows of dorsals enlarged; 14-18 longitudinal rows of

dorsolaterals; 9-12 longitudinal rows of ventrals 19

16-21 gulars between angles of jaws; vertebral rows of
dorsals not enlarged; 20-32 longitudinal rows of
dorsolaterals; 14-29 longitudinal rows of ventrals. ... 20

1 See also the recently described C. c. minor from Matjesfontein, C. P., said to differ from
c. cordylus in having 24-36 longitudinal rows of dorsals, 16 rows of ventrals, only 4-6 femoral
pores, and smaller size.

14 bulletin: museum of comparative zoology

19. Gulars small, the anterior not enlarged; dorsals in 30
transverse rows; ventrals obtusely keeled, in 30 trans-
verse rows; range: Southern Tanganyika Territory. . .• ukingensis

(p. 51)
Gulars very large, a few anterior ones moderately en-
larged followed by a zone of much smaller ones; dor-
sals in 16-19 transverse rows; ventrals smooth in,
20-22 transverse rows; range: Little Namaqualand. . . macropholis

(p. 52)

20. Dorsolaterals in 20-22 longitudinal rows and 15-16
transverse rows; range: Little Namaqualand cataphractus

(p. 53)
Dorsolaterals in 24-32 longitudinal rows and 30-32
transverse rows 21

21. Ventrals in 14 longitudinal rows; range: Central South

West Africa pustulatus

(p. 57)
Ventrals in 18 longitudinal rows; range: Southern

South West Africa namaquensis

tp- 58)

22. Supranasals absent; range: South West Africa campbelli

(p. 59)
Supranasals present; range: South West Africa 23

23. 10-19 femoral pores; upper posterior femorals scarcely
larger or more spinose than the anterior; whorls of
large scales at middle of tail separated by whorls of
smaller ones; a dark streak on side of neck; range:

South West Africa south of Aus p. polyzonus

(p. 60)
5-8 femoral pores; upper posterior femorals much
larger and more strongly spinose than the anterior;
whorls of large scales at middle of tail diminishing
gradually towards tip of tail ; no dark streak on side of

neck; range: South West Africa north of Aus p. jordani

(p. 64)

loveridge: African lizards

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16 bulletin: museum of comparative zoology

CORDYLUS GIGANTEUS Smith

Plate 1, figs. 1-2

1844. Cordylus giganteus A. Smith, 111. Zool. S. Africa, Rept., pis. xxxv-

xxxvi: "Quathlamba" i.e. Drakensberg Mountains. Orange Free
State.
1938. Gorham & Ivy, p. 179.

1845. Zonurus derbianus Gray, Cat. Lizards Brit. Mus., pp. 48, 270: South

Africa.

1890a. Muller, p. 286.

1897. Bateman, p. 106.

1885e. Zonurus giganteus Boulenger, p. 253.

1890b. Muller, p. 699.

1893a. Boettger, p. 66.

1897. Bateman, p. 106.

1898. Sclater, p. 103.

1901. Gadow, p. 536, fig. 135.

1909a. Hewitt, p. 35.

1910b. Boulenger, p. 467.

1910. Ditmars, p. 155, pi. xxxi, fig. 2.
1910a. Hewitt, p. 60.

1911. Gilchrist, p. 230.
1911b. Hewitt, p. 47.

1913. Broom, p. 548, pi. lxxiv.

1913. Boulenger, E.G., p. 72, pl-

1913. Hewitt & Power, p. 153.

1913a. Werner, p. 108, pi. -

1914a. Hewitt, p. 240, map.

1920. Hewitt, pp. 91, 93.

1921a. Dam, p. 242.

1925b. Flower, p. 945.

1929. Rose, p. 100.

1930. FitzSimons, V., p. 29.
1930. Power, p. 14.

1930. Schmidt, p. 153.

1932. FitzSimons, F. W., p. 212.

1935b. FitzSimons, V., p. 349.

1935. Lawrence, p. 44.

1937a. FitzSimons, V., p. 266.

1937. Lawrence, p. 109, fig.

Names. Giant Girdle-tail (English); sonkyker or ouvolk (Afrikaans).

Description. Head slightly depressed; head shields strongly rugose;

rostral in contact with the frontonasal, which is longer than broad, as

loveridge: African lizards 17

long as broad, or shorter than broad; nostril pierced in the postero-
inferior corner of a nasal which is not much swollen; a loreal; a pre-
ocular; median subocular not descending to the lip; fourth or fifth
upper labial smallest, fifth not higher than others ; prefrontals forming
a suture; frontal hexagonal; postfrontals longer than broad; inter-
parietal on a line with, and as large as, the anterior parietals, forming
a suture with the postfrontals; posterior parietals much larger than
the anterior; 4 large, striated, occipital spines, the inner shorter and
slightly curved inwards, the outer largest and slightly curved inwards,
or straight, or directed obliquely outwards, as long as, or longer than,
the three anterior supraoculars; temporals large, rugose, keeled, the
hindmost subtriangular, slightly curved and bluntly pointed ; sides of
neck with large, sharp, slightly curved or erect spines ;gulars small,
the anterior irregularly enlarged, the median slightly imbricate,
smooth, or the lateral obtusely keeled; collar scales large, keeled,
lanceolate, mucronate.

Dorsals subquadrangular, rugose, strongly keeled, sharply mucro-
nate, the vertebral row often somewhat smaller; laterals keeled,
spinose, separated by granular interspaces; ventrals subquadrangular,
the median smooth and strongly imbricate, the lateral obtusely keeled
and shortly mucronate; scales below both fore and hind limbs smooth;
tail with whorls of large, rugose, strongly keeled but not serrate,
spinose scales above and on sides, the lateral spines longest; sub-
caudals elongate, narrow, smooth or obtusely keeled.

For characters common to all species, ■ 9; for scale and pore

counts, see statistical table on p. 15. I g giganteus all scales

are said to be more strongly imbricate and all spines less developed.

Color. Above, head dark brown, lip to ear yellow; back yellowish
clouded with dark brown. Below, gular region and belly whitish or
yellowish, uniform.

That of the young, according to Smith : Above, h&ad anteriorly light
yellowish brown clouded with liver brown ; temples liver brown irregu-
larly spotted with greenish white; posterior part of head, back and
sides intermediate between umber and liver brown, the back, limbs,
and tail irregularly barred with narrow yellow cross-bands; tail-tip
orange red. Below, creamy yellow, except tail-tip which is orange red.
For detailed description of a still earlier stage, and coloured plate of
mother and young, see Broom (1913).

Size. Total length of a ^ type (? derbianvs) 382 (202 + 180) mm.
(Boulenger, 1885e). Smith's measurements of 8 + 7 inches ( = 204
+ 179) mm., for a cT cotype, is rejected by FitzSimons (1937a) who

18 bulletin: museum of comparative zoology

remeasured it as 351 (191 + 160) mm. Total length of a 9 , 365
(195 + 170) mm. (FitzSimons, 1937a). There is nothing to support
the statement by Rose (1929) that the species attains "nearly two
feet," though it appears to have been copied by later authors.

Anatomy. The so-called branchial arch is discussed by Hewitt
(1920), the gall bladder by Gorham and Ivy (1938).

Longevity. Four years, six months, and nineteen days, is the longest
of six records furnished by Flower (1925b).

Diet. In captivity, cockroaches, mealworms, tiny frogs, baby mice,
and raw meat (Bateman, 1897).

Parasites. No mites were found on those examined by Lawrence
(1935).

Defence. F. W. FitzSimons (1932) records that when a hungry mole
snake (Pseudaspis cana) was presented with one of these spiny
lizards, the serpent stalked, seized, and began to constrict it with three
coils; for a time it tightened its coils, then, suddenly relaxing them,
dashed away.

Similarly a captive puff adder (Bitis arietans), having seized a girdle-
tailed lizard by the head, began to engulf it in leisurely fashion. The
lizard, however, which had been quiescent up to this point, began to
slew round and round and to belabour the snake so effectively with its
spinous tail that the viper was glad to disgorge its prey and retire
from the contest.

Habits. According to V. FitzSimons (1935b), these big lowland
lizards are fairly common in the open flat country east of Kroonstad,
where they may be seen sunning on small mounds or anthills. When
disturbed they disappeared into burrows — apparently those of
gerbils.

Its custom, when basking, of carrying its head high and facing the
sun, has won for it the name of "sun-gazer" among the Dutch (Rose).

Apparently unused to water in their rocky haunts, these lizards
would scuttle into a moat "swim frantically for a few seconds, then
sink like a stone to the bottom," there to remain until drowned
(F. W. FitzSimons, 1932).

In Europe captive girdle-tails require artificial heat of from 70° to
85° F., except during the summer months (Bateman, 1897).

Localities. Transvaal: Heidelberg; Paardekop Station, Stand-
erton District; Schaapplaats, Vereeniging District; Zandspruit,
Wakkerstroom District. Orange Free State: Bethlehem; Bloem-
fontein; Lower western slopes Drakensberg Mountains; Geluk
Farm, 20 miles west of Kroonstad; Harrismith; Hoopstad; Kroon-

loveridge: African lizards 19

stad; Ventersburg. Cape Province: Colesberg (over s.w. border
from Orange Free State).

Range. Southern Transvaal, Orange Free State and adjacent areas
in Cape Province and Basutoland.

Cordylus warreni warreni (Boulenger)

Plate 1, fig. 3

1908b. Zonurus warreni Boulenger, Ann. Natal Govt. Mus., 1, pp. 224, 232

pi. xxxv : Ubombo, Zululand.
1909a. Hewitt, p. 36.
1910b. Boulenger, p. 468.
1910a. Hewitt, p. 60.
1930. FitzSimons, p. 30.

1930. Power, p. 14.

1935. Lawrence, p. 43.

1937. Lawrence, p. 111.

Description. Head slightly depressed; head shields rugose; rostral
narrowly in contact with the frontonasal, which is as long as broad;
nostril pierced in the postero-inferior corner of a nasal which is not
much swollen; a loreal; a preocular; median subocular descending to
the lip between the third and fourth or fourth and fifth upper labials ;
fourth, fifth, or sixth upper labial smallest, fifth not higher than others ;
prefrontals forming a broad or narrow suture, or separated; post-
frontals shorter than broad; interparietal enclosed between 2 pairs of
parietals ; posterior parietals slightly larger than the anterior ; 6 rugose,
pointed occipitals; temporals rugose, keeled, the hindmost subtri-
angular, slightly curved and bluntly pointed; sides of neck with large,
sharp, erect spines; gulars small, the anterior irregularly enlarged, the
median not imbricate, smooth, or the lateral obtusely keeled; collar
scales enlarged, keeled, the lateral mucronate.

Dorsals subquadrangular, rugose, strongly keeled, lateral shortly
mucronate, the two vertebral rows reduced; laterals keeled, spinose,
separated by granular interspaces ; ventrals quadrangular, the median
smooth (in females) or obtusely keeled (in males), not, or but slightly
imbricate, the lateral obtusely keeled and shortly mucronate; scales
below fore limbs keeled, below hind limbs smooth or obtusely keeled;
tail with whorls of large, rugose, strongly keeled but not serrate,
spinose scales, alternating with whorls of smaller scales above and
below, the lateral spines longest.

20 bulletin: museum of comparative zoology

For characters common to all species, see definition on p. 9, for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown, lip to ear yellow; back brown, flecked
with black-edged, yellow spots forming more or less regular transverse
series. Below, gular region whitish with brown infuscations; belly yel-
lowish white, uniform.

Size. Total length of cT type, 270 (110 + 160) mm.; of a topotypic
9 (M.C.Z. 21442), 267+ (122 4- 145+) mm.

Remarks. None of the citations given above, excepting that of
Lawrence, contribute to our knowledge.

Breeding. In March and November ova are small (M.C.Z. 21442,
41881).

Diet. Thirty-two Eristalis maggots, each about 20 mm. long, and
fragments of a large snail's shell in one; millipedes and ants in another;
a large grasshopper in a third (A.L.).

Parasites. Mites (Zonurobia debilipes) described from this species
by Lawrence (1935). Nematodes (Thubunaea sp., probably T. agamae
Sandground, and Heterakidae, probably Spinicauda sp.), and trema-
todes (Mesococlium sp.) were taken from topotypic lizards in the
Museum of Comparative Zoology. I am indebted to Messrs Allen
Mcintosh and J. T. Lucker for making the identifications.

Localities. Zululand: Ubombo.

Range. Zululand.

CORDYLUS WARRENI BARBERTONENSIS (van Dam)

1921a. Zonurus barbertonensis van Dam, Ann. Transvaal Mus., 7, p. 240»

pi. iii: Barberton, Transvaal.
1930. FitzSimons, p. 29.

1930. Power, p. 14.

1935. Lawrence, p. 43.

Description. Head slightly depressed; head shields rugose; rostral in
contact with the frontonasal, which is as long as broad, or shorter tha n
broad; nostril pierced in the postero-inferior corner of a nasal which is
not much swollen; a loreal; a preocular; median subocular descending
to the lip between the fourth and fifth upper labials; fourth or fifth
upper labial smallest, fifth not higher than others; prefrontals forming
a suture; postfrontals shorter than broad; interparietal enclosed be-
tween 2 pairs of parietals ; posterior parietals slightly larger than the
anterior; 6 rugose, pointed occipitals; temporals rugose, keeled, the

. loveridge: African lizards 21

hindmost subtriangular, slightly curved and bluntly pointed; sides of
neck with large, sharp, erect spines ; gulars small, the anterior irregu-
larly enlarged, the median not imbricate, smooth, or the lateral ob-
tusely keeled; collar scales enlarged, keeled, the lateral mucronate.

Dorsals subquadrangular, rugose, strongly keeled, lateral shortly
mucronate, the two vertebral rows reduced; laterals keeled, spinose,
separated by granular interspaces ; ventrals quadrangular, the median
smooth, not, or but slightly imbricate, the lateral obtusely keeled
and shortly mucronate; scales below fore limbs keeled, below hind
limbs smooth or obtusely keeled; tail with whorls of large, rugose,
strongly keeled but not serrate, spinose scales, alternating with whorls
of smaller scales above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9, for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown flecked with yellow, lip to ear yellow;
back, blackish brown flecked with yellow spots forming more or less
regular transverse series. Below, gular region whitish with brown in-
fuscations; belly brownish variegated with yellow anteriorly and on
the flanks.

Size. Total length of tf type, 312 (138 + 174) mm.

Remarks. Its describer states that barbertonensis differs from breyeri
"in size of the occipital spines, and number of transverse series of
dorsal scales." While this is correct, it will be seen from the statistical
table that the latter difference is bridged by intermediate forms.
There are other differences separating this, the best marked, race from
breyeri. I proposed treating barbertonensis and allied forms as races
of breyeri, but follow FitzSimons (1943) in regarding them as forms of
warreni.

Parasites. Mites (Zonurobia transvaaleiisis) were described from this
race by Lawrence.

Localities. Transvaal: Barberton.

Range. Transvaal (Barberton only).

CORDYLUS WARRENI PERKOENSIS (FitzSimons)

Plate 2, fig. 1

1930. Zonurus vandami perkoensis FitzSimons, Ann. Transvaal Mus., 14, pp.
27, 30, figs. 8-9: Perkoe Farm, near Olifants River, eastern Transvaal.
1935. Lawrence, p. 43.

Description. Head slightly depressed; head shields posterior o
ocular region rugose; rostral in contact with the frontonasal, whirl is

22 bulletin: museum of comparative zoology

longer than broad; nostril pierced in the postero-inferior corner of a
nasal which is not much swollen; a loreal; a preocular; median sub-
ocular descending1 to the lip between the fourth and fifth upper labials ;
fifth upper labial smallest, fifth not higher than others; prefrontals
forming a suture; postfrontals shorter than broad; interparietal en-
closed between 2 pairs of parietals or largely on a line with the anterior
parietals, forming, or nearly forming, a suture with the postfrontals;
posterior parietals slightly larger than the anterior; 4-5 rugose, pointed
occipitals; temporals rugose, keeled, the hindmost subtriangular,
slightly curved and bluntly pointed; sides of neck with large, sharp,
erect spines; gulars small, the anterior irregularly enlarged, the median
not imbricate, smooth collar scales not enlarged mesially, faintly
keeled, the lateral keeled and mucronate.

Dorsals subquadrangular to subcircular, finely rugose, strongly
keeled, lateral shortly mucronate, the two vertebral rows reduced;
laterals keeled, spinose, separated by granular interspaces; ventrals
quadrangular, the median smooth, not or but slightly imbricate, the
lateral keeled and mucronate, the 2-3 outermost rows much smaller
and almost spinose; scales below forelimbs keeled and mucronate, below
thigh obtusely keeled and shortly mucronate, below tibia strongly
keeled and mucronate; tail with whorls of large, rugose, strongly
keeled but not serrate spinose scales, alternating with whorls of smaller
scales above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9, for
scale and pore counts, see statistical table on p. 15.

Color. Above, head and back dark brown, uniform except for a few
isolated yellow flecks on flanks and tail. Below, gular region yellowish
white with brown inf uscations ; belly mummy brown, uniform except
for a series of short, irregular, transverse bars of yellow at sides and
some traces of same on limbs ; tail with a few scattered flecks of yellow-
ish white.

Size. Total length of 9 type, 276 (126 + 150) mm.

Remarks. Known only from two females. Its describer states that
perkoensis differs from yandami as follows: "head wider and deeper (as
clearly shown in the figures); greater arching of the supraoccipital
ridge; well marked suborbital ridge; short contact of the prefrontals;
colouring on the whole more uniform." As I have seen neither of the
types, I can only add that the grounds for separation appear slen-
der. FitzSimons (1943) continues to regard perkoensis as a race of
vandami.

Breeding. During first week in November (late spring) both females

LOVERIDGE: AFRICAN LIZARDS 23

held eggs, those in the paratype in an early stage, those in the type
containing well-formed embryos curled about the yolk.

Parasites. Mites (Zonurobia transvaalensis) were described from this
race by Lawrence.

Temperament. Gravid females were excessively shy, which, in con-
junction with the inacessibility of their retreats, made them difficult
to capture.

Habitat. Living at an altitude of about 5000 feet among huge rock
masses which had broken away from the slopes above.

Localities. Transvaal: Perkoe Farm near Olifants River.

Range. Transvaal (Eastern only).

CORDYLUS WARRENI VANDAMI (FitzSimOns)

1930. Zonurus vandami FitzSimons, Ann. Transvaal Mus., 14, pp. 25, 30,

figs. 6-7: Gravelotte, near Leydsdorp, eastern Transvaal.
1935. Lawrence, p. 43.

Description. Head slightly depressed; head shields rugose; rostral in
contact with the frontonasal, which is longer than broad or as long as
broad; nostril pierced in the postero-inferior corner of a nasal which is
not much swollen; a loreal; a preocular; median subocular descending
to the lip between the third and fourth or fourth and fifth upper
labials; fourth or fifth upper labial smallest, fifth not higher than
others; prefrontals forming a suture; postfrontals longer than broad;
interparietal enclosed between 2 pairs of parietals ; posterior parietals
slightly larger than the anterior; 4-8 rugose, pointed occipitals; tem-
porals rugose, keeled, the hindmost subtriangular, slightly curved and
bluntly pointed; sides of neck with large, sharp, erect spines; gulars
small, the anterior irregularly enlarged, the median not imbricate,
smooth, or the lateral obtusely keeled; collar scales enlarged, keeled,
the lateral mucronate.

Dorsals subquadrangular, rugose, strongly keeled, lateral shortly
mucronate, the two vertebral rows reduced; laterals keeled, spinose,
separated by granular interspaces ; ventrals quadrangular, not, or but
slightly imbricate, the lateral obtusely keeled but not shortly mucro-
nate; scales below fore limbs keeled, below hind limbs smooth or ob-
tusely keeled; tail with whorls of large, rugose, strongly keeled but
not serrate, spinose scales, alternating with whorls of smaller scales
above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9, for
scale and pore counts, see statistical table on p. 15.

24 bulletin: museum of comparative zoology

Color. Above, head and back dark brown to sepia, a yellow, ir-
regular, transverse bar on nape, four others on body, one at root of
tail; similar yellow barring on limbs and tail. Below, yellowish with
brown inf uscations ; tail broadly banded with yellow.

Size. Total length of cf paratype (T.M. 7410), 273 (155 + 118)
mm.; of type 9 (T.M. 7407), 287 (155 + 132) mm.

Remarks. Its describer states that vandami differs from barbcrto-
nensis by its slightly narrower and more depressed head, greater
rugosity of head shields, and 4 occipitals. The paratype series of 8
lizards, however, was comprised of individuals with 4, 5, 6 (M.C.Z.
41876), and even 8 occipitals, the last, as pointed out by FitzSimons,
resulting from the subdivision of the original 4, this character, there-
fore, cannot be considered of diagnostic value, though used as a key
character by FitzSimons (1943) who continues to regard it as a full
species. In so large a lizard the alleged difference of 25 mm. in size
can not have much significance.

Parasites. Mites (Zonurobia transvaalensis) were described from
this race by Lawrence.

Habitat. Occurs between 4000 and 5000 feet on the northeastern
spur of the Drakensberg.

Localities. Transvaal: Leydsdorp District: Gravelotte; Malta;
Skelem.

Range. Transvaal (Drakensberg Mountains).

CORDYLUS WARRENI DEPRESSUS (FitzSimons)

1930. Zonurus barbertonensis deprcssus FitzSimons, Ann. Transvaal Mus., 14,
pp. 24, 29: Newgate Farm, near Louis Trichardt, Zoutpansberg
Mountains, northern Transvaal.

1935. Lawrence, p. 43.

Description. Head slightly depressed; head shields rugose; rostral
in contact with the frontonasal, which is longer than broad; nostril
pierced in the postero-inferior corner of a nasal which is not much
swollen; a loreal; a preocular; median subocular descending to the lip
between the fourth and fifth upper labials ; fifth upper labial smallest,
fifth not higher than others ; prefrontals forming a suture ; postfrontals
longer than broad; interparietal on a line with the anterior parietals,
forming a suture with the postfrontals; posterior parietals slightly
larger than the anterior; 6 rugose, pointed occipitals ; temporals rugose,
keeled, the hindmost subtriangular, slightly curved and bluntly

loveridge: African lizards 25

pointed ; sides of neck with large, sharp, erect spines ; gulars small, the
anterior irregularly enlarged, the median not imbricate, smooth, or the
lateral obtusely keeled; collar scales enlarged, keeled, the lateral
mucronate.

Dorsals subquadrangular, rugose, strongly keeled, lateral scarcely
mucronate, the two vertebral rows reduced; laterals keeled, spinose,
separated by granular interspaces ; ventrals quadrangular, smooth, not
or but slightly imbricate, the lateral obtusely keeled but not shortly
mucronate; scales below fore limbs keeled, below hind limbs smooth
or obtusely keeled; tail with whorls of large, rugose, strongly keeled
but not serrate, spinose scales, alternating with whorls of smaller scales
above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9, for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown flecked with yellow; back brown with
yellow spots forming more or less regular transverse series. Below,
yellowish with brown infuscations.

Size. Total length of 9 paratype (M.C.Z. 41869), 250+ (120 +
130+) mm. Both paratype cf and type 9 had the same body length
but the tails in all six of the type series were damaged.

Remarks. Its describer states that depressus differs from barberto-
nensis in having the "head considerably more depressed and rugose:
occipital spines not so elongate and subequal : dorsal scales larger and
more rugose: lateral spines on tail shorter and sturdier, directed out-
wards at greater angle: scales on palms of hands and soles of feet
larger and thus fewer in number."

I must confess that, having compared our aged and worn paratype
of depressus with a topotype of barbcrtonensis, the only differences that
I can detect are those affecting the dorsals and possibly some slight
difference in scales on "palms" and soles.

Breeding. Early in July (midwinter) three of the four females held
eggs, the oviduct of one containing "two large undeveloped eggs."
(FitzSimons).

Diet. All were emaciated, probably due to being taken in midwinter.

Parasites. Mites (Zonurobia transvaalensis) were described from this
race by Lawrence.

Habitat. Occurs between 4800 to 5000 feet on the Zoutpansberg, in
or near rock crevices caused by weathering.

Localities. Transvaal: Newgate Farm near Louis Trichardt.

Bnnge. Transvaal (Zoutpansberg Mountains).

26 bulletin: museum of comparative zoology

CoRDYLUS WARRENI LAEVIGATUS (FitzSimons)

Plate 2, fig. 2

1933. Zonurus laevigatus FitzSimons, Ann. Transvaal Mus., 15, p. 276, figs.

1-2: Entabeni, Zoutpansberg Mountains, northern Transvaal.
1935. Lawrence, p. 43.

Description. Head slightly depressed; head shields slightly rugose;
rostral in contact with the frontonasal, which is as long as broad;
nostril pierced in the postero-inferior corner of a nasal which is not
much swollen; a loreal; a preocular; median subocular descending to
the lip between the fourth and fifth upper labials; fifth upper labial
smallest and not higher than others; prefrontals forming a suture ;post-
f rentals longer than broad; interparietal on a line with the anterior
parietals, forming a suture with the postf rentals ; posterior parietals
much larger than the anterior; 6 rugose, pointed occipitals; temporals
rugose, keeled, the hindmost subtriangular, slightly curved and bluntly
pointed; sides of neck with a few large, sharp, erect spines; gulars
small, the anterior irregularly enlarged, the median not imbricate,
smooth, or the lateral almost imperceptibly keeled; median collar
scales enlarged, smooth, the lateral keeled, mucronate.

Dorsals subquadrangular, finely rugose, keeled, the two vertebral
rows reduced; laterals keeled, spinose, separated by granular inter-
spaces ; ventrals quadrangular, the median smooth, not imbricate, the
lateral obtusely keeled and shortly mucronate; scales below fore
limbs keeled, below thigh smooth, below tibia, obtusely keeled and
mucronate; tail with whorls of large, rugose, strongly keeled but not
serrate, spinose scales, alternating with whorls of smaller scales above
and below, the lateral spines longest.

For characters common to all species, see definition on p. 9, for
scale and pore counts, see statistical table on p. 15.

Color. Above, head and body dark brown sparsely spotted with
yellow. Below, slightly paler, gular region with darker infuscations.

Size. Total length of 9 type (T.M. 14229) 226 (116+ 110) mm.

Remarks. Its describer states that laevigatus may be readily dis-
tinguished from barbertonensis, vandami, and breyeri by "its much
slighter build and pronounced reduction of the rugosity and spinosity
of the scales generally." Having seen neither of the females on which
this form was based, I can only add that the grounds of separation
appear slender. FitzSimons (1943) continues to regard it as a full
species.

loveridge: African lizards 27

Parasites. Mites (Zonurobia transvaalensis) were described from this
race by Lawrence.

Habitat. Occurs between 5000 and 6000 feet on the Zoutpansberg,
beneath loose stones and in rock crevices.

Localities. Transvaal: Entabeni.

Range. Transvaal (Zoutpansberg Mountains).

CORDYLUS WARRENI BREYERI (van Dam)

Plate 2, fig. 3

1921a. Zonurus breyeri van Dam, Ann. Transvaal Mus., 7, p. 239, pis. i-ii:
Geelhoutkop Farm, about 45 miles north of Nylstroom, Waterberg
District, northwestern Transvaal.

1930. FitzSimons, p. 29.

1930. Power, p. 14.

1935. Lawrence, p. 44.

1937. Lawrence, p. 111.

Description. Head slightly depressed; head shields rugose; rostral
in contact with the frontonasal, which is as long as broad; nostril
pierced in the postero-inferior corner of a nasal which is not much
swollen; a loreal; a preocular; median subocular descending to the lip
between the fourth and fifth upper labials; fifth upper labial smallest,
fifth not higher than others ; prefrontals forming a suture ; postfrontals
about as long as broad; interparietal on a line with the anterior
parietals, forming a suture with, or separated from, the postfrontals;
posterior parietals slightly larger than the anterior; 6 rugose, pointed
occipitals; temporals rugose, keeled, the hindmost subtriangular,
slightly curved and bluntly pointed; sides of neck with large, sharp,
erect spines ; gulars small, the anterior irregularly enlarged, the median
not imbricate, smooth, or the lateral obtusely keeled; collar scales
enlarged, the lateral keeled and mucronate.

Dorsals subquadrangular, rugose, strongly keeled, lateral shortly
mucronate, the two vertebral rows reduced; laterals keeled, spinose,
separated by granular interspaces ; ventrals quadrangular, the median
smooth, not or but slightly imbricate, the lateral obtusely keeled but
not imbricate; scales below fore limbs keeled, below thigh smooth or
obtusely keeled, below tibia, keeled; tail with whorls of large, rugose,
strongly keeled but not serrate spinose scales, alternating with whorls
of smaller scales above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9, for
scale and pore counts, see statistical table on p. 15.

28 bulletin: museum of comparative zoology

Color. Above, head brownish black; back brown; flanks yellowish
brown; tail grayish brown. Below, slate gray; tail slightly lighter than
above.

Size. Total length of 9 type (T.M. 3769), 275 (120 + 115) mm.

Remarks. Its describer contrasted breyeri with giganteus from which
it differs greatly in many respects, specifically in the presence of alter-
nating whorls of smaller scales on the tail in the entire warrcni group.

The typical form is known only from the adult type, three subadult
lizards, and six foetal young. The latter [circa 90 mm.) are said to
differ from the adult in the head shields being smooth ; occipitals only
keeled, not sharply pointed ; median as well as lateral dorsals strongly
keeled; smaller whorls of caudal scales distinctly visible from above,
but not on the sides and below where they are hidden by the spines.

Breeding. In January the gravid type was found to contain 6 young,
the largest measuring 90 mm.

Parasites. Mites (Zonurobia circularis latior) were described from
this race by Lawrence.

Habitat. Occurs at about 5000 feet in Waterberg District, among
rocks.

Localities. Transvaal: Geelhoutkop Farm.

Range. Transvaal (Waterberg Mountains).

Cordylus caeruleopunctatus (Methuen & Hewitt)

Plate 3, fig. 3

1913c. Zonurus caeruleopunctatus Methuen & Hewitt, Trans. Roy. Soc. S.

Africa, 3, p. 110: Buff el's Nek, between Knysna and Avontuur,

Cape Province.
1914a. Hewitt, p. 239.
1925. Essex, p. 339.

1928a. Essex (1927), p. 932.
1930. FitzSimons, p. 29.

1930. Power, p. 13, pi. ii, fig. 2.

1935. Lawrence, p. 44.

1937. Lawrence, pp. 109, 111.

Description.. Head slightly depressed; head shields smooth ante-
riorly, slightly rugose posteriorly; rostral in contact with, or separated
from, the frontonasal, which is shorter than broad ; nostril pierced in
the lower centre of a nasal which is not much swollen; a loreal; a

loveridge: African lizards 29

preocular; median subocular descending to the lip between the fourth
and fifth upper labials; fourth upper labial lowest, fifth higher than
others except first; prefrontals forming a suture, or separated; post-
frontals as long as broad; interparietal enclosed between 2 pairs of
parietals ; posterior parietals much larger than the anterior ; no occipi-
tals; anterior nuchals very small; temporals large, rugose, keeled,
without spines; sides of neck with keeled scales; gulars moderate, the
anterior irregularly enlarged, the median not imbricate, smooth;
collar scales larger but undifferentiated.

Dorsals elongate, strongly keeled, a few shortly mucronate, posteri-
orly serrate, the two vertebral series enlarged ; laterals elongate, keeled ;
ventrals quadrangular, smooth, not or but slightly imbricate; scales
below fore limbs keeled, below hind limbs smooth or obtusely keeled;
tail with whorls of large, strongly keeled, slightly mucronate and
serrate scales above and below, the subcaudals being long, narrow,
pentagonal, keeled.

For characters common to all species, see definition on p. 9, for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown; back blackish brown variegated ante-
riorly with irregular, longitudinal, fawn-colored stripes ; sides of head
and flanks flecked with bright blue. Below, gular region orange (white
in alcohol) ; belly greenish white to bluish; tail brownish.

Size. Total length of type (T.M. 1920) 150+ (65 + 85+) mm., of
another (from Power Siding) 182 (72 + 110) mm.

Remarks. The describers suggest that this lizard is related to war-
reni and capensis (now in Pseudocordylus) . Power (1930) thinks to
cordylus. It differs from all other members of the genus in lacking
occipitals and in the greatly reduced nuchals, both characters appar-
ently demonstrating affinities with Pseudocordylus with which it also
agrees in lacking osteoderms.

Parasites. Mites (Ixodiderma inverta and Zonurobia sanguined) were
found by Lawrence.

Temperament. When basking in the sun this lizard is extremely
difficult to catch. Of thirty lizards counted on a mile-long wall only
three were captured (Essex). Its eyesight appears abnormally keen
making it difficult to approach even within shooting distance of one
of these lizards (Lawrence).

Habitat. Among rocks on a mountain within the Cape Flora vege-
tational zone, no forest being present.

Localities. Cape Province : Buffet's Nek between Avontuur and
Knysna; Montagu Pass, Outeniqua Mountains; Power Siding,

30 bulletin: museum of comparative zoology

halfway up Montagu Pass; Prince Alfred's Pass, Outeniqua
Mountains; Silver River between George and the Wilderness.
Range. Cape Province (Outeniqua Mountains and vicinity).

Cordylus vittifer (Reichenow)

1887. Zonurus vittifer Reichenow, Zool. Anz., 10, p. 372: Transvaal.

1891a. Matschie, p. 606.

1894e. Boulenger, p. 724.

1898. Sclater, p. 104.

1908b. Boulenger, p. 224.

1909a. Hewitt, pp. 31, 37.

1910a. Hewitt, p. 56.

1911b. Hewitt, p. 74.

1911b. Sternfeld, p. 401.

1911c. Sternfeld, p. 419.

191 Id. Sternfeld, p. 21, fig. 20.

1914a. Hewitt, p. 239.

1921a. Dam, p. 242.

1927c. Power, p. 407.

1930. FitzSimons, p. 30.

1935. Broom, p. 20, fig. 5b.

1935. Lawrence, p. 44.

1889. Zonurus cordylus Boettger (not Linn6), p. 287.

1910b. Boulenger (part), p. 468.

1907b. Zonurus cordylus var. vittifer Roux, p. 418.

1908. Odhner, p. 3.

1930. Power, p. 17.

1931. Power, pp. 41, 48. (as vettifer also).

1910b. Zonurus tropidogaster Boulenger, Ann. S. African Mus., 5, pp. 468,.

495: Barberton, Transvaal.
1911b. Hewitt, p. 47.
1930. Power, p. 15.

Further citations of "vittifer" will be found under cordylus angolcnsis
and c. tropidosternum.

Description. Head much depressed; all head shields strongly rugose;
rostral separated from the frontonasal, which is longer than broad, as
long as broad, or sometimes absent1; nostril pierced in the postero-
inferior corner of a large nasal which is not much swollen; a loreal
present or absent2; a preocular; the median subocular not descending
to the lip; fourth upper labial lowest, fifth highest; prefrontals forming

1 In type of Iropidogasler only.

2 For example fused with preocular in M.C.Z. 41879-80.

loveridge: African lizards 31

a suture, or separated; postfrontals longer than broad, as long as
broad, or shorter than broad; interparietal enclosed between 2 pairs
of parietals or on a line with the anterior parietals separated from or
forming a suture with the postfrontals; posterior parietals much larger
than the anterior; 4-6 rugose, subequal occipitals; anterior row of
nuchals twice as large as the second; temporals large, rugose, slightly
keeled, without spines; sides of neck with keeled, spinose scales; gulars
small, the anterior irregularly enlarged, the median slightly imbricate,
smooth, or the laterals only obtusely keeled and even mucronate1;
collar scales larger but undifferentiated.

Dorsals elongate, rugose, keeled, very shortly mucronate, not or
but slightly serrate, those on the vertebral line not or but rarely
differentiated; laterals keeled, serrate, spinose; ventrals quadrangular,
smooth or keeled2, not or but slightly imbricate; scales below fore
limbs keeled, below hind limbs smooth or slightly keeled; tail with
whorls of large, strongly striate, keeled, serrate, spinose scales above
and below, the lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown ; back yellowish, yellowish brown, reddish
brown, uniform or variegated or spotted with darker, a yellow verte-
bral line present or absent. Below, whitish, uniform. Color of young
richer than that of adults. ,

Size. Total length of largest & (M.CZ. 12416) 172+ (83 + 89+)
mm. (ex. Rhoodeplaat) ; and largest 9 (M.CZ. 21364) 160+ (82+78+)
mm. (ex. Doornkop). The type (Berlin Mus.) was only 112 (55 + 57)
mm., while the type of tropidogaster (Brit. Mus.?) was still smaller,
having a length from snout to anus of only 48 mm.

Remarks. Z. tropidogaster Boulenger was based on a young lizard
which lacked a frontonasal and possessed keeled ventrals. The latter
condition has long been known to occur in occasional specimens of
vittifcr (Roux, 1907b) and such as a specimen from Zululand (M.CZ.
45416) or Marieskop, Transvaal (M.CZ. 41877), so that I entirely
concur with Hewitt's (1911b, p. 147) suggestion that it is nothing but
a synonym of vittifcr. Such a view is reinforced by the occurrence of
typical vittifcr at Barberton, type locality of tropidogaster.

Tornier (1896), impressed by the similarities between vittifcr and
cordylus synonymized the former with the latter. Roux (1907b) on
account of stable differences revived it as a variety. Hewitt (1909a)

i In M.CZ. 41877.

2 See Remarks.

32 bulletin: museum of comparative zoology

rightly restored it to full specific rank on account of the constantly
longer nasal suture, greatly enlarged first row of nuchals, and more
numerous dorsals.

With regard to the latter, it is clear from the context that Roux's
(1907b) counts of longitudinal dorsal rows do not include the laterals,
whereas Sternfeld (1911b), in common with most recent authors, does
include them. Had this been realized much unnecessary discussion
might have been avoided. Monard's (1937b) suggestion that the
Angolan cordylus of Bocage are referable to vittifer is untenable.

Anatomy. Broom (1935) discusses the temporal region of the skull.

Parasites. Mites (Zonurobia circidaris latior) have been described
from this species by Lawrence.

Habitat. Living under stones and in rocky crevices of kopjes (van
Dam) chiefly in the low veld portions of the Transvaal (Hewitt), and
one found at a height of four feet from the ground in a small tree at
Lobatsi (Power).

Localities. Bechuanaland : Lobatsi (fide Power). Transvaal:
Arnhemburg; Barberton; Belfast; Botschabelo near Middleburg;
Carolina; Doornkop near Belfast; Entabeni; Frederikstad; Haen-
ertsburg; Johannesburg; Koster; Krugersdorp; Linokana; Maka-
pan; Marico; Mariepskop ; Metlepetsi River; Middleburg District;
Mphome (Mpoma), Zoutpansberg; Olifants River District; Orange
Grove near Johannesburg; Pietersburg; Potgietersrust; Pretoria Dis-
trict; Roodeplaat; Rustenburg District; Selati; Shilowane; Wakker-
stroom; Waterberg; Witwatersrand; Woodbush. Zululand:
Mkusi River; Ubombo; Umfolosi Rivers junction. Natal: Lady-
smith; Reitvlei, Umvoti; Weenen.

Range. Transvaal and adjacent areas in Bechuanaland, Swaziland,
Zululand, and Natal. (Power (1930) includes: "Albany Division,
Cape Province." It would seem probable that this was based on
Hewitt's (1909a) record from Teafontein, near Grahamstown; a
record which was later repudiated by Hewitt (1911b). Angola, based
on Monard's suggestion, (vide Remarks supra), is also rejected.

Cordylus cordylus rivae (Boulenger)

1896b. Zonurus rivae Boulenger, 1897, Ann. Mus. Civ. Stor. Nat. Genova (2),

17, p. 8: Giacorsa, Gallaland, Ethiopia.
1897g. Boulenger, p. 278.

Description. Head longer than broad; rostral separated from the
frontonasal, which is longer than broad, nostril pierced in the "centre"

loveridge: African lizards 33

of a nasal which is not much swollen; a loreal; a preocular; prefrontals
separated by frontonasal forming a suture with the frontal ; temporals
large, without spines; gulars small, keeled.

Dorsals strongly keeled, shortly mucronate ; median ventrals smooth ;
the lateral obtusely keeled; preanals subequal (!); tails with whorls of
large, strongly keeled, spinose scales above and below.

This scanty description is adapted from the original which might be
consulted for further details; for scale and pore counts see statistical
table on p. 15.

Color. Above, reddish brown, darker brown along the middle of the
back and on the sides. Below, reddish brown.

Size. Total length of young holotype, 79 (39 + 40) mm. Length of
head 13 mm.; width of head 10 mm.; length of body 26 mm.; of fore
limb 15 mm.; of hind limb 19 mm.

Localities. Ethiopia: Gallaland: Giacorsa.

Range. Ethiopia (known only from the type locality).

CORDYLUS CORDYLUS TROPIDOSTERNUM (Cope)

Plate 3, figs. 2a-b

1869. Zonurus tropidosternum Cope, Proc. Amer. Philos. Soc, 11, p. 169:

"Madagascar" (presumed error for Mozambique).

1881c. Boettger, p. 528.

1885e. Boulenger, p. 254.

1896. Tornier, p. 31.

1897. Tornier, p. 64.

1898. Tornier, p. 285.
1900b. Tornier, p. 590.
1909. Mocquard, p. 4.
1913. Boettger, pp. 360, 361.
1913c. Nieden, p. 71.
1920a. Loveridge, p. 143.
1923d. Loveridge, p. 849.
1923h. Loveridge, p. 947.
1924b. Loveridge, p. 10.
1928. Cott, p. 953.

1936J . Loveridge, p. 296.

1937f. Loveridge, pp. 492, 495.

1939b. FitzSimons, p. 29.

1942e. Loveridge, p. 330.

1889. Zonurus frenatus Pfeffer, Jahrb. Hamburg, Wiss. Anst. 6, p. 6:

Mhonda, Tanganyika Territory.
1893. Pfeffer, p. 73, pi. i, figs. 1-2.

34 bulletin: museum of comparative zoology

1896. Zonurus spec. ? cordylus, ? vittifer Tornier, p. 35 (Potue).

1897. Tornier, p. 64.

1897e. Zonurus cordylus Boulenger (not Linne), p. 800 (Nyasaland).

1898. Tornier, p. 285, fig. 4.
1898. Johnston, p. 361.
1902b. Tornier, p. 590.
1934. Pitman, p. 304.

1934a. Zonurus parkeri Cott, Proc. Zool. Soc. London, p. 151, pi. ii: Ama-

tongas, Mozambique.
1937d. Cordylus tropidosternum Mertens, p. 5.

Names. Eastern Girdle-tail (English); kivmambvsi (Swahili: Tor-
nier); chicologolo (Mwera: Loveridge).

Description. Head slightly depressed; head shields strongly rugose;
rostral in contact with, or separated from, the frontonasal, which is
longer than broad; nostril pierced in the postero-inferior corner of a
large nasal' which is not much swollen; a loreal; a preocular; the larger
subocular not descending to the lip; fourth upper labial lowest, fifth
highest; prefrontals forming a suture or separated; postf rentals longer
than broad or as long as broad ; interparietal enclosed between 2 pairs
of parietals; posterior parietals slightly larger than the anterior; 3-4
rugose, subequal occipitals; temporals rugose, with or without keels,
without spines; sides of neck with keeled, spinose scales; gulars small,
the anterior irregularly enlarged, the median slightly imbricate, keeled;
collar scales larger, sometimes mucronate.

Dorsals slightly elongate, rugose, strongly keeled, shortly mucro-
nate, posteriorly serrate, those on the vertebral line not or but slightly
differentiated; laterals keeled, spinose, scarcely separated by minute
granular interspaces; ventrals quadrangular, smooth or keeled, not or
but slightly imbricate; scales below fore and hind limbs slightly keeled;
tail with whorls of large, strongly striate, keeled, serrate (in adults,
but noticeably in young), spinose scales above and below, the lateral
spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head dark brown, lip to ear usually yellow; back
yellowish, yellowish brown, reddish brown, grayish brown, blackish
brown or rich brown, uniform or clouded, variegated, or spotted with
darker; a broad blackish streak on side of neck from above tympanum
to forearm, sometimes persisting as a distinct (Rhodesia) or ill-defined
(Tanganyika) lateral band. Below, uniform greenish, grayish, yel-
lowish, or reddish (stained by laterite soil) white.

loveridge: African lizards 35

Size. Total length of <? , 180 (90 + 90) mm., and 9 , 170 (95 + 75)
mm., both from Morogoro. The head and body length of the types are :
92 mm. {tropidosternum), 70 mm. (Jrenatus), and 92 mm. (parkeri), the
tails being damaged in the two largest.

Remarks. The type, formerly Museum of the Essex Institute No.
500, now Mus. Comp. Zool. 5742, is apparently a cf but it is somewhat
macerated and the viscera have been removed. Mocquard (1909, p. 4)
was correct in eliminating tropidosternum from the Malagasy herpeto-
fauna. Hewitt (1910a) also discusses the matter, but takes a contrary
view. If actually taken on the island, its presence may be readily ex-
plained by the extensive dhow trade across the Mozambique channel,
this species being subject to transportation in hollow logs which have
been cut for fuel.

In this connection it is interesting to note that Cope's type bears
a closer resemblance to the series from Birchenough Bridge, Southern
Rhodesia, than it does to Tanganyika lizards. I have spent much time
in endeavouring to find other than color characters by which to
separate the Tanganyika lizards (for which the name jrenatus would
be available), but the differences appear too slight to justify such
action. I might say, however, that parkeri of Mozambique, which
allegedly differed from tropidosternum, is definitely a synonym of that
race. Elsewhere I (1936j) have discussed in detail the relative lengths
of fingers and toes and other variable characters employed by Cott for
the separation.

In Cott's paratype of parkeri, though not in the type, and in five
of the eight specimens taken at Birchenough Bridge by FitzSimons,
the prefrontals were separated by the frontonasal being in contact with
the frontal. Tornier's (1896) error of dividing Tanganyika material on
the basis of rostral being in contact with, or separated from, the fronto-
nasal by a suture of the nasals, was later corrected by Nieden (1913c).
More recently Mertens (1937d)t has invited attention to minor differ-
ences exhibited by his six lizards from Matete Woods.

Breeding. On July 28, at Makindu, I took a 9 which held 4 large
eggs. Tornier found 5 embryos in a 9 , apparently from Dar es Salaam
but without date.

Diet. Each of four stomachs examined held termites, a glowworm
fell from the mouth of a fifth, while a captive lizard fed readily on
small grasshoppers.

Parasites. The nematode worm {Oochoristica zonuri) described as
from this species, actually came from a Gerrhosaurus m. major, the
error in labeling it in 1918 was mine.

36 bulletin: museum of comparative zoology

Enemies. Their spines do not offer perfect protection for I recovered
one from the stomach of a Bare-faced Hawk (Gymnogenys t. typicus),
while another was caught and eaten by a young galago (Galago c.
panganiensis) which had been temporarily put in the vivarium.

Habitat. Coastal zone and upland savanna, where they are found
upon hollow trees into whose interiors they retreat and from which
it is difficult to dislodge them. One was actually brought into camp in
a hollow log — in which she had remained while it was being chopped
down — and did not even show herself when the log was roughly flung
down. Three were taken from holes in the base of a wall. Two others,
after torrential downpours, were found in roadside gutters in a half-
drowned condition, having evidently been washed out of some retreat.
One was caught running over papers on the table in my tent. In
East Africa the species may generally be considered scarce except
perhaps at Morogoro where eleven were taken during a year.

FitzSimons (1939b), who took eight specimens at Birchenough
Bridge, found them living in the rotted-out cavities of mopane trees.

Localities. Kenya Colony: Sokoki Forest. Tanganyika Terri-
tory: Dar es Salaam; Kakoma; Kipera; Makindu, Msiha
River; Matete; Mhonda; Morogoro; Msimba; Nchingidi, Rondo
Plateau; Pentambili; Potue, Usambara district; Rufigi; Tendaguru;
Unyika; Usaramo. Mozambique: Amatongas. Nyasaland.
Southern Rhodesia: Birchenough Bridge.

Range. Kenya Colony (near Malindi) south through Tanganyika,
Mozambique, and Nyasaland to Southern Rhodesia (where it meets
with C. c. jonesii and C. c. rhodesianus) .

Cordylus cordylus jonesii (Boulenger)

Plate 3, fig. 3

1891d. Zonurus Jonesii Boulenger, Ann. Mag. Nat. Hist. (6) 7, p. 417:

Murchison Range, northeast Transvaal.

1894e. Boulenger, p. 724.

1895. Jeude, p. 228 (as Johnesii).

1898. Sclater, p. 103.

1907j. • Boulenger, p. 484.

1907b. Roux, p. 420.

1909b. Chubb, p. 35.

1909a. Hewitt (part), pp. 31, 36 (omit Steynsburg and Vitenhage).

1910b. Boulenger, p. 468.

1910a. Hewitt, pp. 56, 60.

LOVERIDGE: AFRICAN LIZARDS 37

1911b. Hewitt, p. 47.

1913. Hewitt & Power, p. 153.

1914a. Hewitt, p. 239.

1921a. Dam, p. 243.

1927c. Power, p. 407.

1930. FitzSimons, p. 29.

1934. Pitman, p. 304.

1935. Lawrence, p. 44.

1892a. Zonurus cordylus Boulenger (not Linn6), p. 174.
1909a. Chubb, p. 593.

1930. Zonurus cordylus jonesii Power, p. 16, pi. i, fig. 3.

1931. Power, p. 48.
1933a. Hewitt, p. 48.
1935b. FitzSimons, p. 349.

Description. Head slightly depressed; all head shields strongly
rugose; rostral rarely in contact with, usually separated from, the
frontonasal, which is longer than broad or as long as broad; nostril
pierced in the lower center of a large nasal which is not much swollen ;
a loreal present or absent; a preocular; the larger subocular not de-
scending to the lip; third or fourth upper labial lowest, fourth or fifth
highest; prefrontals forming a suture or sepaiated; postfrontals longer
than broad or as long* as broad; interparietal enclosed betwTeen 2 pairs
of parietals; posterior parietals slightly larger than the anterior; 4
rugose, subequal occipitals; temporals rugose, without keels or spines;
sides of neck with keeled, spinose scales; gulars small, the anterior
irregularly enlarged, the median slightly imbricate, smooth or feebly
keeled; collar scales larger but undifferentiated.

Dorsals slightly elongate, rugose, strongly keeled, shortly mucronate,
posteriorly serrate, those on the vertebral line not or but rarely dif-
ferentiated; laterals keeled, spinose; ventrals quadrangular, smooth,
not or but slightly imbricate; scales below fore limbs keeled, below
hind limbs smooth; tail with whorls of large, strongly striate, keeled,
serrate, spinose scales above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown, lip to ear yellow ; back yellowish brown,
olive brown, or reddish brown, uniform or spotted with darker; from
tympanum to groin a black, brown, or reddish lateral band. Below,
whitish, uniform.

Size. Total length of c? type (Brit. Mus.) 118+ (67 -f 51+) mm.; a
perfect tf (M.C.Z. 14212), 124 (70 + 54) mm.; length from snout to

38 bulletin: museum of comparative zoology

anus of largest <? (FitzSimons) and 9 (M.C.Z. 20990), 73 mm., both
having lost their tail tips.

Remarks. Hewitt (1909a), furnishing actual measurements, shows
that the heads of adult c. jonesii are much less depressed than those
of c. cordylus, but adds that the distinction is not so obvious in young
individuals. In c. jonesii the head shields are strikingly rugose in
comparison with the much smoother ones of c. cordylus.

Hewitt's (1933a) suggestion, however, that Bulawayo lizards should
probably be referred to c. rhodesianus, is not approved by FitzSimons
(1935b) nor confirmed by our single Bulawayo specimen.

Parasites. Mites (Zonurobia circularis var.) were found by
Lawrence.

Habitat. Found among the cracks and crevices of dolomite kopjes
(Power). Beneath the bark of a dead tree and within the hollow trunk
of a living one (van Dam). Under loose bark of tree growing in
Kalahari sand veld; among dead wood of mopane forest; on a fence in
the middle of Bulawayo (FitzSimons).

Localities. Mozambique: Mozambo, Limpopo River. South-
ern Rhodesia: Bulawayo; Empandeni; Lundi River; Matopo
Hills. Bechuanaland : Devondale; Kuke to Molepole; Lobatsi;
Mmoove 42 miles from Serowe; Palapye; Titumi (Totomi) near
Bushman Mine. Transvaal1: Bridgewatef; Crocodile-Komati
River junction; Geelhoutkop; Griffin Mine near Leydsdorp;
Haenertsburg; Hectorspruit; Hornsnek; Leydsdorp; Limpopo
River; Louis Trichardt; Lydenburg; Magalakwin River; Maiepo c.
25 miles N. of Gravelotte; Makoetsi River; Murchison Range;
Olifants River District; Pongola River; Potgietersrust; Pretoria
District; Rustenburg; Selati; Shiny c. 28 miles E. of Gravelotte;
Silwane; Waterberg District; Wilhanshohe; Zoutpansberg Dis-
trict. Cape Province: Kimberly (introduced).

Range. Southern Mozambique and Southern Rhodesia (Bulawayo
south) and Bechuanaland (eastern Kalahari) through Transvaal. Also
Cape Province (at Kimberly where it has been artificially introduced).

Cordylus cordylus angolensis (Bocage)

?1869b. Zonurus griseus Peters (not Cuvier), p. 659 (Damaraland).
?1885e. Zonurus cordylus Boulenger (part, not Linne), p. 256 (Damaraland).
?1895a. Bocage, p. 24 and footnote p. 25 (Angola).

1 Specimens from Johannesburg and Krugersdorp listed under cordylus by Boulenger (1910b),
are all vittifer according to FitzSimons (letter of March, 1943).

loveridge: African lizards 39

?1897b. Boulenger, p. 277 (Aruwimi, Belgian Congo).

?1911d. Sternfeld, p. 21 (Damaraland).

1895a. Zonurus angolensis Bocage, Herp. Angola et Congo, p. 25: Caconda,
Angola.

1930. Power, p. 16.

1937b. Monard, p. 61.

?1937b. Zonurus vittifer Monard (not Reichenow), p. 61.

Description. Nostril pierced in the postero-inferior corner of a nasal
which is not much swollen; a loreal; a preocular; prefrontals forming
a suture or separated ; gulars noticeably smaller than those of cordylus,
imbricate, smooth or keeled.

For scale and pore counts see statistical table on p. 15.

Color. Above, head and back brown, a double series of small, ir-
regular, whitish spots along the back; tail barred with brown. Below,
whitish with darker infuscations.

Size. Total length of cf type, 152+ (74 + 78+) mm.

Remarks. This form is known to me only from Bocage's description,
based on characters which are variable in typical cordylus; in a foot-
note Bocage adds that he has two other Angolan specimens lacking
precise locality, which he considers are quite typical cordylus. The
scanty description furnished above, together with figures in the sta-
tistical table, are composite of the data furnished by Bocage for all three
(cordylus + angolensis) of his Angolan lizards.

Peters (1869b) merely lists griseus without comment so it is im-
possible to say what he had. Boulenger's (1885e) Damaraland male
would presumably be of the same form as Peters', while the later
(1910b) example from Walfish Bay was collected by Nightingale.
Hewitt (1911b) casts doubt on this record for Nightingale was also
credited with taking Oedura africana in the same locality though the
species is otherwise unknown from South West Africa.

Boulenger's (1897b) lizard from Aruwimi may be angolensis or else
an undescribed form, it certainly would not be typical cordylus.
Power (1930) remarks that angolensis is relegated to the synonymy of
C. c. jonesii in the Zoological Record, this disposition is improbable
but should receive attention when specimens are available. Monard
(1937b) refers the two Bocage ("cordylus") without locality to vittifer.

Localities. ? South West Africa: Damaraland. Angola:
Caconda. ? Belgian Congo : Aruwimi River.

Range. ? Northern South West Africa through Angola to the north-
central Belgian Congo?

40 bulletin: museum of comparative zoology

CORDYLUS CORDYLUS RHODESIANUS (Hewitt)

1933a. Zonurus cordylus rhodesianus Hewitt, Occ. Papers Rhodesian Mus.,

p. 48, pi. ix, fig. 3: Monte Cassino, Macheke, Southern Rhodesia.
1935. Lawrence, p. 43.

1939b. FitzSimons, p. 30.

Description. Head much depressed; head shields strongly rugose;
rostral separated from the frontonasal, which is longer than broad or
as long as broad; nostril pierced in the lower centre of a large nasal
which is not much swollen; a loreal; a preocular; median subocular not
descending to the lip; fourth upper labial lowest, fifth highest; pre-
frontals forming a suture or separated; postfrontals as long as broad
or shorter than broad; interparietal enclosed between 2 pairs of
parietals; posterior parietals much larger than the anterior; 4-6
rugose ; subequal occipitals ; temporals rugose, keeled, without spines ;
sides of neck with keeled, spinose scales ; gulars moderate, the anterior
irregularly enlarged, the median slightly imbricate, smooth, the lateral
elongate; collar scales larger but undifferentiated.

Dorsals slightly elongate, rugose, strongly keeled, shortly mucro-
nate, posteriorly serrate, those on the vertebral line slightly smaller;
laterals keeled, serrate, spinose ; ventrals quadrangular, smooth not or
but slightly imbricate; scales below fore limbs keeled, below hind
limbs smooth or slightly keeled; tail with whorls of large, strongly
striate, keeled, serrate spinose scales above and below, the lateral
spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown, lip to ear yellow; back yellowish or olive
brown variegated, spotted, or longitudinally streaked, with darker,
these markings tending to form irregular cross-bars in young; tail
uniform olive brown. Below, whitish, uniform.

Size. Length of c?1 type (Albany Mus.) from snout to anus, 82 mm. ;
of a 9 (T.M. 18526) from snout to anus, 80 mm., both having injured
tails; total length of a young one (M.C.Z. 44422), 69 (38 + 31) mm.
Both of latter being from Vumba Mountain.

Remarks. As indicated by its describer, this form occupies a some-
what intermediate position between vittifer and c. jonesii, agreeing with
vittifer in its much depressed head, and in having the anterior row of
nuchals slightly more enlarged than is the case with any other race of
the cordylus group. It differs from c. jonesii in that its anterior pair
of sublabials form a long (not short) median suture, and, in lacking a

loveridge: African lizards 41

dark lateral band. It differs from c. cordylvs (but scarcely or not from
jonesii) in that its anterior gulars form a slightly enlarged group, yet
scarcely so enlarged as to be called sublinguals, while the majority of
the gulars are longitudinally elongate (not transversely broadened).

Originally based on 8 lizards, one of which is now M.C.Z. 33448,
this form was not again reported until 10, four of which are M.C.Z.
44419-22, were captured by FitzSimons on Vumba Mountain.

Habitat. Occurs at 5000 feet on Vumba Mountain, among rocks.

Localities. Southern Rhodesia: Monte Cassino; Macheke;
Triashill Mission, Rusape; Vumba Mountain.

Range. Southern Rhodesia (C. c. jonesii also occurs but in the south-
west at Bulawayo).

CORDYLUS CORDYLUS LAWRENCI (FitzSimons)

Plate 4, fig. 1

1937d. Zonurus peersi Hewitt (not Hewitt, 1932), p. 208.
1939a. Zonurus lawrenci FitzSimons, Ann. Transvaal Mus., 20, p. 7, figs.
2-4: Lekkersing, Richtersveld, Little Namaqualand.

Description. Head much depressed, longer than broad; head shields
strongly rugose; rostral separated from the frontonasal, which is
shorter than broad; nostril pierced near the centre of a large nasal
which is not much swollen ; a loreal ; a preocular ; median subocular not
descending to the lip; fourth upper labial lowest, fifth highest; pre-
frontals forming a suture; postfrontals as long as broad or shorter
than broad ; interparietal enclosed between 2 pairs of parietals ; poste-
rior parietals subequal to the anterior; 4 strongly keeled, unequal
occipitals; temporals strongly keeled, the hindmost serrated posteri-
orly and almost covering ear; sides of neck with keeled, spinose scales;
gulars large, the anterior irregularly enlarged the median slightly
imbricate, smooth, or the laterals feebly keeled.

Dorsals squarish, not rugose, strongly keeled, shortly mucronate,
posteriorly finely serrate, the two vertebral rows scarcely enlarged ;
laterals in vicinity of midbody as large as the dorsals, keeled, strongly
serrate, spinose; ventrals quadrangular, smooth, not or but slightly
imbricate; scales below fore limbs keeled, below thigh smooth or
slightly keeled, below tibia keeled and mucronate ; tail with whorls of
large, strongly keeled, serrate, spinose scales above and below.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

42 BULLETIN : .MUSEUM OF COMPARATIVE ZOOLOGY

Color. Above, head black, posteriorly flecked with yellow; back
dark brown, anteriorly flecked with yellow; flanks, limbs, and tail
paler than dorsum. Below, gular region grayish, reticulately spotted
with blackish; belly grayish; tail straw yellow.

Size. Total length of 9 type (S.A.M. 18553), 151 (67 + 84) mm.

Localities. Cape Province: Little Namaqualand: Lekkersing.

Range. Little Namaqualand, Cape Province (known only from the
type).

CORDYLUS CORDYLUS TASMANI (Power)

Plate 4, fig. 2

1909a. Zo,nurus jonesii Hewitt (part, not Boulenger), p. 36.

1930. Zonurus cordylus tasmani Power, Ann. Transvaal Mus., 14, pp. 12,

16, pi. i, fig. 4: Dunbrody, Uitenhage Division, Cape Province.
1930. FitzSimons, p. 29.

1935. Lawrence, p. 43.

1937e. Hewitt, p. 30.

Description. Head much depressed; head shields strongly rugose;
rostral separated from the frontonasal, which is longer than broad or
shorter than broad; nostril pierced in the postero-inferior corner of a
large nasal which is not much swollen; a loreal; a preocular; median
subocular not descending to the lip; fourth upper labial lowest, fifth
highest; prefrontals forming a suture or separated; postfrontals as
long as broad or shorter than broad; interparietal enclosed between 2
pairs of parietals; posterior parietals slightly larger than the anterior;
5-6 rugose, subequal or irregular occipitals; temporals rugose, keeled,
without spines; sides of neck with keeled, spinose scales; gulars mod-
erate and large, the anterior irregularly enlarged, the median slightly
imbricate, smooth, the lateral strongly keeled; collar scales larger
but undifferentiated.

Dorsals squarish, rugose, strongly keeled, shortly mucronate, poste-
riorly serrate, the two vertebral rows enlarged; laterals in vicinity of
midbody as large as the dorsals, keeled, serrate, spinose; ventrals
quadrangular, smooth, not or but slightly imbricate; scales below fore
limbs keeled, below hind limbs smooth or slightly keeled; tails with
whorls of large, strongly striate, keeled, serrate, spinose scales above
and below, the lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown, lip to ear yellow; back yellowish brown

loveridge: African lizards 43

or olive brown, spotted or longitudinally streaked with black or with
irregular blackish cross-bars. Below, whitish or olive yellow, uniform.

Size. Total length of c? cotype (M.C.Z. 31572), 136 (70 + 66) mm. ;
of 9 cotype (M.C.Z. 27122), 136+ (78 + 58+) mm., tail-tip missing.

Remarks. Its describer compares this race with c. jonesii with which
it was said to agree in everything except larger laterals ; it differs also
in its more depressed head and in having 3 suboculars. Actually, as
pointed out by Hewitt (1937d), it is more closely related to c. cordylus
as one might expect from its distribution as a pocket within the range
of the typical form.

This intermediate position is emphasized by the specimens from
Steynsburg and Uitenhage in the Albany Museum, first referred to
c. jonesii by Hewitt (1909a), and later (1911b) thought to be young
c. cordylus, but which, according to Power (1930, p. 12) are more
probably c. tasmani, and are therefore listed as such below.

Localities. Cape Province: Uitenhage Division: Dunbrody,
Sunday's River; ? Steynsburg; Uitenhage.

Range. Uitenhage Division of Cape Province (surrounded by
typical C. c. cordylus.)

Cordylus cordylus minor FitzSimons

1943. Cordylus cordylus minor FitzSimons, Lizards of South Africa, p. 458:
Just north of Matjesfontein on road to Sutherland, Cape Province,
Union of South Africa.

Description. Head strongly depressed; head shields finely rugose
throughout; rostral separated from the frontonasal, which is broader
than long; nostril pierced in the postero-inferior corner of nasal, which
is not swollen; a loreal; a preocular; 3 suboculars, the median not
descending to the lip; 6 upper labials, sixth with a strongly compressed
keel; 4 supraoculars, fourth smallest; 3 supraciliaries ; no supranasals;
prefrontals in contact; frontal longer than broad, broadest anteriorly;
interparietal enclosed between 2 pairs of parietals, which are subequal
or posterior pair slightly smaller than anterior; 6 (sometimes reduced
and irregular) occipitals similar in size and shape to adjacent nuchals ;
temporals rugose and sharply keeled; "lower posterior temporal spine
strongly compressed, with a sharp projecting edge"; sides of neck with
keeled, spinose scales; 5 lower labials, posterior keeled, bordered by a
row of 5 large shields ; gulars moderate, the anterior irregularly en-
larged, the median imbricate, smooth.

44 bulletin: museum of comparative zoology

Dorsals squarish, rugose, strongly keeled, not or but shortly mu-
cronate mesially, more strongly so dorsolaterally, posteriorly feebly
serrate, and forming 24-26 longitudinal rows (inclusive of laterals on
flanks) and 27-28 transverse rows ; laterals subequal to dorsals ; ventrals
quadrangular, smooth, forming 16 regular longitudinal and 24-26
transverse rows; 2 enlarged preanals; limbs above with large, keeled,
spinose, imbricate scales; 4-6 femoral pores with 1-2 rows of swollen,
glandular scales anteriorly; tail with whorls of large, striate, strongly
keeled, serrate, spinose scales above and below, the dorsolateral spines
longest.

Color. Above, head dark brown to blackish faintly speckled with
dull yellow; back and tail dull olive brown sparsely spotted with black.
Below, grayish white tinged with olive yellow posteriorly and on tail.

Size. Total length of rf1 cotype (Tvl. Mus. 19563) 139 (65 +
74) mm.

Remarks. Known to me only from original description (based on six
cotypes) which should be consulted for further details. Said to differ
from c. cordylus in being "smaller in size, head and body more strongly
depressed, head shields finely rugose throughout, body scales smaller
and more strongly keeled and spinose, fewer femoral pores and sub-
digital lamellae." That is 10-14 (usually 10-13) lamellae under fourth
toe as against 13-17 in c. cordylus. In scale counts this new form
apparently approaches the pustidatus-7iamaquensis-campbelli group.

Localities. Known only from the type.

Range. Vicinity of Matjesfontein, Cape Province.

Cordylus cordylus cordylus (Linne)
Plate 4, fig. 3

1758. Lacerta cordylus Linne, Syst. Nat. ed. 10, 1, p. 202: Africa.

1760. Linn6 ed. 12, 1, p. 361: "Africa; Asia" (error).

1900. Andersson, p. 7.

1768. Cordylus versus Laurenti, Syn. Rept., p. 52: Africa.

1802d. Stellio cordylus Daudin, p. 8.

1829. Cordylus dorsalis Cuvier, Regne Animal, ed. 2, 2, p. 33: Africa.

1829. Cordylus griseus Cuvier, Regne Animal, ed. 2, 2, p. 33: Africa.

1838a. Smith, A., p. 31.

1843. Smith, A., pi. xxviii, figs. 2-3; pi. xxx, figs. 8-8b.

1884a. Rochebrune, p. 107 (omit Senegambia as error).

1838. Zonurus vertebralis Gray, Ann. Nat. Hist., 1, p. 388: Cape of Good

Hope.

1839. Zonurus griseus Dumeril & Bibron, p. 350.
1845. Zonurus cataphractus Gray (not Boie), p. 47.

loveridge: African lizards 45

1845. Zonurus cordylus Gray, p. 47.

1867a. Steindachner, p. 42.

1885e. Boulenger (part), p. 256 (omit Damaraland?).

1893a. Boettger, p. 66.

1897. Bateman, p. 107.

1898. Jeude, p. 21.

1898. Werner (1896-7), p. 140.

1898. Sclater, p. 103.

1907b. Roux (part), p. 417 (omit niger localities).

1909a. Hewitt, p. 37.

1910b. Boulenger (part), p. 468 (omit Transvaal, Natal and S. W. Africa

records).

1910a. Hewitt, pp. 60, 71.

1910a. ?Werner, p. 324 (see Remarks under C. namaquensis) .

1911. Gilchrist, p. 230, fig. 15b.

1911b. Hewitt, p. 47.

1913. Hewitt & Power, p. 153.

1913a. Werner, p. 107, fig.

1914a. Hewitt, p. 239.

1916. Andersson, p. 39.

1920. Hewitt, p. 93.

1921a. Dam, p. 242.

1925. Essex, p. 338.

1925b. Flower, p. 945

1926b. Rose, p. 492.

1927b. Hewitt, p. 452.

1928. Cott, pp. 927, 928.
1928a. Essex (1927), p. 931.

1929. Rose, p. 97, fig. 62.
1931. Mann, pp. 390, 397, 399.
1934a. Cott, p. 153.

1935. Lawrence, p. 43.

1937e. Hewitt, p. 29, pis. viii, fig. 4; x, fig. 4.
1937. Lawrence, p. 107, fig. 1.

1926b. Zonurus cordylus flavus Rose, Ann. S. African Mus., 20, p. 492:
Robben Island, Cape Province, Union of South Africa.

1930. Zonurus cordylus cordylus FitzSimons, p. 29.
1930. Power (part), p. 16 (omit Umvoti).
1936h. Loveridge, p. 58.

Further citations of "cordylus" and "griseus" will be found under
vittifer, c. tropidosternum, c. jonesii, c. angolensis, c. niger, ukingensis,
and cataphr actus.

Native names. Cape Girdle-tail (English); klip salamander (local
misnomer) ; uroqotyeni.

46 bulletin: museum of comparative zoology

Description. Head much depressed; head shields slightly (anteri-
orly) or strongly (posteriorly) rugose; rostral 23^ (M.C.Z. 21570) to 4
(M.C.Z. 1940) times as broad as high, in contact with, or separated
from, the frontonasal, which is longer than, or shorter than, broad,
and sometimes divided (M.C.Z. 21567, 21570); nostril pierced in the
postero-inferior corner of a moderate nasal which is not, or scarcely,
swollen; a loreal; a preocular; 3 suboculars, the median not or but
rarely descending to the lip between the fourth and fifth upper labials;
5-6 upper labials, fourth lowest, fifth highest; 3-4 supraoculars, the
anterior longest, second broadest; 3 supraciliaries ; no supranasals; pre-
frontals forming a broad or narrow suture, or separated; frontal
pentagonal or hexagonal ; postf rentals as long as broad or shorter than
broad; interparietal enclosed between 2 pairs of parietals; posterior
parietals slightly larger than the anterior; 4-5 rugose, subequal
occipitals; temporals smooth or slightly rugose, keeled, without
spines; side of neck with keeled, spinose scales; mental large; 5-6
lower labials, posterior largest and keeled, bordered by a row of 5
large shields; gulars moderate and large, the anterior irregularly
enlarged, the median slightly imbricate, smooth, the lateral keeled,
in 16-20 rows between angles of mandibles; collar scales larger but
undifferentiated.

Dorsals squarish, slightly rugose, strongly keeled, shortly mucronate,
posteriorly serrate, and forming 16-20 longitudinal rows (inclusive of
laterals on flanks) and 25-29 transverse rows from occiput to base of
tail ; laterals like dorsals, keeled, serrate, spinose ; a lateral fold ; ventrals
quadrangular, smooth not or but slightly imbricate, forming 12,
rarely 10 or 14, regular longitudinal and 23-30 transverse rows be-
tween collar and anals; 2, rarely 4, enlarged preanals; limbs above
with large, keeled, spinose, imbricate scales, below those on fore limbs
keeled, on hind limbs smooth or slightly keeled; 6-91 femoral pores;
tail with whorls of large, striate, strongly keeled, serrate, spinose
scales above and below, the lateral spines longest.

Color. Above, head brown2; back yellowish olive, yellowish brown,
olive brown, reddish brown, or blackish brown, uniform or variegated
with darker, sometimes an irregular cream-colored vertebral line and
a reddish lateral band present. The young are brown, or reddish
brown, flecked with white. Below, greenish or yellowish white,
uniform.

1 10-11 in a "South Africa" specimen (which agrees more nearly with namaquensis) , fide
Werner (1910a).

'Sometimes black, fide Rose (1926a).

loveridge: African lizards 47

Size. Total length of largest cotype (Stockholm Mus.), 170 (85 +
85) mm.; length of d" (M.C.Z. 21571) 166+ (84 + 82+) mm., of 9
(M.C.Z. 21567), 158+ (84 + 74+) mm., their tails being injured, both
surpassed by an unsexed specimen (Brit. Mus.) of 174 (84 -f- 90) mm.

Remarks. Lacerta cordyhis was based on lizards figured in Seba, 1,
pi. Ixxxiv, figs. 3-4 from Africa and Cape of Good Hope respectively,
and Seba, 2, pi. lxii, fig. 5 from Africa, called Lacerta nigra by Seba so
probably the basis of what is now known as C. c. niger Cuvier. This
can be settled as the probability of the five specimens now in the Royal
Swedish Museum at Stockholm being the five cotypes from Drotting-
holm Museum is discussed by Andersson (1900).

Van Dam (1921a, p. 242) presents a table of characters in which
c. cordylus differs from vittifer, though not all are valid. Later Hewitt
pointed out that a median groove on the anterior portion of the frontal,
almost always present in c. cordyhis, is lacking in c. niger.

Anatomy. In a discussion of the branchial arch in lizards, Hewitt
(1920) states that the epibranchials are present and that the hyoid has
no relation with the ear. Mann (1931) remarks upon the structure of
the eyes, which were described as being "reddish brown" by Sir A.
Smith (1843).

Longevity. Three years, six months, ten days, in the London
Zoological Gardens (Flower).

Breeding. Rose states that a single young one, two inches in length,
is produced at a birth, but there are two ova, each measuring about
18 x 10 mm., in a 9 (M.C.Z. 21410) from Grahamstown (N.D.).

Diet. Beetles, cockchafers, cockroaches, crickets and locusts
(Hewitt). Though mainly insectivorous, this girdle-tail may at times
be carnivorous, one having been seen to eat a young skink (Mabnya
sp.), others to take scraps of preserved meat from a discarded tin,
crumbs of currant cake, scraps of orange pulp, and even devour lichen
from the rocks (Essex and Rose).

Parasites. Mites (Scaphothrix convexa and Zonurobia cordylensis)
have been described from this race by Lawrence.

Enemies. One in stomach of a secretary bird (Andersson); "often
a prey of kestrels" (Hewitt).

Habitat. The depressed form of this species facilitates its retreat
into the crevices formed by cracks in the round boulders of dolerite
abundant on the sides of the flat -topped mountains in the vicinity of
Mortimer (Cott). In open country found beneath stones and rocks
near which it basks until disturbed ; on being alarmed it scuttles head
first into its retreat so that the tail forms a protection for the body

48 bulletin: museum of comparative zoology

(Hewitt). At times it lives, like Gerrhosanrus, in holes in the ground
(Essex). Also occurs in dry logs and hollow stumps (Pannel in Hewitt)
and will dash over or into shallow pools when frightened (Cronwright
in Hewitt).

Localities.1 Cape Province: Albany; Alexandria; Amatola
Mountains; Bain's Kloof; Beaufort West; Bedford District; Bush-
man's River mouth; Cala; Cape St. Francis; Cape Town; East
London; Fort Beaufort; French Kraal, Gaus Bay; Frenchhoek
(Franschhoek) ; Gaika's Kop; George; Grahamstown; Hermanus;
Hogsback; Hottentot's Holland Mountain; Houwhoek; Kalk Bay;
Katberg summit; Kingwilliamstown;Kleinpoort near Committees;
Kokstad; Knysna; Middleburg; Mitchel's Pass; Molteno; Mon-
tagu; Mortimer; Mossel Bay; Mount Ayliff; Mvenyane near
Cedarville; Paarl; Philipstown; Port Alfred; Port Elizabeth;
Queenstown; Robben Island; Sir Lowry'sPass; Somerset Strand;
St. Croix Island; Stellenbosch; Steynsburg; Stutterheim; Swellen-
dam; Tsomo, Transkei; Tulbagh District; Willowmore; Wynberg;
Zuurberg, Alexandria Division.

Range. Chiefly coastal strip of Cape Province from Robben Island
to East Pondoland. An isolated subspecies (c. tasmani) occurs within
this range at Dunbrody just north of Port Elizabeth.

cordylus cordylus niger Cuvier
Plate 5, fig. 1

1735. Lacerta nigra Seba, Rerum naturalium Thesauri, 2, p. 62, pi. lxii,

fig. 5: Africa.
1829. Cordylus niger Cuvier, Regne Animal, ed. 2, 2, p. 33: Africa.
1907b. Zonurus cordylus Roux (part, not Linne), p. 417.
1926b. Zonurus cordylus niger Rose, p. 492.

1929. Rose, p. 97, fig. 61.

1930. Zonurus cordylus atrus Power, Ann. Transvaal Mus., 14, pp. 11, 16,

pi. i, fig. 2: Cape Peninsula, Cape Province.

Description. Head much depressed; head shields smooth or only
slightly rugose; rostral separated from the frontonasal, which is as
long as broad or shorter than broad; nostril pierced in the postero-

1 The Uitenhage record of Hewitt (1911b) is removed to c. tasmani by Power (1930), but
apparently Hewitt (1937b) does not concur with Steynsburg being included in the transfer.
Boulenger's (1910b) records of Johannesburg and Krugersdorp, Transvaal, and Umvoti, Natnl
are all vittifer according to FitzSimons (letter of March, 1943).

loveridge: African lizards 49

inferior corner of a moderate nasal which is not swollen; a loreal; a
preocular; median subocular descending to the lip between the fourth
and fifth upper labials; prefrontals forming a suture or separated;
interparietal enclosed between 2 pairs of parietals ; posterior parietals
slightly larger than the anterior; 4, rarely 3, 5, or 6 smooth or slightly
rugose, subequal occipitals ; temporals smooth or slightly rugose, with-
out spines ; sides of neck with keeled, spinose scales ; gulars moderate
and large, the anterior irregularly enlarged, the median slightly im-
bricate, smooth; collar scales larger but undifferentiated.

Dorsals squarish, slightly rugose, strongly keeled, shortly mucro-
nate ; posteriorly serrate ; laterals like dorsals, keeled, serrate, spinose ;
ventrals quadrangular, smooth, not or but slightly imbricate; scales
below fore limbs keeled, below hind limbs smooth or slightly keeled;
tail with whorls of large, striate, strongly keeled, serrate, spinose scales
above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, dull black, uniform. Below, black or gray, in young
greenish white.

Size. Total length of rf> (M.C.Z. 21555),' 150 (67 + 83) mm., of 9
(M.C.Z. 21573), 175 (85 + 90) mm.

Remarks. Apart from slightly smaller size, and color1, niger differs
from c. cordylus not only in the points mentioned in the key but in
certain average characters such as averaging 10 (rarely 12, never 14)
longitudinal rows of ventrals; head scales more or less smooth (not
strongly rugose posteriorly) ; in having the scales of the occipital row
smooth, rectangular, and similar to other head shields; in the vertebral
series of scales being strongly keeled (not smooth).

Breeding. A single uniformly black young one produced at a birth
(Rose).

Localities. Cape Province: Cape Peninsula: Cape Point;
Cape Town; Muizenberg; Table Mountain (Range Cottage).

Range. Cape Peninsula (southern end from Cape Point to Muizen-
berg, meeting with the typical form in about equal numbers at Lion's
Head).

1 For causes of melanism in lizards see Parker, H. W., 1935, Proc. Zool. Soc. London, pp.
137.142.

50 bulletin: museum of comparative zoology

CORDYLUS PEERSI (Hewitt)

Plate 5, fig. 2

1930. Zonurus peersi FitzSimons, p. 30 {nomen nudum).

1932. Hewitt, Ann. Natal Mus., 7, p. 116, photo: Garies, Little Narnaqua-

land, Cape Province, Union of South Africa.
1935. Lawrence, p. 43.

1937. Lawrence, p. 110.

1938. FitzSimons, p. 192.

Description. Head much depressed; head shields very strongly
rugose; rostral separated from the frontonasal (by a granule in one
co type), which is as long as broad or shorter than broad ; nostril pierced
in the postero-inferior corner of a large nasal which is swollen ; a loreal ;
a preocular ; median subocular descending to the lip between the fourth
and fifth upper labials; fourth upper labial lowest, fifth highest; pre-
frontals forming a suture, rarely separated; postfrontals as long as
broad; interparietal enclosed between 2 pairs of parietals; posterior
parietals subequal to the anterior; 4 rugose (but not. keeled), subequal
occipitals; temporals rugose, strongly keeled, without spines; sides of
neck with keeled, spinose scales; gulars moderate and large, the an-
terior not irregularly enlarged, the median slightly imbricate, smooth
or obtusely keeled; collar scales larger but undifferentiated.

Dorsals squarish, slightly rugose, strongly keeled, towards sides
shortly mucronate, posteriorly serrate; laterals like dorsals, keeled,
serrate, spinose; ventrals quadrangular, smooth, not or but slightly
imbricate; scales below fore limbs keeled, below hind limbs smooth
or slightly keeled; tail with whorls of large, striate, strongly keeled,
serrate, spinose scales above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, black, uniform. Below, black, uniform except for
the femoral pores and lemon yellow callose patches on the femur of
males.

Size. Total length of a d*1 (T.M. 18090), 169 (78 + 91) mm.; of a
9 (T.M. 18088), 178 (81 + 97) mm.

Remarks. Its describer distinguishes peersi from c. niger, the only
other entirely black member of the genus, by its swollen nasal which
is never (1937d) so elongate as in cordylus and its races; by the gulars
which are uniform anteriorly (except for a few lateral ones) and pass
gradually into the larger posterior; by the stronger keeling of tem-

loveridge: African lizards 51

porals and dorsals; and by the band of minute scaling in the lateral
fold.

Parasites. Mites (Scaphothrix conversa and Zonurobia polyzonensis)
have been described from this species by Lawrence.

Temperament. Appears to be less timid than other members of the
genus (FitzSimons).

Habitat. About a mile outside Garies, Peers found these lizards
beneath weathered granite, saucer-like flakes which, on being levered
up, were apt to slide down and consequently rendered the capture of
the lizards more difficult. At Nieuwerust, about 90 miles south of
Garies, and separated from it by a vast stoneless plain, others were
encountered on kopjes where they were living under fragments of the
great granite boulders. FitzSimons found peer si occupying the upper
slopes of kopjes while p. polyzonus occurred on the lower.

Localities. Cape Province: Little Namaqualand: Garies;
Kamieskroon; Nieuwerust.

Range. Little Namaqualand, Cape Province.

Cordylus ukingensis (Loveridge)

1900b. Zonurus cordylus Tornier (not Linne), p. 590.

1932a. Zonurus ukingensis Loveridge, Bull. Mus. Comp. Zool., 72, p. 378,

pi. iii, fig. 2: Tandala, Ukinga, Tanganyika Territory.
1933h. Loveridge, p. 301, pi. iii, fig. 2.
1937f. Loveridge, p. 495.

Description. Head slightly depressed; rostral separated from the
frontonasal, which is as long as broad; nostril pierced in the postero-
inferior corner of a very large nasal1 which is not much swollen; no
loreal; a preocular; median subocular not descending to the lip; fifth
upper labial smallest, fifth not higher than others; prefrontals forming
a suture; postfrontals longer than broad; interparietal on a line with
the anterior parietals, in contact with the postfrontals; posterior
parietals slightly larger than the anterior; 6 keeled occipitals; tem-
porals rugose, keeled, without spines; sides of neck with keeled,
spinose scales; gulars small, the anterior not enlarged, the median
slightly imbricate, all strongly keeled and mucronate; collar scales
large, lanceolate, and mucronate.

Dorsals squarish, rugose, strongly keeled, shortly mucronate, poste-
riorly serrate, the two vertebral rows enlarged; laterals like dorsals

1 Tornier remarks that in one of his three specimens the nasal is divided on the right side
to form a prae- and post-nasal as he terms them, on the left it is entire.

52 bulletin: museum of comparative zoology

but smaller, keeled, spinose; ventrals quadrangular, keeled, not or but
slightly imbricate, the lateral shortly mucronate; scales below fore
and hind limbs keeled; tail with whorls of large, striate, strongly
keeled, serrate, spinose scales above and below, dorsal and lateral
spines subequal in length.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head dark brown; back sepia brown variegated with
ochraceous brown; flanks flecked with lighter. Below, white, faintly
mottled with gray.

Size. Total length of cT1 type (M.C.Z. 30761), 87 (54 + 33) mm.

Remarks. Diagnostic characters distinguishing ukingensis from c.
cordylus and c. jonesii have been given in detail (1932a), and for
c. tropidosternum (1933h) also. This dwarf form is known only from
the type, and three specimens in the Berlin Museum.

Localities. Tanganyika Territory: Iringa; Kuthu steppe;
Mbowu River, Unyika; Tandala, Ukinga.

Range. Southern highlands of Tanganyika Territory (i.e. mountains
at north end of Lake Nyasa).

Cordylus macropholis (Boulenger)

1910b. Zonurus macropholis Boulenger, Ann. S. African Mus., 5, p. 494:

Little Namaqualand, Cape Province, Union of South Africa.
1930. Power, p. 16.

1933a. Power, p. 215.
1935. Lawrence, p. 44.

1936h. Loveridge, p. 58.
1937. Lawrence, p. 109.

Description. Head slightly depressed; head shields strongly rugose;
rostral separated from the frontonasal, which is shorter than broad;
nostril pierced in the postero-inferior corner of a large nasal which is
not much swollen; a loreal; a preocular; median subocular not de-
scending to the lip; fifth upper labial longest, not or but slightly
higher than others; prefrontals forming a suture; postfrontals longer
than broad or as long as broad ; interparietal on a line with the anterior
parietals, in contact with, or separated from, the postfrontals ; posterior
parietals smaller than the anterior; 4-5 rugose occipitals; temporals
rugose, keeled, without spines but those of hind row pointed, projecting
over ear; sides of neck with keeled, spinose scales; gulars large, a few
anterior moderate followed by smaller, slightly imbricate, all strongly
keeled and mucronate; collar scales large, keeled, mucronate.

loveridge: African lizards 53

Dorsals squarish, rugose, strongly keeled, shortly mucronate, poste-
riorly serrate, the two vertebral rows enlarged; laterals like dorsals
but slightly smaller, keeled, serrate, spinose; ventrals quadrangular,
smooth, strongly imbricate, the lateral strongly keeled and shortly
mucronate; scales below fore limbs keeled, below hind limbs smooth
or slightly keeled; tail with whorls of large, rugose, strongly keeled,
serrate, spinose scales above and below, dorsal and lateral spines
subequal in length.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head and back olive brown irregularly blotched with
blackish brown. Below, pale olive or brownish.

Size. Total length of type, 130 (68 + 62) mm. ; of a cf (Field Mus.),
119 (67 + 52) mm.

Remarks. For twenty-three years known only from the type, then
Power obtained a series. The scale termed "postnasal" by Power
(1933a) is that usually regarded as a loreal, similarly his "loreal" is
the preocular.

Habitat. Never having been taken in rock crevices, this dwarf
species is suspected of living in burrows (Lawrence).

Localities. Cape Province: Little Namaqualand: Kleinzee.

Range. Little Namaqualand, Cape Province.

CORDYLUS CATAPHRACTUS BoiE

Plate 5, fig. 3
Plate 6, figs. 1-2

1828. Cordylus cataphractus Boie, Nova Acta Acad. Leop. -Carol., 14, pt. 1,

p. 140: South Africa.
1843. Smith, A., pi. xxix, pi. xxx, figs. 9-9a-9b.

1831b. Zonurus Cataphractus Gray, p. 63.
1839. Dumeril & Bibron, p. 355.

1885e. Boulenger, p. 255.
1887b. Boettger, p. 143.
1898. Sclater, p. 103.

1907b. Roux, p. 417.
1909a. Hewitt, pp. 33, 36.
1910b. Boulenger, p. 468.
191 Id. Sternfeld, p. 21.
1914a. Hewitt, p. 240.
1929. Rose, pp. 100, 218, figs. 63-64.

54 bulletin: museum of comparative zoology

1930. FitzSimons, p. 29.

1930. Peers, p. 402, photos. 1-4, pis. i-ii.

1930. Power, p. 15.

1930. Schmidt, p. 154, photo.

1931. Mann, pp. 390, 397.
1931. Popp. p. 191, photos.
1931. Senfft, p. 73, photos.
1933b. Noble & Bradley, p. 75.
1935a. FitzSimons, p. 533.
1935. Lawrence, p. 44.

1937. Lawrence, p. 110, fig. 3.

1938. ' FitzSimons, p. 191.

1834. Zonurus cordylus Schlegel (not Linne), p. 211, pi. vii, figs. 3-3c.
1938a. Cordylus nebidosus Smith, Mag. Nat. Hist. (2), 2, p. 31: South Africa.

Native names. Armadillo lizard (English: Rose); blinkoog (Dutch:
Peers).

Description. Head much depressed; head shields strongly rugose;
rostral separated from the frontonasal, which is longer than broad;
nostril pierced in the centre of a very large nasal which is much
swollen; a small loreal present or absent; a preocular; median subocular
not descending to the lip; third or fourth upper labial smallest, fifth
frequently higher than others ; prefrontals well separated ; postfrontals
longer than broad or as long as broad; interparietal on a line with the
anterior parietals, in contact with the postfrontals ; posterior parietals
much larger than the anterior; 5-6 rugose occiptals, the outermost
pointed and directed obliquely backwards ; temporal region very promi-
nent, temporals large, rugose, keeled, without spines, but those of
hind row pointed, projecting over ear ; sides of neck with keeled, spinose
scales; gulars small, the anterior irregularly enlarged, the median not
or but slightly imbricate, smooth, or the lateral feebly keeled; collar
scales large, mostly smooth, imbricate.

Dorsals elongate, rugose, strongly keeled, shortly mucronate, posteri-
orly serrate, those on the vertebral line regular or irregular; laterals
like dorsals but more spiny, keeled, serrate, spinose; ventrals elongate
or quadrangular, all smooth, not or but slightly imbricate; scales
below fore limbs keeled, below hind limbs smooth or some keeled; tail
with whorls of large, rugose, strongly keeled, serrate, spinose scales
above and below, the lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head and back usually yellowish brown, occasionally
chocolate brown, uniform, or a piebald arrangement of these colors.

loveridge: African lizards 55

Young lizards are predominantly brown. Below, gular region light
yellow vermiculated, streaked, or spotted with black; belly yellowish,
clouded with greenish black or dark brown spots or stripes.

Transference of three armadillo lizards from their arid habitat to
the moister and colder climate of the Cape Peninsula, resulted in
striking color changes, but whether humidity, temperature, or food
was the principal factor is not clear. A young lizard, two-and-a-half
inches long, became a very light creamy yellow and grew larger than
the adults. The latter, originally yellow and brown, changed to a
lighter hue of greenish yellow, most apparent on their heads; mean-
while the brown of their backs became blotchy, not fading into yellow
(Peers).
' Size. Total length of d1 (T.M. 15970), 211 (103 + 108) mm.

Remarks. According to Peers, about 20% of these lizards have the
dorsal rows interrupted on the vertebral line instead of forming regular
rows ; such a variation, as well as normal and intermediate conditions,
being found in members of a family group, there appears to be no
geographical significance attached to it (Peers). Moreover it is to be
found in several other species.

The best account of the bionomics of this lizard is that of Peers, to
whose article I am indebted for much of the information given below.

Anatomy. The iris and pupil have been described by Mann.

Sexual dimorphism and dichromatism. The head of the male is
broader, his habit more robust, his gular markings brighter than those
of his mate, around whom he prances, his head held high, the better to
display his brightly marked throat (Peers).

Breeding. Takes place about October when one or two young are
produced, these rupturing the enveloping membranes immediately
after parturition. (Peers), but in Germany a captive, 200 mm. female
gave birth to a 100 mm. young one on February 2nd or 3rd, according
to Popp, who furnishes a photograph of a pair in coitu.

Diet. When termites appear at the onset of the rains, armadillo
lizards gorge upon them; the fat derived from this prey tides them
over the succeeding weeks of wet and cold. Beetles and grasshoppers,
whose movements are quickly observed by these bright-eyed lizards,
are captured after a short rush which rarely fails (Peers) .

In captivity young lizards will take maggots, other soft-skinned
larvae, and flies. As they grow older, smooth caterpillars, beetles,
crickets, but above all grasshoppers; on the latter, which seem most
appreciated, they will feed to repletion. It is true that at first they
will take mealworms, holding them in their jaws until dead, but these

56 bulletin: museum of comparative zoology

appear to disagree with them for they are frequently regurgitated,
while after a time the lizard will refuse them. Earthworms were also
rejected. When some other lizards (apparently Lacerta. A.L.) were
temporarily placed in their case, they were seized by head or middle,
crunched and eaten by the carnivorous armadillo lizards. Senfft also
fed his specimens with raw egg, brains, and a few drops of vigatol but
failed to keep them alive (Popp, Senfft et al).

Enemies. Mongoose, musihonde and veld rats are a constant
menace to the armadillo lizards which survive attacks by keeping close
to their rocky crevices.

Defence. On gaining its rocky retreat an armadillo lizard takes full
advantage of the prominent keeled temporals and its lateral spinosities
by insinuating itself so firmly into the crevice that removal of the
boulders is often necessary before the creature can be captured. When
exposed in this manner, or if the little reptile has been intercepted
before gaining its rocky refuge, the armadillo lizard coils up and takes
its tail in its mouth. The strongly spinose and plated tail and limbs
thus afford protection to the others wise vulnerable under parts. Once
this posture has been assumed the lizard will permit itself to be rolled
around rather than uncoil, nor will it do so as long as it senses danger
or sees movement in its immediate vicinity. The tail plays so impor-
tant a role that it is never dropped as a means of defence, nor is it
readily detached, but if part is lost regeneration is slow and the re-
produced appendage apparently never attains the full dimensions. For
further details see Peers' account.

Temperament. Relatively slow moving (Rose). In captivity at
least the young are somewhat more active than the sluggish adults
(Schmidt). On account of this sluggishness armadillo lizards are ex-
tremely easy to catch, readily tamed, and make the most confiding
pets (FitzSimons). Consequently in the years 1930-1931 the market
in Germany was flooded with them and accounts such as those of
Senfft and Popp furnish many details of their care in vivaria, the
necessity for sun-lamps, questions of humidity, and maintenance of
temperatures of from 25-28° C. by day, 15-18° by night, etc.

Habitat. Found only in the smooth sandstone rocks; granite out-
crops appear unsuitable, possibly because granite fractures less regu-
larly and deeply than does the sandstone. The presence of these
lizards in a crack is usually betrayed by undigested calcareous or
chitinous fragments of insects from the excrement, such deposits being
considerable where the crevice has been long occupied. Each fissure
is inhabited by one family only, intrusion by an outsider being fiercely

loveridge: African lizards 57

resisted; however a family may consist of as many as eight individuals,
i.e. the original pair of adults and their resulting offspring of several
seasons.

A nocturnal gecko (Pachydactylus b. bibronii) seems to be the only
other lizard permitted to share the family retreat, perhaps because
no competition arises with regard to food. Though Agama a. atra and
Cordylus p. polyzonus may be found on the same kopjes, there is no
association with the armadillo lizard.

Localities. Cape Province: Bitterfontein; Calvinia District;
Clanwilliam District; Hondeklip Bay; Jackalswater to Orange
River; Kamaggas; Malmesbury; Matjesfontein1; Port Nolloth;
Soebatsfontein; 32 miles west of Springbok along the Kamaggas
Road.

Range. Western Cape Province to the southwestern corner of Little
Namaqualand, i.e. a coastal strip of about 100 miles south of the
Orange River and inland for about 150 miles1.

Cordylus pustulatus (Peters)
Plate 6, fig. 3

1862a. Zonurus pustulatus Peters, Monatsb. Akad. Wiss. Berlin, p. 19: Nue

Barmen, South West Africa.
1885e. Boulenger, p. 258.
1909a. Hewitt, p. 36.
1910b. Boulenger, p. 469.
1911b. Sternfeld, p. 401, fig. 1.
191 Id. Sternfeld, p. 22.
1930. Power, p. 15.

1935. Lawrence, p. 44.

Description. Head much depressed; head shields rugose; rostral
separated from the frontonasal ; nostril pierced in the lower centre of
a large nasal which is swollen; a loreal; a preocular; median subocular
not descending to the lip; fifth upper labial smallest, fifth not higher
than others; prefrontals forming a suture; temporals moderate, rugose,
keeled, without spines; gulars small, almost granular.

Dorsals and other scales said to resemble those of polyzonus. This
brief description is based on Peters, scanty one together with data de-
rived from Sternfeld's side view of the head, a drawing which may
not be too exact in details such as the number of labials, etc.

1 Boulenger's (1910b) Matjesfontein record has been questioned by Power (1930) as being
much too far east. However Dr. V. FitzSimons' assures me that caiaphraHus is plentiful at
Matjesfontein.

58 bulletin: museum of comparative zoology

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head and back olive brown spotted with darker brown
and yellow, with a tendency for the latter to form an indistinct dorsal
line. Below, brownish yellow.

Size. Total length of a cotype (Berlin Mus.), 195 (82 + 113) mm.;
and of one of Sternfeld's specimens, 170 (73 -f- 97) mm.

Remarks. Misled by Peters' comparing his cotypes with polyzonus,
Boulenger (1885e), followed by Power (1930) assumed that pustulatus
had supranasals. However, Sternfeld (1911b), with two additional
lizards from "South West Africa", reexamined the cotypes and stated
that they, as well as his specimens, lacked supranasals and were more
nearly related to cordylus. He invited attention to the suture from
nostril to first labial which was displayed by all. Mertens (1937b, p. 8),
commenting on a pustulatus in the Senckenberg Museum, confirmed
Sternfeld's findings. FitzSimons (1943) treats pustulatus as a race of
cordylus, but the large number of longitudinal dorsal scales (see sta-
tistical table) seem to preclude this.

Localities. South West Africa: Hereroland: "Neu Barmen" =
Otjimbingue (fide Sternfeld).

Range. South West Africa.

Cordylus namaquensis (Methuen & Hewitt)

1910a. tZonurus cordylus Werner (not Linne), p. 324.

1914b. Zonurus namaquensis Methuen & Hewitt, Ann. Transvaal Mus., 4,

p. 137: Great Karas Mountains, South West Africa.
1930. FitzSimons, p. 29.

1930. Power, p. 15.

1935. Lawrence, p. 43.

1938. Fitzimons, pp. 190, 191.

Description. Head much depressed; head shields rugose except for
two hindmost supraoculars; rostral rarely in contact with, usually
separated from, the frontonasal, which is shorter than broad; nostril
pierced in the lower centre of a very large nasal which is much swollen ;
a loreal; a preocular; median subocular not descending to the lip; pre-
frontals forming a suture; postfrontals as long as broad; interparietal
on a line with the anterior parietals, in contact with the postfrontals;
posterior parietals slightly larger than the anterior; 6 keeled occipitals;
temporals moderate, rugose, keeled, without spines1, but those of hind

1 Hewitt says spines moderate or poorly developed.

loveridge: African lizards 59

row pointed, projecting over ear; sides of neck with keeled, spinose
scales; gulars small, the anterior irregularly enlarged, the median
slightly imbricate, smooth, or the lateral feebly keeled; collar scales
large, only the lateral lanceolate and mucronate.

Dorsals elongate, scarcely rugose, moderately keeled, neither mucro-
nate nor serrate, those on the vertebral line regular or irregular;
laterals like dorsals but more spiny, keeled, serrate, spinose; ventrals
quadrangular, smooth, not or but slightly imbricate ; scales below fore
limbs keeled, below hind limbs smooth; tail with whorls of large,
striate, strongly keeled, serrate, spinose scales above and below, the
lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown, heavily spotted, a dark streak from
nostril through eye to ear or fore arm, a narrower streak from posterior
corner of eye along upper edge of temporals, lower labials edged with
darker; back light to dark chestnut brown variegated with darker
brown; two dark brown lateral bands sometimes present. Below,
grayish white to light muddy brown; tail light brown.

Size. Total length of type (T.M. 3163), 170 (81 + 89) mm.

Remarks. One wonders if Werner's (1910a, p. 324) cordylus from
"S. Africa" with 10-11 femoral pores and color of namaquensis is not
that species with which it agrees in all of the data published.

Parasites. Mites (Zomirobia montana) were found by Lawrence.

Habitat. Occurs at 6200 feet (Hewitt) as well as on lower slopes of
mountain (FitzSimons) and in rocky crevices of a river bed.

Localities. South West Africa: Great Karas Mountains: —
Kochena Farm; Narudas Sud Farm; Sandmund; and summit of a
mountain near Wasserfall.

Range. Great Karas Mountains and vicinity, South West Africa.

Cordylus campbelli (FitzSimons)

1938. Zonurus campbelli FitzSimons, Ann. Transvaal Mus., 19, p. 189:
Barby Farm, 10-20 miles east of Helmeringshausen, South West
Africa.

Description. Head much depressed; head shields rugose; rostral
separated from the frontonasal, which is longer than broad or as long
as broad; nostril directed upwards and outwards in a nasal which is
much swollen and slightly tubular ; lower eyelid with a semitransparent
disk; prefrontals forming a suture; postfrontals as long as broad; inter-

60 bulletin: museum of comparative zoology

parietal enclosed between 2 pairs of parietals; posterior parietals
slightly larger than the anterior; 6 keeled, rugose occipitals; temporals
moderate, rugose, keeled, without spines, but those of hind row ob-
tusely pointed and flattened, projecting over ear; sides of neck with
keeled, spinose scales; gulars small, the anterior irregularly enlarged,
the median slightly imbricate, smooth, or the lateral feebly keeled;
collar scales large, only the lateral lanceolate and mucronate.

Dorsals elongate, scarcely rugose, obtusely keeled, neither mucro-
nate nor serrate, those on 'the vertebral line irregular; laterals like
dorsals but more spiny, keeled, serrate, spinose; ventrals quadrangular,
smooth, slightly imbricate; scales below forelimbs keeled; below hind
limbs smooth, serrately pointed, imbricate; tail with whorls of large,
striate, strongly keeled, serrate, spinose scales above and below, the
lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head chestnut brown, labial region lighter; back
chestnut brown spotted and irregularly crossbarred with darker brown,
a broad vertebral band or chain bearing pale yellowish spots; tail
chestnut brown more or less barred with dark brown. Below, grayish
white, gular region with rusty infuscations; forelimbs straw yellow;
callose patch anterior to, as well as, femoral pores of male, pale yellow.

Size. Total length of d" type (T.M. 17635), 164 (76 + 88) mm.; of
paratype 9 (T.M. 17637), 162+ (79 + 83+) mm., tail-tip being lost.

Habitat. The type series were taken in rock fissures among outcrops
on valley slopes of Barby Farm, which is northwest of Keetmanshoop
and the Great Karas Mountains, Great Namaqualand.

Localities. South West Africa: Great Namaqualand: Barby
Farm.

Range. Great Namaqualand, South West Africa.

CORDYLUS POLYZONUS POLYZONUS Smith

Plate 6, fig. 4
Plate 7, fig. 1

1838a. Cordylus polyzonus A. Smith, Mag. Nat. Hist. (2), 2, p. 31: Cape of

Good Hope.
1843. Smith, A., pi. xxviii, fig. 1; xxx, figs. 7-7b: North of Orange River

and within the Colony, i.e. Cape Province.
1845. Gray, p. 47.

1839. Zonurus polyzonus Dumeril & Bibron, p. 357.

loveridge: African lizards

1885e. Boulenger (part), p. 257.

1887b. Boettger (part), p. 143.

1888b. Fischer, p. 12.

1893a. Boettger (part), p. 66.

1896c. Bocage, p. 117.

1898. Sclater, p. 103.

1903e. Boulenger, p. 217.

1905h. Boulenger, p. 253.

1907b. Roux, p. 421.

1909a. Hewitt (part), p. 36 (omit Walfish Bay).

1910b. Boulenger, p. 469 (omit Natal as error).

1910a. Werner (part), p. 325 (omit S. Hereroland only).

1911b. Hewitt, p. 47.

1911b. Sternfeld (part), p. 402.

1911d. Sternfeld (part), p. 22, fig. 21 (omit Rehoboth, etc.).

1913. Hewitt & Power, p. 154.

1913a. Werner, p. 108.

1914b. Methuen & Hewitt (part), p. 137.

1918. Power, p. 264.

1928. Cott, p. 953.

1929. Rose, pp. 100, 218.

1930. FitzSimons (part), p. 29 (omit Okahandja).
1930. Power, p. 14.

1933a. Power, p. 215.

1934a. Cott, p. 153.

1935a. FitzSimons, p. 534.

1935. Lawrence, p. 43.

1936. Parker, p. 134.
1937a. FitzSimons, p. 266.
1937e. Hewitt, p. 30, pi. x, fig. 7.

1937. Lawrence, p. 109.

1938. FitzSimons, p. 192.
1940c. Scortecci, p. 7.

Further citations of "polyzonus" will be found under p. jordani.

Names. Karroo Girdle-tail (English); likkwanakkedis (Kogopa).

Description. Head much depressed; head shields rugose; rostral
separated from the frontonasal, which is as long as broad or shorter
than broad; nostril pierced in the postero-superior corner of a very
small nasal which is much swollen; a loreal (rarely fused with first
labial), a preocular; lower eyelid with a semitransparent disk; median
subocular descending to the lip between the fourth and fifth, or fifth
and sixth, upper labials; fourth upper labial lowest, fifth higher than
others; supranasals forming a suture; prefrontals forming a suture;

62 bulletin: museum of comparative zoology

postfrontals as long as broad or shorter than broad; interparietal en-
closed between 2 pairs of parietals; posterior parietals slightly larger
than the anterior ; 5-6 rugose occipitals ; temporals moderate (slightly-
smaller than in -p. jordani), rugose, keeled, without spines, but those
of hind row pointed, projecting over ear; sides of neck with short,
keeled, spinose scales; gulars small or moderate, the anterior irregu-
larly, or not, enlarged, the median slightly imbricate, smooth, or the
lateral feebly keeled; collar scales slightly enlarged, undifferentiated.

Dorsals elongate, rugose, slightly or strongly keeled, neither mu-
cronate nor serrate, those on the vertebral line regular or irregular;
laterals like dorsals but more strongly keeled, serrate, spinose; ventrals
quadrangular, smooth, not or but slightly imbricate, the lateral some-
times slightly keeled and shortly mucronate; scales below fore limbs
keeled, below hind limbs smooth or slightly keeled; tail with whorls
of large, rugose, strongly keeled, serrate, spinose scales above and
below, the lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Very variable, scarcely two the same. Above, head brown;
back yellowish brown, olive brown, chestnut brown, or blackish brown,
uniform, variegated, or blotched with darker, the irregularly shaped
spots tending to form two longitudinal lines; a blackish streak on side
of neck; tail yellowish brown sparingly spotted with umber brown.
Young are olive chequered with black and white, their tails barred
brown and white. Below, whitish, yellowish, brownish or reddish,
uniform or clouded with darker spots and stripes.

Power (1918) writes of the dark dorsal coloring harmonizing per-
fectly with the environment, but the "brilliant red" gular ventral sur-
face possessed by both sexes throughout the year, displayed to ad-
vantage as they sit erect, is immediately concealed at the approach
of danger by the lizard pressing its chin close to the rock. From this
he concludes that the ventral pigmentation has no significance as a
warning color. Later (1933a), he records De Aar lizards as being
"very dark purple, almost black" below, those at Orange River Sta-
tion, as being "brick red sprinkled with very dark purple" etc. Hewitt
(1937e), however, speaks of adults which are almost black in the
breeding season. For further variations consult FitzSimons (1935a;
1938).

Remarks. FitzSimons (1937a) offers comments on three cotypes in
the British Museum. Scortecci (1940c) remarks on scale formation in
this species, contrasting it with that of other families.

loveridge: African lizards 63

Size. Total length of a tf1 (T.M. 18206), 237+ (116 + 121+) mm.,
and of an unsexed specimen (T.M. 15343), 266 (114 -f- 152) mm.

Sexual dimorphism. Femoral pores are present in the males only to
any appreciable degree of development.

Breeding. Mating takes place early in September and the young are
born in January (presumably in Cape Province) according to Hewitt
and Power. However four fully developed young were present in
uteri of a female taken in April at Kubub, South West Africa
(Werner).

Diet. Beetles, locusts and other insects, but one contained a gaudy
and nauseous grasshopper (Zonocercus elegans), another grass stalks
(Hewitt & Power).

Parasites. Mites (Ixodiderma inverta; Scaphothrix convexa; Zonu-
robia polyzonensis) were found by Lawrence ; while nematodes are said
to infest this species at times according to Hewitt and Power.

Defence. The tail is employed to shield the body as soon as the rock
shelter is reached, but the tail, though brittle, is not parted with unless
the lizard is subjected to very rough handling (Hewitt & Power).

Temperament. Alert and active (FitzSimons). Shy and timid
though occasionally attempting to bite when tormented (Hewitt &
Power) .

Hibernation. This is neither long nor deep (Hewitt & Power).

Habits. Apparently polygamous for nine females accompanied by
only a single male were found in one locality (Hewitt & Power).

Habitat. This common species occurs in rocky situations over a
wide area, often in the vicinity of human habitations. They like to
bask in the fierce summer heat with belly pressed to the rock but fore-
part raised, the head and neck being almost vertical. In this position
a lizard will remain for hours, only turning its head in response to
sounds or to watch some movement in the vicinity. On hot days they
face the sun, but on cold ones prefer to expose their backs to its warm-
ing rays. If disturbed each lizard quickly retires beneath somcboulder
or into its particular crevice, for they exhibit a marked attachment to
the same retreat.

Localities. Bechuanaland Protectorate : Kuruman. Orange
Free State: Bloemfontein; Boshof; Jacobsdal; Smithfield. Cape
Province: Albert; Aliwal North; Barkly West; Belmont; Bredas-
dorp; Britstown; Bros Pan; Burghersdorp; Calvinia; Cape Town;
Clanwilliam; Colesberg; Cradock; De Aar; Deelfontein; Faure-
smith; Fort Richmond near Herbert; Fourteen Streams; Garies —
near; Graaff-Reinet ; Hanover; Hay; Herbert; Hoetjes Bay; Hope-

64 bulletin: museum of comparative zoology

town1; Jackalswater; Kamaggas; Kenhardt; Kimberly; Klaver;
Klipfontein; Knysna; Kubroos; Lambert's Bay; Lekkersing; Little
Namaqualand; Malmesbury; Matjesfontein; Middleburg; Morti-
mer; O'okiep to Springbok; Orange River Mouth; Orange River
Station; Philipstown; Pof adder; Port Nolloth; Prieska; Riet Pan;
Rooidam; Rosmead; Rust-en-Vrede; Soebatsfontein; Steinkopf;
Steynsburg; Steytlerville; Strydenburg; Touws River; Uitkyk;
Upington; Van Rhynsdorp; Van Wyksvlei; Vredendal; Worcester.
South West Africa: Aus; Aus to Bethany (Bethanien); Barby
Farm; Great Karas Mountains; Great Namaqualand; Kakamas;
Kolmanskop (M.C.Z.); Kubub; Kuibis; Luderitz Bay (Angra
Pequena) ; Prince of Wales Bay ; Sinclair Mine (? subsp) ; Warmbad.
Range. 2Southern Bechuanaland, Orange Free State, and Cape
Province west to southern South West Africa in Great Karas Moun-
tains and at Aus and Barby (where it meets with the northern race — •
jordani).

Cordylus polyzonus jordani (Parker)

1862a. Zonurus -polyzonus Peters (not Smith), p. 18.

1869b. Peters, p. 660.

1885e. Boulenger (part), p. 257.

1887b. Boettger (part), p. 143.

1890b. Miiller, p. 699.

1893a. Boettger (part), p. 66 (Damaraland only).

1894a. Boettger, p. 89.

1907. ' Schultz, p. 188, fig.

1909a. Hewitt (part), p. 36 (Walfish Bay record).

1910a. Werner (part), p. 325 (Hereroland only).

1911b. Sternfeld (part), p. 402 (2 specimens lacking black mark).

1911d. Sternfeld (part), p. 22 (Rehoboth; Damaraland; Hereroland).

1914. Methuen & Hewitt (part), p. 137.

1915c. Werner, p. 338.

1930. FitzSimons (part), p. 29, (Okahandja, record only).

1936c. Zonurus jordani Parker, Novit. Zool., 40, p. 133: Hoffnung, near

Windhoek, South West Africa.

1938. FitzSimons, p. 191.

1937b. Cordylus jordani Mertens, p. 8.

Description. Head much depressed; head shields rugose; rostral
separated from the frontonasal, which is as long as broad or shorter

1 The Irene record was subsequently corrected by Hewitt (1911b).

2 The Natal record is doubted by both Boulenger and Power, and is quite out of the question
according to FitzSimons (letter).

loveridge: African lizards C5

than broad; nostril pierced in the postero-superior corner of a very
small nasal which is much swollen; a loreal; a preocular; lower eyelid
with a semitransparent disk; median subocular descending to the lip
between the fourth and fifth, or fifth and sixth, upper labials, fourth
or fifth upper labial lowest, fifth or sixth higher than others; supra-
nasals forming a suture; prefrontals forming a suture ; postfrontals as
long as broad; interparietal enclosed between 2 pairs of parietals;
posterior parietals slightly larger than the anterior; 6 rugose occipitals;
temporals large (slightly larger than in p. polyzonus), keeled, without
spines, but those of hind row pointed, projecting over ear; sides of
neck with short, keeled, spinose scales; gulars small, the anterior ir-r
regularly enlarged, the median slightly imbricate, smooth, or the lateral
feebly keeled; collar scales slightly enlarged, undifferentiated.

Dorsals elongate, slightly rugose, moderately keeled, neither mu-
cronate nor serrate, those on the vertebral line regular or irregular;
laterals like dorsals but more strongly keeled, serrate, spinose ; ventrals
quadrangular, smooth, not or but slightly imbricate, the lateral some-
times slightly keeled and shortly mucronate; scales below fore limbs
keeled, below hind limbs smooth or slightly keeled; tail with whorls
of large, rugose, strongly keeled, serrate, spinose scales above and
below, the lateral spines longest.

For characters common to all species, see definition on p. 9; for
scale and pore counts, see statistical table on p. 15.

Color. Above, head brown or yellow; back yellowish, pale brown, or
olive brown, uniform (in old lizards) or variegated and barred with
darker; no black streak on side of neck; flanks yellowish to pale olive.
Below, whitish or pale straw, uniform, or gular region longitudinally
vermiculated with gray; tail yellowish or brownish.

Size. Total length of & (T.M. 17468), 251 (125 -f 126) mm.; of
9 (T.M. 17496), 255 (127 + 128) mm. Length of 9 holotype (Brit.
Mus.) from snout to vent was 111 mm., the tail-tip lost.

Remarks. The describer, in addition to the characters utilized in
the key for separating this form from p. polyzonus, mentions the
larger temporals of jordani — a valid character but difficult to appre-
ciate without comparative material — and the number of dorsals in
transverse row, the latter no longer holds. FitzSimons (1938) says
that in polyzonus there a^e usually 6 large sublabials beneath the lower
jaw, whereas in his twenty-one jordani there are but 5. In our M.C.Z.
material of p. polyzonus both 5 and 6 are so common that I doubt the
value of such an average character for taxonomic purposes.

FitzSimons (1935a) reports that specimens from Aus — which is in

66 bulletin: museum of comparative zoology

the area where one would expect to encounter intergrades — with 12-16
femoral pores, i.e. p. polyzonus, have the intensive caudal keeling, and
spinosity which one would expect of p. jordani, thus offering further
justification of our treating jordani as a race of polyzonus.

Localities. South West Africa : Aus — 15 miles east of ; Barby
Farm; Damaraland; Helmeringhausen; Hereroland; Hoffnung;
Karub; Keetmanshoop; Kobos; Kraikluft; Kraikluft to Sandmund;
Kraikluft to Alt Wasserfall; Neu Barmen; Neudamm Farm;
Okahandja; Okosongomingo; Otjosongombe; Rehoboth; Usakos;
Walfish Bay; Waterberg; Windhoek (Windhuk).

Range. South West Africa from Great Karas Mountains, Aus and
Barby Farm, Great Namaqualand (where the typical form also occurs)
north to Herreroland.

Genus Pseudocordylus

1838a. Hemicordylus A. Smith, Mag. Nat. Hist. (2) 2, p. 32 (type capensis

Smith).
1838a. Pseudocordylus A. Smith, Mag. Nat. Hist. (2) 2, p. 32 {type montanus

Smith = microlepidotus Cuvier).

Head and body depressed; limbs well developed; tail moderate.
Head shields regular; nostril pierced in a nasal or between nasal and
first labial; eyelids well developed; ear-opening large; 4 parietals; sides
of neck covered with granules; a collar fold, attached mesially, at
least indicated; dorsals usually small, soft or nodular, usually inter-
mixed with granules, in more or less regular series which, if extending
to occiput, are much reduced on nape if not granular; ventrals large,
quadrangular or subtriangular, juxtaposed, smooth, forming longi-
tudinal and transverse series; femoral pores present in both sexes;
digits slightly keeled inferiorly; tail spinose.

Those species which I have examined appear to have the following
characteristics in common, consequently these have been omitted
from the specific descriptions.

Head longer than broad; head shields more or less smooth anteri-
orly, rugose posteriorly; rostral at least twice as broad as high; nasal
scarcely swollen; postnasal present only as an aberration; a loreal;
a preocular; subocular descending to the lip between two labials; fifth
upper labial not higher than the others (except in figure of m'.fasciatus) ;
anterior supraocular longest, the second (sometimes first and second
in robertsi) broadest; prefrontals forming a suture (sometimes separated

loveridge: African lizards 67

in capensis and robertsi) ; frontal hexagonal, slightly broader anteriorly;
interparietal enclosed between 2 pairs of parietals (sometimes not in
capensis); posterior parietals much (sometimes only slightly in ro-
bertsi) larger than the anterior; occipitals much reduced or absent;
mental rather large ; lower labials bordered by a row of 5 large shields ;
lateral collar scales enlarged but otherwise undifferentiated; a slight
lateral fold; a pair of enlarged preanals; limbs covered above with
large, keeled, more or less spinose, imbricate or subimbricate scales;
tail with whorls of large, alternating with whorls of smaller, rugose,
keeled, mucronate scales, the lateral spines longest, subcaudals long
and narrow.

Range. Union of South Africa south of 24°S.

Remarks. The transference of capensis (and robertsi) from Cordylus
to this group to which they appear more nearly related, for reasons
stated elsewhere (p. 71), raises the question as to whether the sub-
genus Hemicordylus, which has paragraph precedence, should not
be employed in preference to the former subgenus Pseudocordylus. As,
however, this is not mandatory under the International Code of
Nomenclature, I prefer to use the name Pseudocordylus, not merely on
account of its having been raised to generic status by Gray (1845)
and consequently long-standing in usage, but also because it is not im-
probable that later it may be found advisable to accord generic status
to the group of annectant species at present characterized only by
capensis, robertsi and langi.

Key to the Species

1. Flanks entirely covered with minute granules or at most
some widely separated, small, subcorneal tubercles differ-
ing greatly from the enlarged dorsals 2

Flanks entirely covered with granule-surrounded nodu-
lar scales, or scales only, which are a continuation of
the dorsals 4

2. Back entirely covered with strongly keeled dorsals
forming 10-12 longitudinal rows; ventrals in 8 longi-
tudinal rows; range: southern Cape Province between

False and Mossel Bays capensis

(p. 70)
Back, or vertebral region only, covered with feebly
keeled, or smooth, dorsals of which at most (in robertsi
only) only one or two dorso-lateral rows are strongly
keeled 3

68 bulletin: museum of comparative zoology

3. Slightly enlarged dorsals form 12-20 irregular, ill-
defined, longitudinal rows covering entire back; ventrals "
in 8 longitudinal rows; enlarged temporals 10-13; range:

Van Rhynsdorp District, Cape Province robertsi

(p. 71)
Slightly enlarged dorsals form 6-8 irregular, ill-defined,
longitudinal rows in vertebral region; ventrals in 10
longitudinal rows; enlarged temporals 5-6; range:
Drakensberg of Basutoland to eastern Cape Province
(possibly southwestern Transvaal if Doornkop referable) langi

(p. 73)

4. Temporals of the upper row enlarged and vertically

elongate 5

Temporals of the upper row relatively small and polygo-
nal, at most but one or two vertically elongate 6

5. Median gulars mostly elongate like the lateral; range:
Mountains of southern Transvaal and Orange Free

State south to Drakensberg, Natal m. mclanotus1

(p. 75)
Median gulars more or less squarish, not even slightly
elongate like the lateral; range: Mountains of western

Cape Province m. namaquensis

(p. 78)

6. Enlarged temporals about 8-11; median gulars slightly
elongate like the lateral ; range : Mountains of the Cape

Peninsula, Cape Province m. microlepidotus

(p. 79)
Enlarged temporals about 16-17; median gulars more
or less squarish, not even slightly elongate like the lat-
eral ; range : Mountains of eastern Cape Province m. fasciatus

(p. 80)

includes subviridis (A. Smith) and s. transvaalensis (FitzSimons).

loveridge: African lizards

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capensis

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m. microlepidotus

m. fasciaius

70 bulletin: museum of comparative zoology

PsEUDOCORDYLUS CAPENSIS (Smith)

Plate 7, figs. 2-3

1838a. Cordylus (Hemicordylus) Capensis A. Smith, Mag. Nat. Hist. (2), 2,

p. 32: Cape of Good Hope.

1843. Smith, A., pi. xxvii, fig. 2; pi. xxx, figs. 6-6b : Hottentot's Holland

Mountain, Cape Province.

1839. Zonurus Capensis Dumeril & Bibron, p. 360

1885e. Boulenger, p. 258.

1898. Sclater, p. 103.

1909a. Hewitt, p. 36.

1910b. Boulenger, p. 469.

1910a. Hewitt, p. 60.

1914a. Hewitt, p. 239.

1930. Power, p. 15.

1935. Lawrence, p. 44.

1937a. FitzSimons, p. 266.

1937d. Hewitt, p. 206.

1937. Lawrence, p. 110.

Description. Rostral separated from the frontonasal, which is
longer than broad or as long as broad; nostril pierced in the lower
centre of a nasal; median subocular descending to the lip between the
fifth and sixth upper labials ; postfrontals as long as broad; interparietal
enclosed between two pairs of parietals or on a line with the anterior
parietals, forming a suture with the postfrontals; posterior parietals
much larger than the anterior; no occipitals; temporals of upper row
vertically elongate; gulars smooth, a few anterior ones irregularly en-
larged, the median more or less squarish.

Dorsum covered with large, squarish, unequal-sized, strongly keeled
scales, more or less intermixed with minute granules, forming 10-12
longitudinal dorsal and 43^44 transverse series from nape to base of
tail, the 2 vertebral rows largest and separated on the vertebral line
by flattish granules of irregular shape; flanks covered with small sub-
circular granules; scales below fore limbs smooth, obtusely keeled, or
granular, below hind limbs smooth or obtusely keeled.

For characters common to all species, see definition on p. 66, for
scale and pore counts, see statistical table on p. 69.

Color. Above, head black or brown, back and sides black or purplish
brown with, or without, seven to eight fine, transverse white lines on
nape and back. Below, black, or throat and belly dull brownish red;
soles of feet pale yellowish brown ; tail grayish.

loveridge: African lizards 71

Size. Total length of type (?9 ), first (1838a) given as iy2 inches,
later (1843) expressed as 9 unc. 9 lines, is correctly rendered by
Boulenger (1885e) as 264 (108 + 156) mm.

Remarks. FitzSimons considers that the discrepancy in size clearly
indicates the use of two specimens, on the other hand misprints re-
sulting from Smith's handwriting are not infrequent in "Illustrations."
Smith himself says (1843) that the only example which he had seen
was taken by Mr. Ford of the 49th Regiment and had been deposited
in the Army Medical Museum at Fort Pitt, Chatham. This is ob-
viously the specimen seen by FitzSimons in the British Museum,
having been received by them from the Army Medical College.

No other specimens were recorded until Hewitt (1937d) discussed
eight which he had received from localities to the northwest and east
of the type locality. His findings are embodied in the redescription
given above. It is he who suggested the subspecific relationship with
robertsi which I have hesitated to adopt.

The transference of capensis from Cordylus to Pseudocordylus is
because it agrees with the latter in having the neck covered with
granules instead of scales, and is so closely related to robertsi, whose
dorsals lack osteoderms, that I have no misgivings in effecting the
transfer. Smith himself recognized the annectant character of capensis
by erecting the subgenus Hemicordylus for its reception; my reasons
for not using that name instead of Pseudocordylus are discussed under
the generic heading.

Localities. Cape Province : Hottentot's Holland Mountain east
of Stellenbosch; Jonkershoek near Stellenbosch; Langebergen near
Swellendam.

Range. Cape Province (Between False and Mossel Bays).

Pseudocordylus robertsi (van Dam)

Plate 8, fig. 1

1921a. Zonurus robertsi van Dam, Ann. Transvaal Mus., 7, p. 241, pi. iv:

Klaver, van Rhynsdorp District, Cape Province.

1930. FitzSimons, p. 30.

1930. Power, pp. 13, 15, fig. 1, pi. ii, fig. 1.

1935a. FitzSimons, p. 532.

1935. Lawrence, p. 44.

1937d. Hewitt, p. 207.

1937. Lawrence, p. 110.

72 bulletin: museum of comparative zoology

Description. Rostral separated from the frontonasal, which is longer
than broad or as long as broad; nostril pierced in the lower centre of a
nasal, often between nasal and first labial; median snbocular descend-
ing to the lip between the fourth and fifth or fifth and sixth upper
labials; postfrontals longer than broad or as long as broad; inter-
parietal enclosed between 2 pairs of parietals; posterior parietals
slightly or much larger than the anterior; no occipitals; temporals of
upper row vertically elongate; gulars smooth, a few anterior ones ir-
regularly enlarged, the median more or less squarish.

Dorsum covered with small, soft, subquadrangular, feebly or
strongly keeled scales, sometimes intermixed with granules, forming
12-20 irregular (rarely regular) longitudinal and 42-64 transverse
series from nape to base of tail, the 2 vertebral rows, and sometimes 2
dorsolateral rows, enlarged; flanks covered with small, subcircular
granules ; scales below forelimbs smooth, obtusely keeled, or granular,
below hind limbs smooth or obtusely keeled.

For characters common to all species, see definition on p. 66, for
scale and pore counts, see statistical table on p. 69.

Color. Above, head and back dark brown or blackish with scattered
yellowish spots which sometimes tend to form cross-bars on the dor-
sum. Below, bluish white to blue gray, the gular region dark blue in
males whose callose patches of femoral scales and pores are yellowish
or dirty white.

Size. Total length of tf type (T.M. 3747), 267 (98 + 169) mm.,
exceeded in body length by a cf (T.M. 15656), 264 (104 + 160) mm.;
length of a 9 topotype (M.C.Z. 41875), 213 (85 + 128) mm.

Remarks. I had already transferred robertsi to Pseudocordylus prior
to reading Power's (1930) remarks on the subject. Despite FitzSimons
(1935a) statement, I can find no osteoderms by dissection and thus
there appears no reason for retaining robertsi in Cordylus from all other
species of which it differs in having the neck and flanks covered with
granules. From the description of capensis I imagined that robertsi
should be treated as a subspecies of it, as has been done by FitzSimons
(1943, p. 438). However on receipt of a specimen of capensis and
making direct comparison with a paratype of robertsi, such action
appears unjustified. The nasal variation figured by Power seems of
no significance and can be matched by members of either genus.

Parasites. Mites (Zonurobia subquadrata) described from this
species by Lawrence, are most closely related to the species found on
Cordylus cacrideopunctatus, a point that may have some significance.

Temperament. Shy and difficult to approach (FitzSimons).

loveridge; African lizards 73

Habitat. Found, together with Agama a. atra, on the precipitous
edges of the escarpment overlooking the low country towards Van
Rhynsdorp (FitzSimons), and among the sandstone rocks which form
the "tables" of the Table Mountain sandstone near Clanwilliam and
Van Rhynsdrop (Lawrence).

Localities. Cape Province: Klaver; Van Rhyn's Pass between
Nieuwoudtville and Van Rhynsdorp.

Range. Cape Province (Van Rhynsdorp District).

PSEUDOCORDYLUS LANGI spec. nOV.

The following references •probably relate to this form, which in
FitzSimons (1943, pp. 467-469) is included in his "s. subviridis."

1910b. Pseudocordylus microlepidotus Boulenger (part, not Cuvier), p. 469

(Morija, Basutoland).
1925. Essex, p. 338 (Amatola Mountains, Cape Province).

1925b. Hewitt, p. 356.
1928a. Essex, 1927, p. 932.

1927a. P(seudocordylus) subviridis Hewitt (not A. Smith), p. 392.
1935. Lawrence, p. 44.

1937e. Hewitt, p. 31.

Name. Lesser Caiman Lizard (Hewitt).

Type. Museum of Comparative Zoology, No. 46835, an adult or
subadult cf from Mont-aux-Sources, Drakensberg, Basutoland, col-
lected by Herbert Lang, November, 1930.

Paratypes. Transvaal Museum, Nos. 13846-7, 13849-50 from same
locality as type at 11,000 feet; Nos. 2531, 2533 from Drakensberg on
Basutoland side; No. 20992 from Drakensburg near Underberg at
6,000 feet; No. 21063 from Drakensberg near Kokstad.

Diagnosis. Differs from P. capensis and robcrtsi in the feeble devel-
opment of enlarged dorsal scales which are confined to the vertebral
region. Differs from subviridis, with which it has been confused, and
all races of microlepidotus in having its flanks covered by homogeneous
granules.

Description. Rostral separated from the frontonasal, which is
shorter than broad; nostril pierced in the lower centre of a nasal;
median subocular descending to the lip between the fourth and fifth
labials ; postfrontals as long as broad; upper row of temporals vertically
elongate; gulars smooth, a few anterior ones slightly enlarged, the
median elongated like the lateral.

Dorsum with an irregularly broad vertebral band of small, squarish,

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smooth, juxtaposed scales, of which there are 6-8 in longitudinal series,
flanked by still smaller, unequal granules similar to those on flanks
which scarcely decrease in size towards the lowest part; below fore-
limb covered with granules; below hind limb anteriorly large smooth
scales merging gradually into granules posteriorly. Type with 14
femoral pores.

Agreeing with characters common to all species as defined on p. 66;
for scale and pore counts, see p. 69.

Color. In alcohol. Abo ve, head grayish olive variegated with sepia ;
tip of snout to ear horn-yellow vertically striped with sepia on the
lips ; back olive brown with a vertebral series of coalescing pale spots
which unite with others to form transverse series. Below, lower labials
horny -gray indistinctly blotched with brown ; throat, chest, belly, and
beneath limbs plumbeous; neck, soles of feet, and tail horn-gray.

Size. Total length of type & (M.C.Z. 46835), 221 (90 + 131) mm.

Remarks. I have not seen the Paratypes which are included on the
basis of information kindly supplied by Mr. V. FitzSimons. Appar-
ently lizards somewhat intermediate in character between langi and
subviridis but nearer latter, occur at 7,000 feet on Mont aux Sources,
for of Tvl. Mus. Nos. 13851-3 FitzSimons writes: "the scales on flanks
are better developed, keeled and closer together, though still well
separated by granular interspaces."

It should be added that all information below is based on the as-
sumption that the lizards referred to in the above citations are truly
langi.

Diet. Twice observed eating lichen from the rocks (Essex).

Parasites. Mites (Ixodiderma inverta and I. pilosa) described from
this form by Lawrence.

Temperament. An active and wary lizard (Hewitt).

Habitat. From 11,000 feet down to 7,000 feet and lower at some
points of Mont-aux-Sources, under rocks, but possibly a burrower'
according to Essex, who found one at the terminus of a sixteen-foot
burrow in soft soil near the summit of Katberg.

Localities.1 Basutoland: Drakensberg near Kokstad; Mont-
aux-Sources; Morija; Nemahadi Police Camp; near Underberg at
6,000 feet. Cape Province: Amatola Mountains; Great Winter-
berg; Hogsback; Katberg; Malutsenyane; Thaba Putsua, Ra-
baneng Pass; Ugie.

Range2. Basutoland to eastern Cape Province.

1 As all localities, except those of type and paratypes, are taken from the literature they
should be regarded -with reserve.

2 Unless referable to P. m. melanolus, the specimens from Doornkop, near Belfast, Transvaal,
mentioned by Hewitt, should be added.

LOVERIDGE: AFRICAN LIZARDS 75

PSEUDOCORDYLUS MICROLEPIDOTUS MELANOTUS (Smith)

Plate 8, figs. 2-3
Plate 9, figs. 1-3

1838a. Cordylus (Pseudocordylus) melanotus A. Smith, Mag. Nat. Hist. (2),
2, p. 32: "Cape of Good Hope," later given as Hills near Orange
River east of "Phillopolis," i.e. Phillipolis, Orange Free State.

1838a. Cordylus (Pseudocordylus) sub-viridis A. Smith, Mag. Nat. Hist. (2),
2, p. 33: "Cape of Good Hope," later given as Mountains behind
Kaffirland and Natal.

1843. Cordylus microlepidotus vars. A. Smith, pi. xxv, figs. A-B; pi. xxvi;
pi. xxx, figs. 3-3b (melanotus) and 4-4b (subviridis).

1891a. Pseudocordylus microlepidotus Matschie (not Cuvier), p. 606.

1905h. Boulenger, p. 253.

1907b. Roux, p. 422.

1908b. Boulenger, p. 224.

1909a. Hewitt (part), p. 37 (Transvaal, Orange Free State, Natal).

1910b. Boulenger (part), p. 469 (Natal only).

1920. Hewitt, p. 93.

1937a. Pseucordylus microlepidotus subviridis FitzSimons, p. 266.

1943. Pseudocordylus subviridis subviridis FitzSimons (part, rest langi),
p. 467.

1943. Pseudocordylus subviridis transvaalensis FitzSimons (? part, see Re-
marks below), Lizards of South Africa, p. 469: Woodbush, Pieters-
burg District, northern Transvaal.

Further citations of "subviridis" will be found under langi.

Description. Rostral separated from, or rarely in contact with, the
frontonasal, which is as long as (M.C.Z. 46837), or shorter than, broad
or longitudinally divided, sometimes with an azygous scale between it
and the prefrontals (M.C.Z. 21274) or the nasals (M.C.Z. 21443);
nostril pierced in the lower centre or postero-inferior corner of a nasal ;
median subocular descending to the lip between the fouth and fifth or
fifth and sixth upper labials; postfrontals as long as broad or shorter
than broad ; upper row of temporals vertically elongate ; gulars smooth,
a few anterior ones irregularly enlarged, the median usually moderately
or strongly elongated like the lateral.

Dorsum covered with small, flat or raised, subcircular or squarish,
smooth or keeled, granular scales, forming more or less regular longi-
tudinal and transverse series, those of the vertebral region smallest
and more closely juxtaposed than the larger dorsolateral, which are
separated by minute granules; scales on flanks subcircular or sub-

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triangular, decreasing in size on lowest part of flanks and separated
horizontally by granules interspersed with larger, flat, subcircular
scales (melanotus) or each vertical series of scales juxtaposed (sub-
viridis), the larger scales obtusely keeled (young) or smooth (adult);
scales below fore limbs obtusely keeled or granular, below hind limbs
smooth or obtusely keeled.

For characters common to all species, see definition on p. 66, for
scale and pore counts, see statistical table on p. 69.

Color. Above, head brown variegated with yellow, tip of snout and
lip to ear horn-yellow ; back of male yellowish brown, brown, or black,
with numerous spots in longitudinal series ; back of female chequered
with black and with series of elongated yellowish spots ; both sexes with
faint traces of obsolescent cross-bars descending to the flanks where
they break up to form vertical bars; flanks yellow or orange yellow
tinged with vermilion in male ; sides of neck with one or two large black
spots; limbs and tail handsomely barred or vermiculated with black.
Below, gular region of male deep blue (sometimes extending up on to
sides of head), of female yellowish infuscated with gray; belly greenish
yellow tinged with vermilion; tail with dark cross-bars, the tip en-
tirely black.

Size. Total length of a cf (M.C.Z. 14240), 273+ (120 + 153+) mm.,
from Forbes Reef, Swaziland, of a young 9 (M.C.Z. 21274), 188 (88 +
100) mm., from Kastrol Nek, southeastern Transvaal.

Total length of & (M.C.Z. 14241), 217 (97 + 120) mm., and 9
(M.C.Z. 21443), 177 (88 + 89) mm., both from Giants Castle, Natal.

Remarks. Unfortunately the types of melanotus and subviridis are
lost (FitzSimons, 1937a) and the present disposition must be regarded
only as tentative. The precise status and ranges of the forms of this
difficult group will not be settled until some South African herpetolo-
gist is able and willing to assemble all the material from the South
African museums and subject them to intensive comparative study.

FitzSimons (1943, p. 464) places melanotus in the synonymy of
microlepidotus which has small temporals. This is obviously wrong for
Sir A. Smith's figure of melanotus (3b) shows it as having vertically
elongate temporals like subviridis (4b). What FitzSimons calls sub-
viridis (1943, p. 467) is a composite of melanotus, subviridis and langi
spec. nov. That the name subviridis cannot be applied to the granular
flanked langi (which is more closely related to robertsi than to any
microlepidotus) is clear from Smith's statement that the flanks of
subviridis are covered with keeled scales.

FitzSimons' figures of subviridis (figs. 373-374) from Giant's Castle

loveridge: African lizards 77

undoubtedly represent that form, which I am inclined to think may
be separable as a southeast race on the basis of the almost continguous,
vertical (not horizontal) juxtaposition of the lateral scales. In the
more northern form {melanotus + transvaalensis) such scales are sepa-
rated both vertically and horizontally by granules and with or without
small, scattered, subcircular scales. Where the two forms merge it is
impossible for me to say, and instead of speculating I prefer to treat
both as melanotus for the difference may not prove to be constant when
a large series is studied.
FitzSimons (1943, p. 463) separates two forms as follows :

A single row of large vertically elongate temporals ; lowermost tem-
poral spine moderately projecting in males s. subviridis

Two rows of temporals, the upper vertically elongate and much
larger than the subhexagonal lower; lowermost temporal spine
feebly projecting and only bluntly pointed . . . . s. transvaalensis

Unfortunately these characters fail to separate our material in ac-
cordance with the supposed ranges. In the matter of temporals, not
only do lizards from the same locality answer to both, but the tempo-
rals on one side of the head of an individual may correspond to the
definition of subviridis, on the other to transvaalensis.

As for the degree of bluntness of the "temporal spine" i.e. lowest
anteauricular scale, its condition appears to be affected by age and
the degree of wear to which it has been subjected in rocky crevices.

There is, however, a substantial size difference of about 80 mm. in
total length as between the southern and extreme northern forms.
FitzSimons gives the following:

d1 255 (110 + 145) mm. from Giant's Castle, Natal.

cT 327 (151 + 176) mm. from Woodbush, northern Transvaal.
The seven specimens in the Museum of Comparative Zoology come
from localities indicated by an asterisk below.

Localities. Transvaal: Doornkop near Belfast; *Kastrol Nek
Farm, Wakkerstroom; Pretoria District; *Selati (Paratype of
transvaalensis); Zoutpansberg District. Swaziland: *Forbes Reef.
Natal: Balgowan; *Giant's Castle, Drakensberg; mountains be-
hind Kaffirland; Umvoti.

Range. Transvaal south through Swaziland to Natal, east to Orange
Free State and eastern Cape Province.

* In Museum of Comparative Zoology.

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PSEUDOCORDYLUS MICROLEPIDOTUS NAMAQUENSIS Hewitt

1903e. ? Pseudocordylus microlepidotus Boulenger (not Cuvier), p. 217.

1909a. Hewitt (part), p. 37 (Richmond District record only).

1927a. Pseudocordylus microlepidotus namaquensis Hewitt, Rec. Albany Mus.

3, p. 392, pi. xxii, fig. 1 : Namaqualand.
1935. Lawrence, p. 44.

Description. Rostral separated from the frontonasal; upper row of
temporals slightly vertically elongate ; gulars smooth, the median more
or less squarish or subcircular.

Dorsum covered with small, subequal, keeled, striate, nodular
scales, forming more or less regular longitudinal and transverse series,
those of the vertebral region smallest.

For characters common to all species, see definition on p. 66, for
scale and pore counts, see statistical table on p. 69.

Color. Pattern not readily distinguishable. Below, gular region
without infuscations.

Size. Length of type (S.A.M. 872) from snout to anus, 127 mm.

Remarks. Known to me only from the sketchy description and in-
distinct figure of the original citation, based on four specimens in the
South African Museum. To these I have tentatively added, on geo-
graphical grounds only, Boulenger's (1903e) Deelfontein, Richmond
District, record which was repeated by Hewitt (1909a) as microlepi-
dotus at a time when none of the races were recognized. Whether
namaquensis deserves recognition is uncertain though geographically
probable.

Localities. Cape Province: Beaufort West; Deelfontein, 25
miles w. of Victoria West1; Namaqualand (whether Little or Great
not known).

Range. Western Cape Province (west to "Namaqualand").

Pseudocordylus microlepidotus microlepidotus (Cuvier)

Plate 10, figs. 1-2

1735. Lacerta Africana elegantissima Seba, Rerum naturalium Thesauri

2, p. 62, pi. lxii, fig. 6: Africa.
1829. Cordylus microlepidotus Cuvier, Regne Animal, ed. 2, 2, p. 33:

(based on Seba's plate).
1829-44. Guerin, Icon. Regne Animal, 1, Rept., pi. vi, figs. 1-la.

1 See remarks.

loveridge: African lizards 79

1931b. Gray, p. 119.

1843. Smith, A., (part), pi. xxiv, fig. 1 (= A of text), pi. xxx, figs. 1-la:

Table Mountain near Cape Town, Cape Province.

1831b. Zonurus microlepidotus Gray, p. 63.

1834. Schlegel, p. 216 (pi. is captioned Wittii).

1839. Dumeril & Bibron, p. 361. (omit Sierra Leone as error).

1834. Zonurus wittii Schlegel, pi. vii, figs, la-lc (but text refers to

microlepidotus from) South Point of Africa.

1838a. Cordylus (Pseudocordylus) montanus A. Smith, Mag. Nat. Hist. (2),

2, p. 32: Cape of Good Hope.

1845. Pseudocordylus microlepidotus Gray, p. 49.

1884a. Rochebrune, p. 107 (omit Senegambia as error).

1885e. Boulenger (part), p. 259.

1898. Sclater, p. 104.

1898. Werner (1896-7), p. 140.

1910b. Boulenger (part), p. 469. (Cape Town only).

1910a. Hewitt, p. 71.

1925b. Flower, p. 945.

1926b. Rose, p. 492.

1927a. Hewitt, p. 390, pi. xxiii, fig. 3.

1929. Rose, pp. 102, 106, figs. 67-68.

1935. Lawrence, p. 44.

1931. Zonurus pseudocordylus (in error) Mann, pp. 390, 398.

1939. Popp, p. 263.

Name. Crag Lizard (English: Rose).

Description. Rostral separated from, or rarely in contact with, the
frontonasal, which is shorter or longer than broad; nostril pierced in
the lower centre or postero-inferior corner of a nasal ; median subocular
descending to the lip between the fourth and fifth upper labials ; post-
frontals as long as broad; a few median and posterior temporals
slightly vertically elongate; gulars smooth, the median more or less
slightly elongate like the lateral.

Dorsum covered with small, flat or raised subequal, subcircular,
smooth or obtusely keeled nodular scales, separated by minute gran-
ules, forming more or less regular longitudinal and transverse series,
those of the vertebral region smallest; scales on flanks subequal, not
decreasing in size appreciably, smooth or obtusely keeled; scales below
forelimbs smooth, keeled or granular, below hind limbs smooth or
granular.

For characters common to all species, see definition on p. 66, for
scale and pore counts, see statistical table on p. 69.

Color. Above, head brownish red, lips and sides of head pale brown-

80 bulletin: museum of comparative zoology

ish; back blackish or brownish red with series of ochre yellow spots
which tend to form 7-8 cross-bars descending to the flank; where they
may break up; sides of neck with one or two large blackish spots;
limbs barred with yellow; tail narrowly streaked with yellow. Below,
gular region black (Rose) or blue (Smith); belly dirty gray (Rose),
yellowish or orange (Smith) tinged with red. Eyes chestnut brown
(Smith).

Size. Total length 279 (221 + 58) mm. (Rose).

Remarks. I concur with Hewitt (1927a) that montanus was almost
certainly the form (figured by Seba) on which Cuvier based his micro-
lepidotus (without description).

Breeding. Four young are produced (Rose, fide Hewitt, 1937e).

Longevity. Two years, nine months, seven days (Flower).

Diet. This insectivorous lizard will eat even the spinous wingless
locusts known as korenkrekels and at times vary their diet by taking
lichen from the nearby rocks (Rose).

Parasites. Mites (Lrodidcrma inverta and Zonurobia semilunaris)
have been described from this form by Lawrence.

Enemies. One, when seized by a hawk, freed itself by discarding its
tail, but was killed by the fall to the rocks below (Rose).

Temperament. When first captured this lizard can, and will, inflict
a severe bite. In captivity it soon becomes docile, however, and will
even take food from the fingers (Rose).

Habitat. Confined to the steeper crags from whose deep clefts it
can be secured only by a noose at the end of a long wire (Rose).

Localities. Cape Province: Table Mountain and hills near
Cape Town.

Range. Cape Province (mountains of Cape Peninsula).

PSEUDOCORDYLUS MICROLEPIDOTUS FASCIATUS (Smith)

Plate 10, figs. 3-4
Plate 11, figs. 1-2

1838a. Cordylus (Pseudocordylus) fasciatus A. Smith, Mag. Nat. Hist. (2),

2, p. 32: Cape of Good Hope.
1843. Smith, A., pi. xxvii, fig. 1; pi. xxx, figs. 5-5b: Rocky Hills near

Grahamstown, Cape Province.
1838a. Cordylus (Pseudocordylus) Algoensis A. Smith, Mag. Nat. Hist. (2),

2, p. 32: Cape of Good Hope.
1843. Cordylus microlepidotus A. Smith, (not Cuvier) pi. xxiv, fig. 2; pi.

xxx, figs. 2-2b: Algoa Bay, Cape Province.

loveridge: African lizards 81

1927a. Pseudocordylus fasciatus Hewitt, p. 391.

1935. Lawrence, p. 44.

1937e. Hewitt, p. 31, pi. x, fig. 1.

1937a. Pseudocordylus microlepidotus fasciatus FitzSimons, p. 266.

Names. Crag Lizard or caiman (English); kaaiman, or sometimes
klip-lekkevan (Afrikaans); uroqotyeni (Hottentot; = always on the
stone). All taken from Hewitt (1937e).

Description. Rostral separated from, or rarely in contact with, the
frontonasal, which is as long as broad; nostril pierced in the lower
centre or postero-inferior corner of a nasal ; median subocular descend-
ing to the lip between the fourth and fifth upper labials ; postf rontals
as long as broad or longer than broad; temporals polygonal, not ver-
tically elongate ; gulars smooth, a few anterior ones irregularly enlarged,
the median more or less squarish.

Dorsum covered with small, raised, subcircular or subtriangular,
feebly keeled (young) or smooth (adult) nodular scales, separated by
minute granules, forming more or less regular longitudinal and trans-
verse series, those of the vertebral region smallest; scales on flanks
subequal, not decreasing in size appreciably, obtusely keeled; scales
below fore limbs smooth or striate, below hind limbs smooth.

For characters common to all species, see definition on p. 66, for
scale and pore counts, see statistical table on p. 69.

Color. Above, head browoi variegated with yellow, lip to ear yellow
or more or less tinged with red; back blackish brown, brown, or red-
dish brown, with 6-10 series of pale yellow spots1 or bands2; sides of
neck with one or two large blackish spots ; flanks yellow, orange yellow,
or brick red, without vertical bars, at most the dorsal bands encroach-
ing but slightly; tail irregularly spotted with yellow. Below, yellowish
tinged with reddish.

Size. Total length of d" (M.C.Z. 33449), 304+ (144 4-160+)
mm., tail-tip missing.

Remarks. FitzSimons (1937a) was unable to locate the type. It is
possible that algoensis (including Matschie, 1891a) may prove to be
distinct and have to be removed from the synonymy.

Anatomy. Hewitt (1920) discusses the so-called branchial arch.

Diet. Beetles, crickets, grasshoppers, wood-lice, snails, and even
small lizards (Essex in Hewitt, 1937e).

Parasites. Mites (Ixodiderma inverta 'and Zonurobia semilunaris)
were found on this form by Lawrence.

1 Only 1 or 2 orange spots on Abbotsbury males, whose females are more or less cross-barred.

2 Bands complete in Butterworth lizards (Hewitt).

82 bulletin: museum of comparative zoology

Temperament. An active but shy lizard which keeps close to its
rocky retreat, basking with forepart upraised, the head and chin held
high.

Habitat. Occurs at 5600 feet at Abbotsbury among the rocks on
mountain slopes (Hewitt).

Localities. Cape Province: Abbotsbury near Lett's Kraal;
Algoa Bayx(?); Butterworth; Coetzee's Berg near Pearston; Dor-
drecht; Grahamstown — rocky hills in vicinity; Schurfteberg,
Somerset East District; Tsomo.

Range. Cape Province (eastern part).

Genus Platysaurus

1844. Platysaurus A. Smith, 111. Zool. S. Africa, Rept., footnote to pi. xi
(type capensis Smith).

Head and body depressed; limbs well developed; tail moderate.
Head shields regular; nostril pierced in the nasal2; eyelids well de-
veloped, the lower with a transparent disk; ear -opening large; 4
parietals; a collar fold, attached mesially, at least indicated; dorsals
granular; sides of neck covered with granules; ventrals moderate,
quadrangular, juxtaposed, imbricate, forming longitudinal and trans-
verse series; femoral pores present in both sexes though sometimes
scarcely distinguishable in females; digits slightly keeled inferiorly;
tail with or without spines.

The seven forms which I have examined appear to have the follow-
ing characteristics in common, consequently these have been omitted
from the specific descriptions.

Head much longer than broad ; head shields more or less smooth or
slightly roughened posteriorly; rostral once and a half to twice and a
half as broad as high; nasal not or scarcely swollen; a postnasal; a
loreal; a preocular; 4 suboculars of which one or two descend to the
lip between the labials; 4 supraoculars, the anterior longest, the
second broadest; prefrontals forming a suture; frontal hexagonal,
broader anteriorly; posterior parietals much larger than the anterior;
mental rather large ; lower labials bordered by a row of 5 large shields ;
collar scales enlarged but otherwise undifferentiated.

Dorsal granules small, flat or but slightly convex; median preanals
more or less enlarged; limbs below with smooth (some keeled in

1 Unless algoensis proves to be a recognizable form. Abbotsbury lizards are not quite typical
according to Hewitt (1927a).

s Nol between "nasal and first labial" as stated by Boulenger, the nostril being actually
separated from labial and postnasal by a narrow rim.

loveridge: African lizards 83

g. guttatus and g. rhodesianus) scales which on tibia form a series of
from 6-10 large plates; claws of the adpressed hind limb reach to
axilla or shoulder, rarely neck, in guttatus and its races, to neck (9 )
or temporal region (cf) in capensis, there being a tendency to shorter
limb length in females; tail with whorls of elongate, quadrangular,
rugose or keeled scales, the lateral with strongest keels, the subcaudals
smooth.

Range. South Africa, on east north to the Zambezi, i.e. south of
16° S.

Remarks. An examination of the Statistical Synopsis of Variation
will reveal the homogeneity of this group, rendering the compilation of
a Key extremely difficult. Even so I imagine that when large rseries
of certain forms are available apparent differences in scale-counts will
overlap still more. I should like to see a Key based solely on the strik-
ing color differences of these various montane forms.

Key to the Species

1. A large occipital wedged between the posterior parietals;
sides of neck covered with small and enlarged granules;
granules on flanks distinctly enlarged ; scales covering limbs
above mostly rugose and strongly keeled; adpressed hind
limb reaches axilla or shoulder (except in guttatus fitz-
simonsi where it attains neck); range: southeastern Africa 2

No occipital, at most 4-6 small scales mesially behind
parietals; sides of neck covered with small subuniform
granules; granules on flanks minute or but slightly en-
larged; scales covering limbs above smooth or feebly
keeled; adpressed hind limb reaches neck or temporal "
region; range southwestern South Africa 8

2. Adult length from snout to anus more than 90 mm 3

Adult length from snout to anus less than 80 mm 5

3. Rostral usually in contact with frontonasal; inter-parietal
usually separated from occipital; collar scales 6-7; ventral
plates in 16-20 longitudinal and 39-45 transverse rows
{from collar); length from snout to anus of adults usually

less than 100 mm. ; range: northern Transvaal \ . . . g. guttatus

(p. 89)
Rostral usually separated from frontonasal; inter-parietal
usually in contact with occipital; collar scales 7-12; ventral
plates in 18-26 longitudinal and 40-48 transverse rows
(from collar) 4

84 bulletin: museum of comparative zoology

4 Supraciliaries 4 ; enlarged temporals in 2 rows, upper largest ;
chin and throat very light blue without markings, belly and
limbs below not uniformly dark blue; adult length from
snout to anus over 100 mm.; range: Southern Rhodesia
and Bechuanaland Protectorate g. rhodesianus

(p. 86)
Supraciliaries 5; enlarged temporals in 3 rows, median
largest; chin and throat to chest of adult male entirely black
spotted with pale blue, belly and limbs below uniformly
dark blue; adult length from snout to anus 91 mm.; range:
Lydenburg, eastern Transvaal g. fitzsimonsi

(p. 88)

5. Some dorsals in the vertebral series very slightly enlarged;

lower surface in both sexes dark 6

Dorsals subuniform; lower surface in both sexes light but
males with dark patches on belly 7

6. Interparietal forming a suture with a small occipital which
may be broken up; spines on tibia towards heel poorly de-
veloped, on lateral caudals not at all; chin and throat of
male gray green with black markings; edges of light lines
on dorsum of female sharply defined; range: Tete,
Mozambique g. torquatus

(p. 91)
Interparietal forming a suture with an occipital as large as
itself; spines prominent both on tibia towards heel and on
lateral caudals; chin and throat of male black; edges of light
lines on dorsum of female blurred; range: Barberton, Trans-
vaal and Ubombo, Zuhiland g. wilhelmi

(p. 92)

7. Males above, olive brown to reddish brown with light spots
on dorsum arranged in longitudinal lines; in females such
light spots are arranged in longitudinal lines also, but no
black spots present on abdomen; flanks buff; range: Water-
berg, Transvaal g- minor

(p. 93)
Males above, dull green to bluish green with light spots on
dorsum scattered; in females such light spots are absent or
only present posteriorly; but irregular black spots are
present on abdomen; flanks dull green to bluish green;

range : eastern face of Drakensberg, Transvaal g. orientalis

(p. 94)

8. Range: Little Namaqualand and Victoria West, Cape
Province north to South West Africa capensis

(p. 96)

loveridge: African lizards

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Species

race

g. rhodcsianus
g. fitzsimonsi
g. guttatus
g. torquatus
g. wilhebni
g. minor
g. orientalis
capensis

86 bulletin: museum of comparative zoology

Platysaurus guttatus rhodesianus FitzSimons

1902b. Platysaurus guttatus Boulenger (not A. Smith), p. 16.
1909a. Chubb, p. 593.
1909b. Chubb, p. 35.
1909a. Hewitt (part), pp. 29, 38.
1913. Hewitt & Power, p. 154.

1930. FitzSimons (part), p. 31.

1932. Hewitt (part), p. 118.

1935b. FitzSimons, p. 350.
1939b. FitzSimons, p. 31.

1910b. Platysaurus capensis Boulenger (part), p. 469.
1934. Pitman, p. 304.

1941. Platysaurus guttatus rhodesianus FitzSimons, Ann. Transvaal Mus.,
20, p. 279: Vumba Mountain, Southern Rhodesia.

Description. Rostral sometimes in contact with, though more
usually separated from, the frontonasal, sometimes by an azygous
scale; frontonasal as long as broad or shorter than broad; second or
second and third subocular descending to the lip; interparietal large,
pentagonal, rarely diamond-shaped, rarely enclosed between 2 pairs
of parietals, usually forming a suture with an occipital; enlarged tem-
porals in 2 longitudinal rows, upper largest; sides of neck covered with
granules, both small and enlarged; gulars small, elongate, those on
the median line more or less enlarged and squarish; granules on
flanks enlarged; limbs above with granules and rugose, strongly
keeled scales.

For characters common to all species, see definition on p. 82, for
scale and pore counts, see statistical table on p. 85.

Color, c? . Above, head and back dull bluish green with three light
longitudinal lines usually discernible on head, indicated or disappear-
ing on back in old males, which may be almost uniform, lacking the
light spotting displayed on back and limbs of young; flanks terra cotta
in life; base of tail dull reddish yellow or yellowish green passing dis-
tally to green. Below, chin and gular region light blue (yellow to
greenish yellow in young) ; a black gular collar sometimes extending to
chest (much reduced and narrower in young); belly anteriorly terra
cotta (in life) (blue in young), posteriorly dark grayish blue to black;
limbs blue to dark blue; tail tangerine (in life) or straw color (in
alcohol), passing distally to yellow, yellowish green, and finally green
(yellow at base passing to green in young).

In life, according to FitzSimons, from whom this description is

loveridge: African lizards 87

largely adapted, very young males, though otherwise marked much
like the females, display a terra cotta colored patch on the chest.

9 . Above, head and back dark brown or black with three sharply
defined white to yellowish longitudinal lines, between which are rarely
(Matopos), more usually are not, pale spots; flanks and limbs with
pale spots; tail whitish with a dusky median line anteriorly. Below,
bluish white to bluish gray, chin and gular region sometimes exhibiting
a faint bluish tinge; belly anteriorly sometimes pinkish, middle of
belly usually with a large bluish black patch or spotted with black;
base of tail sometimes pinkish, otherwise straw to whitish.

Size. Total length of type c? (T.M. 18528), 272 (105 + 167) mm.,
and paratype 9 (T.M. 18663), 241 (92 + 149) mm.

Remarks. FitzSimons (1935b) shows that in 8 of his 14, and (1939b)
in 9 of his 31 specimens (which later became the type series), the
rostral is in contact with the frontonasal, while in 4 out of 14 and 9
out of 31 the interparietal is separated from the occipital. Hewitt
(1909a) found that 1 out of 10 lizards displayed an interparietal form-
ing a wide suture with the occipital.

Parasites. Mites (Zonurobia circularis typicus) described by Law-
rence from Salisbury "guttatus".

Habitat. On the granite hills about Zimbabwe this lizard is found
in association with Agama kirkii, Mabuya q. margaritifer, and Gerrho-
saurus v. validus, elsewhere, though found in association with mar-
garitifer in the bush veld, the latter was not observed on the lower
slopes of Vumba Mountain where P. g. rhodesianus was encountered,
and from where the Museum of Comparative Zoology has a good
series of topotypes collected by Mr. V. FitzSimons at the same time
as the type.

Localities. Southern Rhodesia: Bikita; Bindura; Changadzi
River; Chilimansi; Chishagwasha; Devuli River; Driefontein;
Empandeni; Gwanda; Hunyani River; Importuni District; Insiza;
Khami River; Livingstone; Lomagundi District; Matabeleland;
Matopos Hills; Mazoe; Mtoko; Mount Silozi; Penahlonga; Plum-
tree; Salisbury District; Strathmore near Bulawayo; Vumba
Mountain; World's View; Zimbabwe. Bechuanaland Protecto-
rate: Tsessebee.

Range. Southern Rhodesia extending into northeast Bechuanaland
Protectorate. (As yet there is no evidence of the occurrence of any
Platysaurus in Northern Rhodesia so that Pitman's (1934) suggestion
that capensis may occur there, lacks foundation, for the genus appears
to be found south of the Zambezi only).

88 bulletin: museum of comparative zoology

Platysaurus guttatus fitzsimonsi subspec. nov.

Type. Museum of Comparative Zoology, No. 8982, an adult cf
from Lydenburg, Transvaal, received in exchange from the British
Museum, 1913.

Diagnosis. As given in preceding key to the races of guttatus.
To which one might add that from g. wilhelmi, the only other member
of the genus whose male has the whole undersurface either black or
blue, fitzsimonsi differs in the pale blue spotting of its chin and jaws,
and the very dark blue (instead of light blue) undersurface of body
and limbs. Also in its three rows of temporals; smooth (not strongly
keeled) plates on the upper anterior surface of hind limb; subuniform
character of dorsal granules; more numerous transverse rows of ven-
trals ; larger size, etc. etc.

Description. Head much longer than broad; rostral twice as broad
as high, separated from the frontonasal, which is shorter than broad
nostril pierced in the postero-inferior corner of the nasal ; a postnasal
a loreal; a preocular; 4 suboculars, the second descending to the lip
4 upper labials anterior to subocular; 4 supraoculars, the anterior
longest, the second broadest; 5 supraciliaries; prefrontals forming a
suture; frontal hexagonal, broader anteriorly; a pair of postf rontals ;
interparietal large, pentagonal, forming a suture with an occipital;
posterior parietals much larger than the anterior; enlarged temporals
in three longitudinal rows, the median largest; sides of neck covered
with granules, both small and enlarged; mental rather large; 4-5 lower
labials, bordered by a row of 5 large shields; gulars small, elongate,
those on the median line slightly enlarged and squarish; collar scales
10, the ill-defined collar fold attached mesially.

Dorsum covered with small, flat, granular scales, subequal; granules
on flank enlarged; ventrals moderate, squarish or slightly broader
than long, smooth, juxtaposed, forming 20 longitudinal and 45 trans-
verse rows (from collar); median preanals enlarged; limbs above with
granules and rugose, strongly keeled scales, limbs below with enlarged
smooth scales which on tibia form a series of 7 large plates; claws of
adpressed hind limb reach to neck; 20-21 femoral pores; tail with
whorls of elongate, quadrangular, rugose or keeled scales, keels on the
lateral strongest, only some of the subcaudals smooth.

Color, c? Above, head dark blue (? green in life) with three light
longitudinal lines; back pale blue (? green in life) uniform or with a
few faintly discernible light spots; limbs pale blue, the hind limbs

loveridge: African lizards

brown posteriorly; tail straw yellow (? red in life). Below, chin and
gular region deep black flecked with light blue, merging with black of
collar; belly and limbs uniformly bright ultramarine blue; tail straw
yellow.

Size. Total length of type cf (M.C.Z. 8982), 226 (91 + 135) mm.

Remarks. This appears to be the most southern form of the larger
races of guttatus. The only possible reference to it in the literature may
be that of Hewitt (part, 1932, p. 119) where he writes of guttatus
ranging as far south as White River, Barberton District.

Localities. Transvaal: Lydenburg (possibly also White River)

Range. Eastern Transvaal (Lydenburg District).

Platysaurus guttatus guttatus Smith

1849. Platysaurus guttatus A. Smith, 111. Zool. S. Africa, Rept. App., p. 8:

Limpopo River near the Tropic of Capricorn.
1882a. Peters, p. 52.
1885e. Boulenger, p. 262.
1898. Sclater, p. 104.

1907j. Boulenger, p. 484.
1909a. Hewitt (part), pp. 29, 38.
1910. Hewitt, pi. i, fig. 1 (for 1909a).

1911b. Hewitt, p. 48.
1930. FitzSimons (part), p. 31.

1935. Broom, p. 20, fig. 5c.

1935. Lawrence, p. 44.

1937a. FitzSimons, p. 266.
1938. Gorham & Ivy, p. 179.

1891a. Platysaurus intermedins Matschie, Zool. Jahrb. Syst., 6, p. 606:

Mphome, Zoutpansberg District, Transvaal.
1911c. Sternfeld, p. 419 (Mpoma).
1910b. Platysaurus capensis Boulenger (part, not Smith), p. 469 (Pietersburg

only.

Further citations of "guttatus" will be found under its various races
and capensis.

Description. Rostral usually in contact with, though sometimes
separated from, the frontonasal, which is shorter than broad ; second
and third subocular descending to the lip; interparietal large, diamond-
shaped, rarely pentagonal, enclosed between 2 pairs of parietals, rarely
forming a suture with an occipital, which may be transversely divided;
enlarged temporals in 2 longitudinal rows, upper largest; sides of neck

90 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

covered with granules, both small and enlarged; gulars small, elongate,
those on the median line much enlarged, squarish or polygonal;
granules on flanks enlarged; limbs above with granules and rugose,
strongly keeled scales.

For characters common to all species, see definition on p. 82, for
scale and pore counts, see statistical table on p. 85.

Color, d*. Above, head brown; back bluish green to pale brownish
red, with or without three light longitudinal lines, between which are
irregular longitudinal series of light spots; flanks like back with faint
spotting; base of tail pale orange or yellowish with or without a dusky
median line. Below, chin and gular region light blue with or without
blackish, or deep purplish brown, markings; a black gular collar; belly
anteriorly pale blue, posteriorly dark blue; hind limbs blue though
posteriorly like tail, straw yellow.

9 . Above, head and back olive brown with three rather poorly
defined light longitudinal lines, between which are two longitudinal
series of pale spots ; flanks and limbs with pale spots ; tail straw yellow,
anteriorly with a dusky median line. Below, bluish white to blue;
chin and gular region tinged with blue and marked with black; belly
anteriorly pale blue, middle of belly usually with a large dark bluish
black patch; tail straw yellow.

Size. Total length of c? (M.C.Z. 21231), 217 (84 + 133) mm., ex-
ceeded in length from snout to anus by one of 105 mm. from the same
locality— Woodbush; length of type 9 (Brit. Mus.), 165 (71 + 94)
mm., exceeded in length from snout to anus by one of 87 mm. (Bou-
lenger, 1907J).

Anatomy. The temporal region of, the skull is discussed by Broom,
the gall bladder by Gorham and Ivy.

Parasites. Mites (Zomirobia circularis typicus) recorded from this
race by Lawrence.

Localities. Transvaal Blyde River; D wars River; Gerlachshoop,
Elands River; Gravelotte; Haenertsburg; Jachtsdrift; Koedoes
River; Letaba Drift; Leydsdorp; Limpopo River near Tropic of
Capricorn; Mokeetsi; Mphome or Mpoma; Nwanedsi River;
Pietersburg; Shilowane; Woodbush; Zoutpansberg. (Hewitt's
(1932) record of White River, Barberton District, is tentatively re-
moved to the race fitzsimonsi His (1909a) record from Victoria
West, Cape Province was based on a capensis Mr. FitzSimons
informs me.

Range. Northern Transvaal (Zoutpansberg region).

loveridge: African lizards 91

Platysaurus guttatus torquatus Peters
Plate 11, fig. 3

1854. Platysaurus capensis Peters (not A. Smith), p. 616.

1855. Peters, p. 45.

1854. Platysaurus guttatus Peters (not A. Smith), p. 616.

1855. Peters, p. 45.

1879a. Platysaurus torquatus Peters, Sitz. Ges. naturf. Freunde Berlin, p.

10: Tete, Mozambique.
1882a. Peters, p. 52, pi. ixa.
1885e. Boulenger, p. 262.
1896a. Bocage, p. 98.

1907j. Boulenger, p. 484 (comments on). .
1909a. Hewitt, pp. 31, 38.
1910a, Hewitt, p. 60.

Native name. Biinio (at Tete: Peters).

Description. Rostral in contact with, or separated from, the fronto-
nasal, which is shorter than broad; second and third subocular de-
scending to the lip; interparietal large, diamond-shaped, forming a
suture with an occipital which may be broken up; enlarged temporals
in 2 longitudinal rows, upper largest; sides of neck covered with gran-
ules both small and enlarged; gulars small, elongate, those on the
median line slightly enlarged and squarish; granules on flanks en-
larged; limbs above with granules and rugose, strongly keeled scales.

For characters common to all species, see definition on p. 82, for
scale and pore counts, see statistical table on p. 85.

Color, c?. Above, head and back black with three light (golden
yellow in life) longitudinal lines on head, the lateral disappearing on
back, flanking the vertebral line is a regular longitudinal series of
light spots; flanks malachite green spotted with lighter; limbs dark
brown spotted with rusty yellow; tail pale orange yellow (red in life).
Below, chin and gular region gray green with black markings ; a black
gular collar; tail yellow (red in life).

9 . Above, head and back brownish black with three sharply de-
fined, light, longitudinal lines between which are a longitudinal series
of light spots; flanks grayish or brownish green; tail greenish gray
(not red) with a median dusky line anteriorly. Below, chin and gular
region bluish green; (?) a black gular collar; belly gray ; tail gray brown.

Size. Total length of cotype 'd> (Berlin Mus.), 185 (72 + 113) mm.,
of cotype 9 (M.C.Z. 37203), 161 (68 + 93) mm.

92 bulletin: museum of comparative zoology

Remarks. It is just as well that Peters (1882a) described this lizard
in great detail for no other specimens of it have been taken. He points
out that in 1854 he referred the males to guttatus, the females to
capensis: twenty-five years later he used the same material as types
of torquatus.

Boulenger (1885e), doubtless misled by Peter's figure for he had no
material, erred in saying that the gulars on the median line were not
enlarged for Peters definitely says that two rows are enlarged (two,
or possibly three in our cotype), though his figured specimen shows
but slight enlargement and that only in the basal part. Hewitt (1909a)
is equally mistaken in suggesting that torquatus can be separated from
capensis on the grounds that the former has two and the latter only
one median row enlarged, for in most of the forms much variation is
displayed in this character.

Boulenger (1907j) points out that guttatus has from 70-90 dorsal
granules in longitudinal series while torquatus has from 90-100. This
is a character worthy of study in all the forms and one to which I have
paid no attention.

Anatomy. Utilising C. c. cordylus for comparative purposes, Peters
(1882a) discusses the viscera, skeleton, and skull of torquatus, stating
that the latter has 20 teeth in the upper, 21 in the lower jaw; that the
pupil is round, and that no lateral fold is present in life.

Habits. He claims that it is difficult to capture owing to the agility
with which this lizard seeks shelter in the rock crevices, from which it
can be taken only by removal of the rocks.

Habitat. It is to be found on the carbonaceous sandstone rocks
between the Caruera Mountains and Tete, also a few miles north and
southwest of Tete.

Localities. Mozambique: Tete, south bank of the Zambezi.

Range. Mozambique.

Platysaurus guttatus wilhelmi Hewitt

1908b. Platysaurus guttatus Boulenger (not A. Smith), p. 224.

1909a. Platysaurus wilhelmi Hewitt, Ann. Transvaal Mus., 2, p. 29, Nelspruit,

Barberton District, Transvaal.
1910. Hewitt, pi. i, fig.*2 (for 1909a).

1911b. Hewitt, p. 48.
1930. FitzSimons (part), p. 32.

1932. Hewitt (part), p. 119.

1935. Lawrence, p. 44.

Further citations of "wilhelmi" will be found under g. orientalis.

loveridge: African lizards 93

Description. Rostral in contact with the frontonasal, which is as
long as broad or shorter than broad; second subocular descending to
the lip; interparietal large, pentagonal, forming a suture with an
occipital; enlarged temporals in 2 longitudinal rows, upper largest;
sides of neck covered with granules, both small and enlarged; gulars
small, elongate, those on the median line much enlarged and squarish;
granules on flanks enlarged; limbs above with granules and rugose,
strongly keeled scales.

For characters common to all species, see definition on p. 82, for
scale and pore counts, see statistical table on p. 85.

Color, c?. Above, head brown with three light longitudinal lines;
back dull greenish with ill-defined light spots; flanks greenish, uni-
form; limbs dark variegated with lighter; tail straw yellow (red in
life). Below, chin to chest deep black; belly ultramarine with some
black mesially; limbs ultramarine; tail straw yellow (red in life).

9 . Above, head and back brownish black with three rather poorly
defined light longitudinal lines, between which are a single, or posteri-
orly double, series of light spots; flanks and limbs with pale spots; tail
terra cotta with a dusky median line anteriorly. Below, black or
blackish gray; tail gray brown (pinkish in life).

Size. Total length of cotype cf (T.M. ), 194 (82 + 112) mm.,

and 9 (T.M. ), 184 (66 + 118) mm.

Remarks. Hewitt (1909a) differentiated the adult male wilhelmi
from guttatus by reason of its alleged broader head and heterogeneous
dorsal granules which, in the female also, show some enlargement along
the vertebral and dorsolateral lines. The character of the occipital
scale he (1911b) later dropped from the diagnosis. It is more likely to
be confused with P. g . fitzsimonsi under which race the points of differ-
ence are enumerated.

Parasites. Mites (Zonurobia circularis transvaalensis) are recorded
by Lawrence.

Localities. Transvaal: Nelspruit; Queen's River (both in Bar-
berton district). Zululand: Ubombo.

Range. Eastern Transvaal south to Zululand.

Platysaurus guttatus minor FitzSimons

1930. Platysaurus guttatus minor FitzSimons, Ann. Transvaal Mus., 14,
pp. 30, 31, fig. 10: Vygeboompoort, Waterberg District, northern
Transvaal.

1935. Lawrence, p. 44.

Description. Rostral separated from the frontonasal, which is

shorter than broad; second or second and third subocular descending

94 bulletin: museum of comparative zoology

to the lip; interparietal large, diamond-shaped or pentagonal, some-
times separating the anterior parietals, sometimes anteriorly in con-
tact with a small azygous scale, or posteriorly forming a suture with
an occipital; enlarged temporals in 2 longitudinal rows, upper largest;
sides of neck covered with granules, both small and enlarged; gulars
small, elongate, those on the median line enlarged, squarish or irregu-
lar ; granules on flanks enlarged ; limbs above with granules and rugose,
strongly keeled scales.

For characters common to all species, see definition on p. 82, for
scale and pore counts, see statistical table on p. 85.

Color, cf. Above, head and back olive green with three light longi-
tudinal lines, indistinct on head and tending to break up or disappear
on back, between the lines are longitudinal series of light spots ; flanks
buff; tail terra cotta. Below, chin to collar vivid blue; belly dark
Prussian blue extending on to base of tail, rest of tail salmon buff.

9 . Above, head and back dark olive brown with three sharply de-
fined light longitudinal lines, between which are one or two longi-
tudinal series of light spots; flanks and limbs with pale spots; tail terra
cotta with a broad, dusky, median line anteriorly. Below, chin, gular
region, chest and belly whitish (pale blue in life), uniform; tail gray
brown.

Size. Total length of cotype cT (TM. 2109), 158 (65 + 93) mm.
of cotype 9 (T.M. 2367), 146 (61 + 85) mm.

Remarks. Of 135 specimens examined by FitzSimons not one ex-
ceeded 160 mm. in total length, the race was based on 73 specimens of
which four are now M.C.Z. 41882-5. Of these 73 apparently about 44
possessed anterior parietals which were separated by the interparietal.
The form is a perfectly good montane race though no great reliance
should be placed on the scale characters originally cited by Fitz-
Simons, for most will be found to vary widely within certain limits.

Parasites. Mites (Zonurobia circularis spiniventer) were described
from this form by Lawrence.

Localities. Transvaal: Geelhoutkop; Modder Nek; Nylstroom;
Palala River; Vgeboompoort.

Range. Transvaal (Waterberg District).

Platysaurus guttatus orientalis FitzSimons

1930. Platysaurus wilhehni FitzSimons (part, not Hewitt), p. 32.

1932. Hewitt (part, not Hewitt of 1909a), p. 119.

1941. Platysaurus minor orientalis FitzSimons, Ann. Transvaal Mus., 20,

p. 280: Sekororo on lower slopes of Drakensberg about 40 miles

from Leydsdorp, Transvaal.

loveridge: African lizards 95

Description. Known to me only from the original description, which
was based on three males and a female, and took the form of con-
trasting with minor. These differences are given without comment.

Snout more sharply pointed; head more swollen in temporal region
and in general a little larger in proportion to the body; granules in
temporal region in 2-3 longitudinal rows (as against 3-4 in minor);
gular scales across throat between last large sublabial on either side
19-22 (as against 16-18 in minor); ventrals in 20-22 longitudinal (as
against 16-18 in minor) and 381 transverse (as against 34-381 in minor)
rows; femoral pores 14-20 (as against 14-17 in minor) preceded by
modified glandular scales arranged in 2-3 rows (as against a single row
in minor) ; scales on forearm and tibia more strongly keeled and spinose,
especially on heel where the spines are long and strongly pointed;
upper caudals obtusely keeled (more or less smooth in minor), lateral
caudals strongly keeled and sharply spinose (moderately keeled and
not or but bluntly spinose in minor).

For characters common to all other races which I have examined,
see definition on p. 82, for scale and pore counts, see statistical table
on p. 85.

Color c? . Above, head and back bluish green to dull green with
three light longitudinal lines on head only; back with small irregularly
scattered light spots; flanks dull green to bluish green. Below, chin
and gular region blue passing to dark blue or bluish black on belly.

9 . Above, head and back very dark brown or black with three
sharply defined light longitudinal lines, between which are no light
spots except for a few posteriorly. Below, bluish white with scattered
irregular black spots.

Size. Total length of type c? (T.M. 4527), 180 (65 + 115) mm.

Localities. Transvaal : Dientje Farm, near Vaalhoek, Pilgrim's
Rest District; Perkoe Farm, near Olif ant's River; Sekororo, 40
miles south of Leydsdorp.

Range. Transvaal (eastern slopes of Drakensberg).

1 Counting from collar to within three or four rows of preanals, in contrast to method em-
ployed by FitzSimons.

96 bulletin: museum of comparative zoology

Platysaurus capensis Smith

1844. Platysaurus capensis A. Smith, 111. Zool. S. Africa, Rept. pi. xl:
Great Namaqualand, South West Africa.

1845. Gray, p. 49.
1885e. Boulenger, p. 261.
1898. Sclater, p. 104.
1909a. Hewitt, pp. 31, 38.
1910b. Boulenger (part), p. 469.
1911b. Hewitt, p. 48.

1911b. Sternfeld, p. 403.

191 Id. Sternfeld, p. 23.

1932. Hewitt, p. 118.

1935a. FitzSimons, p. 534.

1935. Lawrence, p. 44.

1937a. FitzSimons, p. 266.

1938. FitzSimons, p. 193.

1909a. Platysaurus guttatus Hewitt (part, not Smith), pp. 31, 38 (Victoria
West record).

Further citations of "capensis" will be found under g. rhodesianus,
g. guttatus and g. torquatus.

Description. Rostral rarely in contact with, usually separated from,
the frontonasal, sometimes by an azygous scale; frontonasal as long
as broad, sometimes with an azygous scale posteriorly; second or
second and third subocular descending to the lip; sometimes a pre-
frontal also descending to the lip between loreal and preocular; inter-
parietal large, diamond-shaped, enclosed between 2 pairs of parietals;
occipitals broken up or absent; enlarged temporals in 2 longitudinal
rows, upper largest; sides of neck covered with small granules only;
gulars small, elongate, those on the median line more or less enlarged
and squarish; granules on flanks minute or but slightly enlarged;
limbs above with granules and smooth or feebly keeled scales.

For characters common to all species, see definition on p. 82, for
scale and pore counts, see statistical table on p. 85.

Color, cf- Above, head and back bright green anteriorly passing to
dull orange on posterior third, with or without three light longitudinal
lines on head which tend to disappear on back; between the lines some
ill-defined light spots may be present or absent; flanks greenish blue
and orange; tail yellow ringed with brown, a dusky median line an-
teriorly. Below, chin and gular region bright blue; belly anteriorly
dark blue, in middle black, posteriorly whitish; limbs and tail pale
straw yellow (grenadine in life), the latter ringed with darker.

loveridge: African lizards 97

9 . Above, head and back very dark brown' with three well-defined
light longitudinal lines, between which there are no light spots; flanks
and limbs with obsolete pale spots ; tail yellow alternately ringed with
pale brown, a dusky median line anteriorly. Below, whitish, belly
with or without pale blue and pinkish suffusions and a small black
patch; tail straw yellow ringed with gray brown.

The above descriptions are adapted largely from FitzSimons (1935a),
for more details of color variation see the same author for 1938.

Size. Total length of tf (T.M. 15883), 207 (78 + 129) mm., of
type 9 (Brit. Mus.), 197 (77 + 120) mm.

Remarks. FitzSimons (1937a) has reexamined and confirmed the
identity of the 9 holotype in the British Museum. Boulenger (1910b)
was in error in synonymizing guttatus with this species, a fact soon
pointed out by Hewitt (1911b) who, in 1932, listed many distinguish-
ing characters though some have not stood the test of larger series.
Peters (1882a) states that the dorsal granules are in 80-84 rows.

Parasites. Mites (Zonurobia circularis capensis) were described from
this species by Lawrence.

Habits. These extremely active lizards dart up vertical rock sur-
faces with the greatest ease (FitzSimons) to seek shelter beneath flat
stones or in crevices from which they are difficult to dislodge on account
of the use they make of their strong claws (Smith).

Habitat. In Great Namaqualand found always on rocky precipices
(Smith), but in Little Namaqualand they appear to be confined to
the rock and boulder-strewn river beds where great numbers may be
seen in favoured localities (FitzSimons).

Localities. Cape Province: Aughrabies Falls, Orange River;
Garies to Kamiesberg; Goodhouse; Kamieskroon; Kuboos; Lek-
kersing; Victoria West (Sclater, 1898, and Hewitt (1909a) but as
gidtatus.). South West Africa: Churutabis; Great Namaqualand.

Range. Cape Province (Victoria West and Little Namaqualand) to
South West Africa.

98 bulletin: museum of comparative zoology

Genus Chamaesaura

1801. Chamaesaura1 Schneider (part), Hist. Amphib., 2, pp. 205, 210 (type
restricted to anguina Linne).

1820. Monodactylus Merrem, Versuch Syst. Amphib., p. 76 (type anguina
Linn6).

1826. Chamaesaura Fitzinger, Neue Class. Rept., pp. 18, 59 (type anguina
Linne).

1832. Cricochalcis Wiegmann, Handbuch der Zool., p. 185 (no type desig-
nated but aena described though not named).

1862. Mancus Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 339 (type
macrolepis Cope).

For further synonymy see Boulenger, 1885e, Cat. Lizards Brit.
Mus., 2, p. 263.

Head elongate; body serpentiform ; limbs short, rudimentary, or fore
limbs absent; tail extremely long. Head shields regular, strongly
striated; nostril pierced in the postero-inferior corner of the nasal;
eyelids well developed, the lower scaly; ear-opening moderate; 4
parietals; no collar fold; dorsals larger than ventrals, otherwise both
uniform, lanceolate, strongly keeled, strongly imbricate, forming longi-
tudinal and transverse series; femoral pores present in both sexes
though sometimes scarcely distinguishable in females; tail without
spines.

The five known forms, all of which I have examined, have the fol-
lowing characteristics in common, consequently these have been
omitted from the specific descriptions.

Rostral 3-42 times as broad as high; a large subocular bordering the
lip with 3 (rarely 2 or 4) labials anterior to it; 3 supraoculars, the
anterior longest, the second broadest; 3 supraciliaries, the anterior
longest; frontal hexagonal or heptagonal; a pair of postfrontals which
are subequal to the 4 parietals; temporals covered by imbricate,
keeled scales; 4 (rarely 3 or 5) lower labials, bordered by a row of 4
(rarely 5 in anguina) chin shields.

Range. South, East, and Central Africa to northern Uganda, i.e.
to 4° N.

1 Sherborn, 1925, Index Animalium, p. 1209, is wrong in citing Chamaesaurus, and Fitz-
Simons, 1943, p. 409, in giving the date as 1799.

2 Five for aenea according to Boulenger (1885e, p. 263), seems improbable and does not agree
with M.C.Z. material.

loveridge: African lizards 99

Remarks. An extraordinary homogeneous group except for the
stages in limb reduction from the four-limbed, pentadactyle aenea to
the two-limbed, monodactyle macrolepis, for which Cope proposed
his genus Mancus solely on the absence of fore limbs.

Key to the Species

1. Midbody scale-rows 28; dorsals in 46 transverse rows from
parietals to above anus; prefrontals forming a suture; fore
and hind limbs pentadactyle; range: Cape Province north to
Orange Free State and Transvaal aenea

(p. 101)
Midbody scale-rows 26 or less; dorsals in 38-40 transverse
rows from parietals to above anus; prefrontals separated by
frontonasal forming a suture with frontal; fore and hind
limbs, if present, monodactyle or didactyle 2

2. Fore limbs, though rudimentary and scale-like, divided or

undivided 3

Fore limbs minute or absent 4

3. Fore limb longer, its length being contained from 1 to \\i
times in the distance from end of snout to anterior corner of

orbit; range: eastern Cape Province, Natal and Zululand a. anguina

(p. 102)
Fore limb shorter, its length being contained from 1 K to 1 %
times in the distance from end of snout to anterior corner of
orbit; range: eastern Belgian Congo and western Tanganyika

north to Kenya and Uganda a. tenuior

(p. 105)

4. Midbody scale-rows 24-26; fore limb minute; range: Angola

east to Nyasaland and southern Tanganyika miopropus

(p. 107)
Midbody scale-rows 22; fore limb absent; range: Natal and

Zululand north to the Transvaal macrolepsis

(p. 108)

100

bulletin: museum of comparative zoology

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loveridge: African lizards 101

Chamaesaura aenea (Fitzinger)

1843. Cricochalcis aenea (Wiegmann) Fitzinger, Syst. Rept., p. 21: Africa

1880. Chamaesaura miodactyla Giinther, Ann. Mag. Nat. Hist. (5), 6, p. 235,

fig: "Peri" i.e. Pirie Bush, Cape Province.

1885e. Chamaesaura aenea Boulenger, p. 263.
1898. Sclater, p. 104.

1898. Werner (1896-7), p. 140.

1901. Gadow, p. 537.

1907b. Roux, p. 422.
1908b. Boulenger, p. 224.
1909a. Hewitt, pp. 34, 37.
1910b. Boulenger, p. 470.
1910a. Hewitt, pp. 56, 71.
1911b. Hewitt, p. 47.
1913a. Werner, p. 109.

1922. Duerden & Essex, p. 269, fig. 1.

1923. Duerden, p. 362.

1928a. Essex (1927), p. 883, figs. 1, 4, 7-14.

1935. Lawrence, p. 44.

1937e. Hewitt, p. 32, pi. x, fig. 3.

1938. Gorham & Ivy, p. 179.

1898. Chamaesaura macrolepis Werner (1896-7) (not Cope"), p. 141.

Nome. Transvaal Snake-lizard (English: Hewitt).

Description. Rcstral in contact with, or separated from, the fronto-
nasal; two superposed lcreals (= a loreal over a postnasal); a pre-
ocular; prefrontals forming a suture; anterior pair of parietals sepa-
rated by an interparietal, posterior pair by an occipital-; fore and
hind limbs well-developed, pentadactyle, clawed.

For characters common to all species, see definition on p. 88, for
scale and pore counts, see statistical table on p. 101.

Color. Above, head and back dark brown with three (vertebral and
dorsolateral) light yellow or gray, black-edged, longitudinal lines;
flanks straw color with two or three longitudinal series of dark spots
or a reddish brown lateral band. Below, whitish or greenish white.

Size. Total length of d" (M.C.Z. 14204), 397 (95 -f 302) mm., from
Belfast; of the type of miodactyla (Brit.Mus.), 320+ (90 + 230+) mm .,
the tail being regenerated.

Thus the tail may comprise nearly three-quarters of the total
length. Two 460 mm. adults had fore limbs of 11 mm. long, hind
limbs of 14 and 16 mm. respectively (Essex).

Remarks. Boulenger (1885e) places the nostril in the antero-inferior

102 bulletin: museum of comparative zoology

portion of the nasal, this is rejected for postero-inferior, for it does not
differ in this respect from other members of the genus a position
which some might prefer to describe as in the lower centre.

Schmidt (1919) sees no reason for Werner (1898) having referred
three Natal lizards to macrolepis, two at least had four limbs and on
one they were pentadactyle, apparently three species are involved!

Anatomy. Duerden & Essex (1922) discuss the skeletal structure
of the limbs; Gorham and Ivy (1938) deal with the gall bladder.

Habitat. Grass veld uplands.

Localities. Transvaal: Belfast (M.C.Z.); Lydenburg District;
Middleburg District; Modderfontein; Pretoria District; Stander-
ton District; Zoutpansberg District. Natal. Orange Free State.
Gape Province: near Cathcart, Amatola Mountains; Drakens-
berg; Fenella Falls, Tarkastad District; Oakville Farm, Elliot Dis-
trict; Pirie (Peri) Bush.

Range. Transvaal south through Orange Free State to eastern Cape
Province and Natal.

Chamaesaura anguina anguina (Linne)
Plate 12, figs, lb, 2-3

1735. Vermis serpentiformis Seba, Rerum naturalium Thesauri, 2, p. 70,

pi. lxviii, figs. 7-8: Cape of Good Hope.
1758. Lacerta anguina Linne, Syst. Nat., ed. 10, 1, p. 210: Cape of

Good Hope, i.e. Union of South Africa.
1760. Linn6, ed. 12, 1, p. 371.

1768. Chalcides pinnata Laurenti, Syn. Rept., p. 64: Cape of Good Hope.
1795. Chalcida anguina Meyer, p. 31.
1801. Cha?naesaura Anguinea (sic) Schneider, p. 210.
1802d. Seps monodactylus Daudin, Hist. Nat. Rept., 4, p. 342, pi. lvii, fig.

1 : Cape of Good Hope.
1811. Chalcides monodactylus Oppel, p. 45.
1820. Monodactylus anguinus Merrem, p. 76: Africa.

1838. Chamaesaura anguina Gray, p. 392.

1839. Dumeril & Bibron, p. 441.
1845. Gray, p. 61.

1849. Smith, A., App., p. 10.

1851. Gravenhorst, p. 309, pi. xxx.

1867a. Steindachner, p. 42.

1885e. Boulenger, p. 264.

1889. Boettger, p. 292.

1890a. Muller, p. 286.

1893a. Boettger, p. 66.

loveridge: African lizards

103

1898. Sclater, p. 104.

1898. Werner (1896-7), p. 140.

1905h. Boulenger, p. 253.

1907b. Roux, p. 423.

1908b. Boulenger, p. 224.

1909a. Hewitt, pp. 34, 37.

1910b. Boulenger, p. 470.

1911b. Hewitt, p. 47.

1913a. Werner, p. 109, fig.

1922. Duerden & Essex, p. 270, figs. 2-3.

1923. Duerden, p. 362.
1925. Essex, p. 339.
1926b. Rose, p. 493.
1927b. Hewitt, p. 454.

1928a. Essex (1927), pp. 884, 932, figs. 2, 5, 15, 16, 17.

1929. Rose, pp. 113, 125, fig. 82.

?1933m. Witte, p. 72 (Elisabethville, B. C, so removed to C. a. tenuiof).

1935. Lawrence, p. 44.

1937e. Hewitt, p. 32, pi. x, figs. 2, 6.
1890d. Chamaesaura didactyla Boulenger Proc. Zool. Soc. London, pp. 78,

82, pi. xi, fig. 1 : South Africa.

1898. Sclater, p. 104.
1898. Chamaesaura macrolepis Werner (1896-7) (not Cope), p. 141.

For an extensive bibliography for the century prior to 1839, see
Dumeril & Bibron, 1839, Erpet. Gen., 5, p. 441.

Native names. Cape Snake-lizard (English : Hewitt) ; sweepslang or
pootjie slang (Afrikaans, but latter not even generic: Hewitt); un-
ombatamb' eza?itsi (Kaffir: Hewitt).

Description. Rostral in contact with, or separated from, the fronto-
nasal; a postnasal; a single or two superposed loreals; a preocular;
prefrontals separated by the frontonasal being in contact with the
frontal; anterior, and sometimes the posterior, pairs of parietals sep-
arated by a large interparietal, or the posterior by one or two small
scales, or posterior pair in contact posteriorly; fore and hind limbs
styliform, undivided, or hind limbs divided, terminating in minute
claws, or clawless.

For characters common to all species, see definition on p. 98, for
scale and pore counts, see statistical table on p. 101.

Color. Above, head and back dark brown, uniform or latter laterally
edged with black forming a pair of dark longitudinal lines; a light
vertebral line; flanks straw color with or without a narrow, longi-
tudinal, white lateral band. Below, whitish or golden yellow.

104 bulletin: museum of comparative zoology

Size. Total length of type of didactyla (Brit. Mus.), 530 (110 + 420)
mm., of a 9 (M.C.Z. 21430), 467 (115 + 352) mm.

Remarks. As Hewitt (1909a) and so many authors have pointed
out, the variability of this species in respect to its limbs being mono-
dactyle or didactyle is unquestionable. Roux (1907b) comments on
one lizard with a didactyle and two-clawed right fore limb whose other
three feet were monodactyle, another had clawed fore limbs but claw-
less hind limbs. Whether clawlessness in members of this genus im-
plies that the claws have not been developed or have been lost, is a
matter which requires investigation.

Anatomy. Duerden & Essex (1922) and Essex (1928) remark that
the fore and hind limbs average 6 and 8 mm. respectively and usually
terminate in a single claw, and discuss limb reduction generally.

Breeding. In February, 4 or 5 young are born when about six
inches in length (Rose).

Habits. Even in its slowest movements this active lizard makes no
use of its limbs so far as can be seen, the forepart of the body being
raised clear of the ground. Its passage through grass is too swift and
smooth for the eye to follow (Rose). So snake-like were the motions
of one which was encountered gliding over a rock in rocky country,
that it was mistaken for a Psammophis crucifer (Essex).

Habitat. Occurs chiefly among grass or stones near streams or the
sea coast in humid localities (A. Smith).

Localities. Zllluland: Umfolosi Station (fide Boulenger, 1905h).
Natal: Durban (fide Boulenger, 1910b). Cape Province1: Bains
Kloof; Bathurst; Caledon Division; Cape Peninsula; Cape Town;
Coldspring; Grahamstown; Kaffraria; Kei Road; Kentani;Knysna;
Maclear; Mquanduli; Muizenberg; New Brentingville; Ngqeleni;
Pondoland; Port Alfred; Port Elizabeth; Range Cottage, Table
Mountain; Schoonster's Drift; Swellendam; Tokai; West Hill near
Grahamstown; Worcester. (Witte's (1933m) record from Elisa-
bethville, Belgian Congo is tentatively transferred to the northern
race tenuior. As C. macrolepis is the common species of Natal
and Zululand it might be advisable to check the identifications of
1905 and 1910 listed above).

Range. Transvaal, Swaziland, and Zululand south to Natal west
across Cape Province to Little Namaqualand.

'The Irene record of Hewitt was later corrected by him (1911b) to Tokai; that of Little
Namaqualand (Boulenger, 1910b) is obviously wrong locality data, for FitzSimons (in lift.)
informs me that no Chamaesaura occurs there.

loveridge: African lizards

105

Chamaesaura anguina tenuior Gunther

1895. Chamaesaura tenuior Gunther, Ann. Mag. Nat. Hist. (6), 15, p. 524,

pi. xxi, fig. 6: Kampala, Uganda.

1896. Tornier, p. 37.

1897. Tornier, p. 64.
1898c. Boulenger, p. 915.

1902d. Boulenger, p. 445 (as taennior).

1902b. Tornier, p. 582.

1909. Peracca, p. 166 (as taenuior).

1912c. Sternfeld, p. 209.

1913c. Nieden, p. 74.

1923a. Loveridge, p. 18.

1923d. Loveridge, p. 850.

1924b. Loveridge, p. 10.

1929h. Loveridge, p. 59.

1930b. Barbour & Loveridge, p. 788.

1936J. Loveridge, p. 297.

1939. Someren, p. 157, pi. A, fig. 2.

1942e. Loveridge, p. 330.

1899c. Chamaesaura annectens Boulenger, Proc. Zool. Soc. London, p. 97:

Ravine Station, Mau Mountains, 7500 ft., Kenya Colony.

1900b. Tornier, p. 590.

1902d. Boulenger, p. 445.

1908c. Boulenger, p. 5.

1909. Peracca, p. 166.

1911c. Boulenger, p. 162.

1913c. Nieden, p. 74.

1923d. Loveridge, p. 850 (as annectans).

1924b. Loveridge, p. 10 (as annectans).

?1933m. Chamaesaura anguina Witte (not Linn6), p. 72.

Native names. Mugoye (Ragoli); shikoye (Tereki), nyarunyansi
(Toro and Wamba).

Description. Rostral in contact with1, or separated from, the fronto-
nasal; a postnasal; a loreal; a preocular; prefrontals separated by the
frontonasal being in contact with the frontal; anterior and posterior
pairs of parietals separated by a large interparietal, or the posterior
by one or two small scales ; fore and hind limbs styliform, undivided, or
hind limbs divided, terminating in minute claws, or clawless.

For characters common to all species, see definition on p. 98, for
scale and pore counts, see statistical table on p. 101.

1 Tornier (1896) states that this condition obtains in a 9 from Kavirondo, but that while
two of her offspring agree in this respect, in the other embryos the rostral is separated from the
frontonasal.

106 bulletin: museum of comparative zoology

Color. Above, head and back pale or dark brown, latter laterally
edged with black forming a pair of dark longitudinal lines; a light
vertebral line present or absent ; flanks straw color with or without a nar-
row, longitudinal, white lateral band. Below, whitish or pinkish white.

Size. Total length of type of tenuior (Brit. Mus.), 570 (114 4- 456)
mm., of type of annectens (Brit. Mus.), 450+ (150 + 300+) mm., tail
reproduced, of a 9 (M.C.Z. 41101), 637 (135 -f 502) mm.

It is interesting to note that in very young embryos the tail is little
more than twice the length of head and body, while in adults it is as
much as 3.7 times.

Remarks. Giinther stated that the fore limb was about equal to the
orbit in length, and that the hind limb was not quite twice as long,
actually the former's length is contained from once and a half to once
and three quarters in the distance from end of snout to anterior corner
of orbit, thus differing from typical anguina of Southeast Africa where
the hind limb about equals or is contained once and a quarter times
in the distance from end of snout to anterior corner of orbit. In colora-
tion and other respects there seems little to differentiate the two so I
prefer to regard tenuior as a northern race.

My reasons for uniting annectens with tenuior were given (1929h)
in detail. In a Kaimosi series both monodactyle and didactyle lizards
occurred, some having 24, others 26, midbody scale rows. Nor can
Kaimosi be regarded as the meeting place of two forms for apparently
the same variation occurs throughout its range.

Breeding. On February 12, at Kaimosi, five females were gravid,
two examined held 9 and 10 embryos respectively, one of the latter
measured 117 (35 + 82) mm., another lizard held only 6 embryos, one
measuring 77 (25 + 52) mm. In June or July, in mountains northwest
of Lake Tanganyika, Sternfeld records embryos, the viviparous nature
of this race being first recorded by Tornier (1896). On December 12,
at Fort Portal, a 138 mm. 9 held embryos (Loveridge).

Diet. Observed capturing flies and moths, a captive lizard took
nymphal grasshoppers (van Someren).

Enemies. Recovered from stomach of Bare-faced Hawk {Gymno-
genys t. typicus ) by van Someren.

Habits. These serpentiform lizards delight to bask on tufts of dry
grass into which they dive, or from which they slide to the ground, on
being disturbed. If approached quietly, however, it was possible to
sweep them up in a butterfly net. Wakamba, who showed no fear of
skinks such as Mabuya striata, hesitated to catch tenuior on account of
their serpentiform appearance (van Someren).

loveridge: African lizards 107

Habitat. Grasslands from 3000 to 8000 feet.

Localities. Uganda: Fort Portal; Kabulamuliro; Kacheliba;
Kampala; Kasiba; Kitende, west of Mbarara; Mabira Forest;
Mitiana; Sesse Islands. Kenya Colony: Chuyulu Hills; Gilgil
(M.C.Z.); Kaimosi; Kossowo; Kwa Raschuongo in Kavirondo;
Loita Plains; Mount Kenya; Ravine Station; Ugowe Bay, Kavi-
rondo Gulf; Yala River. Tanganyika Territory : Kakutta (1902b)
later spelled Kalkutta (1913c) is probably a locality in the western
Usambara Mountains where Eggel collected. Belgian Congo:
? Elizabethville; Mountains northwest of Lake Tanganyika.

Range. Kenya Colony and northeastern Tanganyika Territory west
to Uganda and eastern Belgian Congo.

Chamaesaura miopropus Boulenger

1894e. Chamaesaura miopropus Boulenger, Proc. Zool. Soc. London, pp. 724

732: "Fwambo" i.e. Fuambo, Nyasaland.

1900b. Tornier, p. 590.

1911. Sternfeld & Nieden, p. 385.

1913c. Nieden, p. 74.

1923a. Loveridge, p. 18.

1923d. Loveridge, p. 850.

1924b. Loveridge, p. 10.

1933h. Loveridge, p. 302.

1933. Schmidt, p. 10.

1937f. Loveridge, p. 498.

1895a. Chamaesaura macrolepis Bocage (not Cope), p. 25.

1937b. Monard, p. 61.

Native names. Nyoka lusagalla (Hehe); nunduswa (Kinga); nombo
(at Galanga: Bocage).

Description. Rostral separated from the frontonasal; a postnasal;
an elevated loreal; a preocular; prefrontals separated by the fronto-
nasal being in contact with the frontal; anterior and posterior pairs of
parietals separated by a large interparietal ; fore limbs a minute clawed
vestige; hind limbs styliform, undivided, clawed.

For characters common to all species, see definition on p. 98, for
scale and pore counts, see statistical table on p. 101.

Color. Above, head and back pale brown, uniform except for minute
black flecks, or like tenuior with or without a narrow, longitudinal,
white lateral band. Below, whitish or pinkish white.

Size. Total length of type (Brit. Mus.), 550 (120 + 430) mm., of
a <? (M.C.Z. 30765), 443 (87 + 356) mm., of a 9 (M.CZ. 30764),
455 (103 + 349) mm.

108 bulletin: museum of comparative zoology

Remarks. Schmidt (1933) has suggested the probability that the
two Angolan lizards identified as macrolcpis by Bocage are more prob-
ably miopropus. Bocage's remarks about the minute fore limbs makes
this practically certain. One wonders whether the Elisabeth ville reptile
referred to anguina by Witte (1933m), which I have tentatively placed
under a. tenuior, might not possibly be a miopropus also. The high
labial count given by me (1933h) for some Tanganyika miopropus was
due to the inclusion of the subocular and small scales over the gape.

Breeding. On February 8, at Ipemi, a 9 held several small ova; on
April 30, at Igale, another held a single round egg measuring 7 mm. in
diameter.

Diet. Black field crickets, grasshoppers, caterpillars and apparently
a beetle larva.

Habits. Fond of sunning itself on tussocks of dry grass into which it
vanishes with great speed when disturbed.

Habitat. In long dry grass at high altitudes.

Localities. Tanganyika Territory: Livingstone Mountains;
near Manitete, Unyika; Poroto Mountains — Igale Pass; Ukinga
Mountains — Ihenye and Tandala; Uzungwe Mountains — Da-
baga and Ipemi. Nyasaland: Fuambo (Fwambo). Angola:
Caconda; Chitau; Galanga (see Remarks).

Range. Southern Tanganyika Territory and Nyasaland. Angola.

Chamaesaura macrolepis (Cope)

1862. Mancus macrolepis Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 339:

Natal.

1885e. Chamaesaura macrolepis Boulenger, p. 264.

1898. Sclater, p. 104.

1898. Werner (1896-7; part), p. 141.

1908b. Boulenger, p. 224.

1908. Odhner, p. 3.

1909a. Hewitt, pp. 34, 37.

1910b. Boulenger, p. 470.

1910a. Hewitt, p. 60.

1911b. Hewitt, p. 48.

1913a. Werner, p. 109.

1919. Schmidt, p. 607.

1922. Duerden & Essex, p. 273, fig. 4.

1923. Duerden, p. 362.
1927b. Hewitt, p. 454.

1928a. Essex (1927), p. 884, figs. 3, 6, 18, 19.

1935. Chamaesaura macropholis (sic) Lawrence, p. 44 (lapsus).

loveridge: African lizards 109

Further citations of macrolepis will be found under aenea, a.
anguina, and miopropus.

Description. Rostral separated from the frontonasal, which may be
transversely divided; a postnasal; an elevated loreal1; a preocular; pre-
frontals separated by the frontonasal being in contact with the frontal;
anterior, and usually the posterior, pair of parietals separated by a
large interparietal; fore limbs absent, hind limbs styliform, undivided,
terminating in a minute claw, or clawless.

For characters common to all species, see definition on p. 98, for
scale and pore counts, see statistical table on p. 101.

Color. Above, head and back pale brown, a pair of dark, longitu-
dinal dorsal lines; flanks straw color with two longitudinal series of
dark dots. Below, whitish.

Size. Total length of c? (M.C.Z. 16162), 519 (124 + 395) mm.
from Zululand, of unsexed type (A.N.S.P. 9709), 638 (138 + 500)
mm. (fide Schmidt).

Remarks. Schmidt (1919) describes in detail Philadelphia Academy
No. 9709 as type, designating No. 9708 as paratype. Former iY.N.S.P.
9713 (now M.C.Z. 17736) is labeled cotype, actually, however, Cope
writes in the singular of "this lizard."

Anatomy. Duerden and Essex (1922) discuss the skeletal structure
of the limbs, also limb reduction has been studied by Essex who figures
different degrees of it in the bones of the hind limb of this species.

Diet. Small orthoptera and spider (in M.C.Z. specimen).

Localities.2 Transvaal: White River, Lydenburg District.
Zululand: Dukuduku (Indukuduku); Mseleni; Somkele (Mich.
Mus.). Natal: Durban; Lower Tugela River; Pietermaritzburg.

Range2. Southeastern Transvaal, Swaziland, and Zululand south
to Natal.

1 Schmidt writes of two loreals, merely a difference in designation, the anterior being here
regarded as a postnasal.

2 Bocage (1895a) records from Angola are referred to miopropus.

110 bulletin: museum of comparative zoology

BIBLIOQRAPHY

of 180 works mentioning African CORDYLIDAE from 1735-1942,

but no attempt has been made to complete those prior to 1880.

Titles of papers are omitted for sake of brevity.

Andersson, L. G.

1900. Bihang K. Svenska Vet.-Akad. Handl., 26, No. 1, pp. 1-29.
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Barbour, T. & Loveridge, A.

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1930b. in Strong, R., "Report of the Harvard African Expedition upon
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Bateman, G. C.

1897. "The Vivarium." (London), pp. 1-424, figs. 1-98, pi. — .

BOCAQE, J. V. B. DU

1895a. "Herp6tologie d'Angola et du Congo." (Lisboa), pp. i-xx + 1-203,

pis. i-xix.
1896a. Jorn, Sci. Lisboa (2), 4, No. 14, pp. 65-104, pis. i-ii.
1896c. Jorn. Sci. Lisboa (2), 4, No. 14, pp. 115-120.

BOETTGER, O.

1881c. Abhand. Senckenberg. Naturf. Ges., 12, pp. 435-558, pis. i-v.

1887b. Ber. Senckenberg. Naturf. Ges., pp. 135-173, pi. v.

1889. Ber. Senckenberg. Naturf. Ges., pp. 267-316.

1893a. "Katalog der Reptilien-Sammlung in Museum der Senckenberg-

ischen naturforschenden Gesellschaft in Frankfurt-am-Main."

(Frankfurt a. M.), pp. i-ix + 1-140.
1894a. Ber. Senckenberg. Naturf. Ges., pp. 88-93.
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3, pp. 269-376, pis. xxiii-xxx.

Boie, F.

1828. Nova Acta Acad. Caes. Leop.-Carol., 14, pp. 139-142.

BOTTLENGER, E. G.

1929. "Reptiles and Batrachians." (London), pp. i-xiv + 1-278,
78 pis.

BOTJLENGER, G. A.

1884a. Ann. Mag. Nat. Hist. (5), 14, pp. 117-122.

1885e. "Catalogue of the Lizards in the British Museum (Natural
History)," ed. 2 (London), 2, pp. i-xiii + 1-497, pis. i-xxiv.

loveridge: African lizards 111

1890d. Proc. Zool. Soc. London, pp. 77-87, pis. viii-xi.

1891d. Ann. Mag. Nat. Hist. (6), 7, p. 417.

1892a. in Distant, W. L. "A Naturalist in the Transvaal." (London),

pp. 174-176, fig.
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1897b. Ann. Mag. Nat. Hist. (6), 19, pp. 276-281, figs. -.
1897e. Proc. Zool. Soc. London, pp. 800-803, pi. xlvi.
1897g. Ann. Mus. Civ. Stor. Nat. Genova (2), 17, pp. 275-280.
1898c. Proc. Zool. Soc. London, pp. 912-923, pis. lv-lvii.
1899c. Proc. Zool. Soc. London, pp. 96-98, pi. x.
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445-447.
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1905h. Proc. Zool. Soc. London, pp. 248-255.

1907J. Proc. Zool. Soc. London, pp. 478-487, figs. 140-141, pis. xxi-xxii.
1908b. Ann. Natal Govt. Mus., 1, pp. 219-235, figs. 1-3, pis. xxv-xxvi.
1908c. Ann. Mus. Civ. Stor. Nat. Genova (3), 4, pp. 5-7, fig. -.
1910b. Ann. S. African Mus., 5, pp. 455-538.
1911c. Ann. Mus. Civ. Stor. Nat. Genova (3), 6, pp. 161-169.

Broom, R. /

1913. Proc. Zool. Soc. London, p. 548, pi. lxxiv.
1935. Ann. Transvaal Mus., 18, pp. 13-31, figs. 1-3.

Camp, C. L.

1923. Bull. Am. Mus. Nat. Hist., 48, pp. 289-481, figs. A-L + 1-112.

Chubb, E. C.

1909a. Proc. Zool. Soc. London, pp. 590-597.

1909b. Rhodesia Mus. Bulawayo, 8th. Ann. Rep., pp. 34-36.

Cope, E. D.

1862. Proc. Acad. Nat. Sci. Philadelphia, pp. 337-344.
1869. Proc. Amer. Philos. Soc, 11, pp. 147-169.

Cott, H. B.

1928. Proc. Zool. Soc. London, pp. 923-961, pis. i-iv.

1934a. Proc. Zool. Soc. London, pp. 145-173, figs. 1-3, pis. i-iii.

Cuvier, G. L. C. F. D.

1829. "Le Regne Animal distribue' d'apres son organisation pour servir
de base a l'Histoire naturelle des Animaux et d'introduction a
l'Anatomie Comparee. 2, Les Reptiles." (Paris), ed. 2, pp.
i-xvi + 1-406.

Dam, see van Dam

112 bulletin: museum of comparative zoology

Daudin, F. M.

1802d. "Histoire naturelle, generale et particuliere, des Reptiles." 4, pp.
1-397, pis. xlvi-lviii.

DlTMARS, R. L.

1910. "Reptiles of the World." (London), pp. i-xi + 1-373, pis. i-lxxxix.

DUERDEN, J. E.

1923. S. African Journ. Sci., 20, pp. 257-265 (largely a synopsis of 1922
paper; see below).

Duerden, J. E. & Essex, R.

1922. S. African Journ. Sci., 19, pp. 269-275.

DUMERIL, A. M. C. & BlBRON, G.

1839. "Erpetologie Generale ou Histoire Naturelle complete des Rep-
tiles." (Paris), i-viii + 1-855, pis. 1-lviii.

Essex, R.

1925. Rec. Albany Mus., 3, pp. 332-342, figs. 1-8.

1928a. Proc. Zool. Soc. London, 1927, pp. 879-945, figs. 1-86, pis. i-viii.

Fischer, J. G.

1888b. Jahrb. Hamburg. Wiss. Anst., 5, pp. 11-17, pi. ii.

Fitzinger, L. J. F. J.

1826. "Neue Classification der Reptilien . . . Zoologischen Museum's

zu Wien." (Wien), pp. i-viii + 1-66, pi. i.
1843. "Systema Reptilium . . . Fasiculus primus: Amblyglossae (Con-
spectus geographicus)." (Vindobonae), pp. 1-106.

FitzSimons, F. W.

1932. "Snakes." (London), pp. i-xiv + 1-286, pis.

FitzSimons, V.

1930. Ann. Transvaal Mus., 14, pp. 20-48, figs. 1-29, pi.

1933. Ann. Transvaal Mus., 15, pp. 273-280, figs. 1-6, pi.
1935a. Ann. Transvaal Mus., 15, pp. 519-550.

1935b. Ann. Transvaal Mus., 16, pp. 295-397, figs. 1-30, pis. x-xi.

1937a. Ann. Transvaal Mus., 17, pp. 259-274, pi. x.

1938. Ann. Transvaal Mus., 19, pp. 153-209, map, pis. ii-iv.

1939a. Ann. Transvaal Mus., 20, pp. 5-16, figs. 1-17.

1939b. Ann. Transvaal Mus., 20, pp. 17-46.

1941. Ann. Transvaal Mus., 20, pp. 273-281, figs. 1-8, pi. xii.

1943. Transvaal Mus. Mem. 1. "The Lizards of South Africa" (Pretoria),

pp. i-xv + 1-528, figs. 1-384, pis. i-xxiv, map. (received too late

for general citation in this revision).

Flower, S. S.

1925b. Proc. Zool. Soc. London, pp. 911-981.

loveridge: African lizards 113

Gadow, H.

1901. in Cambridge Natural History, "Amphibia and Reptiles." (Lon-
don), 8, pp. i-xiii + 1-668, figs. 1-181, map. (reprinted 1920).

Gilchrist, J. D. F.

1911. "South African Zoology." (Cape Town). Nonvidi.

Gorham, F. W. & Ivy, A. C.

1938. Field Mus. Nat. Hist. Zool. Series, 22, pp. 159-213.

Gravenhorst, J. L. C.

1851. Nova Acta Acad. Caes. Leop.-Carol., 23, 1, pp. 291-394, pis.
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1831b. "Synopsis of the Species of the Class Reptilia." pp. 1-110, in

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1838. Ann. Nat. Hist., 1, pp. 274-283 + 388-394.
1845. "Catalogue of the Specimens of Lizards in the Collection of the

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Gronovius, L. T.

1763. "Zoophylacium Grovianum exhibiens Animalia Quadrupeda, Am-
phibia, ..." etc. (Lugduni Bataviorum), pp. i-vi + 1-380, pis.
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Guerin, Meneville (F. E. Guerin-Meneville)

1829- "Iconographie du Regne Animal de G. Cuvier, ou representa-
1841. tion d'apres Nature de l'une des especes les plus remarquables, et
souvent non encore figures de chaque genre d'animaux. Avec un
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GtJNTHER, A.

1880. Ann. Mag. Nat. Hist. (5), 6, pp. 234-238, figs. -
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Hewitt, J.

1909a. Ann. Transvaal Mus., 2, pp. 29-40, pi. -.

1910a. Ann. Transvaal Mus., 2, pp. 50-71.

1911b. Ann. Transvaal Mus., 3, pp. 42-55.

1914a. S. African Journ. Sci., 10, pp. 238-253, maps.

1920. Trans. Roy. Soc. S. Africa, 7, pp. 91-93, figs. -.

1925b. Rec. Albany Mus., 3, pp. 343-368, figs. 1-4 + 3, pis. xv-xix.

1927a. Rec. Albany Mus., 3, pp. 371-415, figs. 1-2, pis. xx-xxiv.

1927b. S. African Journ. Sci., 24, pp. 452-456.

1932. Ann. Natal Mus., 7, pp. 105-128, fig. -, pi. vi.

1933a. Occ. Papers Rhodesian Mus., 2, pp. 45-50, pi. ix.

114 bulletin: museum of comparative zoology

1937d. Ann. Natal Mus., 8, pp. 199-209, pi. xii.

1937e. "A Guide to the Vertebrate Fauna of the Eastern Cape Province,
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Hewitt, J. & Power, J. H.

1913. Trans. Roy. Soc. S. Africa, 3, pp. 147-176.

Jeude, T. W. van L. de

1895. Notes Leyden Mus., 16, pp. 227-230.

1898. Mus. Hist. Nat. Pays-Bas, 10, part 2, pp. 1-54 + 1-52 + 1-11.

Laurenti, J. N.

1768. "Austriaci Viennensis Specimen Medicum, exhibiens Synopsis
Reptilium Emendatam cum Experimentis circa Venena et Anti-
dota Reptilium Austriacorum." (Vienna), pp. 1-216.

Lawrence, R. F.

1935. Parasitology (Cambridge, Eng.), 27, pp. 1-45, figs. 1-22.
1937. Cape Naturalist (Cape Town), 1, pp. 107-113, figs. 1-4.

Linne, Carl von

1758. "Systema Naturae." (Holmiae), ed. 10, 1, pp. 1-824.

1760. "Systema Naturae." (Halae Magdeburgicae), ed. 12, 1, pp. 1-532.

Loveridge, A.

1920a. Proc. Zool. Soc. London, pp. 131-167.

1923a. Journ. Tanganyika Civil Serv. Assoc, pp. 10-28.

1923d. Proc. Zool. Soc. London, pp. 841-863.

1923h. Proc. Zool. Soc. London, pp. 935-969.

1924b. Journ. E. Africa & Uganda Nat. Hist. Soc, Spec. Suppl. No. 3,

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1929h. Bull. U. S. Nat. Mus., No. 151, pp. 1-135, pi. i.

1932a. Bull. Mus. Comp. Zool., 72, pp. 375-387.

1933h. Bull. Mus. Comp. Zool., 74, pp. 197-416, pis. i-iii.

19361-l. Field Mus. Nat. Hist., Zool. Series, 22, pp. 1-111.

1936j. Bull. Mus. Comp. Zool., 79, pp. 209-337, pis. i-ix.

1937f. Bull. Mus. Comp. Zool., 79, pp. 481-541, pis. i-iv.

1942e. Bull. Mus. Comp. Zool., 91, pp. 235-374, pis. i-vi.

Mann, I.

1931. Trans. Zool. Soc. London, 21, pp. 355-412, figs. 34-35, pis. xx-xxiv.

Matschie, P.

1891a. Zool. Jahrb Syst., 5, pp. 605-611.

Merrem, B.

1820. "Versuch eines Systems der Amphibien. Tentamen systematis
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loveridge: African lizards 115

Mertens, R.

1937b. Abhand. Senckenberg. Naturf. Ges., No. 435, pp. 1-23.
1937d. Veroften. Deutschen Kolon.-Ubersee-Mus. Bremen, 2, pp. 1-9.

Methtten, P. A. & Hewitt, J.

1913c. Trans. Roy. Soc. S. Africa, 3, pp. 107-111, pi. vii.
1914b. Ann. Transvaal Mus., 4, pp. 118-145, figs. 14-16, pi. xiv.

Meyer, F. A. A.

1795. "Synopsis Reptilium, novam ipsorum sistens Generum Methodum
nee non Gottingensium huius Ordinis Animalium Enumera-
tionem." (Gottingae), pp. 1-32.

Mocquard, F.

1909. Nouv. Arch. Mus. Paris (5), 1, pp. 1-110.

MONARD, A.

1937b. Arqu. Museu Bocage, Lisboa, 8, pp. 1-154, figs. 1-3.

Muller, F.

1890a. Verh. Naturf. Ges. Basel, 8, pp. 249-296, pis. i-iii.
1890b. Verh. Naturf. Ges. Basel, 8, pp. 685-705, pi. x.

NlEDEN, F.

1913c. Mitt. Zool. Mus. Berlin, 7, pp. 51-100.

Noble, G. K. & Bradley, H. T.

1933b. Ann. New York Acad. Sci., 35, pp. 25-100.

Odhner, T.

1908. Arkiv. Zool., 4, No. 18, pp. 1-7.

Oppel, M.

1811. "Die Ordnungen, Familien und Gattungen der Reptilien als
Prodrom einer Naturgeschichte derselben." (Munchen), pp. 1-87,

Parker, H. W.

1936c. Novit. Zool. (Tring), 40, pp. 115-146, figs. 1-2.

Peers, B.

1930. S. African Journ. Nat. Hist. (Pretoria), 6, pp. 402-411, photos.

Perraca, M. G.

1909. in Abruzzi, "II Ruwenzori." (Milano), 1, pp. 165-180.

Peters, W. C. H.

1854. Montasb. Akad. Wiss. Berlin, pp. 614-628.

1855. Arch. Naturg., 21, pp. 43-58.

1862a. Monatsb. Akad. Wiss. Berlin, pp. 15-26.
1862c. Abh. Berlin Akad., pp. 165-225, pis. i-iii.
1869b. Ofver. Kongl. Vetensk.-Akad. Forh., pp. 657-662.
1879a. Sitz. Ges. Naturf. Freunde Berlin, pp. 10-11.

116 bulletin: museum of comparative zoology

1882a. "Naturwissenschaftliche Reise nach Mossambique . . . Zoologie,
3, Amphibien." (Berlin), pp. i-xv + 1-191, pis. i-xxx.

Pfeffer, G.

1889. Jahrb. Hamburg. Wiss. Anst., 6, pp. 1-36.

1893. Jahrb. Hamburg. Wiss. Anst., 10, pp. 71-105, pis. i-ii.

Pitman, C. R. S.

1934. "Report on a Faunal Survey of Northern Rhodesia." (Living-
stone), pp. i-xii + 1-500 + Index i-xxxii, maps A-K.

Popp, B.

1931. Aquarium (Berlin), pp. 191-194, photos.

1939. Wochenschr. Aquar. Terra. (Braunschweig), 36, pp. 263-264,
photos.

Power, J. H.

1918. S. African Journ. Sci., 14, pp. 263-271.

1927c. Trans. Roy. Soc. S. Africa, 14, pp. 405-422, fig. 1, pis. xviii-xxii,
map.

1930. Ann. Transvaal Mus., 14, pp. 11-19, fig. 1, pis. i-ii.

1931. Trans. Roy. Soc. S. Africa, 20, pp. 39-49, figs. 1-2.
1933a. Trans. Roy. Soc. S. Africa, 21, pp. 211-220, map.

Reichenow, A.

1887. Zool. Anz., 10, pp. 369-372.

RoCHEBRUNE, A. T. DE

1884a. "Faune de la Senegambie. Reptiles.'' (Paris), pp. 1-221, pis. i-xx.

Rose, W.

1926b. Ann. S. African Mus., 20, pp. 491-494.

1929. "Veld and Vlei." (Cape Town), pp. i-xxiii + 1-240, figs., photos
1-125.

Roux, J.

1907b. Zool. Jahrb. Syst., 25, pp. 403-444, pis. xiv-xv.

SCHLEGEL, H.

1834. Tijdschr. Nat. Gesch. Phys., 1, pp. 203-221, pi. vii.

Schmidt, K. P.

1919. Bull. Amer. Mus. Nat. Hist., 39, pp. 385-624, figs. 1-27, pis.
vii-xxxii.

1933. Ann. Carnegie Mus., 22, pp. 1-15, figs. 1-2, pis. i-ii.

Schmidt, P.

1930. Aquarium (Berlin), pp. 153-155, photos.

Schneider, J. G.

1801. "Historiae Amphibiorum naturalis et literariae." (Jena), pp.
1-364, pis. i-ii.

loveridge: African lizards 117

SCLATER, W. L.

1898. Ann. S. African Mus., 1, pp. 95-108, pi. v.

SCORTECCI, G.

1940c. Atti. Soc. Ital. Sci. Nat. (Milano), 79, pp. 1-10, figs. 1-2, pis. i-ii.

Seba, A.

1735. "Locupletissimi Rerum Naturalium Thesauri Accurata Descriptio,
et Iconibus Artificiosissimis Expressio, per Universam Physices
Historiam," etc. (Amstelaedami), 2,pp.i-xxx + 1-154, pls.i-cxiv.

Senfft, W.

1931. Aquarium (Berlin), pp. 73-75, photos.

Smith, A.

1838a. Mag. Nat. Hist. (2), 2, pp. 31-33.

1843. "Illustrations of the Zoology of South Africa . . . Reptiles."

1849. (London), pis. i-lxxviii., App. pp. 1-28.

SOMEREN, See VAN SoMEREN

Steindachner, F.

1867a. in "Reise der Osterreichischen Fregatte Novara um die Erde in
den Jahren 1857, 1859, unter den Befehlen des Commodore
B. von Wullerstorf-Urbair. Zool." 1, pp. 1-98, pis. i-iii.

Sternfeld, R.

1911b. Mitt. Zool. Mus. Berlin, 5, pp. 393-411.
1911c. Mitt. Zool. Mus. Berlin, 5, pp. 413-420.
191 Id. in "Die Fauna der Deutschen Kolonien (Berlin), 4, part 2, pp.

i-iv + 1-65, figs. 1-76, map.
1912c. in "Wiss. Ergeb. der Deutschen Zentral-Afrika-Expedition 1907-

1908." (Leipzig, 1913), 4, pp. 197-279, figs. 1-4, pis. vi-ix.

Sternfeld, R. & Nieden, F.

1911. Mitt. Zool. Mus. Berlin, 5, pp. 383-385.

TORNIER, G.

1896. "Die Kriechthiere Deutsch-Ost-Afrikas. Beitrage zur Systematik
und Descendenzlehre." (Berlin), pp. i-xiii + 1-164, figs. 1-11,
pis. i-v.

1897. Arch. Naturg., 63, 1, pp. 63-66.

1898. in Werther, C. W. "Die mittleren Hochlander des nordlichen
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1900b. Zool. Jahrb. Syst., 13, pp. 579-618, figs. A-H.
1902b. Zool. Jahrb. Syst., 15, pp. 578-590, figs. -.

van Dam, G. P.

1921a. Ann. Transvaal Mus., 7, pp. 239-243, pis. i-iv.

118 bulletin: museum of comparative zoology

i
VAN SOMEREN, V. G. L.

1939. Journ. E. Africa & Uganda Nat. Hist. Soc, 14, pp. 155-160.

Werner, F.

1898. Jahrb. Abhand. Natur. Magdeburg (1896-7), pp. 139-148.
1910a. Denks. Med -Nat. Ges. (Jena), 16, pp. 279-370, figs. 1-15, pis.

vi-xi.
1913a. in Brehm, A. "Tierleben." (Leipzig), ed. 5, 4, pp. i-xvi + 1-598,

figs, plates, maps.
1915c. in Michaelsen, W. "Beitrage sur Kenntnis der Land- und Sus-

wasserfauna Deutsch-Slidwestafrikas. III." (Hamburg), pp. 325-

376, pi. vii.

WlEGMANN, A. F. A. & RUTHE, J. F.

1832. "Handbuch der Zoologie." (Berlin), pp. i-vi + 1-621.

WlTTE, G. F. DE

1933m. Ann. Mus. Congo Beige, Zool. (1), 3, pp. 55-98, pis. i-iv.

PLATES

PLATE 1

Loveridqe — African Lizards

PLATE 1

Fig. 1. Cordylus giganteus (Type after Smith).
Fig. 2. Cordylus giganteus (Type after Smith).
Fig. 3. Cordylus w. warreni (Type after Boulenger).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 1.

■-■ Hi

wpy^

2 i

PLATE 2

Lovebidqe — African Lizards

PLATE 2

Fig. 1. Cordylus warreni perkoensis (Type after FitzSimons).
Fig. 2. Cordylus warreni laevigatus (Type after FitzSimons).
Fig. 3. Cordylus warreni breyeri (Type after van Dam).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 2.

' I lr

PLATE 3

Loveridge — African Lizards

PLATE 3

Fig. 1. Cordylus caeruleopunctatus (after Power).
Fig. 2. Cordylus cordylus tropidosternum (Cope).

a. Type of frenatus Pfeffer.

b. Type of parkeri Cott.

Fig. 3. Cordylus c. jonesii (after Power).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 3.

PLATE 4

Loveridqe — African Lizards

PLATE 4

Fig. 1. Cordylus cordylus lawrencei (Type after FitzSimons).
Fig. 2. Cordylus cordylus tasmani (Cotype after Power).
Fig. 3. Cordylus cordylus cordylus (after Smith as griseus).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 4.

?» rX<

•>

Y MM

. rS :

- 3$^

PLATE 5

Loveridgb — African Lizards

PLATE 5

Fig. 1. Cordylus cordylus niger (Cotype of atrus Power).
Fig. 2. C. peersi (Type after Hewitt).
Fig. 3. C. cataphractus (after Smith).

BULL. MUS. COMP. ZOOL.

Loverid3E. African Lizards. Plate 5.

^jGABM'-m,

>* ■

PLATE 6

Loveeidge — African Lizards

PLATE 6
Fig. 1. Cordylus cataphractus (after Smith).
Fig. 2. C. cataphractus in defensive attitude (after Rose).
Fig. 3. C. pustulatus (after Sternfeld).
Fig. 4. C. p. polyzonus (Type after Smith).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 6 .

>•»

4fc>

■*» «.

£*-*-*■ ■'

tare

PLATE 7

Loveridge — African Lizards

PLATE 7

Fig. 1. Cordylus p. polyzonus (Type after Smith).
Fig. 2. Pseudocordylus capensis (Type after Smith).
Fig. 3. P. capensis (Type after Smith).

1ULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 7.

'.'.V < .J: A

PLATE 8

Loveridge — African Lizard =

PLATE 8

Fig. 1. Pseudocordylus robertsi (after Power).

Fig. 2. P. microlepidotus subviridis (Co type after Smith).

Fig. 3. P. m. subviridis (Cotype after Smith).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 8.

'"' -"-N

', '

■' in ■•-

leU

PLATE 9

Loveridge — African Lizards

PLATE 9

Fig. 1. Pseudocordylus microlepidokis melanotus (cf Cotype after Smith).
Fig. 2. P. m. melanotus (d1 Cotype after Smith).
Fig. 3. P. m. melanotus ( 9 Cotype after Smith).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 9.

'>V! .. .).■

***>

,'3

PLATE 10

Loveridge — African Lizards

PLATE 10

Fig. 1. Pseudocordylus m. microlepidotus (Type of montanus Smith).

Fig. 2. P. m. microlepidotus (Type of montanus Smith).

Fig. 3. P. m. fasciatus (Type after Smith).

Fig. 4. P. m. fasciatus (Type of algoensis Smith).

BULL. MUS. COMP. ZOOL

Loveridge. African Lizards. Plate 10.

It l! \ I

.v*vV

■

rv-

v-i-^i*.

* w& >

PLATE 11

Loveridge — African Lizards

PLATE 11

Fig. 1. Pseudocordylus m. fasciatus (Type of algoensis Smith).

Fig. 2. P. m. fasciatus (Type after Smith).

Fig. 3. Platysaurus guttatus torquatus (Cotype after Peters).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 11.

'•*-.-.

PLATE 12

Loveridge — African Lizards

PLATE 12

Fig. 1. Toe reduction in Chamaesaura (after Boulenger).

a. C. aenea (Fitzinger).

b. C. anguina anguina (Type of didactyla Boulenger).

c. C. macrolepis (Cope).

Fig. 2. C. a. anguina (Type of didactyla Boulenger).
Fig. 3. C. a. anguina (after Rose).

BULL. MUS. COMP. ZOOL.

Loveridge. African Lizards. Plate 12.

- ■:>

Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
- Vol. XCV, No. 2

UEC 28 1944*

FURTHER REVISIONS OF AFRICAN SNAKE GENERA

By Arthur Loveridge

CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM

December, 1944

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIV, and Vol. XCV, No. 1 and 2 have
appeared and of the Memoirs, Vol. I to LVI.

These publications are issued in numbers at irregular intervals.
Each number of the Bulletin and of the Memoirs is sold separately.
A price list of the publications of the Museum will be sent upon ap-
plication to the Director of the Museum of Comparative Zoology,
Cambridge, Massachusetts.

After 1941 no more Memoirs are to be published.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. XCV, No. 2

FURTHER REVISIONS OF AFRICAN SNAKE GENERA

By Arthur Loveridge

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

December, 1944

g& Zoology %>

' DEC 281944

No. 2 — Further Revisions of African Snake Genera
By Arthur Loveridge

The miscellaneous genera here assembled are not quite so diverse
as may appear at first sight, the majority of them being sylvicoline
and largely associated with rain forest. In fact the uniformly black
species or races of the four subfossorial genera are sufficiently similar
as to be difficult to differentiate in the field, while some have actually
been mistaken for burrowing vipers (Atractaspis) by several herpeto-
Iogists of note. The new monotypic genus is erected for a species which
occupies a somewhat intermediate position between Naja and Elap-
soidea, having been referred to the former by Werner and Nieden, to
the latter by Boulenger and de Witte.

These revisions, with the exception of the last two, were under-
taken four years ago during preparation of a report (1942, Bull. Mus.
Comp. Zool., 91, pp. 237-373) on the 81 species, or races, of snakes
collected during the course of an expedition sponsored by the John
Simon Guggenheim Memorial Foundation, to which this Museum is
grateful for much of the comparative material which has made these
studies possible. There remained, however, certain species or races
unrepresented in any American museum, besides questions regarding
types which could be satisfactorily settled only by reference to Euro-
pean museums. With this object in view the revisions were put on one
side, but as another three years may elapse before conditions are
sufficiently settled to deal with such matters, I have thought it better
to delay their publication no longer, though the synoptical keys to
some were included in the aforementioned report.

Since the Uganda Journal is inaccessible to many herpetologists, I
have deviated from usual practice and furnished both page reference
to Pitman's articles in the Journal (1935-1 938a) as well as the repagin-
ation as they appeared in book form (1938b) of his "Guide to the
Snakes of Uganda," such being followed in parenthesis by the word
'reprint.' In such cases it is listed under the name or spelling as
finally adopted in the concluding parts.

In all other respects the procedure adopted is similar to that out-
lined in the earlier of half-a-dozen similar revisions published in this
Bulletin during recent years (1939-1944). In each I have attempted
to assemble all pertinent data and information published since 1880
after allocating it to its subspecies as here recognized. Where an
author furnishes a locality record without scale-counts or other data

122 bulletin: museum of comparative zoology

which would assist in checking his identification, the locality in ques-
tion is listed under the subspecies within whose range it occurs, usually
with footnote or other comment. The generic definitions are substan-
tially those of Boulenger (1894a, 1896d) modified or expanded to in-
clude the findings of C. M. Bogert and others.

The opportunity is taken of thanking both Messrs C. M. Bogert
(American Museum) and V. FitzSimons (Transvaal Museum) for all
the trouble they have taken to supply me with data of specimens in
their care, the latter furnishing scale counts of thirty-one Elapsoidea,
while the former and Messrs K. P. Schmidt and C. H. Pope (Chicago
Natural History Museum) leave me indebted for the loan of material.

Index to the Species Recognized
AGLYPHA Page

*Rhamnophis batesii (Boulenger) 125

*Rhamnophis aethiopissa aethiopissa Gunther •. 126

*Rhamnophis aethiopissa ituriensis Schmidt 128

*Rhamnophis aethiopissa elgonensis Loveridge 129

Thrasops occidentalis Parker % 131

*Thrasops flavigularis (Hallowell) 132

*Thrasops jacksonii jacksonii Gunther 134

*Thrasops jacksonii schmidti Loveridge 137

*Duberria tutrix abyssinica (Boulenger) i 139

*Duberria tutrix shirana (Boulenger) 142

"Duberria tutrix tutrix (Linnaeus) 144

Duberria variegata (Peters) T47

OPISTHOGLYPHA

*Thelotornis kirtlandii kirtlandii (Hallowell) 149

*Thelotornis kirtlandii capensis Smith 154

*Calamelaps concolor (Smith) 161

Calamelaps unicolor polylepis Bocage 162

*Calamelaps unicolor warreni Boulenger 163

*Calamelaps unicolor unicolor (Reinhardt) 165

Calamelaps unicolor feae Boulenger 168

*Miodon acanthias (Reinhardt) 170

Miodon gabonensis gabonensis (Dumeril) 172

*Miodon gabonensis notatus (Peters) 173

*Miodon gabonensis collaris (Peters) •. . . 175

* Miodon gabonensis christyi Boulenger 178

* Miodon gabonensis graueri Sternfeld 180

loveridge: African snakes 123

AGLYPHA+OPISTHOGLYPHA

* Aparallactus lincatus (Peters) 183

* Aparallactus modestus (Gtinther) 186

Aparallactus liddiardae Parker 191

* Aparallactus concolor (Fischer) 192

* Aparallactus lunulatus (Peters) 195

* Aparallactus jacksonii (Gtinther) 197

* Aparallactus werneri Boulenger 199

* Aparallactus turneri Loveridge 201

Aparallactus capensis bocagii Boulenger 202

* Aparallactus capensis capensis Smith 205

* Aparallactus capensis uluguruensis Barbour & Loveridge 210

Aparallactus nigriceps (Peters) 212

PROTEROGLYPHA

Elapsoidea sundevallii laticincta (Werner) 216

* Elapsoidea sundevallii decosteri Boulenger 217

*Elapsoidea sundevallii semiannulata Bocage 220

*Elapsoidea sundevallii giintherii Bocage 222

* Elapsoidea sundevallii nigra Gtinther 225

* Elapsoidea sundevallii sundevallii (Smith) 228

*Elapsoidea sundevallii fitzsimonsi subsp. nov 229

*Paranaja gen. nov 231

Paranaja multicincta multicincta (Werner) 233

'Paranaja multicincta anomala (Sternfeld) 234

* Represented in the collections of the Museum of Comparative Zoology: examples of species
or forms without asterisk are earnestly desired.

124 bulletin: museum of comparative zoology

Genus RHAMNOPHIS

1862b. Rhamnophis Gunther, Ann. Mag. Nat. Hist. (3), 9, p. 129, pi. x

(type aethiopissa Gunther).
1864b. Crypsidomus Gunther, Proc. Zool. Soc. London, p. 309 (substitute

name).

Maxillary teeth 17-35, followed after an interspace by 3 enlarged
ones; anterior mandibular teeth slightly enlarged. Head rather short,
distinct from neck; eye very large, with round pupil; nasal divided or
semi-divided; a loreal; a preocular1. Body compressed; scales oblique,
narrow, smooth, with apical pits, in 13-19 rows of which the vertebral
is distinctly enlarged; ventrals rounded or obtusely keeled laterally.
Tail long; subcaudals in two rows.

Range. Africa in forested areas of equatorial belt from French
Guinea to western Kenya Colony.

Remarks. The substitute name Crypsidomus was proposed by
Gunther in the mistaken belief that his Rhamnophis, 1862, was pre-
occupied by Rhamphiophis Peters, 1854. It is by no means sure that
Rhamnophis should not be united with Thrasops Hallowell, 1857, with
which it has been confused, and from which it is barely separable.

The above description is largely that of Boulenger (1896d, p. 632)
after exclusion of the keeled-scaled Thrasops jacksonii, and extension
of the maxillary tooth count resulting from the inclusion of batesii,
together with notes on its dentition kindly supplied me by Bogert in
1940.

Key to the Species

1. Midbody scale-rows 13; anal entire; subcaudals less than 116; maxillary

teeth 30 +3 to 35 +3 batesii

(p. 125)

Midbody scale-rows 15—19; anal divided; subcaudals more than 116;

maxillary teeth 17+3 to 20+3 .*. .2

2. Midbody scale-rows' 17, very rarely 15, 16, or 19; lower postocular in con-

tact with 3 upper labials; range: French Guinea east to the western

Belgian Congo a. aethiopissa

(p. 126)

Midbody scale-rows 15, very rarely 17; lower postocular in contact with

2 or 3 upper labials; range; eastern Belgian Congo to western Kenya

Colony 3

3. Subcaudals 134-150; normally upper labials 8, sometimes 7; range: Ituri

region of eastern Belgian Congo a. ituriensis

(p. 128)

1 Rarely 2, fide Werner.

loveridge: African snakes 125

Subcaudals 117-138; normally upper labials 7, sometimes 6 or 8; range:

western Uganda to western Kenya Colony a. elgonensis

(p. 129)

Rhamnophis batesii (Boulenger)

1908a. Thrasops batesii Boulenger, Ann. Mag. Nat. Hist. (8), 2, p. 93: Akok

and Efulen, French Cameroon.
1909b. Sternfeld, p. 16.
1919b. Boulenger, p. 283.
1929a. Werner, p. 97.
1923. Rhamnophis batesii Schmidt, p. 83, fig. 5.

Description. Rostral twice as broad as deep, just visible from above ;
nasal divided or semidivided; internasals as broad as long, about as
long as the prefrontals ; frontal bell-shaped, once and a half to twice as
long as broad (in the middle), longer than its distance from the end of
the snout, as long as, or a little longer or shorter than, the parietals,
as broad as a supraocular; loreal longer than deep; preocular 1, in
contact with or separated from the frontal ; eye very large, its diameter
much greater than its distance from the mouth, postoculars 3, the
lowest in contact with 3 upper labials ; temporals 1, or 1 + 1 and occipi-
tal ; occipitals 4 ; upper labials 7 or 8, the fourth and fifth or fifth and
sixth entering the orbit; 4-6 lower labials in contact with the anterior
sublinguals, which are about as long as the median pair. Midbody
scales in 13 rows, smooth, oblique, the vertebral row enlarged; ventrals
163-177; anal entire; subcaudals 91-114 pairs.

Dentition. Boulenger found about 30+3 maxillary teeth, Bogert —
for Ituri specimens ■ — 35+3, the enlarged posterior ones separated by
a diastema from the subequal anterior teeth.

Color. Above, pale broAvn or pea green, each scale edged or striped
with black and spotted with lighter which may form irregular or
interrupted crossbands; labials edged with black posteriorly. Below,
white anteriorly with scattered black spots, brown or olive posteriorly
blotched with darker.

For further notes on color, see Schmidt (1923, p. 84).

Size. Total length of tf (M.C.Z. 38393), 1217 (827 + 390) mm. from
Batouri district, Cameroon, surpassed by that of an unsexed cotype
(Brit. Mus.) of 1800 (1450 + 350 mm.) '

Localities. French Cameroon: Akok; Batouri district; Efulen;
Ja River district, Belgian Congo: Gamangui; Niapu.

Range. French Cameroon east to the Ituri region, Belgian Congo.

126

bulletin: museum of comparative zoology

Rhamnophis aethiopissa aethiopissa Gunther

1862b. Rhamnophis aethiopissa Gunther, Ann. Mag. Nat. Hist. (3), 9 p.

129, pi. x: West Africa.

1884a. Rochebrune, p. 176, pi. xix, fig. 1 (ignored).

1875a. Rhamnophis aethiops Peters, p. 199.

1876a. Peters, p. 119.

1896d. Boulenger, p. 632.

1897b. Mocquard, p. 13.

1897. Sjostedt, p. 35.

1897b. Werner, p. 399.

1898a. Werner, p. 210.

1899a. Werner, p. 138.

1900b. Boulenger, p. 453.

1902a. Werner, p. 344.

1906i. Boulenger, p. 213.

1906. Johnston, p. 832.

1909. Gendre, p. cvi (as Rhaumophis).

1915a. Boulenger, p. 207.

1917. Chabanaud, p. 372.

1917b. Chabanaud, p. 11.

1919b. Boulenger, p. 284.

1919g. Boulenger, p. 23.

1922. Aylmer, p. 15.

1927d. Witte, p. 324.

1933f. Angel, p. 118, figs. 43-43a (not 44-44a; captions transposed).

1933m. Witte, p. 90.

1888a. Crypsidomus aethiops Boettger, p. 64.

1890. Crypsidomus aethiopissa Biittikofer, p. 478.

1891b. Chrysidomus aethiops Matschie, p. 616.

1892. Zenker, p.. 183.

1893c. Matschie, p. 212.

1901. Thrasops splendens Andersons, Bihang Till K. Svenska Vet.-Akad.

Handl., 27, No. 5, p. 11, pi. i, fig. 8: Bibundi and Mapanja, British

Cameroon.

1908a. Thrasops aethiops Sternfeld, pp. 409, 425.

1908b. Sternfeld, pp. 215, 230.

1909a. Sternfeld, p. 15.

1909b. Sternfeld, p. 17.

1910. Muller, p. 602.

1911. Lampe, p. 194.
1929a. Werner, p. 97.

1941e. Rhamnophis aethiopissa aethiopissa Loveridge, p. 121.

Further citations of 'aethiops' will be found under a. ituriensis.

loveridge: African snakes 127

Description. Rostral twice as broad as deep, visible from above;
nasal divided or semidivided; internasals as broad as long, about as
long as the prefrontals; frontal bell-shaped, twice as long as broad (in
the middle), as long as, or slightly longer than, its distance from the
end of the snout, as long as, or a little shorter than, the parietals, as
broad as a supraocular; loreal longer than deep; preocular 1, rarely 2/
in contact with or separated from the frontal; eye very large, its diam-
eter much greater than its distance from the mouth; postoculars 2,
rarely 3 or 4,1 the lower in contact with 3 upper labials; temporals 1,
or 1 + 1 and occipital (or 1 + 2, fide Boulenger); occipitals 2, usually
present; upper labials 8, the fourth and fifth, rarely fourth, fifth and
sixth, entering the orbit; 4 or 5 lower labials in contact with the
anterior sublinguals, which are shorter than the median pair. Midbody
scales in 17, rarely 15 (French Congo, fide Angel) 16 (Liberia, fide
Loveridge) or 19 (French Cameroon, fide Midler) rows, smooth,
oblique, the vertebral row enlarged; ventrals 158-179; anal divided;
subcaudals 139-159 pairs.

Dentition. 19 or 20 + 3 maxillary teeth, the enlarged posterior ones
separated by a diastema from the subequal anterior teeth.

Color. Above, green, each scale heavily edged with black; head pale
olive, the shields usually spotted and edged with black; five broad
black lines on tail. Below, pale olive or yellow with a light line along
either lateral angle; outer ends of ventrals green edged with black;
tail with a narrow, median, black line and edged with black laterally.

Size. Total length of cf (cotype of splendens in Stockholm Museum).
1330 (855 + 475) mm. from British Cameroon; of 9 (cotype Stock,
Mus. 1979) 1470 (950 + 520) mm. from Bibundi, British Cameroon;,
both surpassed by Boulenger's unsexed record of 1500 (970 4- 530)
mm.

Remarks. Angel (1933f, p. 118) has synonymized ituriensis with
aethiopissa on account of the occurrence in the French Congo of a
specimen with 15 midbody scale-rows. The incidence is so slight,
however (circa 3%) as not to invalidate the recognition of ituriensis
as a race within whose range 10% may have 17 midbody scale-rows. In
passing, it might be pointed out that the captions under the figures of
T. jacksonii and R. acthiops have been transposed.

Localities. Senegal ? (fide Rochebrune). French Guinea:
Ditinn (Ditiun). Sierra Leone. Liberia: Harbel. Gold Coast:
Accra. Togo: Adele (Bismarckburg) ; Kete; Misahohe. Nigeria:
Lagos. British Cameroon: Bibundi; Buea; Isongo; Mapanja;

lfide Werner.

128 bulletin: museum of comparative zoology

Victoria. French Cameroon: Batouri district; Bipindi; Dehane;
Dibongo near Edea; Longji; Yaundeland. Fernando Po. French
Congo: Lambarene; Ogowe. Belgian Congo: Banziville; Makaia
Ntete; Umangi.

Range. French Guinea east to the western Belgian Congo.

Rhamnophis aethiopissa ituriensis Schmidt

1919g. Rhamnophis aethiops Boulenger (part), p. 23.

1934a. Schwetz, pp. 381, 383.

1923. Rhamnophis ituriensis Schmidt, Bull. Am. Mus. Nat. Hist., 49, p.

81, fig. 4: Niapu, Belgian Congo.

1933m. Witte, p. 90.

1936. Pitman (part), p. 49 (but not plates).

1938b. Pitman (part), p. 104 (but not plates, reprint).

1929a. Thrasops ituriensis Werner, p. 97.

Further citations of 'ituriensis' will be found under a. elgonensis.

Description. Rostral about twice as broad as deep, just visible from
above; nasal divided; internasals as broad as long, about as long as
the prefrontals; frontal bell-shaped, once and three-quarters as long as
broad (in the middle), as long as its distance from the end of the snout,
slightly shorter than the parietals, as broad as a supraocular; loreal
longer than deep; preocular 1, in contact with or separated from the
frontal; eye very large, its diameter much greater than its distance
from the mouth; postoculars 2, rarely 3 or 4, the lower in contact with
2 upper labials ; temporals 1, or 1 -f 1 and occipital; occipitals 2, rarely
3 ; upper labials 7, rarely 6 or 8, the fourth and fifth entering the orbit ;
4 or 5 lower labials in contact with the anterior sublinguals, which are
shorter than the median pair. Midbody scales in 15, rarely 17, rows,
smooth, oblique, the vertebral row enlarged; ventrals 159-172; anal
divided; subcaudals 134-150 pairs.

Color. Above, green (bluish black in alcohol), each scale edged or
striped with black and spotted or striped with lighter; five broad
black lines on tail. Below, chin and throat yellowish, rest olive with a
light line along either lateral angle, a black spot either adjacent to this
line or at the outer end of the ventrals; tail with a narrow, median,
black line and edged with black laterally.

Size. Total length of type & (A.M.N.H. 12505), 1305 (865 + 440)
mm. of paratype 9 (A.M.N.H.), 1290 (832 -f- 458) mm. both from
Niapu.

loveridge: African snakes 129

Parasites. Hemogregarines reported from a Stanleyville snake by
Schwetz.

Localities. Belgian Congo: Gamangui; Medje; Niangara;
Niapu; Stanleyville. (Reported from Uganda in error by Pitman).

Range. Eastern Belgian Congo.

Rhamnophis aethiopissa elgonensis Loveridge

1916a. Thrasops rothschildi Loveridge (not Mocquard), pp. 79, 84.

1923e. Loveridge (part), p. 879 (Yala specimens only).

1929h. Rhamnophis aethiopissa elgonensis Loveridge, Bull. U. S. Nat. Mus.

161, p. 24: Yala (= Lukosa) River, Kenya Colony.
1937f. Loveridge, p. 502.
1938a. Pitman, p. 210.
1938b. Pitman, p. 308 (reprint).
1942e. Loveridge, p. 276.

1936. Rhamnophis ituriensis Pitman (part), p. 49, pi. vi. fig. 6, pi. T, fig. 3.
1938a. Pitman, pp. 210, 231.
1938b. Pitman, pp. 104-106, 308, 329, pis. as above (reprint).

Description. Rostral once and two-thirds to twice as broad as deep,
visible from above; nasal divided (its posterior half fused with loreal
only in the type specimen) ; internasals broader than long (in young)
or as broad as long (in adults), about as long as the prefrontals; frontal
bell-shaped, once and a half as long as broad (in the middle), as long
as (adult) or much longer than (young) its distance from the end of
the snout, noticeably shorter than the parietals, much broader than a
supraocular; loreal longer than deep; preocular 1, in contact with or
separated from the frontal; eye very large, its diameter much greater
than its distance from the mouth; postoculars 2, the lower in contact
with 2 upper labials; temporals 1, or 1 + 1, or 1 + 1 and occipital;
occipitals 2; upper labials 7, rarely 5, 6 or 8, the fourth and fifth or
third and fourth entering the orbit; 4 or 5 lower labials in contact with
the anterior sublinguals, which are shorter than the median pair.
Midbody scales in 15 rows, smooth, oblique, the vertebral row enlarged ;
ventrals 154-164; anal divided; subcaudals 117-138 pairs.

Color. Above, green, each scale edged with black; head olive, uni-
form or the shields edged with black, upper labials pale green anteri-
orly and yellowish posteriorly in their upper portion, blue below;
five broad black lines on tail. Below, ventrals greenish flecked with
white, a brown line along either lateral angle, outer ends of ventrals

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pale olive green, tail with a median dusky line flanked by irregular
dark flecks.

The above was based on notes made in the field of a snake from
Kibale Forest. For further notes on color, see Pitman (1938b, p. 105,
and col. pi. T, fig. 3) under name of "iturie?isis".

Size. Total length of d" 1255 (825 + 430) mm. from Budongo
Forest (Pitman); of type 9 (M.C.Z. 18198), 1147 (760 + 387 mm.)
from Yala River.

Localities. Uganda: Budongo Forest; Kajansi Forest; Kibale
Forest; Mabira Forest. Kenya Colony: Kaimosi; Kakamega;
Yala River.

The last three localities are really one, Kaimosi being only an hour's
walk from the Yala River flowing through the Kakamega Forest,
which is an outlier of the Elgon Forest. The species has not as yet
been taken on Mount Elgon.

Range. Western Uganda to western Kenya Colony.

Genus Thrasops

1857. Thrasops Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p. 67 (type
Dendrophis fiavigularis Hallowell).

Maxillary teeth 17-18, followed after an interspace by 3-4 enlarged
ones; anterior mandibular teeth slightly enlarged. Head rather short,
distinct from neck; eye large, with round pupil; nasal divided, semi-
divided, or entire; a loreal; 1 or 2 preoculars. Body more or less com-
pressed; scales oblique, strongly imbricate, smooth in young, at least
the median ones keeled in adults, with apical pits, in 13-21 rows;
ventrals rounded or with a discontinuous lateral keel. Tail long;
subcaudals in two rows.

Range. Africa in forested areas of equatorial belt from French
Guinea east to central Kenya Colony.

Remarks. It is possible that the so-closely related genus Rhamnophis
Giinther, 1862, may ultimately have to be united with Thrasops.
Schmidt (1923, p. 83), however, states that in R. a. ituriensis the
hemipenis has four large spines about its base, the remainder being
calyculate, while in T. j. jacksonii the hemipenis Is heavily spinose on
one side for its entire length. The hemipenes of both T. j. jacksonii
and T. fiavigularis have been described in detail by Bogert (1940,
pp. 58, 59) who finds them almost identical in structure.

loveridge: African snakes 131

Key to the Species

1. Three labials in contact with the lower postocular; midbody scale-rows

15-19; ventrals 175-187; subcaudals 120-140; range: French Guinea

east to Togo occidentalis

(p. 131)

Two, very rarely 3, labials in contact with the lowest postocular; range:

Nigeria east to Kenya 2

2. Midbody scale-rows 13-15; the dorsals much longer than the ventrals;

range: Nigeria south to Cabinda and western Belgian Congo .fiavigularis

(p. 132)

Midbody scale-rows 17-21 ; the dorsals not or but slightly longer than the

ventrals; range: central Belgian Congo east to Kenya Colony 3

3. Midbody scale-rows 19, rarely 17 or 21; ventrals 187-211; range: central

Belgian Congo east to western Tanganyika Territory and western Kenya

Colony j. jacksonii

(p. 134)
Midbody scale-rows 17; ventrals 170-178; range: Mount Kenya to Nairobi

in Central Kenya Colony j. schmidti

(p. 137)

Thrasops occidentalis Parker

1894a. Thraso-ps fiavigularis Boulenger (part), p. 105.
1906. Johnston, p. 832.

1909. Gendre, p. cvi.

1915a. Boulenger (part), p. 206.
1919b. Boulenger (part), p. 283.
1922. Aylmer, pp. 15, 19.

1929a. Werner (part), p. 97.
1933f. Angel (part), p. 115.

1908b. Thrasops jacksoni Sternfeld (not Gunther), pp. 215, 230, figs, 2-3.
1909a. Sternfeld, p. 16, figs. 21-22.
1929a. Werner (part), p. 98.

1915a. Rhamnophis jacksonii Boulenger (part), p. 207.
1915c. Boulenger (part), p. 624.
1919b. Boulenger (part), p. 284.
1919g. Boulenger (part), p. 23.
1922. Aylmer, p. 15.

1940a. Thrasops occidentalis Parker, Ann. Mag. Nat. Hist. (11), 5, p. 273,
figs. 1 and 2a: Axim, Gold Coast.

Description. Rostral subquadrangular, a little broader than deep,
visible from above; nasal divided; internasals as long as the prefront-

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als ; frontal slightly longer than broad, longer than its distance from the
rostral (in adult), longer than its distance from the end of snout (in
young), slightly shorter than the parietals; no enlarged occipitals;
loreal present; preocular 1, separated from the frontal; eye large, its
diameter much greater than its distance from the mouth; postoculars
3, lowest in contact with 3 upper labials; temporals 1 + 1; upper
labials 8, rarely 7, the fourth and fifth entering the orbit; 4 lower labials
in contact with the anterior sublinguals, which are shorter than the
posterior. Midbody scales in 15-19 rows, not longer than the ventrals,
at least the median rows keeled in adults though all may be smooth in
young; ventrals 175-187; anal divided; subcaudals 120-140 pairs.

Color. Above, in adults, black; in young, head and neck olive, sides
of head whitish, the sutures between the scales picked out in black,
dorsum chequered with subrectangular black and yellow spots arranged
in 6 to 8 longitudinal rows. Below, in adults, chin and throat straw
colored, otherwise dark olive; in young, chin and throat whitish, belly
black, the ventrals with alternately two or three transversely oval
yellow spots ; subcaudals yellow margined with black.

Size. Total length of paratype d71 (B.M. 66.1.28.6), 1165 (670 +
495) mm. from Sierra Leone; of 9 holotype (B.M. 1911.6.30.2),
1085 (682 + 403) mm. from Axim, Gold Coast.

Remarks. In the absence of material, I have based the above de-
scription on that of Parker, supplemented by such additional data as
is to be found in the literature cited above.

Localities. French Guinea: Ditinn (Diteien); Labe; Yambering.
Sierra Leone. Liberia: Monrovia. Gold Coast: Axim; Dunkwa.
Togo: Misahohe.

Range. French Guinea east to Togo (Rochebrune's (1884a, pp. 174,
176) records of niger and flarigularis from Senegambia are questionable
and omitted).

Thrasops flavigularis (Hallowell)

1852b. Dendrophis flavigularis Hallowell, Proc. Acad. Nat. Sci. Philadelphia,

p. 205: "Liberia" later corrected to Gaboon.
1857. Hallowell, p. 66.

1857. Thrasops flavigularis Hallowell, p. 67.
1877c. Peters, p. 615.

1888a. Boettger, p. 63 (inc. var. pustulate/,).
1889. Boettger, p. 279 (inc. var. pustulata).

1889. Hesse, p. 267.

1894a. Boulenger (part), pp. 105, 358.

loveridge: African snakes 133

1895a. Bocage, p. 97.

1897b. Boulenger, p. 278.

1897. Sjostedt, p. 35.

1898. Boettger, p. 59.
1898a. Werner, p. 208.
1899a. Werner, p. 138.
1900b. Boulenger. p. 453.
1902a. Werner, p. 344.
1905f. Boulenger, p. 185.
1906L Boulenger, p. 213.
1908a. Sternfeld, pp. 408, 425.
1909b. Sternfeld, p. 16, figs, 18-19.
1911. Lampe, p. 194.

1915a. Boulenger (part), p. 206.

1919b. Boulenger (part), p. 283.

1927d. Witte, p. 324.

1929a. Werner (part), p. 97.

1933L Angel (part), p. 115.

1933m. Witte, p. 90.

1938b. Mertens, p. 47.

1940. Bogert, p. 58.

1940a. Mertens, p. 241.

1940a. Parker, p. 271, fig. 2b.

1872a. Hapsidophrys niger Giinther, Ann. Mag. Nat. Hist. (4), 9, p. 25:
Gaboon.

1875a. Thrasops pustulatus Buchholz & Peters, Monatsb. Akad. Wiss. Ber-
lin, p. 199: Mungo, British Cameroon.

1876a. Peters, p. 119.

For other citations see occidentalis with which it has been long
confused.

Native name. Mduma (near Banana, fide Hesse).

Description. Rostral subquadrangular, about once and a third to
once and a half as broad as deep (in the middle), visible from above;
nasal divided, semidivided or entire; internasals broader than long (in
young) or as broad as long (in adults), about as long as the prefrontals;
frontal once and two-thirds to twice as long as broad (in the middle),
as long as its distance from the rostral (adult) or longer than its dis-
tance from the end of the snout (young), as long as the parietals; as
broad as, or narrower than, a supraocular; loreal present; preocular 1,
rarely 2, separated from the frontal; eye large, its diameter much
greater than its distance from the mouth ; postoculars 3, the lowest in
contact with 2, very rarely 3, upper labials; temporals 1 + 1; upper

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labials 8, rarely 9, the fourth and fifth or rarely fifth and sixth entering
the orbit; 3-5 lower labials in contact with the anterior sublinguals,
which are shorter than the posterior. Midbody scales in 13 or 15 rows,
much longer than the ventrals, at least the median dorsals keeled in
adults, all smooth in young; ventrals 196-215; anal divided; subcaud-
als 128-146 pairs.

Anatomy. Both dentition and hemipenes are discussed under the
generic definition.

Color. Above, in adults, uniformly black with a silken lustre; in
half-grown or young, dark brown or dark olive, the neck yellow with
black-tipped scales and orange on the sides, dorsum chequered with
black and yellow spots, the former predominating. Below, in adults,
black, though paler than dorsum, uniform, or the throat yellowish,
grayish, or brownish white; in half-grown or young, chequered black
and yellow on belly, with roundish yellow spots disposed alternately
on the inner and outer part of successive scales on belly and tail.

Boettger (1889, p. 279) also furnishes detailed color descriptions.

Size. Total length of unsexed record 2000 (1440 + 560) mm. from
Isongo (fide Lampe); of 9 (type of niger, Brit. Mus.), 1552 (1120 +
432) mm. from Gaboon.

Diet. Mammal remains in a Metet snake (Bogert) ; a chameleon (C.
gracilis etiennei) in a Povo Nemlao reptile. (Boettger).

Habitat. Arboreal in primary forests.

Localities. Nigeria. British Cameroon: Bibundi; Bota; Buea;
Isongo; Mungo; Tiko; Victoria. French Cameroon: Bipindi;
Dehane; Ebolowa; Kribi; Metet; Pungo Songo; Sakbayeme;
Yaunde. Fernando Po: Moka. Spanish Guinea. French
Congo: Gaboon; Loango River. Belgian Congo: Bikori (?
Bikoro); Ganda Sundi; Kwango (Kuango) River; "Mayon"; Povo
Nemlao nr. Banana; Temvo nr. Mayumbe; Vista. Cabinda :
Chinchoxo.

Range. Nigeria south to Cabinda and western Belgian Congo
(Records of flavigidaris and jacksonii from west of Nigeria are refer-
able to occidentalis Parker, 1940).

Thrasops jacksonii jacksonii Giinther

1895. Thrasops Jacksonii Giinther, Ann. Mag. Nat.- Hist. (6), 15, p. 528:

Kavirondo, Kenya Colony.
1910a. Sternfeld, p. 20, figs. 18-19.
1923. Schmidt, p. 85. fig. 6.

loveridge: African snakes 135

1924b. Loveridge, p. 5.

1928j. Loveridge, p. 75.

1929a. Werner, p. 98.

1933f. Angel, p. 116, figs. 44-44a (not 43-43a; captions transposed).

1940a. Parker, p. 271, fig. 3.

1896d. Rhamnophis jacksonii Boulenger, p. 632.

1902a. Boulenger, p. 446.

1909. Peracca, p. 172.

1911c. Boulenger, p. 165.

1912. Hobley, p. 49.

1915a. Boulenger (part), p. 207.

1915c. Boulenger (part), p. 624.

1916a. Loveridge, p. 84.

1919b. Boulenger (part), p. 284.

1919g. Boulenger (part), p. 23.

1923e. Loveridge (part), p. 879.

1933m. Witte, p. 90 (but Temvo record should be checked).

1934a. Schwetz, pp. 381, 383.

1934b. Schwetz, p. 24.

1905b. Thrasops Rothschildi Mocquard, Bull. Mus. Hist. Nat. Paris, 11, p.

287: "Afrique orientale anglaise."

1915c. Boulenger, p. 624.

1923e. Loveridge (part), p. 879.

1929a. Werner, p. 97.

1936j. Thrasops jacksonii jacksonii Loveridge, p. 249.

1936. Pitman, p. 52, pi. vii, fig. 1, pi. G, fig. 4.

1937c. Loveridge, p. 274.

1937f. Loveridge, p. 502.

1938a. Pitman, pp. 210, 231.

1938b. Pitman, pp. 39, 107, 308, 329, pis. and figs, as above (reprint).

1940. Bogert, p. 58.

1942e. Loveridge, p. 277.

Further citations of 'jacksonii' and 'rothschildi' will be found under
Rhamnophis a. elgonensis, T. occidentalis and T. j. schmidti.

Native names. Mambala (at Stanleyville, fide Schwetz); wahimbiri
(Wamba and Toro); ntemankima (Ganda); isilukanga (Gishu for
olivaceous halfgrown examples); yakobe (Gishu for black adults).
Probably confused with Dispholidus typus.

Description. Rostral subquadrangular, about once and a quarter to
once and a half as broad as deep (in the middle), visible from above;
nasal divided, semidivided or entire; internasals broader than long (in
young) or as broad as long (in adults), about as long as the prefrontals;

136 bulletin: museum of comparative zoology

frontal once and a third to twice as long as broad (in the middle), as
long as, or longer, or shorter than, the parietals ; as broad as, or broader
than, a supraocular; loreal present; preoculars 1 or 2, separated from,
or in contact with, the frontal; eye large, its diameter much greater
than its distance from the mouth; postoculars 3, rarely 4, the lowest in
contact with 2, very rarely 3, upper labials; temporals 1 + 1; upper
labials 8, rarely 9, the fourth and fifth or rarely fifth and sixth entering
the orbit; 4 or 5 lower labials in contact with the anterior sublinguals,
which are subequal to, or shorter than, the posterior. Midbody scales
in 17 or 19, very rarely 21, rows, not or but scarcely longer than the
ventrals, at least the median dorsals keeled in adults (probably all
smooth in young); ventrals 187-211; anal divided; subcaudals 130-
155 pairs.

Dentition. Maxillary teeth 18, subequal, followed after a diastema
by 3 enlarged ones (Bogert, based on snakes from Kampala and
Lukolela).

Anatomy. The hemipenes are described by Bogert, as follows:
"Everted on the Lukolela specimen, single, sulcus undivided. Greatly
enlarged basal spines decreasing in size distally and merging into
fringed reticulate calyces at the end. The organ appears to be nearly
identical with that of T. flavigvlaris." (vide ante).

Color. Above, in adults, uniformly black with a silken lustre; in
halfgrown or young, dark brown or dark olive, the neck yellow with
black-tipped scales and orange on the sides; dorsum chequered with
black and yellow spots, the former predominating. Below, in adults,
black, though paler than dorsum, uniform, or the throat yellowish,
grayish, or brownish white; in halfgrown or young, chequered black
and yellow on belly, with roundish yellow spots which are sometimes
disposed alternately on the inner and outer part of successive sub-
caudals.

Size. Total length of d" (A.M.N.H. 12288), 1900 (1320 + 580)
mm.; of 9 (A.M.N.H. 12290), 2160 (1550 + 610) mm., both from the
Belgian Congo (fide Schmidt).

Breeding. At Sipi and Butandiga on the western slopes of Mt.
Elgon, four gravid females were taken between December 14, 1933
and January 11, 1934. The number of eggs varied from 7 to 12,
average 9, and in size ranged from 19 x 8 mm. to 35 x 8 mm. (Lover-
idge).

Diet. Mammals, such as the tree rat (Oenomys b. editvs), a bird,
lizard (Agama atricolHs) and chameleons (C. senegalensis, C. b. bitaen-
iatus and C. b. hohnelii) were found in their stomachs (Loveridge).

loveridge: African snakes 137

Parasites. Hemogregarines in Stanleyville snakes (Schwetz), and
ticks on Uganda specimens (Pitman).

Defence. According to Christy this snake distends its neck like a
cobra. If confirmed, it is probably achieved by inflation as is prac-
tised by the Boomslang (Dispholidus typus) to whose black phase
T. j. jacksonii bears so striking a resemblance that it deceives even
herpetologists. The snake described as T. j. mossambicus by Mertens,
was only a Boomslang, whose grooved teeth were overlooked.

Habitat. Found in primary forest, being essentially an arboreal
species.

Localities. Belgian Congo: Albert ville; Avakubi; Bosabangi;
Buta; Diambo; Eala; Kasai basin; Leopold ville; Lukolela; Medje;
Niangara; Temvo (fide Witte); Upper Mulinga on Idjwi Id.,
Lake Kivu. Tanganyika Territory: Kabare near Bukoba.
Uganda: Bundibugyo; Bussu; Butandiga; Entebbe; Fort Portal;
Jinja; Kampala; Kilembe; ? Kitala; Mabira Forest; Sipi; Toro.
Kenya Colony: Kaimosi; Kakamega; Kavirondo; Yala River.

Range. Central Belgian Congo east to western Tanganyika Terri-
tory and western Kenya Colony.

Thrasops jacksonii schmidti Loveridge

1911. Thrasops rothschildi Lonnberg (not Mocquard), p. 22.

1912. Hobley, p. 49.

1928j. Thrasops jacksonii Loveridge (part), p. 75.
1923e. Loveridge, p. 879.

1936f. Thrasops jacksonii schmidti Loveridge, Proc. Biol. Soc. Washington,
49, p. 63: Meru Forest, Mount Kenya, Kenya Colony.

Description. Rostral subquadrangular, about once and a third as
broad as deep (in the middle), visible from above; nasal divided;
internasals broader than long (in half grown), about as long as the pre-
frontals; frontal twice as long as broad (in the middle), longer than its
distance from the rostral (half grown), as long as the parietals, as broad
as a supraocular ; loreal present ; preocular 1 , separated from the frontal ;
eye large, its diameter much greater than its distance from the mouth;
postoculars 3, the lowest in contact with 2 upper labials; temporals
1 + 1 ; upper labials 8, the fourth and fifth entering the orbit; 4 or 5
lower labials in contact with the anterior sublinguals, which are shorter
than the posterior. Midbody scales in 17 rows, not or scarcely longer
than the ventrals, faintly keeled (in halfgrown); ventrals 170-178;
anal divided; subcaudals 140-144 pairs.

138 bulletin: museum of comparative zoology

Color. Above, in adults, uniformly black with a silken lustre; in
half grown type, brownish olive. Below, in adults, black; in half grown
type, grayish white becoming gray beneath tail with a median darker
gray line posteriorly.

Size, Total length of type d" (M.C.Z. 9276), 1065 (700 + 365) mm.
from Meru Forest; of ? 9 (Nairobi Museum), 2255 (1671 + 584) mm.
from Muthaiga. That is to say, almost 73^ feet.

Localities. Kenya Colony: Meru Boma and Forest; Muthaiga,
near Nairobi.

Range. Central Kenya Colony.

Genus Duberria

1826. Duberria Fitzinger (part), Neue Class. Rept., p. 29 (type Coluber

canus Linnaeus "und Consorten").
1830. Homalosoma Wagler, Syst. Amphib., p. 190 (n.n. for Duberria).
1894a. Boulenger, p. 274, fig. 19.

Maxillary short, with 10-12 teeth, subequal; anterior mandibular
teeth longest. Head small, not distinct from neck; eye small, with
round pupil; nasal entire or very rarely semidivided; loreal small or
absent; 1, very rarely 2, preoculars. Body cylindrical, short; scales
smooth, with apical pits, in 15, very rarely 16, rows; ventrals rounded.
Tail short; subcaudals in two rows.

Range. East and Central Africa from Ethiopia and Uganda (chiefly
in montane grasslands of equatorial belt) southwards to the Cape (but
not recorded from Angola and South West Africa).

Remarks. Boulenger (1894a), who should be consulted for further
generic synonymy, correctly followed Cope (1864) in using Pseudaspis
Fitzinger (1843) for Coluber canus Linnaeus (1758) of which it was the
genotype, thus Duberria Fitzinger (1826) was left unused. When pro-
posing Duberria, Fitzinger included among its species Coluber arcti-
ventris Daudin (1803) = Coluber duberria Merrem (1790) = Coluber
lutrix Linnaeus (1758) which consequently becomes the type by
tautonomy of the genus Duberria. Homalosoma was proposed by
Wagler (1830) merely as a substitute name for Duberria, as is shown
by his footnote 4 on p. 190.

Bogert (1940, p. 39) remarks that the anterior sixth of the maxilla
is devoid of teeth or sockets, and that the maxillary teeth are more
widely spaced than is usually the case with snakes. Like Peters (1882a,
pi. xvi, fig. 1), he found 10 maxillary teeth, whereas Boulenger (1894a,
fig. 19) shows 11 and gives from 10 to 12 in the text.

loveridge: African snakes 139

For various comments on the following Key1, see Loveridge (1942e,
pp. 279-280) where the following scale-counts are accepted.
D. variegata Range of ventrals 97-110, subcaudals 25-36.

D. I. tutrix Range of ventrals 120-134, subcaudals 25-51.

D. I. shirana Range of ventrals 126-151, subcaudals 24-46.
D. I. abyssinica Range of ventrals 118-149, subcaudals 17-39. ■

Key to the Species

1. Ventrals 97-110; postoculars 2; a loreal; belly reticulated black and white;

range: Zululand northwards to Inhambane, Mozambique variegata

(p. 147)
Ventrals 118-151 2

2. Postoculars usually 2 (87%, 1 in 13%); a loreal (absent in 7%); belly

yellowish in middle; range: highlands and lowlands of Africa south of the

Zambesi I. tutrix

(p. 144)
Postocular usually single 3

3. Postocular 1 (85%, 2 in 15%); no loreal (100%); belly yellowish in middle

rarely dark; range: highlands around Lake Nyasa and southern Tangan-
yika Territory I. shirana

(p. H2)

Postocular 1 (100%); a loreal (absent in 10%); belly usually very dark,

rarely yellowish in middle; range: highlands of northeastern Tanganyika

Territory and western Belgian Congo, north to Uganda and Ethiopia . . .

I. abyssinica (p. 139)

Duberria lutrix ABYSSINICA (Boulenger)

1870. Homalosoma lutrix (not of Linnaeus) Blanford, p. 458.
1896d. Boulenger, p. 642.

1896. Tornier (part), p. 72 (little use has been made of this reference which

contains misprints).

1897. Tornier, p. 65.
1902a. Boulenger, p. 446.
1902b. Mocquard, p. 406.
1909. Peracca, p. 172.

1910a. Sternfeld (part), p. 22 (part text, not fig.).

1912b. Boulenger, p. 332.

1912b. Sternfeld, p. 385.

1912c. Sternfeld, p. 271.

1 Based on 56 lulrix, 50 shirana, and 53 abyssinica counts.

140

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1915a.

1915c.

1915d.

1918a.

1923e.

1924b.

1925a.

1929h.

1930b.

1933.

1933j.

1933m.

1934a.

1894a.

1908c.
1910a.
1912c.

1933h.

1936h.

1936.

1937d.

1937.

1938a.

1938b.

1940.

Boulenger (part), p. 208.

Boulenger (part), p. 625.

Boulenger (part), p. 650.

Loveridge, p. 332.

Loveridge, p. 880.

Loveridge, p. 5.

Angel, p. 33.

Loveridge, p. 28.

Barbour & Loveridge, p. 787.

Schouteden, p. 236.

Witte, p. 123.

Witte, p. 91.

Schwetz, p. 381 (latrix, misprint).
Homalosoma abyssinicum Boulenger, Cat. Snakes Brit. Mus., 2, p.

276, pi. xiii, fig. 2: Lake Ashangi, Ethiopia.

Sternfeld, pp. 240, 243.

Sternfeld, p. 22.
Homalosoma tutrix var. atriventris Sternfeld, Wiss. Ergebn. Deut.

Zentral-Afrika-Exped. 1907-1908, 4, p. 271: Kissenje = Kisenyi,

Belgian Ruanda.
Duberria lutrix shiranum Loveridge (not Boulenger), p. 241.

Loveridge, p. 34.

Pitman, p. 61, pi. vii, fig. 5, pi. H, fig. 1.

Mertens, p. 7.

Uthmoller, p. 112.

Pitman, pp. 211, 231.

Pitman, pp. 116, 309, 329, pis. as above (reprint).

Bogert (part), p. 39 (Fort Portal specimen only).
Duberria lutrix abyssinicum Bogert, p. 40.

Names. Abyssinian Slug-eater; bulifu (Kiga).

Description. Rostral once and a third to once and three quarters
as broad as deep, visible from above; nasal entire, very rarely
semidivided; internasals broader than long, much shorter or about
as long as the prefrontals; frontal once and a quarter to nearly
twice as long as broad, longer than its distance from the end of the
snout, as long as, or slightly longer or slightly shorter than, the
parietals; once and a quarter to once and two thirds as broad as a
supraocular; loreal small, rarely absent; preocular 1 ; eye moderate,
its diameter greater than its distance from the mouth; postocular 1,
rarely 2; temporals 1+2, rarely 1 -f- 1 ; upper labials 6, very rarely
5 or 7, the third and fourth, or very rarely the third only, or third,
fourth and fifth, entering the orbit; 3, rarely 4, lower labials in con-
tact with the anterior sublinguals, which are subequal with, or

loveridge: African snakes 141

shorter than, the posterior. Midbody scales in 15 (rarely 161)
rows, smooth; ventrals 110-1492; anal entire; subcaudals 17-392
pairs.

Color. Above, dark olive or olive brown or blackish, usually a
fine, more or less continuous, black vertebral line; sides dark,
flecked with white. Below, usually deep gray-black, more or less
variegated with lighter, throat sometimes yellow, this color rarely
continued as a median stripe as far as the anal shield which may be
spotted with yellow.

Size. Total length of d> (M.C.Z. 48354), 332 (281 + 51) mm.
from Mushongero, Lake Mutanda; of 9 (M.C.Z. 34921), 434
(384 -f- 50) mm. Lake Bunyoni.

Sexual dimorphism. Subcaudals of females range from 17-27, in
males from 27-39. The latter figure, however, is an unsexed record
of Sternfeld's, the highest count on an M.C.Z. male being 37.

Breeding. Eight out of thirteen Kigezi females examined between
October and November, by Pitman, held from 6 to 12 eggs. Others
examined in June and July by the same author were gravid. Four
females from Nyakabande, Kigezi, examined on January 27, 1939,
by Loveridge held, respectively, 7 eggs (measuring 12 x 9 mm.),
10 eggs (8x5 mm.), 11 eggs (14 x 7 mm.) and a fourth with large
embryos. At Kabare, Bukoba, January 10, 1923, a female, though
small (290 + 38 mm.), held 10 eggs (12 x 8 mm.). At Lulenga,
Ruanda, March 1, 1927, a 9 held at least two embryos, which
were uniformly plumbeous above, blue-gray below, measuring cf
103 (86 + 17) mm., and 9 100 (85 + 15) mm.

Diet. Slugs in four Nyakabande snakes; eggs (fide Sternfeld) in
stomach of type of atriventris.

Parasites. Worms were observed in Kigezi snakes by Pitman.

Temperament. Pitman (1938b, p. 118) writes: "Its general
demeanour suggests inoffensiveness. I have caught and handled
numerous specimens and very rarely have any attempted to bite.
They are as a rule most docile, placid and friendly, and within a
few moments of capture even the most frightened has become
tame and confiding!" and adds that, though abundant, the Bakiga
do not think it is harmless.

Habitat. Upland country (3,000 to 10,000 feet) with short grassy
tussocks.

i Bogert (1940, p. 39) in a Fort Portal snake.

2 Pitman's (1938b, p. 117) record of 151 ventrals and 46 subcaudals, rejected pending con-
firmation as possibly based on a native's counting.

142 bulletin: museum of comparative zoology

Range. Highlands of Central Lake region northwest through
Uganda to Ethiopia, south in highlands of Kenya and northern
Tanganyika.

Localities. Ethiopia: Gara Mulata; Grau; Lake Ashangi; Webi
Mana. Uganda: Bufundi; Fort Portal; Harutindo; Hoima to
Kampala; Kisolo (Kissoro); Kitagueta; Lake Bunyonyi; Lake
Chahafi; Mityana; Muko; Mushongero; Niwashenya, s.of Kishasha
Valley; Nyakabande. Kenya Colony: ? Aberdare Mountains;
Kinangop Plateau; Meru; Nairobi; Nyeri (Ndjiri). Tanganyika
Territory: Amani; Arusha; Bukoba; Gomberi; Kabare, Bukoba;
Kilema, Kilimanjaro Mountain; Kilimanjaro to Teita; Marungu;
Moshi; Ngorongoro. Belgian Ruanda: Katana, Lake Kivu;
Kisenyi; Lulenga; Volcano region. Belgian Congo: Boundary
Mountains N. W. of Lake Tanganyika; Ngoma; Rutshuru; Stan-
leyville {fide Schwetz).

Duberria lutrix shirana (Boulenger)

1893. Homalosoma lutrix Giinther (not Linnaeus), 1892, p. 555.

1898. Johnston, p. 361a.

1894a. Homalosoma shiranum Boulenger, Cat. Snakes Brit. Mus., 2, p. 276,

pi. xiii, fig. 1 : Shire Highlands, Nyasaland.
1896a. Bocage, p. 103.

1933h. Duberria lutrix shiranum Loveridge, p. 241 (but range wrong).
1934. Pitman, p. 295 (lists only).

1940. Bogert (part), p. 39 (exclude Portal specimen).

Further citations of 'shiranum' will be found under D. I. abys-
sinica.

Names. Shire Slug-eater; nyaluhercka (Kinga); isakani (Nya-
kusa). But both Wakinga and Banyakusa consider this small
snake to be the young of Trimerorhinus t. tritaeniatvs .

Description. Rostral once and a half to twice as broad as deep;
visible from above; nasal entire; internasals broader than long,
shorter or longer or about as long as the prefrontals; frontal once
and a quarter to once and a half as long as broad, longer than its
distance from the end of the snout, as long as, or shorter than, the
parietals; once and a half to twice as broad as a supraocular;
loreal absent, very rarely present; preocular 1; eye moderate, its
diameter greater than its distance from the mouth; postocular 1,
rarely 2; temporals 1+2, rarely 1 + 1 ; upper labials 6, very rarely

loveridge: African snakes 143

7, the third and fourth, or very rarely the second, third and fourth,
or third, fourth and fifth, entering the orbit; 3, rarely 2, lower
labials in contact with the anterior sublinguals, which are subequal
with, or shorter or longer than, the posterior. Midbody scales in
15 rows, smooth; ventrals 122-151; anal entire; subcaudals 24-47,
pairs.

Color. Above, dark olive, olive brown, red brown or black,
usually a fine, more or less continuous, black vertebral line. Below,
sometimes deep gray black more or less variegated with lighter,
sometimes the dorsal coloration extending on to the ventrals but
leaving a median stripe of pale yellow or white down the centre.

Size. Total length of & (M.C.Z. 30184), 379 (300 + 79) mm.
from Mangoto, Ubena Mtns.; of 9 (M.C.Z. 30174), 412 (362 +
50) mm. from Kigogo, Uzungwe Mtns.

Sexual dimorphism. Subcaudals in females range from 25-38, in
males from 40-47. Length of tail included in total length from 4.2
to 5.1 times in males, from 6 to 9 times in females.

Breeding. From Rungwe Mountains, Bogert records females
holding from 6 to 17 eggs, measuring from 8 x 7 to 13 x 6 mm. He
does not observe much correlation between size of snake and the
number of eggs produced; it seemed to me that the number increases
with the size of the snake though doubtless waning with declining
fertility in old individuals. Condensed, my records read: At Kigogo
Uzungwe Mtns., January 23 & 30, 1930, seven females held 8 to
13 eggs, measuring from 9 x 6 to 12 x 8 mm., all the larger being
on the later date. At Madehani, Ukinga Mtns., February 14, 1930,
five females held from 7 to 12 eggs, one batch measured, being
10 x 6 mm. At Mangoto, Ubena Mtns., February 10, 1930, an
evidently recently born young male measured 118 (93 -+- 25) mm.

Diet. Almost exclusively slugs, which are taken by even very
young snakes. The only other food found, together with a slug,
was a 125 mm. D. I. shirana in the stomach of a larger Madehani
snake of the same species.

Parasites. Nematode worms in stomachs and intestines of
Dabaga and Kigogo snakes.

Enemies. Cannibalistic as related above, though it is possible
that the young snake was engaged in swallowing the slug and that
this was the incentive for the larger snake to attack so that the
engulfing of the young snake was only incidental!

Habitat. They like to bask on the grassy tussocks where their
olivaceous color renders them inconspicuous, such tussocks occurred

144 bulletin: museum of comparative zoology

on hillsides where hoeing by natives resulted in uncovering a good
many, for on being disturbed these snakes seek refuge in the loose
soil at the base of the tussocks. Others were found in rich, but
short, grass in the marshlands of highland valleys.

In habitat then, as well as in diet, size, and a somewhat similar
appearance, D. I. shirana fills a niche in the East African fauna similar
to that occupied by Storeria d. dekayi in eastern North America.

Near Kigogo a settler, whose native employees were clearing land
for planting coffee, informed me that "blind snakes" were very
abundant, and that in digging a furrow forty feet in length they had
destroyed over ninety of them! As I found no Typhlops in the Uzungwe
Mountains I concluded that he referred to Duberria. In view of the
great economic value of these snakes in a coffee plantation by reason
of their diet of slugs they deserve protection by all intelligent settlers :
the same applies to Typhlops which subsists almost entirely on ter-
mites with an occasional caterpillar or slug.

Localities. Tanganyika Territory : Dabaga; Ihanganya; Ilolo;
Kigogo; Mangoto; Madehani; Rungwe Mountain; Tandala;
Ugano. (All these localities being in the Matengo, Ubena, Ukinga,
Uzungwe or Rungwe highlands). Nyasaland: Shire highlands.

Range. Highlands of southern Tanganyika Territory and Nyasa-
land.

Duberria lutrix lutrix (Linnaeus)

1735. Hydra zeylanica Seba, Rerum Nat. Thesauri, 2, p. 2, pi. i, fig. 6:

"Ceylon".
1735. Serpens eximia Seba, Rerum Nat. Thesauri, 2, p. 92, pi. lxxxvi, fig. 5:

Africa.
1758. Coluber lutrix Linnaeus, Syst. Nat. ed. 10, 1, p. 216: "Indiis."
1766. Linnaeus, 1, p. 375 (275 misprint).

1788. Gmelin, 1, p. 1086.

1790. Coluber Duberria Merrem, Beytr. Naturg., p. 7, pi. i: No locality

(after Seba).

1801. Elaps Duberria Schneider, 2, p. 297.

1802. Coluber tetragonus Latreille, Hist. Nat. Rept., 4, p. 97: "? France."
1803c. Daudin, p. 207.

1803c. Coluber arctiventris Daudin, Hist. Nat. Rept., 7, p. 221: n.n. for

duberria Merrem.
1820. Kuhl, p. 82.

1804. Coluber erathon Hermann, Observat. Zool., p. 273: "India orientali."
1826. Duberria arctiventris Fitzinger, p. 55.
1830. Homalosoma arctiventris Wagler, p. 191.

loveridge: African snakes 145

1849. Smith, A., App., p. 16.

1837. Calamaria arctivenlris Schlegel, 2, p. 36, pi. i, figs. 24-26.

1854. Homalosoma tutrix Dumeril & Bibron, 7, p. 110.

1858. Gunther, p. 20.

1862. Jan, 2, p. 33.

1865. Jan, livr. 13, pi. iii, fig. 3.

1867a. Steindachner, p. 59.

1884a. Rochebrune, p. 152 (this is erroneous).

1885a. Miiller, p. 142.

1887b. Boettger, p. 156.

1887h. Boulenger, p. 175.

1891a. Matschie, p. 609.

1894a. Boulenger, p. 274.

1896. Tornier (part), p. 72 (Cape material).

1898. Boettger, p. 77.

1898. Jeude, p. 35.

1898. Werner, 1896-7, p. 143.

1902. Lampe & Lindholm, p. 29.

1907a. Roux, p. 77.

1907c. Roux, p. 735.

1908b. Boulenger, p. 229.

1908. Gough, p. 25.

1909b. Chubb, p. 35.

1910b. Boulenger, p. 509.

1910a. Hewitt, p. 57.

1910a. Sternfeld (part), p. 22, fig. 25.

1910b. Sternfeld (part), p. 21, fig. 22.

1912. FitzSimons, F. W., p. 90.

1913. Hewitt & Power, p. 162.
1916. Andersson, p. 40.
1922c. Angel, p. 357.

1929. Rose, p. 152, fig. 97.

1929a. Werner (part), p. 150, fig. 44.

1935. Power, p. 334.

1868. Cyclophis catenatus Theobald, Cat. Rept. Asiatic Soc. Mus., p. 49:

"Simla, India."

1908. Homalosoma shiranum Gough (not of Boulenger), p. 25.

1933h. Duberria lutrix tutrix Loveridge, p. 242.

1 939b. FitzSimons, V. , p. 2 1 .

1940. Bogert, p. 39.

1937e. Duberria lutrix Hewitt, p. 52.

Names. Russet Slug-eater (English); rooislang (Dutch).
Description. Rostral once and a third to nearly twice as broad as
deep, visible from above; nasal entire; internasals broader than long,

146 bulletin: museum of comparative zoology

about as long as the prefrontals; frontal once and a half to twice as
long as broad, longer than its distance from the end of the snout, as
long as, or slightly longer or slightly shorter than, the parietals, once
and a half to twice as broad as a supraocular; loreal small, rarely trans-
versely divided or absent; preocular 1, very rarely 2; eye moderate, its
diameter greater than its distance from the mouth; postoculars 2,
lower sometimes minute, or 1 only; temporals 1+2, rarely 1 + 1 or
1 + 2; upper labials 6, the third and fourth entering the orbit; 3,
rarely 4, lower labials in contact with the anterior sublinguals, which
are subequal with, or longer than, the posterior. Midbody scales in 15
rows, smooth; ventrals 120-144; anal entire, very rarely divided1;
subcaudals 24-51, pairs. (21-46 fide Boulenger).

Color. Above, brick red, reddish brown, pale brown, olive or
yellowish, with or without a vertebral series of fine dark dashes; flanks
gray or plumbeous, usually sharply distinct from dorsal coloring.
Below, white, cream or yellowish, the outer edges of the ventrals gray,
usually flecked or spotted with black.

Size. Total length of d1 (M.C.Z. 42639), 364 (297 + 67) mm.,
from Port St. John; of 9 (M.C.Z. 11921), 377 (322 + 55) mm., from
Cape Town. Both surpassed by Boulenger's unsexed (?cf ) record of 390
(325 + 65) mm.

Sexual dimorphism. In Museum of Comparative Zoology material
the subcaudals of females range from 25-322, in males from 33-49.

Breeding. Viviparous, producing from 3 to 10 young in February
(F. W. FitzSimons).

Diet. Mainly slugs (Rose), also snails, insects and their larvae
(Hewitt).

Enemies. Two in gullet of Secretary Bird (Sagittarius serpentarius)
(Anderson).

Defense. When alarmed, curls up like a roll of tobacco (Hewitt).

Habits. Slow-moving and, though not a burrower, highly secretive,
being found in loose soil about the base of bushes, beneath fallen leaves
and pine needles, and in dry grass (Smith, Rose, Hewitt).

Habitat. Common alike in the coastal districts of Cape Province
and the high plateaux of the interior (Hewitt).

Localities. Mozambique : Rikatla. Southern Rhodesia : Bula-
wayo; Chirinda Forest. Transvaal: Barberton; Belfast; Haenerts-
burg; Irene; Johannesburg; Lydenburg; Mariepskop; Mphome;

1 In Chirinda Forest specimen (Transvaal Mus. 16185), fide FitzSimons (1939b, p. 21).

2 The alleged 9 with 48 recorded by Bogert (1940, p. 39), proved on re-examination to have
been a d\ fide Bogert (letter of 17, VI, 40).

loveridge: African snakes 147

Potchefstroom ; Pretoria; Sabi; Woodbush. Zululand: Melmoth'
Natal: Durban; Hilton Road, Merebank. Orange River Colony
{fide F. W. FitzSimons).Cape Province: Albany; Burghersdorp;
Caledon; Cape Town; Ceres; Grahamstown; East London;
Fransche Kraal; Kalk Bay; Knysna; Little Namaqualand; Malms-
bury; Middleburg; Paarl; Port Alfred; Port Elizabeth; Port St.
John; Sir Lowry's Pass; Stellenbosch; Table Mountain; Tokai.

Range. Africa south of the Zambesi exclusive of South West Africa
and Angola.

Duberria variegata (Peters)

1854. Homalosoma variegatum Peters, Monatsb. Akad. Wiss. Berlin, p. 622.

Inhambane, Mozambique.

1855. Peters, p. 51.

1882a. Peters, p. 107, pi. xvi, fig. 1.

1888d. Boulenger, p. 140.

1894a. Boulenger, p. 276.

1896a. Bocage, p. 93.

1898. Sclater, p. 99.

1908b. Boulenger, p. 229.

1910b. Boulenger, p. 509.

1912. FitzSimons, F. W., p. 90.

1929a. Werner, p. 151.

Description. Rostral broader than deep, visible from above; nasal
entire; internasals slightly longer than the prefrontals; frontal once
and a half as long as broad, longer than its distance from the end of
the snout, as long as the parietals; nearly twice as broad as a supra-
ocular ; loreal small ; preocular 1 ; eye rather larger than in lutrix, its
diameter greater than its distance from the mouth; postoculars 2;
temporals 1 + 2 ; upper labials 6 or 7, the third and fourth or fourth and
fifth entering the orbit; 3 lower labials in contact with the anterior sub-
linguals. Midbody scales in 15 rows, smooth; ventrals 97-110; anal
entire; subcaudals 25-36, pairs.

Color. Above, dark or olive brown with three series of dark brown
spots or irregular lichen-like brownish-white variegation. Below,
reticulated black and white.

Size. Total length 250 (217 + 33) mm.

Sexual dimorphism. Subcaudals in a female were 25, in a male 36.

Remarks. In the absence of material, the foregoing is adapted from
Boulenger (1894a, p. 276).

148 bulletin: museum op comparative zoology

Localities. Mozambique: Delagoa Bay; Inhambane; Lorenzo
Marques. Zululand: Mseleni.

Range. Mozambique to Zululand. (Senegambia in error by Roche-
brune, 1884a, p. 152).

Genus Thelotornis

1849. Thelotornis A. Smith, 111. Zool. S. Africa, 3, App. p. 19 (type capensis

Smith.)
1859. Cladophis A. Dumeril, Arch. Mus. Hist. Nat. Paris, 10, p. 204 (type

Leptophis kirtlandii Hallowell).

For further partial synonymy see Boulenger (1896d, p. 184).

Maxillary teeth 11-17, gradually increasing in size, followed after
an interspace by 2-3 enlarged grooved fangs situated below the pos-
terior border of the eye; anterior mandibular teeth strongly enlarged.
Head distinct from neck, with strong canthus rostralis ; eye large with
horizontal pupil; nasal entire; 2, rarely 1 or 3, loreals; preocular.
Body cylindrical; scales oblique, narrow, slightly keeled, with apical
pits, in 19 rows, of which the vertebral is not enlarged; ventrals
rounded. Tail long, subcaudals in two rows.

Range. Africa south of 15° N., i.e. Portuguese Guinea and northern
South West Africa, east to Italian Somaliland and Natal.

Remarks. Boulenger (1896d, p. 185), from whom the above descrip-
tion is largely taken, with increase in range of maxillary teeth from
Bogert (1940, pp. 69, 71), remarks that the ectopterygoid bone is
forked, the two branches articulating with the maxillary; an arrange-
ment which he states is unique among ophidia. Bogert describes the
hemipenes for both races.

Schmidt (1923, p. 113) points out that: "The depressed and flat
head, with the canthus rostralis distinctly projecting, forming a shal-
low loreal groove, is very characteristic, distinguishing the species at
once from all other African snakes." In the field one is most likely to
mistake it for the slender, vine-like, bush-climbing Psammophis
biseriatus, and I have confused it with the more blunt-headed young of
its near relative, Dispholidus typus, the latter being vinaceous colored
with white labials.

Key to the Races

Rostral and anterior ends of nasals broadly visible from above; crown of head
immaculate, labials more or less immaculate, neck crossbanded; range: Portu-

loveridge: African snakes

149

guese Guinea to northern Angola, east to southern Somaliland and1 central

Tanganyika Territory k. kirtlandii

(p. 149)
Rostral and anterior ends of nasals narrowly visible from above; crown of head
speckled with black, labials heavily speckled with black, neck not crossbanded
though black lateral blotches usually present; range: central Angola and1
northern South West Africa, east to central Tanganyika Territory and Natal.

k. capensis (p. 154)

Thelotornis kirtlandii kirtlandii (Hallowell)

1844. Leptophis Kirtlandii Hallowell, Proc. Acad. Nat. Sci. Philadelphia,

p. 62: Liberia.

1854a. Oxybelis Lecomtei Dumeril & Bibron, Erpet. Gen., 7, p. 821: Gaboon.

1854a. Tragops rufulus Dumeril & Bibron, Erpet. Gen. ,7, p. 827: Senegal.

1854a. Dryophis Kirtlandi Hallowell, p. 100.
1858c. 2Gunther, p. 156.
1863a. Giinther, p. 22.
1866a. Bocage, p. 48.
1869. Jan, livr. 32, pi. vi. fig. 2.

1885d. Muller, p. 684.
1886. Martinez y Saez, p. 339.

1888a. Boettger, p. 65.
1889. Hesse, p. 267.

1889. Mocquard, p. 145.
1893. Prato, p. 13.
1895a. Bocage, p. 119.
1895c. Bocage, p. 13.
1903a. Bocage, p. 44.

1856b. Oxybelis violacea Fischer, Abhand. Nat. Ver. Hamburg, 3, p 91 pi.

ii, fig. 7: Edina, Grand Bassa County, Liberia.

1857. Oxybelis Kirtlandii Hallowell, p. 59.

1859. Cladophis Kirtlandii Dumeril, p. 204, pi. xii, fig. 8.
1874. Reichenow, p. 292.

1867b. Thelotornis kirtlandii Peters, p. 235.
1875a. Peters, p. 199.
1884a. Fischer, p. 11.
1884a. Rochebrune, p. 178 (ignored).

1890. Biittikofer, p. 478.
1892. Matschie, p. 110.

1 It should he home in mind that Angola and Tanganyika are areas of intermediates and
that an occasional lowland specimen in the Voi region of southeast Kenya may preponderate
in capensis attributes.

2 This, and some of the following, were spelt Dryiophis.

150

bulletin: museum of comparative zoology

1893c.

1893.

1893b.

1896d.

1896.

1897b.

1897g.

1897.

1898.

1898a.

1899a.

1900b.

1900.

1901b.

1902a.

1905f.

1906i.

1906.

1907.

1908a.

1908b.

1909a.

1909b.

1910.

1910a.

1911.

1911b.

1911a.

1912.

1913.

1913a.

1915a.

1915c.

1915d.

1916a.

1918a.

1919b.

1919g.

1921a.

1921b.

1922.

1923e.

Matschie, p. 212.

Prato, p. 13.

Stejneger, p. 733.

Boulenger (part), p. 185.

Tornier (part), p. 83.

Boulenger, p. 279.

Sjostedt, p. 35.

Tornier, p. 65.

Boettger, p. 107.

Werner, p. 209.

Werner, p. 140.

Boulenger, p. 454.

Ferreira, p. 52.

Tornier, p. 64.

Boulenger, p. 447.

Werner, p. 345.

Boulenger, p. 185.

Boulenger, p. 214.

Johnston, p. 832.

Lonnberg, p. 16.

Sternfeld, pp. 413, 428.

Sternfeld, pp. 219, 233.

Sternfeld, p. 21, fig. 33.

Sternfeld, p. 21, fig. 28.

Miiller, p. 607.

Sternfeld (part), p. 31, fig. 34.

Lampe, p. 201.

Nieden, p. 442.

Sternfeld, p. 251.

Hobley, p. 52.

Lonnberg & Andersson, p. 4.

Werner, in Brehm, p. 403, pi. viii, fig. 3.

Boulenger (part), p. 213.

Boulenger (part), p. 631.

Boulenfer (part), p. 654.

Loveridge, p. 86.

Loveridge, p. 327.

Boulenger (part), p. 290.

Boulenger, p. 26.

Chabanaud, p. 471.

Chabanaud, p. 525.

Noble, p. 168, fig.

Aylmer, pp. 15, 21.

- 887.

'V"">-'! FK- ■*"> "'■•

Loveridge (part), p.

loveridge: African snakes

151

1923. Schmidt, p. 112, pi. xiv.

1924b. 'Loveridge (part), p. 7.

1925. Werner, 1924, p. 131, fig. 5.

1927. Calabresi, p. 56.

1927c, Power, p. 410.

1927d. Witte, p. 325.

1928c. Barbour & Loveridge, p. 128.

1928g. Loveridge (part), p. 34.

1929h. Loveridge, p. 33.

1930a. Barbour & Loveridge, p. 773.

1933f. Angel (part), p. 165, figs, 62, 62a.

1933m. Witte, p. 94.

1934a. Schwetz, p. 382.

1934c. Scortecci, p. 70, fig. 30.

1936h. Loveridge (part), p. 39.

1936J. Loveridge, p. 265.

1936c. Parker, p. 125.

1937c. Loveridge, p. 277.

1937f. Loveridge, pp. 493, 496, 503.

1937. Pitman (part), p. 242, pi. xi, fig. 5.

1937. Uthmoller, p. 120.
1938a. Pitman, pp. 216, 233.

1938. Uthmoller, p. 45.
1939a. Scortecci, p. 283.

1939c. Scortecci (part), p. 159', figs. 88-89.

1940. Bogert, p. 69, fig. 10.

1942b. Thelotornis kirtlandii capensis Bogert (intermediates), p. 2.

1942e. Thelotornis kirtlandii kirtlandii Loveridge, p. 292.

Further citations of 'kirtlandii will be found under k. capensis.

Names. Western Bird Snake or Vine Snake (English); bokarrabai
(Temne : Sierra Leone : Aylmer) ; mbcija (Wamba : Uganda : Loveridge) ;
mraringa (Teita: Kenya: Loveridge); lukukuru (Kami: Tanganyika:
Loveridge); kawaikukoto (Cazengo region, Angola: Ferreira).

Description. Rostral about once and three-quarters to twice as broad
a9 deep, strongly recurved on snout so broadly visible from above;
nasal entire, its anterior end reaching upper surface of snout; inter-
nasals about as broad as long, as long as or shorter than the prefrontals;
frontal semi-bell-shaped, twice to thrice as long as broad (in the mid-
dle), as broad as, or narrower or broader than, a supraocular, as long
as, or slightly shorter or longer than, its distance from the end of the
snout, as long as, or slightly shorter than, the parietals; loreals 2,
sometimes 1 or 3 (or absent fide Hallowell) ; preocular 1 ; eye very large,

152 bulletin: museum of comparative zoology

its diameter much greater than its distance from the mouth; postocu-
lars 3, rarely 2, the lowest in contact with 2 upper labials ; temporals
1 + 2, very rarely 2 + 2; occipitals 2, separated by 1, 2, or 3 smaller
shields; upper labials 8, rarely 7 or 9, the fourth and fifth, or third,
fourth and fifth, or fifth and sixth, entering the orbit; 4, rarely 3 or 5,
lower labials in contact with the anterior sublinguals, which are usually
much shorter than, though sometimes as long as, the posterior. Mid-
body scales in 19l rows which are narrow, very oblique, and feebly
keeled; ventrals 153-189; anal divided; subcaudals 137-175 pairs.

Color. Above, head green, uniform, lips cream-colored or pink,
uniform (in west) or slightly flecked with black (in east); dorsum
pinkish brown speckled and striated with brown, anteriorly heavily
crossbarred with black. Below, vinaceous, gray or white, speckled or
striated with brown. Iris golden (Reichenow). Tongue bright red
with a black tip (A.L.)

Size. Total length of <?, 1422 (821+601) mm., from Mount
Mbololo, Kenya Colony; total length of 9 , 1478 (919 + 559) mm.
from Morogoro, Tanganyika Territory.

Remarks. Dr. Dunn (7. x. 1940) tells me that the type of kirtlandii
cannot be located in the Academy of Natural Sciences of Philadelphia.
Only in recent times has the Southeastern Bird Snake been definitely
accepted as a recognizable geographical race. In this account an
attempt has been made to carefully allocate all available data to its
correct subspecies.

Dentition. The number of teeth anterior to the three enlarged fangs
vary from 11 to 14 according to Bogert (1940, p. 70) whom see for
further discussion.

Anatomy. For a full description of the hemipenes of four Congo
snakes, in which they extended to the sixth subcaudal only, see Bogert
(1940, p. 70).

Breeding. On October 4, a Nyange 9 held 5 eggs, each measuring
15 x 15 mm. On January 16, a Morogoro 9 laid 8 eggs, each measur-
ing 27 x 15 mm.

Diet. Actually birds seem less frequently an article of diet than
arboreal lizards or snakes. A green snake (Chlorophis carinatus) has
been found in a Lukolela specimen (Bogert); while at Nyange, a
captive Bird Snake ate a Chlorophis neglcctus, Neusterophis o. ulugur-
uensis and Crotaphopeltis h. tornieri, and apparently the same fate
befell an Egg-eater (Dasypcltis s. medici) that shared the cage. At

1 My (1929h, p. 33) count of 15 is erroneous, as also Hallowell's (1854, p. 10) of 13, and
Uthmoller's (1934, p. 120) of 17. Their oblique nature often makes an accurate count difficult.

loveridge: African snakes 153

Bundibugyo an Agama atricollis was recovered from a Bird Snake,
while a Buta specimen was found to have swallowed a skink (Mabuya
m. maculilabris) and two large nestlings of a weaver (Spermophaga).

When the Bird Snake seized the large Tornier's Snake it held on
doggedly, occasionally chewing with its poison fangs. The Tornier's
Snake felt about with its tail for twigs or branches on which to get a
purchase. After eight minutes spent in this way the Bird Snake tried
to swallow and was then observed to be in difficulties; the Tornier's
Snake had hooked its teeth into the mucous membrane of the Bird
Snake's mouth. I intervened and separated them whereupon the
Tornier's Snake tried to make off, but the Bird Snake — which had
itself withdrawn — returned swiftly, seized its victim, and began to
swallow again. The Tornier's Snake, being a large one as I have said,
resulted in an unusually laboured deglutition. In all it took an hour
from the moment when the Tornier's Snake was first seized until the
last of it disappeared.

Parasites. Nematode in a Lukolela snake (Bogert), and fragment of
a cestode in one from Nyange (Loveridge).

Defence. The first line of defence of this remarkably vine-like
reptile is cryptic. Partly lying along a branch about which its tail is
entwined, the Bird Snake projects its anterior third far into space and
so remains rigidly motionless except, perhaps, for an occasional flicker
of its black-tipped, scarlet tongue. The bright green top of its some-
what leaf-shaped head assists in the illusion, for the Bird Snake
furnishes one of the finest examples of cryptic coloring to be found
among African snakes.

When molested, however, prior to lunging, the snake assumes a most
threatening attitude, being able, like the Boomslang, to vertically
inflate its anterior third to a surprising extent. This is made possible
by the cartilaginous rings, which support the trachea, being incomplete
dorsally. The result of this inflation is to accentuate the brighter color-
ing of the distended neck, particularly the broad black crossbands.
Miiller (1910, p. 608), who has given an excellent account of this
behavior, states that the excited snake also extends its strangely
colored tongue to the fullest extent, the shiny black tips closely applied
together, or spread so widely apart as to form an angle of 180°.

Habitat. Though essentially an arboreal species, it seems reasonable
to suppose that even the western form descends to the ground at times
in search of prey.

Localities. Portuguese Guinea. French Guinea: Beyla.
Sierra Leone. Liberia: Edina; Gbanga. Gold Coast: Adjah

154 bulletin: museum of comparative zoology

Bippo; Ashanti; Fantee. Togoland: Misahohe. Nigeria: Ifo,
Ondo Province; Oil River. British Cameroon: Johann Albrecht-
shohe; Victoria. French Cameroon: Bipindi; Bitye; Dibongo
near Edea; Ja River; Jossplatte; Kribi; Longji. Spanish Guinea:
Benito River; Corisco; Elobey district. Fernando Po: Bahia de
S. Carlos. French Congo: Gaboon; Loango Mouth; Loudinia-
Niari. Belgian Congo1: Akenge; Avakubi; Banana; Basongo;
Buta; Dika; Elisabeth ville; Epulu Ferry; Ganda Sundi in May-
umbe; Kanzenze; Kasai Kunungu; Lukolela; Mayon; Mayumbe;
Niangara; Niapu; Nyampoko; Poko; Povo Nemlao; Pove Netonna;
Saidi's Village; Stanleyville; Vube. Angola2: Cazengo; Duque de
Braganca; Quirimbo (For southern localities see T. k. capensis).
Uganda: Budongo Forest; Bundibugyo; Entebbe; Fort Portal;
Jinja; Lutoto Hill in w. Ankole. Italian Somaliland: Belet
Amin; Kismayu; Mofi. Kenya Colony: Jilore; Kilibassi;3 Mt.
Mbololo; Sokoki Forest; Tana River; Taveta; Teita Mtns.; Voi3.
Tanganyika Territory:2 Arusha; Dunda on Kingani River;
Kilimanjaro Mtns. Gomberi and Kibonoto; Marangu; Morogoro;
Tumbanatji; Uleia; Uluguru Mtns. Nyange and Vituri; Usambara
Mtns. Amani, Derema, and Mlalo near Ambangula.

Range. Tropical Africa from Portuguese Guinea to northern Angola,
east through Uganda and Kenya to Italian Somaliland and south to
central Tanganyika Territory.

Thelotornis kirtlandii capensis Smith

1849. Thelotornis capensis A. Smith, 111. Zool. S. Africa, 3, App., p. 19:

Kaffirland and the country towards Port Natal.
1940. Bogert, p. 70, fig. 11.

1854. Oxybelis Lecomtei Peters (not Dumeril & Bibron), p. 623.

1855. Peters, p. 52.

1881b. Dryiophis Oatesii Giinther, in Oates, Matabeleland & Victoria Falls

App., p. 330, pi. D.
1889a. Giinther, p. 337, pi. D. (Dnjophis).
1894a. Giinther, p. 618.
1898. Johnston, p. 361a.

i Some of these in the southern Congo may be referable to the race capensis.

2 For further localities see those listed under the race capensis.

3 Though characters of the Voi and Kilibassi specimens are preponderatingly capensis, crown
of head is immaculate.

loveridge: African snakes

155

1882a.

1890b.

1891a.

1896d.

1896.

1897e.

1897.

1898.

1899a.

1907a.

1907J.

1908b.

1908.

1908b.

1908c.

1909a.

1909b.

1910b.

1910.

1910a.

1910b.

1910c.

1912.

1913.

1915a.

1915c.

1915d.

1915.

1915c.

1919b.

1921a.

1923e.

1924b.

1928.

1928d.

1928g.

1931.

1933f.

1933h,
1934.

Thelotornis Kirtlandii Peters (not Hallowell), p. 131, pi. xix, fig. 2.
Boulenger, p. 93.
Boulenger, p. 307.
Boulenger (part), p. 185.
Tornier (part), p. 83.
Boulenger, p. 801.
Tornier (part), p. 65.
Johnston, p. 361a.
Sclater, p. 100.
Werner, 1896-7, p. 146.
Mocquard, p. 219.
Boulenger, p. 11.
Boulenger, p. 487.

Boulenger, p. 229.
Chubb, p. 221.
Gough, p. 32.

Mocquard, p. 558.

Sternfeld, p. 246.

Chubb, p. 596.

Chubb, p. 36.

Boulenger, p. 515.

Peracca, p. 4.

Sternfeld (part), p. 31.

Sternfeld, p. 29, fig. 33.

Sternfeld, p. 56.

FitzSimons, F. W., p. 126.

Peracca, p. 6.

Hewitt & Power, p. 164.

Boulenger (part), p. 213.

Boulenger (part), p. 631.

Boulenger (part), p. 654.

Breijer, p. 113.

Werner, p. 363.

Boulenger (part), p. 290.

Angel, p. 42.

Loveridge (part), p. 887

Loveridge (part), p. 7.

Cott, p. 953.

Loveridge, p. 56.

Loveridge (part), p. 34.

Monard, p. 106.
Power, p. 48.
Angel (part), p. 165.
Loveridge, p. 257.
Pitman, p. 297.

156 bulletin: museum of comparative zoology

1935. Cott, p. 969.

1935. Cunha, p. 11.

1936h. Loveridge (part), p. 39.

1937a. FitzSimons, V., p. 274.

1937b. Monard (part), pp. 128, 135.

1937a. Parker, p. 630.

1937. Pitman (part), p. 242, col. pi. L, fig. 3.

1939b. FitzSimons, V., p. 23.

1939c. Scortecci (part), p. 159.

1941. Moreau & Pakenham, p. 108.

1887h. Dryophis kirtlandii Boulenger (not Hallowell), p. 177.

1893. Pfeffer, p. 86.

1895. Jeude, p. 229.

1895a. Dryiophis Kirtlandii var. mossambicana Bocage, Herp. Angola Congo,

p. 119: Manica, Mozambique (restricted).

1913. Thelotornis kirtlandi var. capensis Boettger, p. 345.

1927. Theborius Kirtlandii (sic) Wyllie, p. 129.

1937b. Thelotornis kirtlandii capensis Mertens, p. 14.
1942e. Loveridge, p. 294.

, Further citation of 'capensis' will be found under k. kirtlandii.

Names. Southeastern Bird Snake or Vine Snake (English); nondo
(Rungu : Tanganyika) ; lukungu (Nyika : Tanganyika) ; nalakutu (Yao
at Dodoma) ; lukukuti (Yao at Kitaya, Tanganyika) ; lukukutu (Konde :
Tanganyika) ; likukuhi (Mawiha : Tanganyika : all Loveridge) ; injaru-
cucutue (Sena : Mozambique : Peters) ; nharicucuto (Sena : Mozambique :
Cott); cucuta (Quando, Angola: Anchieta); nhocamenha (Bibala,
Angola: Bocage); kalakukwiti (N. Rhodesia: Neave); ukotikoti (Mata-
bele : S. Rhodesia : Chubb) ; vogclvreter slang (Dutch : F. W. FitzSimons).

Description. Rostral about once and a quarter to twice as broad as
deep, strongly recurved on snout so moderately or narrowly visible
from above ; nasal entire, its anterior end not, or but scarcely, reaching
upper surface of snout ; internasals about as broad as long, or longer
than broad, as long as, or shorter or longer than, the prefrontals;
frontal semi-bell-shaped, twice to twice and a half as long as broad (in
the middle), as broad as or narrower than a supraocular, as long as, or
slightly shorter or longer than, its distance from the end of the snout,
as long as or slightly shorter than, the parietals ; loreals 2, sometimes 1 ;
preocular 1 ; eye very large, its diameter much greater than its distance
from the mouth; postoculars 3, rarely 2 or 4, the lowest in contact with
2 upper labials ; temporals 1 + 2; occipitals 2, separated by a smaller
shield; upper labials 8, the fourth and fifth, or rarely the third and

loveridge: African snakes 157

fourth, entering the orbit; 4, rarely 5, lower labials in contact with the
anterior sublinguals, which are usually shorter than the posterior.
Midbody scales in 19 rows which are narrow, very oblique, and feebly
keeled; ventrals 147-170; anal divided; subcaudals 131-166 pairs.

Color. Above, head of young pink, uniform, of adult green or
pinkish brown flecked or speckled with darker, a brownish black-dotted
band passing through the eye, an oblique streak below the eye on the
upper lip which is cream or pink speckled with dark brown or black;
dorsum pinkish brown or gray above, uniform or with blotches and
striations and one or more black streaks on the nape. Below, vinaceous
or gray speckled and striated with brown.

Size. Total length of d\ 1453 (875 + 578) mm. from Nchingidi,
Tanganyika Territory; total length of 9 , 1348+ (933 + 415+) mm.
from Zengeragusu, Tanganyika Territory; both, however, exceeded by
an unsexed specimen of 1470+ (935 + 535+) mm. from Caconda,
Angola.

Remarks. The type of capcnsis cannot now be located (V. Fitz-
Simons, 1937a).

Dentition. The number of teeth anterior to the three enlarged fangs
vary from 11 (Hanha) to 16 (Rungwe) according to Bogert (1940,
p. 71) whom see for further discussion.

Anatomy. For a full description of the hemipenes of two Hanha and
Mlanje snakes, in which they extended to the ninth subcaudal, see
Bogert (1940, p. 71) who also comments on the binocular vision attri-
buted to this species by Walls (1932, p. 69).

Breeding. On December 24, at Zengeragusu, a 9 laid 2 eggs, meas-
uring 38 x 15 and 34 x 14 mm., respectively, which were dry when
found; but for her escape the following day she might have laid more.
Diet. Only one snake of many examined, held feathers, apparently
those of a weaver or finch. The Zengeragusu snake, confined with a
Typhlops s. excentricus, allegedly swallowed the latter, according to
the native in charge of them. Some weeks after her escape she was
located in a tree only two hundred yards from the house, in her stomach
was a recently swallowed chameleon (C. d. dilcpis). Bogert (1940,
p. 71) records finding a chameleon (Brookesia platyceps) in each of
two Rungwe snakes, and two small terrestrial toads in a snake from
Hanha. At Nchingidi two snakes each held a toad (Breviceps mossam-
bicus).

Parasites. Cestode in a Northern Rhodesian snake (Pitman).
Defence. See account under typical form, and also that of Cott
(1935, p. 969) with special reference to its cryptic coloration.

158 bulletin: museum of comparative zoology

Migration. Mr. E. Wyllie, a surveyor who had spent many years in
the veld, states (1927) that: "a number of these snakes" followed two
days later by "a second contingent" passed through his camp on the
Pongola River, Piet Retief District, travelling in a northwesterly
direction. Never before had he witnessed snakes travelling "m masse."

Habitat. Boulenger (1897e) has stated that Bird Snakes occur on
the Nyika Plateau between 6,000 and 7,000 feet. Pitman (1937)
remarks on its occurrence from 1,500 feet in the Zambezi Valley to
5,000 feet on the plateau. He also invites attention to the frequency
with which he has encountered this arboreal reptile upon the ground.
As a savanna species in the East and South, from sea level upwards,
it is obvious that it must make its way from tree to tree by means of
the ground. Bogert (1940, p. 72) discusses at length the alternatives as
to whether Thelotornis originated in the western rain forest and then
spread to the savanna — the most reasonable view it seems to me —
. or else whether capensis represents a stage in the evolution of kirt-
landii. The assumption of the absence of intergrades, however, is
fallacious, being made in the absence of a good series of Tanganyika
snakes.

At Kitungulu I obtained a Bird Snake under rather unusual cir-
cumstances. In a native clearing stood two huge trees about whose
bases were piled quantities of dry grass. As the situation appeared
ideal for cobras, I had the heap ignited. The heat from the flames
rose into the trees though the flames fell far short; towards the end of
the conflagration the Bird Snake dropped from a height of at least
twenty feet. Though I saw it fall I mistook it for a branch till a shout
from one of my "boys" drew my attentoin to the departing snake
which I pursued and captured among the standing maize.

Localities. Tanganyika Territory1: Kitaya; Kitungulu; Kon-
doa Irangi; Lake Victoria; Marangu; Mikindani; Mpwapwa;
Nchingidi; Rungwe Mtn.; Sanya; Uleia; Zengeragusu. Mafia
Island. Mozambique: Cabaceira Id.; Cheringoma Farm,
Inhaminga; Chifumbazi; Delagoa Bay; Fambani; Inguenha,
Maputa; Loangwa Valley; Lumbo; Manica; Massangulo; Ngaza;
Querimba Id.; Quilimane; Sena; Tete. Nyasaland: Lake Nyasa;
Mandala; Mlanje (Milanji); Nyika Plateau; Shire Highlands;
Zomba. Northern Rhodesia: Barotze; Batoka; Broken Hill to
Bwana Mkubwa; Feira district; Loangwa Valley; Mpika; Mumb-
wa; Mwengwa on Kafue River; Namwala; Petauke; Serenje; Upper

1 For further localities Bee those listed under the typical form.

loveridge: African snakes 159

Zambezi. Southern Rhodesia: Bulawayo; Chirinda Forest;
Eldorado; Empandeni; Gwena's, Gwanayaya River ;Iro me; Khami;
Matabeleland; Zambesi. Bechuanaland Protectorate: Lealui;
Lobatsi (reported). Transvaal: Banolierkop; Comati and Croco-
dile Rivers junction; Dwar's River, Zoutpansberg district; Legogot;
Louw's Creek; Malta near Leysdorp; Mariepskap; Mmoouve, 42
miles N. of Serowe; Piet Retief district; Pretoria; Zoutpansberg.
Zulllland: Hlabisa; Kosi Bay; Mseleni; Somkele; Ubombo.
Natal: Country towards Port Natal; Durban; Umvoti River.
Cape Province: Francistown. South West Africa: Gobabis;
Grootfontein. Angola1: Bibala; Caconda; Chimporo; Cunene
River; Hanha; Kuvangu; Quando River; Quilengues; Querimbo;
Quissangue; Vila da Ponte.

Distribution. Africa south of the equator from central Angola and
northern South West Africa, east to central Tanganyika Territory and
Natal.

Folklore. An Myeye of Tabora voiced the belief that the species
was not vicious, but that it would bite if trodden upon and that then
the victim would die in a minute (Loveridge, 1928d). Pitman (1937)
directs attention to local names in Northern Rhodesia such as: "the
little bit of wood which bites," and "he who has been bitten can get
as far as to see the roofs of his village but no farther before he dies."
Wyllie (1927) was told by natives of the Piet Retief district, Trans-
vaal, that this snake was very dangerous, averring that it licks its
victim — ox, goat, or man — instead of striking. This results in the
skin peeling off and the exposed tissue becoming septic leads to the
demise of the victim.

Genus Calamelaps

1849. Choristodon A. Smith (not Jonas), 111. Zool. S. Africa, 3, App. p. 18

(type concolor Smith).
1866a. Calamelaps Gunther, Ann. Mag. Nat. Hist. (3), 18, p. 26 (type

Calamaria unicolor Reinhardt).

Maxillary very short with 3-4 teeth gradually increasing in size,
followed after an interspace by a large grooved fang situated below the
eye; anterior mandibular teeth enlarged. Head small, not distinct
from neck; eye minute, with round pupil; nasal entire2 or divided, in

1 For further localities see those listed under the typical form.

2 Tornier (1901a) states that in young examples of C. u. unicolor the nasal is entire, becoming
semidivided or divided with growth. He mentions one individual in which the nasal is entire
on one side of the head, divided on the other.

160 bulletin: museum of comparative zoology

contact with the rostral; no loreal; no preocular. Body cylindrical;
scales smooth, without pits, in 17-211 rows; ventrals rounded. Tail
very short; obtuse, subcaudals in two rows.

Range. Africa from Portuguese Guinea and Angola east to Kenya
Colony and Natal.

Remarks. Apparently the rostral develops with age, as in Prosymna;
it is not distinguishable from that of Rhinocalamus and should there-
fore be avoided as a key character.

Key to the Species

1. Frontal as long as, or shorter than, its distance from the rostral; temporals

1+1; upper labials 7; fifth lower labial largest; posterior sublinguals as
long as, or almost as long as, the anterior; ventrals 133-148; range

Natal concclor

(p. 161)

Frontal as long as, or longer than, its distance from the end of the snout;

temporal 1 only; upper labials 6 or 5; fourth lower labial largest; posterior

sublinguals often scarcely differentiated, if distinct then much shorter

than the anterior; ventrals 161-194; unknown in Natal 2

2. Midbody scales in 21 rows; range Angola and Transvaal north to

extreme southwestern Tanganyika Territory (at Tukuyu nr. Lake

Nyasa) u. polylepis

(p. 162)
Midbody scales in 19 to 15 rows 3

3. Midbody scales in 19 rows; range Zululand, north along coast to Kenya

Colony (at Ngatana, Tana River), penetrating inland in Transvaal
(to Gravel otte) in Southern Rhodesia; (to Aatosma, and Empandeni
where it meets with polylepis) and in Northern Rhodesia (to Lake

Bangweulu) w. warreni

(p. 163)
Midbody scales in 17 to 15 rows 4

4. Midbody scales in 17 rows; range Tanganyika Territory (Uluguru Mtns.)

north to Kenya Colony (at Peccatoni, fide Boettger) west to Sierra Leone
and "Guinea" u. unicolor

(p. 165)

Midbody scales in 15 rows; range Portuguese Guinea (Rio Cassine, known

only from the type) u. feae

(p. 168)

Sexual dimorphism. The marked dimorphism in the number of sub-
caudals in the races of unicolor is best shown in tabular form. It is

1 Said to be 13 in type of C. concolor but considered erroneous by FitzSimons (1937a).

loveridge: African snakes 161

important to note, however, that the sexes have had to be assumed in
the case of polylepis as authors have not furnished the sex, moreover,
though Boulenger stated that the type offeac was a cf, from the scale-
counts it would appear to be a 9 .

c?&. 9 9. <?<?. 9 9

C. u. polylepis 163-194, 197-212 ventrals ; 27-27, 16-20 subcaudals
C. u. warreni 161-177, 179-209 ventrals ; 26-30, 17-32 subcaudals
C. u. unicolor 164-182, 201-208 ventrals ; 28-38, 21-27 subcaudals
C.u.feae 196, ventrals; 23 subcaudals

Calamelaps concolor (Smith)

1849. Choristodon concolor A. Smith, 111. Zool. S. Africa, 3, App., p. 18:
Kaffirland eastward of Cape Colony, i.e. Natal.

1897h. Boulenger, p. 175.

1896d. Calamelaps concolor Boulenger, p. 246.

1898. Sclater, p. 100.

1910b. Boulenger, p. 516.

1912. FitzSimons, F. W., pp. 127, 128.

1925. Werner, 1924, p. 151.

1937a. FitzSimons, V., p. 263.

1905a. Calamelaps Mironi Mocquard, Bull. Mus. Hist. Nat. Paris, 11, p.
77: Upper Natal.

1925. Werner, 1924, p. 151.

Further citations of "concolor" will be found under u. unicolor and
u. polylepis.

Name. Purple-glossed Snake (English).

Description. Rostral nearly twice as broad as deep, the triangular
portion visible from above included once and a half to two times in its
distance from the frontal; nostril in a semidivided or divided nasal;
internasals much broader than long, their median suture half to two-
thirds the length of that of the prefrontals ; frontal as long as broad (in
the middle), shorter than its distance from the rostral, much shorter
than the parietals; supraocular small; no loreal1; no preocular; eye
small, its diameter only \ialf its distance from the mouth; postocular
small, sometimes fused with the supraocular; temporals 1 + 1; upper
labials 7, the third and fourth entering the orbit, third in contact with
the prefrontal, fifth largest and in contact with the parietal; fourth
lower labial largest; 4 lower labials in contact with the anterior sub-

1 The frenal mentioned by Smith, is the posterior nasal.

162 bulletin: museum of comparative zoology

Unguals, which are slightly longer or shorter than the posterior. Mid-
body scales in 171 rows, smooth; ventrals 1332-148; anal divided; sub-
caudals 32-39.

Color. Above and below uniformly black (fading to brown in alco-
holic specimens exposed to sunlight) except for the ventrals some, or
all, of which may be margined with white posteriorly.

Size. Total length of c? (type of mironi), 442 (367 + 75) mm.,
from Upper Natal; total length of 9 (M.C.Z. 16163), 348 (303 + 45)
mm. from Durban, Natal.

Remarks. The alleged type in the Royal Scottish Museum has been
discussed by FitzSimons (1937a) who finds that it has 17 (instead of
13 as stated by Smith) midbody scale-rows. Smith's misprint resulted
in the description of mironi by Mocquard (1905a) subsequently synon-
ymized by Boulenger (1910b). Chubb's (1909b) record of concolor
from Southern Rhodesia was shown by Hewitt (1913e) to have been
based on a misidentified C. u. polylepis. My own (1928g) reference to
concolor was a lapsus for unicolor.

The third known specimen (M.C.Z. 16163) of this rare snake was
received from the Albany Museum in 1922, identified as Macrelaps
microlepidotus, a species readily distinguishable by its more numerous
midbody scale-rows, which number 25 to 27.

Habitat. A burrowing snake, easily mistaken for the burrowing
viper (Atractaspis bibronii) found in the same locality, Durban.

Localities. Natal: Durban; Kaffraria; Upper Natal.

Range. Natal.

Calamelaps unicolor polylepis Bocage

1873b. Calamelaps polylepis Bocage, Jorn. Sci. Lisboa, 4, p. 216: Dondo,

Angola.
1895a. Bocage, p. 126, pi. ix. fig. 2.
1896d. Boulenger, p. 246.

1897a. Bocage, p. 201 (misprinted Galamelaps).
1901a. Tornier (part), p. 86.
1904. Ferreira, p. 116.

1910a. Sternfeld, p. 32, fig. 36.
1913e. Hewitt, p. 480.
1915a. Boulenger (part), p. 214.
1915c. Boulenger (part), p. 632.

1 Said to be 13 in type, considered erroneous by FitzSimons (1937a).

2 Said to be 134 in type, recounted as 140 by FitzSimons (1937a).

loveridge: African snakes 163

1925. Werner (part), 1924, p. 151.

1937b. Monard, p. 129.

1888b. Calamelaps miolepis Gtinther, Ann. Mag. Nat. Hist. (6), 1, p. 323:

Cape MacLear, Lake Nyasa, Nyasaland.

1891a. Boulenger, p. 307.

1896a. Bocage, p. 103.

1909b. Calamelaps concolor Chubb (not Smith), p. 34.

1939b. Calamelaps unicolor FitzSimons, V. (not Reinhardt), p. 24.

Names. Angolan Purple-glossed Snake (English).

Description. Differs from C. u. warreni only in its more numerous
midbody scale-rows which are 21, smooth; ventrals 163-212; anal
divided; subcaudals 16-27.

Color: As in C. u. warreni.

Size. Total length of supposed d\ 480 (430 + 50) mm. from
Angola (Bocage); total length of supposed 9 , 535 (504 + 31) from
Birchenough Bridge surpassed by another of 690 (. . . + . . .) mm.
from Empandeni, Southern Rhodesia (Hewitt).

Sexual dimorphism. None of the ten specimens recorded in the
literature have been sexed, but they would appear to fall into two
groups, viz. supposed c? c? with ventrals 163-194, subcaudals 27, and
supposed 9 9 with ventrals 200-212, subcaudals 16-20.

Habitat. A burrowing snake, easily mistaken for the burrowing
viper (Atractaspis eo>igica) which sometimes occurs in the same
localities.

Localities. Angola: Cazengo; Dondo; Humbe; Quissangues.
Transvaal: Hectorspruit (but scale-count not given); near Nyl-
stroom; Palmary ville. Southern Rhodesia: Birchenough Bridge;
Empandeni. Nyasaland: Cape MacLear, i.e. Livingstonia. Tan-
ganyika Territory: Tukuyu (Langenburg) at northwest end of
Lake Nyasa.

Range. Angola and (?) Transvaal north to extreme southwestern
Tanganyika Territory.

Calamelaps unicolor warreni Boulenger

?1896a. Calamelaps unicolor Bocage (not Reinhardt), p. 94.

1901a. Tornier (part), p. 86.

1933h. Loveridge (part), p. 260.

1936J. Loveridge (part), p. 267.

1937. Pitman (part), p. 326.

1938a. Pitman (part), p. 217.

1938b. Pitman (part), pp. 40, 179, 315 (reprint).

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?1908c. Calamelaps polylepis Sternfeld (not Bocage), p. 247.

1915a. Boulenger (part), p. 214.

1915c. Boulenger (part), p. 632. .

?1921a. Angel, p. 42.

1923e. Loveridge, p. 889.

1925. Werner (part), 1924, p. 151.

?1934. Pitman, p. 298.

1908b. Calamelaps warreni Boulenger, Ann. Natal Mus., 1, pp. 230, 234,
fig. 3: Kosi Bay, Zululand.

1910b. Boulenger, p. 516.

1912. FitzSimons, F. W., pp. 127, 128.

1912. Hewitt, p. 276.

1913e. Hewitt, p. 480.

1925. Werner, 1924, p. 151.

1928. Cott, p. 953.

1935. Cott, p. 970.

1915a. Calamelaps mellandi Boulenger, Proc. Zool. Soc. London, p. 214:
Chirini Island, Lake Bangweulu, Northern Rhodesia.

1925. Werner, 1924, p. 151.

1934. Pitman, p. 298.

1942e. Calamelaps unicolor warreni Loveridge, p. 295. ,

Names. Eastern Purple-glossed Snake (English) ; ngogoma (Pokomo) ;
mbitu (Makonde, but not specific); nyeresi (Nyakusa).

Description. Rostral once and three quarters to twice as broad as
deep, the triangular portion visible from above as long as, or nearly as
long as, its distance from the frontal; nostril in a semidivided or
divided, rarely entire,1 nasal; internasals much broader than long,
their median suture as long as, or longer than, that of the prefrontals;
frontal once and a third to once and two thirds as long as broad (in
the middle), longer than its distance from the end of the snout, much
shorter than the parietals; supraocular small; no loreal; no preocular;
eye small, its diameter less than half its distance from the mouth;
postocular small, rarely absent2 through fusion with the supraocular;
temporal 1 only; upper labials 6, rarely 5, the third and fourth entering
the orbit, third, rarely second and third2 or only second3, in contact
with the prefrontal, fifth, rarely fourth3, largest and in contact with
the parietal ; fourth lower labial largest ; 4 lower labials in contact with
the anterior sublinguals, which are much longer than the posterior
which are often scarcely differentiated. Midbody scales in 19 rows,
smooth; ventrals 161-203; anal divided; subcaudals 17-30.

1 In type of warreni.

2 In type of warreni.

3 In type of mellandi.

loveridge: African snakes 165

Color. Above and below uniformly iridescent black (fading to brown
in alcoholic specimens exposed to sunlight) or opaque bluish gray when
about to slough.

Size. Total length of <? (M.C.Z. 30399), 475 (430 + 45) mm. from
Mwaya; total length of 9 (M.C.Z. 48428), 670 (630 + 40) mm. from
Mbanja.

Remarks. This form with 19 midbody scale-rows, being intermediate
between unicolor and polylepis was at first united with one, and then
the other, of these forms until Boulenger described warreni in 1908 on
an individual which was not wholly typical. The characters on which
he founded C. mellandi, viz. absence of postocular, second supralabial
in contact with prefrontal and fourth with parietal, also occur spas-
modically in the typical form, which, however, has 17 midbody scale-
rows. I placed mellandi in the synonymy of warreni in 1942.

Sexual dimorphism. cJ^cf with ventrals 161-177, subcaudals 26-30,
and 9 9 with ventrals 179-209, subcaudals 17-22.

Habitat. A burrowing species, readily mistaken for one or other of
the very venomous burrowing vipers (Atractaspis) occurring within its
range. It has been found wandering above ground in the early morning
after a night of rain.

Localities. Zululand : Kosi Bay. Transvaal : Southern Rho-
desia: Empandeni. Northern Rhodesia: Chirini Id., Lake
Bangweulu; Lealui; Sesheke. Mozambique: Caia; Chifumbazi;
Lumbo. Tanganyika Territory: Amboni Estate near Tanga;
Mbanja; Morogoro (? as found bottled in house); Mwaya; Nchin-
gidi; Tanga. Kenya Colony: Ngatana, Tana River.

Range. Zululand and Southern Rhodesia (at Empendeni, where it
meets with polylepis) north to Kenya Colony (at Ngatana.).

Calamelaps unicolor unicolor (Reinhardt)

1843. Calamaria unicolor Reinhardt, K. Danske Vidensk. Selsk. Afhandl.'

10, p. 236, pi. i, figs. 1-3: Gunea ,West Africa.
1862. Amblyodipsas unicolor Jan, p. 41.
1896. Tornier, p. 79.

1877c. Atractaspis Hildebrandtii Peters, Monatsb. Akad. Wiss. Berlin, p. 616,

pi. -, fig. 3: Zanzibar coast.
1896d. Boulenger, p. 512.
1898. Boettger, p. 136.

1901a. Tornier, p. 85.
1912. Hobley, p. 56.

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1913. Boettger, p. 353.

1866a. Calamelaps unicolor Giinther, p. 25.

1893. Pfeffer, p. 77.

1896d. Boulenger, p. 245.

1897. Tornier, p. 65.

1901a. Tornier (part), p. 85.

1902a. Boulenger, p. 447.

1908b. Sternfeld, pp. 219, 233.

1909a. Sternfeld, p. 22.

1910a. Sternfeld, p. 32.

1911c. Boulenger, p. 166.

1915c. Boulenger, p. 632.

1919b. Boulenger, p. 291.

1922. Aylmer, p. 15.

1923. Schmidt, p. 116.
1924b. Loveridge, p. 7.
1925. Werner, 1924, p. 151.
1928c. Barbour & Loveridge, p. 130.
1933f. Angel, p. 170, figs. 64-64a.
1933m. Witte, p. 95.

1 936j . Loveridge (part), p. 267.

1937f. Loveridge, pp. 493, 496.

1937. Pitman (part), p. 326, pi. xii, fig. 2; pi. M, fig. 2.

1938a. Pitman (part), p. 217.

1938b. Pitman (part), pp. 40, 179, 315 & pis. as above (reprint).

1923. Calamelaps niangarae Schmidt, Bull. Am. Mus. Nat. Hist., 49, p.

117, fig. 12: Niangara, Belgian Congo.

1928g. Calamelaps concolor Loveridge (not Smith), p. 41 (lapsus for unicolor.)

A further citation of "anicolor" will be found under u. polylcpis.

Names. Equatorial Purple-glossed Snake (English).

Description. Rostral once and a half to nearly twice as broad as
deep, the triangular portion visible from above as long as, or nearly as
long as, its distance from the frontal; nostril in a semidivided or
divided, rarely entire, nasal; internasals much broader than long,
their median suture as long as, or shorter, or longer1 than the length of
that of the prefrontals; frontal once and a quarter to once and two
thirds as long as broad (in the middle), as long as, or longer than, its
distance from the end of the snout, much shorter than the parietals;
supraocular small; no loreal; no preocular; eye small, its diameter only
half its distance from the mouth; postocular small, rarely absent

1 In types of hildebrandiii and niangarae.

loveridge: African snakes 167

through fusion with the supraocular; temporal 1 only; upper labials 6,
rarely 51, the third and fourth, rarely second and third, entering the
orbit, third, rarely second, in contact with the prefrontal, fifth, rarely
fourth, largest and in contact with the parietal; fourth lower labial
largest; 4 lower labials in contact with the anterior sublinguals, which
are much longer than the posterior, which are often scarcely differen-
tiated. Midbody scales in 17 rows, smooth; ventrals 164-208: anal
divided; subcaudals 21-38.

Color. Above and below uniformly iridescent black (fading to brown
in alcoholic specimens exposed to sunlight) or opaque bluish gray
when about to slough.

Size. Total length of cf (type of niangarae), 414 (366 -f- 48) mm.
from Niangarae; total length of 9 (A.M.N.H.), 722 (672 + 50) mm.
from Faradje.

Remarks. Tornier (1901a) was the first to synonymize Atractaspis
hildcbrandtii with unicolor. Accorinti's (1913, p. 300) record of it
from Eritrea, however, obviously refers to a true viper and not to a
Calamelaps. In 1925 Werner (1924, p. 151) synonymized niangarae, a
species which was thought to differ from unicolor because the median
suture of its internasals was longer (instead of as long as, or shorter)
than that between the prefrontals. In this, however, it agreed with
hildcbrandtii. Its other supposed differences were chiefly those of sex,
and it is interesting to note that Boulenger (1896d) had recorded a
Sierra Leone specimen with exactly the same ventral and subcaudal
counts as had the type of niangarae. Witte (1933m) has since obtained
unicolor at Niangara.

As a result of Tornier's remarks and records, Barbour and Loveridge
(1928c) added polylepis to the synonymy of unicolor, but now, with
additional material, and after a study of all the literature, the true
position begins to emerge of forms occupying definite geographical
areas, though a good deal of overlapping occurs where the western and
eastern forms meet.

Sexual dimorphism, cf cf with 164-182 ventrals, 28-38 subcaudals,
and 9 9 with 201-208 ventrals, 17-21 subcaudals.

Diet. A wolf snake (Lycophidion c. acutirostrc) was disgorged by an
Equatorial Purple-glossed Snake shortly after capture, the prey being
only 20 mm. shorter than the predator. The similarity in the parallel
development of these two blackish, burrowing snakes was striking, the
prey having nine ventrals fewer, and four subcaudals more, than its

1 These rare labial variations in a snake from Mt. Mbololo.

168 bulletin: museum of comparative zoology

vanquisher at Changamwe. A skink (Lygosoma kilimensis) in a Nyange
snake, a snake (Aparallactus werneri) and a caecilian (Boulengerula
boulengeri) in Amani specimens.

Defence. The fact that Peters, overlooking the absence of poison
fangs, described this snake as a viper, and was followed by Boulenger,
Boettger and others who identified fresh material with hildebrandtii.
is sufficient and eloquent proof of the close superficial resemblance
between the two genera, making it essential when capturing Calamelaps
to treat them as if they were indeed dangerous vipers.

Habitat. A burrowing species of which several were taken in clearing
land at Amani, while another was hoed up in a native garden at
Nyange. This race seems to be chiefly associated with .forested, or
recently deforested, areas at altitudes below 3,000 feet.

Localities. Bocage's (1896a) and Sternf eld's (1908c) records for
Mozambique have been arbitrarily transferred to C. u. warreni as other
Mozambique material, whose scales could be counted, were referable
to that race.

Tanganyika Territory: Amani, Usambara Mtns.; Bagamoyo
(Pfeffer, requires checking); Nyange, Uluguru Mtns.; Tanga.
Kenya Colony: Changamwe; Mombasa; Mount Mbololo; Pecca-
toni. Uganda: Bussu; Kampala. Belgian Congo: Faradje;
Niangara. Nigeria: Togo. Gold Coast. Sierra Leone. Guinea.

Range. Tanganyika Territory (Uluguru Mountains) north to Kenya
Colony (atPeccatoni,/?de Boettger) west to Sierra Leone and "Guinea."

Calamelaps unicolor feae Boulenger

1906L Calamelaps feae Boulenger, Ann. Mus. Civ. Stor. Nat. Genova, (3),

2, p. 214, fig. 9: Rio Cassine, Portuguese Guinea.
1919b. Boulenger, p. 291.
1925. Werner, 1924, p. 151.

1933f. Angel, p. 171.

Names. Western Purple-glossed Snake (English).

Description. Rostral once and a half as broad as deep, the triangular
portion visible from above as long as its distance from the frontal;
nostril in a divided nasal; internasals much broader than long, their
median suture longer than that of the prefrontals ; frontal once and a
third as long as broad (in the middle), longer than its distance from the
end of the snout, much shorter than the parietals ; supraocular small ;
no loreal; no preocular; eye small, its diameter only half its distance

loveridge: African snakes 169

from the mouth; postocular small; temporal 1 only; upper labials 5,
the second and third entering the orbit, second in contact with the
prefrontal and posterior nasal, fourth largest and in contact with the
parietal; fourth lower labial largest; 4 lower labials in contact with
the anterior sublinguals, which are much longer than the posterior
which may be scarcely differentiated. Midbody scales in 15 rows,
smooth; ventrals 196; anal divided; subcaudals 23.

Color. Above and below uniformly iridescent black (fading to brown
in alcoholic specimens exposed to sunlight).

Size. Total length of alleged & , 610 (560 4- 50) mm. Type.

Remarks. Known only from the type, said to be a male. The above
description is based on the original and on the figures.

Localities. Portuguese Guinea: Rio Cassine.

Range. Portuguese Guinea.

Genus Miodon

1858. Microsoma Jan (not Mocquart), Revue Mag. Zool. (2), 10, p. 519

(type neuwiedii Jan).

1859. Miodon A. Dumeril, Arch. Mus. Hist. Nat. Paris, 10, p. 206 (type

Elapomorphus gabonensis A. Dumeril).

1860. Urobelus Reinhardt, Vidensk. Meddel. Kjobenhavn, p. 229 (type

acanthias Reinhardt).
1902a. Cynodontophis Werner, Verh. Zool.-Bot. Ges. Wien, 52, p. 346 (type
aemulans Werner = notatus Peters) .

Maxillary very short with 2-4 teeth followed after an interspace by
1-2 large grooved fangs situated in advance of the eye; second and
third, or third and fourth, mandibular teeth enlarged, fang-like.
Head small, not distinct from neck; eye very small, with round pupil;
nasal entire or divided, not in contact with the rostral, the internasal
forming a suture with the first labial; no loreal; 1 preocular. Body
cylindrical; scales smooth, without pits, in 15 rows; ventrals rounded.
Tail very short; subcaudals in two rows.

Range. Africa in forested areas of equatorial belt from Liberia east
to Uganda and western Tanganyika Territory.

Remarks. Microsoma of Jan is antedated by that of Mocquart,
1854 (Ann. Soc. Ent. France (3), 2, p. 737) proposed for a genus of
Diptera. Cynodontophis was originally believed to differ from Miodon
in dentition. Bogert (1940, p. 46) has shown, however, that such differ-
ences were due to fang replacement ; his remarks on the subject should
be consulted. This author also describes the hemipeneal structure.

170 bulletin: museum of comparative zoology

Key to the Species

1. Anal entire; dorsum with pattern of 5 parallel black lines; ventrals 183-

216; subcaudals 16-22; range Sierra Leone to Togoland (or Nigeria, fide

Angel) acanthias

(p. 170)
• Anal divided1 2

2. Dorsum with pattern of 3 parallel black lines; ventrals 219-246; subcaudals

11-21; range Gold Coast to Dahomey (with type allegedly from Gabon,

i.e. French Congo) g. gabonensis

(p. 172)
Dorsum without pattern of parallel black lines 3

3. Dorsum with pattern of 2 parallel series of black spots; ventrals 178-228;

subcaudals 14-19; range French Cameroon south to French Congo

g. notatvs

(p. 173)

Dorsum uniformly dark 4

4. Nape and crown pale fawn with dark mottlings; throat white; ventrals

181-252; subcaudals 15-25; range (Togo1 fide Werner) southeastern
Nigeria south to Angola east to Uganda (west of Ruwenzori) .g. collaris

(p. 175)

Nape and crown of head entirely black, or with a sharply distinct broad

white band across the parietal region 5

5. Top of head iridescent black like nape and dorsum; throat white or black;

ventrals 202-241; subcaudals 15-21; range western Congo (Poko) east
to central Uganda (Mabira Forest) and western Tanganyika, .g. christyi

(p. 178)

Top of head from snout to behind eyes, black, posterior half of head and

nape pure white; ventrals 237-258; subcaudals 13-18; range western

Congo (Idjwi Island, Lake Kivu) to central Uganda (Entebbe) .g. graueri

(p. 180)

Miodon acanthias (Reinhardt)

1860. Urobelus acanthias Reinhardt, Vidensk. Meddel. Kjobenhavn, p. 229

pi. iii: Guinea.
1893c. Matschie, p. 213.

1863. Elapomorphus acanthias Jan, p. 47.
1863. Jan, p. 39.

1865. Jan, livr. 14, pi. iii, fig. 4.

1888b. Giinther (part), p. 323.

1 Allegedly entire in a Congo specimen of M. g. collaris, fide Bocage (1895a, p. 126)

loveridge: African snakes 171

1896d. Miodon acanthias Boulenger, p. 250.

1908b. Sternfeld, pp. 219, 233.

1909a. Sternfeld, p. 22.

1915a. Boulenger, p. 215.

1919b. Boulenger, p. 291.

1922. Aylmer, pp. 15, 22.

1925. Werner, 1924, p. 152, fig. 13.

1930a. Barbour & Loveridge, p. 773.

1933f. Angel, p. 173.

1941e. Loveridge, p. 123.

Name. Five-striped Snake-eater (English).

Description. Rostral broader than deep, just visible from above;
nostril in a divided or entire nasal, separated from rostral ; internasals
as broad as long, two-thirds to three-quarters the length of the pre-
frontals; frontal as long as, or once and two-thirds as long as, broad,
as long as, or longer or shorter than, its distance from the rostral,
much shorter than the parietals, once and a half to twice as broad as a
supraocular; no loreal; preocular 1 ; eye small, its diameter half to two-
thirds its distance from the mouth; postoculars 1-2; temporals 1 + 1;
upper labials 7, the third and fourth entering the orbit; 4 lower labials
in contact with the anterior sublinguals, which are longer than the
posterior. Midbody scales in 15 rows, smooth; ventrals 183-216; anal
entire; subcaudals 16-22 pairs.

Based on original descriptions, the literature, 8 Liberian specimens
in the M.C.Z. and 1 in the U.S.N.M.

Color. Above, head more or less black, snout, upper lip, and an
occipital bar white (reddish in life) ; a black nuchal band; dorsum pale
reddish (Brazil red in life) with five longitudinal black stripes on body,
three on tail, tip of tail white (reddish in life). Below, white (Bitter-
sweet orange in life). The colors in life are from Ridgway and were
based on a Liberian snake as noted by G. M. Allen.

Size. Total length of <? (M.C.Z. 22525), 514 (475 + 39) mm. from
Nickabo; total length of 9 (M.C.Z. 22524), 585 (555 + 30) mm. from
Gbanga.

Sexual dimorphism. Assuming the two types to have been females,
then cfcf have 183-195 ventrals, and 22 subcaudals, and 9 9 have
207-212 ventrals, and 16-18 subcaudals, this being based on five males
and five females.

Breeding. Between April 10-16, at Gibi, a 9 held 4 eggs, each
measuring circa 22 x 7 mm.

Localities. Liberia: Du River; Gbanga; Gibi Si Mountain;

172 bulletin: museum of comparative zoology

Nickabo; Paiata. Gold Coast: Ashanti. Togoland: Misahohe.
Nigeria {fide Angel).

Range. Sierra Leone east to Nigeria.

MlODON GABONENSIS GABONENSIS (Dumeril)

1856c. Elapomorphus gabonensis A. Dumeril, Revue Mag. Zool. (2), 8, p.

468: Gaboon.
1859. Dumeril, A., p. 206, pi. xvi, fig. 2.

1884b. Sauvage, p. 201.

1858. Microsoma neuwiedi Jan, Revue Mag. Zool. (2), 10, p. 519: Christians-

borg, Gold Coast.

1859. Jan, pi. iv.

1866. Elapomorphus (Urobelus) neuwiedi Jan, livr. 15, pi. i, fig. 2.

1884a. Miodon gabonense Rochebrune, p. 153, pi. xvii, fig. 1 : (the figure is of

gabonensis but text is ignored).

1909a. Sternfeld, p. 22.

1896d. Miodon neuwiedii Boulenger, p. 253.

1908b. Sternfeld, pp, 219, 234.

1909a. Sternfeld, p. 22.

1917. Chabanaud, p. 377.

1917b. Chabanaud, p. 12.

1925. Werner, 1924, p. 153.

1933f. Angel, p. 173, figs. 65-65a.

Further citations of 'gabonensis, will be found under g. collaris and
g. christyi.

Name. Three-striped Snake-eater (English).

Description. Rostral slightly broader than deep, just visible from
above; nostril in an entire nasal; internasals as broad as long, slightly
shorter than the prefrontals; frontal as long as, or longer than, broad,
as long as its distance from the rostral, much shorter than the parietals,
nearly twice as broad as a supraocular ; no loreal ; preocular 1 ; eye small,
its diameter two-thirds its distance from the mouth; postoculars 1-2;
temporals 1 + 1 ; upper labials 7, the third and fourth entering the
orbit; 3 or 4 lower labials in contact with the anterior sublinguals,
which are slightly shorter than the posterior. Midbody scales in 15
rows, smooth; ventrals 219-246 {fide Angel); anal divided ;subcaudals
11-21 pairs.

Based on original descriptions and figures; no material seen.

Color. Above, head black; back pale brown with three longitudinal
black stripes; tail black. Below, white.

loveridge: African snakes 173

Size. Total length of type 550 (510 + 40) mm. allegedly from
Gaboon, and (type of ncuwicdii), 172 (162 + 10) mm. from Christians-
borg.

Remarks. Both Dumeril and Jan have figured their reptiles showing
that both have three prominent black stripes running the entire length
of the dorsum. With the exception of Sternfeld and Chabanaud, no
one appears to have received true gabonensis, by which I mean a
striped snake.

Boulenger, when writing the Catalogue of Snakes (1896d), referred
to gabonensis material which I consider identical with collaris, for he
had no representatives of either true (striped) gabonensis or neuwiedi,
though recognizing the latter by trivial characters which are demon-
strably variable in any large series of collaris.

I am uncertain if it is wise to treat collaris as a race of gabonensis
for the ranges appear to overlap, this would be especially the case if
the type of the latter actually came from the Gaboon. On the other
hand to treat as full species snakes which differ only in color and
pattern, would seem retrogressive.

Localities. Gold Coast: Christiansborg. Togo: Klein Popo.
Dahomey: Agouagou. French Congo as Gaboon.

Range. Gold Coast east to Dahomey (and ? French Congo).

Miodon gabonensis notatus (Peters)

1882d. Microsoma noialum Peters, Sitz. Ges. Naturf. Freunde Berlin, p. 127:

No locality.
1887b. Mocquard, p. 64.

1896d. Miodon notatus Boulenger, p. 252.

1897. Sjostedt, p. 35.

1898. Werner, p. 212.
1908a. Sternfeld, pp. 414, 429.
1909b. Sternfeld, p. 22.
1910. Muller, p. 609.
1915a. Boulenger, p. 215.
1919b. Boulenger, p. 292.
1925. Werner, 1924, p. 153.
1940. Bogert, p. 46.

1902a. Cynodontophis aemulans Werner, Verh. Zool.-Bot. Ges. Wien, 62, p.

346: Congo.
1910. Muller, p. 610.

1915a. Boulenger, p. 215.
1923. Schmidt, p. 120.

1925. Werner, 1924, p. 153.

Further citation of 'notatus' will be found under g. collaris.

174 bulletin: museum of comparative zoology

Name. Spotted Snake-eater (English).

Description. Rostral broader than deep, just visible from above;
nostril in a divided nasal, separated from1, or in contact with2, the
rostral; internasals longer than broad, as long as, or two-thirds as long
as, the prefrontals; frontal longer than broad, as long as its distance
from the rostral, much shorter than the parietals, twice as broad as a
supraocular; no loreal; preocular 1 ; eye small, its diameter half to two-
thirds its distance from the mouth; postoculars 1-2; temporals 1 + 1;
upper labials 7, the third and fourth entering the orbit; 3 lower labials
in contact with the anterior sublinguals, which are as long as, or longer
than, the posterior. Midbody scales in 15 rows, smooth; ventrals
178-228; anal divided; subcaudals 14-19 pairs.

Based on original descriptions, the literature, and one specimen.

Color. Above, head and nuchal collar black; back light brown with
a double series of black spots which may be light-edged; tail black.
Below, white (yellow in life?).

Size. Total length, 317 (288 -f 29) mm. from Brazzaville.

Remarks. Andersson (1901) advocated the uniting of notatus with
gabonensis but lacked material of both. Sternfeld (1908a), with
notatus material, objected, though the characters with which he sup-
ported this view have since proved to be variable.

M tiller (1910) referred aemulans to the synonymy of notatus but
. proposed to retain the genus Cynodontophis. Boulenger (1915a)
retained both species and genera, separating Cynodontophis on the
nasal being in contact with the rostral — which is not the case with
our Cameroon specimen (M.C.Z. 14995) and is variable in the M.C.Z.
series of gabonensis collaris.

Schmidt (1923) and Werner (1925) agree with the synonymizing of
the species, but suggest that the genus should be retained. Bogert
(1940), after a thorough study of the dentition in g. collaris, found that
the alleged differences between Miodon and Cynodontophis were due
to fang replacement and that Cynodontophis could not be retained.

Witte (1933m) referred six specimens from Kunungu, Belgian Congo,
to notatus one of which is now M.C.Z. 42957. It is a typical g. collaris
in coloration and it may be safely assumed that the others are the same
for he recorded both gabonensis and collaris from Kunungu in the same
paper.

Sexual dimorphism. None of the material mentioned in the literature
is sexed, a Cameroon 9 in the M.C.Z. has 201 ventrals and 14 sub-
caudals. Males probably have a range from 17-19.

1 In type of notatus.

2 In type of aemulans.

loveridge: African snakes

175

Localities. French Cameroon: Barombi; Bipindi; Dibongo near
Edea ; Kribi ; Sakbayeme. French Congo : Brazzaville ; Cap Lopez.
(Recorded from Belgian Congo: Kunungu by Witte (1933m) in
error) .

Range. French Cameroon and adjacent French Congo.

Miodon gabonensis collaris (Peters)

1862. Elapomorphus gabonicus Jan (not Dumeril), p. 47.

1866. Jan, livr. 15, pi. i, fig. 1.

1865a. Polemon barthii Giinther (not Jan), p. 90.

1881d. Microsoma collare Peters, Sitz. Ges. Naturf. Freunde Berlin, p. 148;

Macange, Cuango = Kwango, French Equatorial Africa.

1887a. Bocage, p. 182.

1895a. Bocage, p. 124, pi. xiv, figs. 1-2.

1895c. Bocage, p. 13.

1903a. Bocage, p. 44.

1887b. Urobelus gabonicus Boulenger, p. 127.

1887b. Microsoma fulvicollis Mocquard, Bull. Soc. Philom. Paris (7), 11, p.

65: Franceville, French Congo.

1888b. Elapomorphus acanthias Giinther (not Reinhardt), p. 323.

1888b. Elapomorphus caeutiens Giinther, Ann. Mag. Nat. Hist. (6), 1, p.

323, pi. xix, fig. B: Cameroon Mountains, British Cameroons.

1896d. Miodon collaris Boulenger, p. 251.

1897b. Mocquard, p. 13.

1900b. Boulenger, p. 454.

1903. Gough, p. 468.

1905c. Boulenger, p. 114.

1905. Ferreira, p. 169.

1908a. Sternfeld, p. 413.

1911. Lampe, p. 202.

1915a. Boulenger, p. 215.

1919b. Boulenger, p. 291.

1923. Schmidt, p. 120.

1933m. Witte, p. 95.

1937b. Monard, p. 129.

1896d. Miodon gabonensis Boulenger (part: no material), p. 252.

1897. Sjostedt, p. 35.

1897b. Werner, p. 400.

1898a. Werner, p. 211.

1899a. Werner, p. 140.

1900b. Boulenger, p. 454.

1901. Andersson, p. 23.

1908a. Sternfeld, pp. 413, 428.

176 bulletin: museum of comparative zoology

1909b. Sternfeld, p. 22.

1910. Muller, p. 609.

1911. Lampe, p. 202.
1915a. Boulenger, p. 215.
1915c. Boulenger, p. 633.
1917. Sternfeld, p. 480.
1919b. Boulenger, p. 291.
1919g. Boulenger, p. 26.
1923. Schmidt, p. 118.
1925. Werner, 1924, p. 153.
1932b. Witte, p. 14.

1933f. Angel, p. 175.

1933. Schouteden, p. 236.

1933m. Witte, p. 95.

1940. Bogert, p. 45.

1908b. Miodon gabonensis var. collaris Sternfeld, pp. 219, 234.

1910. Cynodontophis werneri Muller, Abh. Bayer Akad. Wiss., 2 Kl., 24,

p. 612: Cameroons?

1933m. Miodon notatus Witte (not Peters), p. 95.

1942e. Miodon gabonensis collaris Loveridge, p. 298.

Name. Pale-collared Snake-eater (English).

Description. Rostral broader than deep, just visible from above;
nostril in a semidivided, divided, or entire nasal; internasals as broad
as long, as long as, or almost as long as, the prefrontals; internasal and
prefrontal in contact with, or separated from the labials; frontal
longer than broad, as long as, or shorter than, its distance from the
rostral, much shorter than the parietals, once and a half to twice as
broad as a supraocular; no loreal; preocular 1; eye small, its diameter
half1 to three-quarters its distance from the mouth; postoculars 2,
rarely 1 ; temporals 1 + 1 ; upper labials 7, the third and fourth enter-
ing the orbit; 4, very rarely 3, lower labials in contact with the anterior
sublinguals, which are as long as, or longer than, the posterior. Mid-
body scales in 15 rows, smooth; ventrals 181-252; anal divided2; sub-
caudals 15-25 pairs.

Based on original descriptions, all data in literature, and much
material in A.M.N.H., C.N.H.M. and M.C.Z.

Anatomy. For discussion on dentition and description of hemipenes,
see Bogert (1940).

Color. Above, head and nape pale brown or fulvous with some
black blotches on the snout or crown and below the eye; back bluish
black or slaty gray, the scales edged with darker; tip of tail white.

1 A third to equal according to some authors.
Entire in one Congo snake according to Bocage (1895a, p. 126).

loveridge: African snakes 177

Below, white, except for some dusky markings on the chin and the
dorsal color impinging on the outer (lateral) edges of the ventrals, or
throat white and rest of under surface black (in an Angolan specimen.
M.C.Z. 32593).

Size. Total length of & (A.M.N.H. 12452), 535 (501 + 34) mm.
from Medje; total length of 9 (M.C.Z. 13606), 661 (630 + 31) mm.
from Niapu, but exceeded by an unsexed specimen, though almost cer-
tainly a 9 , recorded as 860 (818 + 42) mm. by Sternfeld, which is
probably the Togo snake of 850 (808 + 42) mm. of Werner (1897b).

Remarks. As indicated under M. g. gabonensis, I am of the opinion
that it may prove possible to separate two forms which have long been
confused, and that the overlap in ranges between the striped gabonensis
and the pale-collared collaris may not prove to be so great as has been
supposed.

Boulenger (1896d) synonymized fulvicollis and caeutiens with the
composite which he called gabonensis. Bogert (1940) would add both
collaris and werneri but he had in mind the composite rather than the
striped snake of Dumeril, for apart from this his remarks form the
most important contribution to our understanding of the genus in
recent times.

Sexual dimorphism. Using the sexed (but unchecked) records in the

literature, cfcf have 200-232 ventrals, and 19-25 subcaudals, while

9 9 have 195-252 ventrals, and 16-19 subcaudals, this being based on

five males and twelve females; 19 subcaudals is correct for both sexes,

being found in M.C.Z. material.

Diet. A snake, 180 mm. in length but with head digested,. in a 230
mm. collaris from the Congo (Schmidt).

Parasites. Two large linguatulids in lungs of a Metet snake (Bogert).

Habitat. Apparently associated with virgin forest.

Localities. Togo (based on a single specimen collected by Butt-
ner, recorded by Werner (1897b) and repeated by Sternfeld
(1908a-b), it may well be regarded as doubtful pending confirma-
tion by fresh material from Togo). Nigeria : Old Calabar. British
Cameroon: Bibundi; Cameroon Mountains; Cape Debundscha;
Isongo; Mapanja; Rio del Ray; Victoria. French Cameroon:
Bipindi; Bitye; Dibongo near Edea; Kribi; Lekungg River;
Lolodorf; Longji; Metet. Fernando Po: Spanish Guinea:
Esong (or Eosung) near Bakossiberge. French Congo: France-
ville; Lambarene. Angola: Cazengo; Golungo Alto; Missao de
Donda; Pungo Adungo; Quindumbo. French Equatorial Africa:
Macange, Kwango. Belgain Congo: Duma; Goma (Ngoma);

178 bulletin: museum of comparative zoology

Kunungu; Medje; Moanda; Mocra; Niangara; Niapu; Nyampoko;
Panga, Aruwimi River; Sandoa; Stanleyville. Uganda: Bundi-
bugyo, northwest of Ruwenzori Mountains.

Range. Togo?, certainly southeastern Nigeria, south to Angola and
east to Uganda (northwest of Ruwenzori Mountains).

Miodon gabonensis christyi Boulenger

1903f. Miodon Christyi Boulenger, Ann. Mag. Nat. Hist. (7), 12, p. 354:
Uganda.

1910a. Sternfeld, p. 34 (as Chrystii).

1911c. Boulenger, p. 166.

1915c. Boulenger, p. 633.

1924b. Loveridge, p. 7.

1925. Werner, 1924, p. 153.

1937f. Loveridge, p. 502.

1937. Pitman, p. 332, pi. xii, fig. 3, col. pi. M, fig. 3.

1938a. Pitman, pp. 217, 233.

1938b. Pitman, pp. 40, 185, 187, 188, 315, 331 and plates as above (re-
print).

1910a. Miodon gabonensis Sternfeld (not Dumeril), p. 34.

1924b. Loveridge, p. 7.

1933h. Loveridge, p. 261.

1936h. Loveridge, p. 40.

1937f. Loveridge, p. 502.

1938a. Pitman, p. 217.

1938b. Pitman, pp. 186, 187, 315 (reprint).

1923. Miodon unicolor Schmidt, Bull. Am. Mus. Nat. Hist., 49, p. 119, fig.
13: Poko, Belgian Congo.

1925. Werner, 1924, p. 153.

Name. Eastern Snake-eater (English).

Description. Rostral broader than deep, just visible from above;
nostril in a divided1 or entire2 nasal; internasals longer than broad,
shorter than the prefrontals; frontal slightly longer than broad, as
long as1, or shorter than2, its distance from the rostral, much shorter
than the parietals, broader than a supraocular; no loreal; preocular 1;
eye small, its diameter half to three-fifths its distance from the mouth;
postoculars 2; temporals 1 + 1, rarely 1 only2; upper labials 7, the
third and fourth entering the orbit, rarely the fifth3 or the seventh2 in
contact with the parietals ; 4 lower labials in contact with the anterior

1 In type of unicolor from Poko.

2 In type of christyi from Uganda.

3 In specimen from Mambawanga Hill.

loveridge: African snakes 179

sublinguals, which are longer than the posterior. Midbody scales in 15
rows, smooth; ventrals 202-241 ; anal divided; subcaudals 15-24 pairs.

Based on original descriptions and figures, data in literature cited,
and three specimens from Congo, Uganda and Tanganyika.

Color. Above, black with an iridescent bluish bloom. Below, uni-
form with dorsum or throat only black or white; ventrals and sub-
caudals dull creamy- or dirty white broadly margined with black later-
ally, or mainly black with their posterior edges mottled with white.

Size. Total length of d\ 661 (617 + 44) mm. from Mambawanga
Hill; total length of type of christyi, allegedly a 9 , 430 (402 + 28)
mm.

Remarks. In the extreme eastern part of its range, M. g. collaris
shows a marked tendency to melanism, several males being uniformly
black like a Calamelaps while other specimens have the black ventrals
mottled with white on the posterior edges (as in unicolor) or have
retained the light ventrals but more or less broadly edged with black
(as in christyi). All, however, would appear to have the head, nape and
dorsum uniformly colored so I have utilized this rather slender distinc-
tion to retain the name christyi for these eastern snakes. Perhaps addi-
tional material will show it to be untenable.

Sternfeld (1910a) had no material of christyi, all his descriptions
being translations into German from Boulenger, a point that Pitman
has not realized in retranslating back to English. While Sternfeld's
text regarding gabonensis is taken from Boulenger, I regard his record
of a specimen from Dar es Salaam with misgiving until it is verified, it
is more probably a Calamelaps. If it is correct it extends the range 400
miles to the east and introduces into the coastal zone a species usually
associated with forested areas.

Sexual dimorphism. Assuming that the type of christyi is a male,
then cf a71 have 202-217 ventrals, and 19-24 subcaudals, and 9 9
have 221-241 ventrals, and 15-18 subcaudals, this being based on
seven (four supposedly so) males and three females.

Diet . The tip of a blind snake's ( Typhlops or Leptotyphlops) tail was
present in the stomach of the Ilolo snake.

Habitat. Associated with forested or recently deforested land such
as at Mbango, where Pitman's natives secured one while clearing
thickets of "lantana" from an old rubber plantation. A burrowing
species occurring at altitudes of 4000 (Mubango) to 4600 (Ilolo) feet.

Localities. Belgian Congo: Mambawanga Hill; Poko. Uganda:
Budongo Forest; Bussu; Katebo, n.w. Lake Victoria; Kilembe, e.
of Ruwenzori Mountains ; Mubango, Mabira Forest. Tanganyika

180 bulletin: museum of comparative zoology

Territory: Ilolo; ? Dar es Salaam (see comments regarding this
record under the heading of Remarks).

Range. Belgian Congo (in extreme east) east to Uganda and Tan-
ganyika Territory (in extreme west).

Miodon gabonensis graueri Sternfeld

1908. Miodon Graueri Sternfeld, Sitz. Ges. Naturf. Freunde Berlin, p. 94:

Entetbe, i.e. Entebbe, Uganda.

1908c. Sternfeld, p. 244, fig.

1910a. Sternfeld, p. 35, fig. 40.

1915c. Boulenger, p. 633.

1924b. Loveridge, p. 7.

1925. Werner, 1924, p. 153.

1937. Pitman, p. 334, pi. xii, fig. 4; col. pi. M, fig. 4.

1938b. Pitman, pp. 40, 187, plates as above (reprint).

1942e. Miodon gabonensis graueri Loveridge, p. 298.

Name. Central Lake Region Snake-eater (English).

Description. Rostral nearly twice as broad as deep, just visible from
above; nostril in a divided nasal; internasals as broad as long, as long
as, or almost as long as, the prefrontals; frontal as long as, or longer
than, broad, as long as, or shorter than, its distance from the rostral,
much shorter than the parietals, once and a half to twice as broad as a
supraocular; no loreal ; preocular 1 ; eye small, its diameter half to two-
thirds its distance from the mouth; postocular 2; temporals 1 + 1;
upper labials 7, the third and fourth entering the orbit ; 4 lower labials
in contact with the anterior sublinguals, which are longer than the
posterior. Midbody scales in 15 rows, smooth; ventrals 237-258; anal
divided; subcaudals 13-18 pairs.

Based on data of type and examination of the three Idjwi snakes.

Color. Above, black with an iridescent bloom; a broad white band
crosses head and nape, extending from the back of the frontal across the
parietals and over three scale-rows on the nape. Below, and on lower
half of outer scale-rows, white.

Size. Total length of rf (Mus. Congo), 325 (310 + 15) mm.; total
length of 9 (M.C.Z. 48432), 370 (358 + 12) mm.; both from Idjwi
Island.

Remarks. The species is known only from the type, apparently a
cf; and three others (tf 9 9 ) which I collected on Idjwi Island in
Lake Kivu, Belgian Congo. The latter is in the general region where
Grauer did so much of his collecting and it seems strange that, if the

loveridge: African snakes 181

type was correctly labeled, no second example has been taken at
Entebbe where so much collecting has been done by Johnston, Degen,
Pitman and others.

Sexual dimorphism. Assuming the type to be a male, then c? c? have
237-238 ventrals, and 16-18 subcaudals, and 9 9 have 254-258
ventrals, and 13 subcaudals, but only two of each sex are known.

Diet. Remains of a blind snake (Typhlops b. lestradei) and an egg,
possibly a lizard's, were recovered from two Idjwi snakes.

Habitat. If the type locality is correct, then the fact that two of the
Idjwi snakes were taken near the lake shore, may indicate a preference
of this race for lakeside habitats. The Idjwi male, however, was taken
while crossing a path in bush (recently deforested) country at an
altitude of about 6000 feet.

Localities. Belgian Congo : Idjwi Island in Lake Ki vu. Uganda :
Entebbe.

Range. Western Belgian Congo east to central Uganda.

Genus Aparallactus

1849. Aparallactus A. Smith, 111. Zool. S. Africa, 3, App. p. 15 (type capensis

Smith).
1849. Elapomorphus A. Smith (not Wiegmann), 111. Zool. S. Africa, 3, App.

p. 16, footnote (type capensis Smith).
1854. Uriechis Peters, Monatsb. Akad. Wiss. Berlin, p. 623 (type lunulatus

Peters).
1859a. Elapops Giinther, Ann. Mag. Nat. Hist. (3), 4, p. 161 (type modestus

Giinther).
1860. Periaspis Cope, Proc. Acad. Nat. Sci. Philadelphia, pp. 241, 266

(type plumbeatra Cope = modestus Giinther).
1863a. Cercocalamus Giinther, Ann. Mag. Nat. Hist. (3), 11, p. 21 (type

collaris Giinther = capensis A. Smith).
1923b. Guyomarchia Angel, Bull. Mus. Hist. Nat. Paris, 29, p. 348 (type

unicolor Angel = modestus Giinther).

Maxillary short with 5-10 small teeth followed with or without an
interspace by 1-2 large grooved or solid fangs situated below the eye;
anterior mandibular teeth longest. Head small, not distinct from neck;
eye very small, with round pupil; nasal entire or divided; no loreal;
1, rarely 2, preoculars. Body cylindrical; scales smooth, without pits,
in 15 rows; ventrals rounded. Tail moderate or short; subcaudals
single.

Range. Africa south of 15° N., i.e. from Portuguese Guinea to
Ethiopia.

182 bulletin: museum of comparative zoology

Remarks. It would appear as if Sir A. Smith, realising that his
Elapomorphus was preoccupied by that of Wiegmann in Fitzinger
(1843 p. 25), sent his description of Aparallactus to be inserted by the
printers. Whatever the circumstances, however, he published the
description of capensis twice (p. 15 and p. 16) under different generic
names.

The argument for merging Elapops with Aparallactus was presented
by Bogert (1940, p. 43) whose studies of the dentition of modestus are
responsible for range extensions incorporated in the above description.

Aparallactus hagemanni Gough was long ago removed from this
genus by Boulenger (1919a) when he referred it to the synonymy of
Polemon bocourti Mocquard.

Werner (1925), who was the last to publish a key to the species of
this genus, listed 25, here reduced to 12!

In studying the genus it was interesting to note that a block of six
western and northern species invariably have seven upper labials of
which the third and fourth enter the orbit, while three eastern species
(werneri, turneri and nigriceps) have six labials with the second and
third entering; three others (c. bocagii, c. capensis and c. uluguruensis)
may present either arrangement though the first appears to be normal.

Synopsis of the Species of Aparallactus

1. A single prefrontal; range French Guinea east to Gold Coast. . . .lineakis

(p. 183)
A pair of prefrontals 2

2. First lower labial in contact with its fellow behind the mental 3

First lower labial not in contact with its fellow behind the mental 9

3. One or two upper labials in contact with a parietal 4

Upper labials separated from parietal by temporals 7

4. Usually two upper labials (fifth and sixth, rarely fifth only or sixth only)

in contact with a parietal; a single temporal (very rarely 1+1); range

Sierra Leone east to Uganda modestus

(p. 186)
A single upper labial (the fifth) in contact with a parietal; temporals

1+1 5'

5. Diameter of eye twice its distance from the mouth; range Togo east to

Nigeria liddiardae

(p. 191)
Diameter of eye less than twice its distance from the mouth 6

loveridge: African snakes 183

6. Preocular usually not in contact with the nasal; range Anglo-Egyptian

Sudan (Lado) east to Eritrea south to northern Tanganyika Territory

(Arusha) concolor

(p. 192)

Preocular in contact with the nasal; range Belgian Congo (Dika) and

Northern Rhodesia, east to Mozambique (Tete), south to Transvaal . . .

lunulatus
(p. 195)

7. Upper labials 7, the third and fourth entering the orbit; range dry uplands

of southern Ethiopia (east of Lake Rudolf) to southeastern Tanganyika

Territory (Rondo Plateau) jacksonii

(p. 197)
Upper labials 6, the second and third entering the orbit 8

8. Ventrals of d71 and 9 141-161; range Tanganyika Territory (montane

forests of Magrotto, Usambara and Uluguru) werneri

(p. 199)
Ventrals of <? and 9 120-139; range Kenya Colony (dry coastal plain

between Lamu and Malindi) burneri

(p. 201)

9. Ventrals 168-191; range Angola and South West Africa east through ?

southeastern Belgian Congo (Elisabethville) to western ? Mozambique

(Chifumbazi) and Transvaal (Kruger Park) c. bocagii

(p. 202)
Ventrals 167 or less 10

10. No light and black nuchal collar, uniformly iridescent plumbeous; size

larger; range virgin forests of coastal Kenya Colony (Ngatana) and
Tanganyika Territory (Magrotto, Usambara, Uluguru Mountains) ....

c. uluguruensis

(p. 210)

A black, light-edged nuchal collar; size smaller 11

11. Ventrals 134-166; subcaudals 35-59; range savanna of Tanganyika Terri-

tory (Ujiji) east to southern Kenya Colony (Mt. Mbololo) south to

Pondoland, and west to Angola (Quindumbo) c. capensis

(p. 205)
Ventrals 108-123; subcaudals 20-35; range Mozambique (Inhambane and

Tete) nigriceps

(p. 212)

Aparallactus lineatus (Peters)

1870c. Uriechis (Metopophis) lineatus Peters, Monatsb. Akad. Wiss. Berlin,

p. 643, pi. i, figs. 3-3c: Keta, Guinea, i.e. Quittah, Gold Coast.
1885d. Miiller, p. 678.

184

bulletin: museum of comparative zoology

1893a. Uricchis anomala Boulenger, Ann. Mag. Nat. Hist. (6), 12, p. 273:

Gold Coast. d\

1895h. Aparallactus anomalus Boulenger, p. 173.

1896d. Boulenger, p. 262, pi. xi, fig. 3.

1919b. Boulenger, p. 293.

1921a. Chabanaud, p. 471.

1921b. Chabanaud, p. 525.

1925. Werner, 1924, p. 158.

1933f. Angel, p. 179.

1895h. Aparallactus lineatus Boulenger, p. 173.

1896d. Boulenger, p. 261.

1919b. Boulenger, p. 293.

1925. Werner, 1924, p. 158.

1933f. Angel, p. 178.

1938d. Loveridge, p. 60.

1897a. Aparallactus niger Boulenger, Ann. Mag. Nat. Hist. (6), 19, p. 154:

Sierra Leone, d".

1919b. Boulenger, p. 293.

1921a. Chabanaud, p. 471.

1921b. Chabanaud, p. 525.

1922. Aylmer, p. 15.

1925. Werner, 1924, p. 158.

1933f. Angel, p. 180.

1908a. Elapops hekrolepis Mocquard, Bull. Mus. Hist. Nat., 14, p. 261:

Assini, Ivory Coast. 9 .

1925. Werner. 1924, p. 158.

1917. Rouleophis Chevalieri Chabanaud, 1916, Bull. Mus. Hist. Nat., 22, p.

379, figs. 22-23: Sampouyara, French Guinea.

1917b. Chabanaud, p. 12.

1921a. Chabanaud, p. 471.

1933f. Angel, p. 181, figs. 67-67a.

Description. Rostral nearly twice as broad as deep, the portion
visible from above one third1 to half2 as long as its distance from the
frontal; nostril in an entire or semidivided nasal; internasals as long
as, or much shorter than, the single prefrontal; frontal once and a
third to once and two thirds as long as broad (in the middle), as long
as, or much longer than, its distance from the end of the snout, much
shorter than the parietals, twice as broad as a supraocular; no loreal;
preocular 1, in contact with the nasal; eye small, its diameter equal to,
or slightly less or greater than, its distance from the mouth; post-
oculars l2 or 21, separated from the only temporal 1, flanking the

1 In type of niger.

- In type of chevalieri.

loveridge: African snakes 185

outer border of the parietal without any enlarged shield below it
posteriorly; upper labials 7, the third and fourth entering the orbit,
sixth largest, fifth and sixth in contact with a parietal; first lower
labial in contact with its fellow behind the mental; 4, rarely 5, lower
labials in contact with the anterior sublinguals, which are as long as,
or slightly longer than, the posterior. Midbody scales in 15 rows,
smooth on females and young males, usually keeled on tail and pos-
terior part of body in adult males; ventrals 151-170; anal entire;
subcaudals SS^O.

Based on all original descriptions and literature, also a#young
striped cf (unquestionably lineatus) from "Guinea" (M.C.Z. 25826),
and two adult plumbeous cf cf (undoubtedly niger) from Sanoquelle,
Liberia (M.C.Z. 43206-7).

Color. Juvenile and Female. Above, olive, with three dark longi-
tudinal lines, each scale of the outer series with a yellow spot. Below,
pinkish white minutely speckled with gray.

Adult. Above, uniformly black. Below, and on lips, white, each
ventral and subcaudal shield edged with black laterally and poste-
riorly and with a median black line, or tendency thereto, on tail.

Size. Total length of cf (type of anomalus), 440 (360 + 80) mm.,
and total length of 9 (type of lineatus), 440 (383 + 57) mm.

The maximum length of 605 (407 + 198) mm. for chevalieri furnished
by Chabanaud (1921a) appears to be erroneous, for Angel (1933f) in
his redescription of the species, gives only 475 (375 + 100) mm. for
an unsexed, but presumably a cf, specimen.

Remarks. Boulenger (1919a) referred Rouleophis chevalieri to the
synonymy of niger, an action with which Chabanaud (1921a) con-
curred. Angel (1933f), however, demurred on the grounds that the
type of niger had 2 postoculars, while the six examples of chevalieri
possess only 1, and for other reasons, which, after a study of the
genus, I consider of even less importance. It is a fact that the type
of niger, which is the most westerly specimen, remains unique in the
possession of 2 postoculars; on the other hand, its nearest ally,
modestus, displays 1 or 2 indiscriminately throughout its range
though in a selected locality one or other condition may be fairly
constant.

Angel (1933f) synonymized Elapops heterolcpis with anomalus;
later I (1938d) placed all four in the synonymy of lineatus for reasons
stated there in considerable detail.

Sexual dimorphism. If one assumes the type of anomalus to have

1 It seems possible that 33 and 35 were based on truncated tails, the next lowest subcaudal
count is 41.

186

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been a 9 , and not a d", then: cfcf have 151-164 ventrals, and 58-60
subcaudals, and 9 9 have 161-170 ventrals, and 331 or 41-52 sub-
caudals.

Localities. French Guinea: Beyla; Sampouyara. Sierra Le-
one. Liberia: Sanoquelle (Sanikole). Ivory Coast: Assini.2
Gold Coast: Quittah (Keta).

Range. French Guinea east to Gold Coast.

1859a.

1860.

1865b.

1875a.

1884a.

1891b.

1893c.

1896d.

1897b.

1897.

1898.

1898a.

1899a.

1900b.

1903.

1906i.

1908a.

1909a.

1909b.

1910.

1911.

1915a.

1919b.

1919g.

1922.

1923.

1925.

1927d.

1933f.

1933m.

1934a.

Aparallactus modestus (Giinther)

1 Elapops modestus Giinther, Ann. Mag. Nat. Hist. (3), 4, p. 161, pi.
iv, fig. C: West Africa.
Cope, p. 566.
Giinther, p. 152.
Peters, p. 198.

Rochebrune, p. 151 (ignored).
Matschie, p. 614.
Matschie, p. 213.
Boulenger, pp. 262, 649.
Mocquard, p. 8.
Sjostedt, p. 35.
Boettger, p. 112.
Werner, p. 210.
Werner, p. 141.
Boulenger, p. 454.
Gough, p. 468.
Boulenger, p. 214.
Sternfeld, pp. 414, 429.
Sternfeld, p. 22, fig. 34.
Sternfeld, p. 22, fig. 29.
Muller, p. 615.
Despax, p. 240.
Boulenger, p. 217.
Boulenger, p. 293.
Boulenger, p. 27.
Aylmer, pp. 15, 22.
Schmidt, p. 121, pi. xvii, fig. 2.
Werner, 1924, p. 158.
Witte, p. 326.
Angel, p. 183, figs. 68-68a.
Witte, p. 96.
Schwetz, p. 381.

1 It seems possible that 33 and 35 were based on truncated tails, the next lowest subcaudal
count is 41.

s Given as Assinie, presumably the town on the Ivory Coast side of the border.

loveridge: African snakes 187

1936h. Loveridge, p. 40.

1937c. Loveridge, p. 278.

1937. Pitman, p. 337, pi. xii, fig. 6, col. pi. N, figs. 1-2.

1938a. Pitman, pp. 218, 233.

1938b. Pitman, pp. 188, 190, 316, 331, plates as above (reprint).

1860. Periaspis plumbeatra Cope, Proc. Acad. Nat. Sci. Philadelphia, p.

242: Liberia.

1860. Elapops plumbeater Cope, p. 566.

1862. Elapops (Calamaria) Petersi Jan, Arch. Zool. Anat. Fisiol., 2, p. 32:

Gold Coast.

1865. Jan, livr. 13, pi. iii, fig. 2.

1885d. Muller, p. 678.

1889. Mocquard, p. 145.

1890b. Muller, p. 692.

1893. Prato, p. 10.

1896. Aparallactus boulengeri Werner, Verh. Zool.-Bot. Ges. Wien, 46, p.

363, pi. vi, figs. 6-6b: Cameroon.

1897. Sjostedt, p. 35 (as boulangeri).

1897b. Aparallactus peraffinis Werner, Verh. Zool.-Bot. Ges. Wien, 47, p.

404, pi. ii, fig. 3: Interior of Cameroon.
1897b. Aparallactus ubangensis Boulenger, Ann. Mag. Nat. Hist. (6), 19, p.

279, fig. : Zongo, Ubangi Rapids, Belgian Congo.
1901g. Boulenger, p. 11, pi. iv, fig. 2. ,
1903b. Bocage, p. 64.
1915a. Boulenger, p. 217.
1925. Werner, 1924, p. 151.

1901g. Aparallactus flavitorques Boulenger, Ann. Musee Congo (1), 2, p. 11,

pi. iv, fig. 3: Lubue, Kasai, Belgian Congo.
1903b. Bocage, p. 64.
1915a. Boulenger, p. 217.
1925. Werner, 1924, p. 151.

1927d. Witte, p. 325.
1933m. Witte, p. 96.
1938b. Pitman, pp. 40, 191.
1902a. Aparallactus dolloi Werner, Verh. Zool.-Bot. Ges. Wien, 52, p. 346:

Banzyville, Ubangi River, Belgian Congo.
1915a. Boulenger, p. 216.
1919b. . Boulenger, p. 292.
1925. Werner, 1924, p. 157.

1902a. Aparallactus congicus Werner, Verh. Zool.-Bot. Ges. Wien, 52, p. 346:

Lingunda, Belgian Congo.
1915a. Boulenger, p. 217.
1925. Werner, 1924, p. 157.

1907c. Aparallactus Batesii Boulenger, Ann. Mag. Nat. Hist. (7), 19, p. 325:

5 miles inland from Kribi, French Cameroon.

188 bulletin: museum of comparative zoology

1908a. Sternfeld, pp. 414, 429.

1909b. Sternfeld. p. 22.

1919b. Boulenger, p. 292.

1925. Werner, 1924, p. 157.

1910c. Aparallactus christyi Boulenger, Ann. Mag. Nat. Hist. (8), 6, p. 512:

Mabira Forest, Chagwe, Uganda.
1915c. Boulenger, p. 634.
1924b. Loveridge, p. 7.
1925. Werner, 1924, p. 158.

1937f. Loveridge, p. 502.
1938b. Pitman, pp. 40, 190, 191.
1939c. Scortecci (part), p. 178 (omit record).
1917. Aparallactus nigrocollaris Chabanaud, 1916, Bull. Mus. Hist. Nat.

Paris, 22, p. 377, figs. 18-19: French Congo.
1919b. Boulenger, p. 293.
1925. Werner, 1924, p. 157.

1938a. Pitman, p. 198, pi. xvii, fig. 5, col. pi. W, fig. 6.
1938b. Pitman; p. 296, plates as above (reprint).
1917. Aparallactus nigrocollaris Roucheti Chabanaud, 1916, Bull. Mus.

Hist. Nat. Paris, 22, p. 378, figs. 20-21 : French Congo.
1919b. Aparallactus roucheti Boulenger, p. 293.
1925. Werner, 1924, p. 157.

1923b. Guyomarchia unicolor Angel, Bull. Mus. Hist. Nat. Paris, 29, p. 348,

figs. 1-4: French Congo (probably from near Sangha).
1924b. Aparallactus Graueri Werner, Sitz. Akad. Wiss. Wien, 133, p. 42:

Beni, Belgian Congo.
1925. Werner, 1924, p. 158.

1940. Aparallactus modestus Bogert, p. 43, fig. 5.
1941e. Loveridge, p. 123.
1942e. Loveridge, p. 299.

Further citations of 'modestus' and 'christyi' will be found under
concolor.

Names. Gray Forest Snake (English) ; mboli (Togo) ; kileba ( Wamba)
Description. Rostral nearly twice as broad as deep, the portion
visible from above half to three quarters as long as its distance from
the frontal; nostril in a divided, rarely semidivided, or entire1 nasal;
internasals as long as, or slightly shorter than, the prefrontals; frontal
once and a third to once and two thirds2 as long as broad (in the mid-
dle), as long as, or much longer than, its distance from the end of the
snout, much shorter than the parietals, twice as broad as a supra-

'In types of boulengeri, nigrocollaris and roucheti.

2In type of peraffinis said to be twice but only two thirds in figure of type.

loveridge: African snakes 189

ocular; no loreal1; preocular 1, in contact with, or rarely separated
from2, the nasal; eye small, its diameter equal to, or slightly less or
slightly greater than, its distance from the mouth; postoculars 1-2,
not in contact with the only temporal; temporal 1, rarely 1 + l3,
flanking the outer border of the parietal with a smaller enlarged shield
below and posterior to it ; upper labials 7, the third and fourth entering
the orbit, sixth largest, fifth, or fifth and sixth, or sixth4 only in contact
with a parietal; first lower labial in contact with its fellow behind
the mental; 4, sometimes 3, lower labials in contact with the anterior
sublinguals, which are as long as, or slightly longer than the posterior.
Midbody scales in 15 rows, smooth; ventrals 134-164; anal entire;
subcaudals 32-51.

Based on the descriptions of all species in the synonymy and litera-
ture after examination of the entire series in the M.C.Z. The following
color description is likewise a composite of adult and young from the
original descriptions.

Color. Above, black, brown, yellow brown, olive gray, or blue gray
with each scale edged with black, uniform, or a more or less faintly
indicated pale, yellow, or black nuchal collar; labials pale or dusky
reddish yellow, more or less blotched with black. Below, except for
outer (lateral) edges of the ventrals which are colored like the back,
orange, yellow, white, gray, or grayish green, uniform or edged with
lighter, or with dusky or black flecks and infuscations, particularly on
tail which may be entirely gray or black beneath. Eye brown (fide
Pitman).

Size. Total length of d\ 457 (377 + 80) mm. from Mabira Forest
.(Pitman, 1937); total length of 9 (M.C.Z. 9253), 559 (486 + 73)
mm. from Lolodorf. Angel mentions 565 mm. for an unsexed snake.
Remarks. The species plumbeatra and petersi were synonymized by
Gunther (1865b), boulengeri and pcraffinis by their own author (1898,
p. 192), Werner, who later (1925) added unicolor. Parker (in Pitman,
1937) decided that flavitorques and christyi were nothing but juvenile
modestus, for Schmidt (1923) had already pointed out the complete
transition from collared juveniles to uniform adults in his series of

'Gough (1903) reports a loreal as present on the right side of a West African snake (pre-
sumably split off from the posterior nasal).

2ln type of nigrocollaris and in M.C.Z. 38961 (Liberia) and on left side of M.C.Z. 29355
(Cameroon).

3 See remarks regarding nigrocollaris under Remarks.

4Werner, in describing graueri, says third and fourth or fourth and fifth, but this appears
to be a lapsus for fourth and fifth and fifth and sixth as he has already stated that the third
only or third and fourth upper labials enter the orbit in the azygous type which possesses
6 labials on one side, 7 on the other.

190 bulletin: museum of comparative zoology

nineteen snakes from the Ituri region. The remaining seven species I
added to the synonymy in 1942e.

A. nigrocollaris from French Congo was founded on two specimens
in which the posterior nasal has fused with the second upper labial,
resulting in the second upper labial being in contact with the prefrontal.
In A. n. roucheti, with same locality and collector, the arrangement is
normal. It seems reasonable to assume that the condition is an aber-
ration for in one of our Liberian specimens (M.C.Z. 38961) the second
upper labial is broadly in contact with the prefrontal, while in another
Liberian snake (M.C.Z. 38962) it is separated. In one Cameroon
reptile (M.C.Z. 9253) the second upper labial is narrowly in contact
on the left side only, in yet another (M.C.Z. 29355) it is the third
upper labial which is narrowly in contact. In view of Pitman having
twice obtained nigrocollaris in Uganda, I suggest that the condition
may crop up in any part of the range from Liberia to Uganda, for the
other point of difference — that of the temporal being subdivided —
also occurs on one side of the head in a Uganda snake.

Angel's (1925) reference to modestus from Kenya Colony, and that
of Loveridge (1929b) quoted by Scortecci (1939c) for a snake taken
between Kenya and Ethiopia which was referred to christyi are now
transferred to concolor after reexamination of both specimens.

Dentition. This has been studied recently by Bogert (1940) as
indicated in the generic diagnosis.

Hemipenis. The hemipenis of modestus has been described in detail
and figured by Bogert (1940).

Sexual dimorphism. Assuming that the Niger specimen mentioned
by Boulenger (1896d) and his type of batesii were males and not
females then cfcf have 135-144 ventrals, and 41-51 subcaudals, while
9 9 have 152-164 ventrals, and 34-44 subcaudals.

Breeding. Without date, at Kribi, a 9 held 7 eggs measuring about
20 x 5 mm., on November 13, at Mabira Forest, a 9 held 7 eggs
averaging about 25 x 8 mm.

Diet. Nothing known !

Temperament. "Extremely placid and makes no attempt to bite
when handled." (Pitman).

Habitat. A forest-dwelling species, taken during clearing operations
or in coffee plantations on previously forested areas.

Localities. Sierra Leone. Liberia. Harbel. Gold Coast.
Togo: Adele (Bismarckburg); Grand Popo; Misahohe; Wegbe.
Nigeria: Lagos; Niger River. British Cameroon: Bonjongo
(? Banjo); Buea; Johann Albrechts Heights. French Cameroon:

loveridge: African snakes 191

Bipindi; Bitye; Dibongo near Edea; Kribi; Ja River; Lolodorf;
Longji; Metet; Mukonje Farm, Mundame; Sakbayeme; Yaunde.
Spanish Guinea: Esong (as Esosung, Bakossiberge). French
Congo: Fernand Vaz; Lambarene; Loudinia-Niari, Niari River;
? near Sangha; Sette Kama (Cette Cama). Belgian Congo:
Avakubi Banzyville; Beni; Kai Bumba; Lingunda; Lubue, Kasai;
Lukolela; Makaia Ntete; Medje; Ngombe, Kasai; Saidi; Stanley-
ville; Temvo; Zongo. Uganda: Bisu; Budongo Forest; Bundi-
bugyo; Mabira Forest; 'Rom Mtn., northeast iVcholi; ^emliki
Valley.

The two Kenya Colony records, as mentioned above, were
based on misidentified examples of concolor.

Range. Equatorial Africa from Sierra Leone east to Uganda.

Aparallactus liddiardae Parker

1908b. Aparallactus bocagii Sternfeld (not Boulenger), pp. 219, 234 (Togo).
1909a. Sternfeld, p. 22.

1933. Aparallactus liddiardae Parker, Ann. Mag. Nat. Hist. (10), 12, p. 545:
Jos, northern Nigeria.

Description. Rostral broader than deep, the portion visible from
above less than half as long as its distance from the frontal; nostril in
a divided nasal ; internasals shorter than the prefrontals ; frontal once
and a half as long as broad (in the middle), much longer than its dis-
tance from the end of the snout, slightly shorter than the parietals; no
loreal; preocular 1, in contact with the nasal; eye small, its diameter
equal to twice its distance from the mouth ; postocular 1, not in contact
with the anterior temporal; temporals 1 + 1 ; upper labials 7, the third
and fourth entering the orbit, fifth in contact with a parietal; first
lower labial in contact with its fellow behind the mental ; 4 lower labials
in contact with anterior sublinguals, which are longer than the pos-
terior. Midbody scales in 15 rows, smooth; ventrals 170 (Togo) to
174 (Nigeria); anal entire; subcaudals 41 (Togo) to 45 (Nigeria).

Based on description of type and data from Sternfeld (1908b).

Color. Above gray, upper surface of the head darker, edges of
parietal and temporal shields stippled with black, ocular region and lip
below the eye, black; a broad black nuchal collar; the third scale-row
on each side and the seven mid-dorsal rows each with a dark spot, the

1 As nigrocollaris.

192

bulletin: museum of comparative zoology

spots of the middle row largest, and forming a narrow line which is
continued forward on to the parietal shields. Below, pale gray.

Size. Total length of type, 80 (44 + 36) mm.

Diet. A relatively large centipede.

Localities. Togo: Sausane Mangu. Northern Nigeria: Jos.

Range. Togo east to Nigeria.

Aparallactus concolor (Fischer)

1884a. Uriechis concolor Fischer, Jahrb. Hamburg. Wiss. Anst., 1, p. 4,

pi. i, fig. 1 : Arusha, Tanganyika Territory.

1888b. Giinther, p. 325.

1895h. Aparallactus concolor Boulenger, p. 172.

1896d. Boulenger, p. 257.

1896e. Boulenger, p. 216.

1896. Tornier, p. 79.

* 1896c. Boulenger, p. 21.
1897g. Boulenger, p. 279.

1897. Tornier, p. 65.

* 1902b. Mocquard, p. 406.
1907. Lonnberg, p. 16.
1908c. Sternfeld, p. 241.
1909d. Boulenger, p. 311.
1910. Lepri, p. 327.
1910a. Sternfeld, p. 36.

1912. Hobley, p. 53.
1912c. Sternfeld, p. 274.

1913. Lonnberg & Andersson, p. 5.
1915c. Boulenger, p. 634.

1915d. Boulenger, p. 655.

1916a. Loveridge (part), p. 86.

1923b. Calabresi, p. 162.

1924b. Loveridge, p. 7.

1925. Werner, 1924, p. 157.

1927. Calabresi, pp. 33, 56.

1929h. Loveridge, p. 34.

1934c. Scortecci, p. 74, fig. 32.

1936J. Loveridge, p. 269.

*1936e. Parker, p. 608.

1937f. Loveridge, pp, 493, 496.

1937. Pitman, p. 335, pi. xii, fig. 5, col. pi. M, fig. 5.

1938a. Pitman, p. 217.

1938b. Pitman, pp. 40, 188, 315, plates as above (reprint).

loveridge: African snakes 193

*1939a. Scortecci, p. 285.

1939c. Scortecci, p. 175 (as A. c. concolor).

1940a. Scortecci, p. 136.

?1896a. Aparallactus lunulatus Boulenger (not Peters), p. 554.

?1928b. Scortecci, p. 306.

1925a. Elapops modestus Angel (not Gunther), p. 36.

*1929h. Aparallactus christyi Loveridge (not Boulenger), p. 34.

*1939c. Scortecci (part), p. 178.

*1931c. Aparallactus concolor boulengeri Scortecci, Atti. Soc. Ital., 70, p. 212:
Villaggio Duca Abruzzi and inland from Mogadiscio, Italian
Somaliland.

* 1934c. Scortecci, p. 75.

*1939c. Scortecci, p. 177, figs. 99-100.

Names. Plumbeous Centipede-eater (English); mowa (Teita, but
generic).

Description. Rostral nearly twice as broad as deep, the portion
visible from above half to two thirds as long as, or even equal to, its
distance from the frontal ; nostril in a divided, semidivided, or entire
nasal (which is fused with a labial on one side of the head in the
cotype 9 of boulengeri) ; internasals much shorter than the prefrontals ;
frontal once and a half to twice as long as broad (in the middle), much
longer than its distance from the end of the snout, as long as, or
slightly shorter than, the parietals; once and two thirds to twice as
broad as a supraocular; no loreal; preocular 1, not, or but rarely1, in
contact with the nasal ; eye small, its diameter equal to, or greater than,
its distance from the mouth; postocular 1, not in contact with the
anterior temporal ; temporals 1 + 1 ; upper labials 7, the third and
fourth entering the orbit, fifth largest and in contact with a parietal;
first lower labial in contact with its fellow behind the mental ; 4 lower
labials in contact with the anterior sublinguals, which are about as
long as the posterior. Midbody scales in 15 rows, smooth; ventrals
1402-1733; anal entire; subcaudals 43-77.

Based on original descriptions, literature, and Kenya material in
the M.C.Z. and U.S.N.M.

'Some or all of these specimens have the preocular in contact with the nasal a character of
boulengeri which may yet prove to be B recognizable race, though the name is preoccupied by
boulengeri Werner, 1896.

1 In Athi Plains specimen (Mocquard) and Kenya-Ethiopian frontier snake (Loveridge), types
of boulengeri, etc.

2 In "lunulalus" from Let Marefia; Calabresi's record of 133 is rejected.

3 In "lunulalus" from Isole; if these two are not concolor then the range is 143—169.

194 bulletin: museum of comparative zoology

Color. Above, uniformly plumbeous. Below, rather paler, sometimes
almost white, (c? and 9 adults. Tanganyika and Kenya. Loveridge).

Above, uniformly brown with a light purplish reflection. Below,
slightly lighter, (adult. Mogadish, Italian Somaliland. Lepri, 1910).

Above, rather pale yellowish, a large transverse black band on hinder
part of head and nape, each dorsal scale edged with brown. Below,
uniformly whitish yellow. ( cf 9 adults. Durgale to Magghiole, Italian
Somaliland. Calabresi, 1927).

Above, pale maroon, a four-scale wide transverse black band on
hinder part of head and nape, pre- and postocular and third or third
and fourth labials black. Below, very pale rosy yellowish. (Neghelli,
Ethiopia. Scortecci, 1940a).

Size. Total length of type, ?cf, 360+ (300 + 60+) mm. from
Arusha; and total length of 9 (M.C.Z. 40711), 520 (420 + 100) mm.
from Voi.

Remarks. Werner's (1908) reference to concolor in the Sudan appears
to have been based on a misidentification, possibly of a Prosymna.
The snake from Kitui, identified by Boulenger, and listed by Loveridge
(1916a) is evidently something else.

Parker (1936e) synonymized boulengeri with concolor the same year
that I (1936J) placed it in the synonymy of uluguruensis! Actually it
is intermediate between the two, agreeing with concolor in the first
lower labial being in contact with its fellow, but agreeing with uluguru-
ensis in having the preocular in contact with the nasal. Of these char-
acters that of the lower labials is more constant and important in the
genus so I admit that I was wrong in my disposition of boulengeri.

I am inclined to think — on account of the extensive range in
variation in ventral and subcaudal counts — that we really are dealing
with two forms and that a race — for which the name boulengeri is not
available — does occur in Italian Somaliland. The question can only
be decided by someone assembling all the concolor material, sexing it,
and seeing whether there is any discernible geographical significance
or correlation between higher scale counts and naso-preocular contact.
The data at my disposal does not reflect it, but perhaps authors have
referred to concolor, without comment, snakes that might have been
assigned to "boulengeri" .

There is an undoubted tendency in northern Kenya and southern
Somaliland for the preocular to be in contact with the nasal. In the
northern part of its range, concolor certainly exhibits a wider variation
in the number of ventrals, which do not go above 158 in the south.
The coloration in that region also appears to be uniformly plumbeous,

loveridge: African snakes 195

thereby differing from many specimens described from Somaliland
and Eritrea.

Diet. A centipede, about as long as a finger and nearly the same
diameter as the snake itself, in Kenya (Sternfeld).

Localities. Anglo-Egyptian Sudan: Lado. Eritrea: Isole1
near Massaua. Ethiopia: Between Ethiopia and Kenya; between
Gara Mulata and Lake Haramaya; Let Marefia3, Shoa; Neghelli.
Italian Somaliland: Balad; Bardera; Belet Amin; Durgale to
Magghiole; Kismayu; between Lugh and Matagoi; Mofi; Moga-
dish; between Obbia and Tobungab; Villa Duca de Abruzzi.
Kenya Colony: Athi Plains; Boran country; Bulessa; Bura;
Lamu Island; Lodwar; Mount Mbololo; Mtito Andei; east of
Tsavo (Izavo); Turkana; Voi. Tanganyika Territory: Arusha;
Usambara.

Range. Anglo-Egyptian Sudan east to Eritrea and south to extreme
northern Tanganyika Territory.

Aparallactus lunulatus (Peters)

1854. Uriechis lunulatus Peters, Monatsb. Akad. Wiss. Berlin, p. 623: Tete>

Mozambique.

1855. Peters, p. 53.

1882a. Peters, p. 113, pi. xviii, fig. 2.

1888b. Gunther, p. 324.

1891a. Boulenger, p. 308.

1896a. Bocage, p. 100.

1895h. Aparallactus lunulatus Boulenger, p. 172.

1896d. Boulenger, p. 258.

1907a. Boulenger, p. 12.

1910a. Sternfeld, p. 36.

1915a. Boulenger, p. 216.

1915c. Boulenger, p. 634.

1917. Sternfeld, p. 481.

1924b. Loveridge, p. 7.

1925. Werner, 1924, p. 158.

1928d. Loveridge, p. 57.

1933m. Witte, p. 96.

1934. Pitman, p. 298.

1937f. Loveridge, p. 496.

1937b. Mertens, p. 14.

Further citations of 'lunulatus' will be found under concolor and c.
bocagii.

i Recorded as lunulatus by Boulenger and Scortecci respectively, have not been reexamined.

196 bulletin: museum of comparative zoology

Names. Blotched-back Centipede-eater (English); bubse (Tete).

Description. Rostral nearly twice as broad as deep, the portion
visible from above one third to half as long as its distance from the
frontal ; nostril in a divided or entire1 nasal ; internasals much shorter
than the prefrontals; frontal once and a half to once and two thirds as
long as broad (in the middle), much longer than its distance from the
end of the snout, as long as, or slightly shorter than, the parietals;
once and two thirds to twice as broad as a supraocular; no loreal;
preocular 1, in contact with the nasal; eye small, its diameter equal to,
or greater than, its distance from the mouth; postocular 1, not in
contact with the anterior temporal ; temporals 1 + 1 ; upper labials 7,
the third and fourth entering the orbit, fifth largest and in contact
with a parietal ; first lower labial in contact with its fellow behind the
mental; 4, sometimes 32, lower labials in contact with the anterior
sublinguals, which are about as long as, or shorter than, the posterior.
Midbody scales in 15 rows, smooth; ventrals 151-1673; anal entire;
subcaudals 48-58.

Based on original description, literature, and two specimens in the
Museum of Comparative Zoology.

Color. Composite. Above, olive, olive gray, or olive green, dark or
light brown, uniform or with a light yellow collar preceding a black
transverse band which may be followed by a series of black bars or
spots; a black fleck below the eye; each scale of back edged with
darker. Below, white or greenish yellow.

Size. Total length of <? (M.C.Z. 23075), 288 (232 + 56) mm. from
Mukwese, being surpassed by the unsexed type, measuring 415 (325
+ 90) mm., and a 485 (385 + 100) mm. specimen from Punda Maria,
Transvaal.

Remarks. The records of Boulenger (1896a) and Scortecci (1928b,
repeated 1939c), and listing of Zavattari (1930b) of two "lunulatus"
from Ethiopia and Eritrea respectively, have been transferred arbi-
trarily to concolor. This has been done not merely because lunulatus
is otherwise unknown north of Tanganyika, but because the scale
counts of both and color (of the one furnished) approximate more
nearly to those of concolor, with which species they appear to have
been identified on the basis of Boulenger 's (1896d) key which fails to
distinguish concolor as now understood.

1 Entire in Victoria Falls specimen (M.C.Z. 21481).

2 Three in Victoria Falls and Mukwese specimens in M. C. Z. Perhaps these are aberrant
c. capensis, and the type and Boulenger's two snakes, which had four, represent true lunulatus
which would then agree with its preceding allies in this character.

3 167 in a Punda Maria, Transvaal, snake (FitzSimon's letter of 18, vii, 1941), otherwise 161.

loveridge: African snakes 197

Indeed it seems highly probable that the eleven recorded, and widely
scattered specimens of lunulatus may prove to be only aberrant c.
capensis in which the first lower labials are in contact behind the
mental — throwbacks to their ancestral condition. This appears to be
the only difference between the two if one excepts the strange coloring
of the type, for subsequent specimens appear to be colored like c.
capensis. In this connection it may be noted also that Peters himself
recorded both species from Tete, and both occur at Victoria Falls.

Localities. Belgian Congo : Dika. Tanganyika Territory:
Duma; Lake Tanganyika; Mukwese near Manyoni; Rufigi River;
Mozambique: Cheringoma Farm, Inhaminga; Tete. Nyasaland:
Lake Nyasa. Northern Rhodesia: Ulungu Mountain west of
Luangwa River; Victoria Falls. Transvaal: Punda Maria.1

Range. Belgian Congo and Northern Rhodesia east to Mozambique,
south to Transvaal.

Aparallactus jacksonii (Giinther)

1888b. Uriechis Jacksonii Giinther, Ann. Mag. Nat. Hist. (6), 1, p. 325, pi

xix, fig. E: Foot of Mount Kilimanjaro, Tanganyika Territory.

1895h. Aparallactus jacksonii Boulenger, p. 172.

1896d. Boulenger, pp. 256, 649.

1896. Tornier, p. 79.
1898a. Boulenger, p. 721.

1897. Tornier, p. 65.
1907. Lonnberg, p. 16.
1910a. Sternfeld, p. 35.
1912. Hobley, p. 53.
1915c. Boulenger, p. 633.
1916a. Loveridge, p. 86.
1916b. Loveridge, p. 122.
1918a. Loveridge, p. 325.
1923e. Loveridge, p. 889.
1924b. Loveridge, p. 7.
1925. Werner, 1924, p. 157.
1928g. Loveridge, p. 41.
1929h. Loveridge, p. 34.
1936. Roux, p. 178.
1937f. Loveridge, p. 496.
1937d. Mertens, p. 8.
1939c. Scortecci, p. 174.
1942e. Loveridge, p. 300.

Further citation of 'jacksonii' will be found under turncri.

167 in a Punda Maria, Transvaal, snake (FitzSimon's letter of 18, vii, 1941), otherwise 161.

198 bulletin: museum of comparative zoology

Names. Jackson's Centipede-eater (English).

Description. Rostral nearly twice as broad as deep, the portion visi-
ble from above one third to half as long as its distance from the
frontal; nostril in a semidivided or entire nasal; internasals much
shorter than the prefrontals ; frontal once and a half to once and two
thirds as long as broad (in the middle), much longer than its distance
from the end of the snout, as long as, or slightly shorter than, the
parietals, once and a half to twice as broad as a supraocular; no
loreal; preocular 1, in contact with the nasal; eye small, its diameter
equal to, or greater than, its distance from the mouth; postoculars 2,
in contact with the anterior temporal ; temporals 1 -f- 1 ; upper labials
7, the third and fourth entering the orbit, sixth largest but not in
contact with a parietal ; first lower labial in contact with, very rarely
separated from1, its fellow behind the mental; 3 lower labials in
contact with the anterior sublinguals, which are about as long as, or
slightly longer than, the posterior. Midbody scales in 15 rows, smooth;
ventrals 134-157; anal entire; subcaudals 33-46.

Based on original description, literature, and eight specimens.

Color. Above, head black, a six-scale wide transverse black band,
edged before and behind by scale-wide bands of bright yellow, on nape ;
back and tail a delicate pinkish brown (in life) or terra cotta with, or
without, a fine black vertebral line, more rarely a lateral series of white
scales bordered with black above and below, which, if confluent, would
form lateral lines. Below, bright yellow (in life) or white.

Size. Total length of d\ 276 (228 + 48) mm. from foot of Mount
Longido; total length of 9 (M.C.Z. 48442), 259 (213 + 46) mm. from
Nchingidi.

Sexual dimorphism, cfcf have 134-144 ventrals, and 35-46 sub-
caudals. 9 9 have 148-157 ventrals, and 33-44 subcaudals.

Temperament. Inoffensive, not attempting to bite.

Habitat. This species favours the hot upland steppe with scattered
acacia forest where, during the rains, I collected half-a-dozen beneath
boulders, stones and logs in the centipede-infested region at the
western foot of Mount Longido.

Elsewhere2 I have recounted how one of these snakes attempted to
cross the face of a recumbent trooper encamped at the foot of Mount
Meru, and was subsequently captured beneath his blankets.

Localities. Ethiopia: Between Dime (? Dima) and Lake Ru-

1 Separated in one of a series of six Longido snakes (U.S.N.M. 62919) .
2 Fauna (Philadelphia), 4, p. 119. December, 1942.

loveridge: African snakes 199

dolf. Kenya Colony: Kell's Farm near Nairobi; Lamu Island1
(!); Lat. O'O, Long. 39E.; Naivasha; "Uganda".2 Tanganyika
Territory: foot of Mt. Kilimanjaro; foot of Mt. Longido; Matete
Bach; foot of Mt. Meru near Ngare Mtoni; Nchingidi, Rondo
Plateau; Ngare na Nyuki; Tanga (!).

Range. Dry uplands from southern Ethiopia to southeastern Tan-
ganyika Territory.

Aparallactus werneri Boulenger

1895h. Aparallactus Werneri Boulenger, Ann. Mag. Nat. Hist. (6), 16, p. 172 •'

Usambara Mountains, Tanganyika Territory.
1896d. Boulenger, p. 257.

1896. Tornier, p. 79.
1897c. Mocquard, p. 123.

1897. Tornier, p. 65.

1898. Boettger, p. 111.
1910a. Sternfeld, p. 35, fig. 41.
1911b. Nieden, p. 442.
1915c. Boulenger, p. 633.
1923e. Loveridge, p. 889.
1924b. Loveridge, p. 7.

1925. Werner, 1924, p. 157.

1926. Werner, p. 248.

1928c. Barbour & Loveridge, p. 131.

1928g. Loveridge, p. 41.

1937f. Loveridge, p. 502.

1942e. Loveridge, p. 301.

Names. Werner's Centipede-eater (English) ; nyolca usambia (Sham-
baa, but applied by them to Neusterophis o. uluguruensis also).

Description. Rostral nearly twice as broad as deep, the portion
visible from above half to two thirds as long as its distance from the
frontal; nostril in an entire (or semidivided3) nasal; internasals shorter
than, or equal to, the prefrontals; frontal once and a third to once
and two thirds as long as broad (in the middle), much longer than its
distance from the end of the snout, as long as, or slightly shorter than,
the parietals; once and two thirds to twice as broad as a supraocular;
no loreal ; preocular 1, in contact with the nasal; eye small, its diameter

'Sternfeld's (1910a) record, more probably an A. turneri.

5 Boulenger's (1896d, p. 649) record, for "Uganda" on this page read Kenya Colony.

3 Fide Tornier, but entire in all our fifty snakes.

200 bulletin: museum of comparative zoology

equal to, or greater than, its distance from the mouth; postoculars 2,
very rarely l1, in contact with the anterior temporal ; temporals 1 + 1 ;
upper labials 6, the second and third entering the orbit, fifth largest
but not in contact with a parietal ; first lower labial not, or but very
rarely2, in contact with its fellow behind the mental ; 3 lower labials in
contact with the anterior sublinguals, which are about as long as, or
slightly longer than, the posterior. Midbody scales in 15 rows, smooth ;
ventrals 141-161; anal entire; subcaudals 32-45.

Based on original description, literature, and over fifty specimens
in the Museum of Comparative Zoology.

Color. Above, head black, a six-scale wide transverse black band,
edged before, and sometimes behind, by scale-wide bands of olive or
yellow, on nape ; back and tail usually olive, rarely pale brown. Below,
throat whitish tinged with yellow, rest of undersurface bright lemon
yellow.

Size. Total length of a cotype, 390 (325+ 65) mm. from Usam-
bara Mountains, and total length of a 9 (M.C.Z.), 354 (295 + 59)
mm. from Amani.

Sexual dimorphism. Most unfortunately, in 1928, I included two
young males in a series of thirty-three females, thus masking the
dimorphism in ventral counts. I have carefully rechecked all extremes
of the series of fifty specimens now available and find that cf cf have
141-151 ventrals, and 35-45 subcaudals, while 9 9 have 154-1613
ventrals, and 32-42 subcaudals.

Breeding. In November, at Amani, 17 out of 31 females held large
well-developed, very elongated eggs, the largest (on 25.xi.26) measur-
ing 39 x 6 mm. Seven snakes held 2 eggs, eight had 3 eggs, and two
had 4 eggs.

Diet. Centipedes in each of four snakes.

Enemies. One was recovered from the stomach of a Calamelaps u.
unicolor.

Habitat. I found these snakes beneath logs, bark and stones, both
within and without the rain forest but chiefly along its edge. Some
were unearthed by hoeing up grass and weeds on the outskirts of the
forest. In fact its habitat is very similar to that of the North Ameri-
can ring-necked snakes (Diadophis spp.) which it so closely resembles
in color, markings and size. A montane species.

1 In only one of a series of fifty snakes.

2 On one side only in two out of fifty snakes.

3 Not 163 as given in Barbour & Loveridge, 1928c, p. 131, recounted as 161.

loveridge: African snakes 201

Localities. Tanganyika Territory: Amani; Baglio; Bumbuli;
Kizerui; Kukulio; Magrotto; Mkarazi; Mt. Lutindi; Nguelo;
Tanga; Vituri.

These places, with the exception of Tanga, are all in one or other
of the four mountains — Magrotto, Pare, Uluguru, Usambara. Moc-
quard (1897c) is responsible for the Tanga record, Tanga is the port
for Usambara which is about forty miles away, and pending confirma-
tion this record for Tanga should be accepted with reserve.

Range. Mountains of eastern Tanganyika Territory.

Aparallactus ttjrneri Loveridge

?1910a. Aparallactus Jacksoni Sternfeld (part, not Giinther), p. 35.
1935c. Aparallactus turneri Loveridge, Bull. Mus. Comp. Zool., 79, p. 9:

Sokoki Forest, near Malindi, Kenya Colony.
1936J. Loveridge, p. 268.
1937f. Loveridge, p. 493.

Description. Rostral nearly twice as broad as deep, the portion
visible from above one third to two thirds as long as its distance from
the frontal ; nostril in an entire nasal ; internasals much shorter than the
prefrontals; frontal once and a half to once and two thirds as long as
broad (in the middle), longer than its distance from the end of the
snout, as long as the parietals; once and two thirds to twice as broad
as a supraocular ; no loreal ; preocular 1 , in contact with the nasal ; eye
small, its diameter greater than its distance from the mouth; post-
oculars 2, rarely 1, in contact with the anterior temporal; temporals
1 + 1 ; upper labials 6, the second and third entering the orbit, fifth
largest but not in contact with a parietal ; first lower labial in contact
with its fellow behind the mental ; 3 lower labials in contact with the
anterior sublinguals, which are about as long as the posterior. Mid-
body scales in 15 rows, smooth; ventrals 120-139; anal entire: sub-
caudals 31^2.

Based on the type series of four males and two females.

Color. Above, head black with white blotches, one anterior, one
posterior, to the eye, the second blotch larger and extending upwards
on to the anterior temporal ; a black transverse band, sometimes edged
before and behind by scale-wide light bands (which may break up
into spots), on the nape, the anterior, being just posterior to the parie-
tals, tends to separate the black of the head from the black of the nape ;
back and tail a pallid, pinkish brown, the edges of each scale darker,

202

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and with, or without, a fine black vertebral line. Below, white,
uniform except for a slight encroachment of the black nape patch in
two downward -pointing patches.

Size. Total length of cf, 202 (167 + 35) mm. from Peccatoni;
total length of 9 , 196 (165 + 31) mm. from near Witu.

Sexual dimorphism, c? cf have 120-129 ventrals, and 33-42 sub-
caudals. 9 9 have 134-139 ventrals, and 31-37 subcaudals.

Habitat. Beneath logs and stones on the sandy coastal plain.

Localities. Kenya Colony: ? Lamu Island1; Mkonumbi; near
Witu; Peccatoni; Sokoki Forest.

Range. Kenya Colony coast between Lamu and Malindi.

Aparallactus capensis bocagii Boulenger

1895a. Uriechis capensis Bocage (part, not Smith), p. 127: Gambos; Novo
Redondo; and Quindumbo, Angola.

1895h Aparallactus guentheri Boulenger (part), p. 172: Angola only.

1895h. Aparallactus bocagii Boulenger, Ann. Mag. Nat. Hist. (6), 16, p. 173:
Angola.

1896d. Boulenger, p. 259.

1905c. Boulenger, p. 114.

1915a. Boulenger, p. 216.

1925. Werner, 1924, p. 158.

1933f. Angel (part), p. 179 (omit Togo from range).

1937b. Monard, p. 129.

1897a. Uriechis Guentherii Bocage, p. 201.

1897a. Uriechis Bocagi Bocage, p. 201.
?1908c. Aparallactus lunulatus Sternfeld (not Peters), p. 247.

1910c. Aparallactus Lubberti Sternfeld, Mitt. Zool. Mus. Berlin, 5, p. 57:
Between Omaruru and Okanjanda (?Okahandja), South West
Africa.

1910b. Sternfeld, p. 30, fig. 35.

1925. Werner, 1924, p. 157 (as luebberti).

?1933m. Aparallactus punctatolineatus Witte, p. 96.
?1937b. Aparallactus nigriceps Mertens (not Peters), p. 14.

Further citation of 'bocagii' will be found under lidiardae.

Names. Angolan Centipede-eater (English).

Description. Rostral nearly twice as broad as deep, the portion
visible from above one third to two thirds as long as its distance from

1 It seems probable that Sternfeld's (1910a) specimen of "jacksonii" from Lamu should be
referred to lurneri which occurs on the opposite mainland at Mkonumbi.

loveridge: African snakes 203

the frontal; nostril in an entire or divided2 nasal; internasals much
shorter than the prefrontals ; frontal once and a half to once and two
thirds as long as broad (in the middle), much longer than its distance
from the end of the snout, as long as, or slightly shorter than, the
parietals; once and two thirds to twice as broad as a supraocular; no
Ioreal; preocular 1, in contact with, rarely separated from1, the nasal;
eye small, its diameter equal to, or greater than, its distance from the
mouth; postocular 1, not in contact with the anterior temporal;
temporals 1 + 1, rarely 0 + 1 ; upper labials 7, rarely 62, the third and
fourth, rarely second and third2, entering the orbit, fifth, rarely fourth,
largest, and fifth, or fourth and fifth in contact with a parietal ; first
lower labial not in contact with its fellow behind the mental ; 3 lower
labials in contact with the anterior sublinguals, which are as long as,
or slightly longer than, the posterior. Midbody scales in 15 rows,
smooth; ventrals3 168-191 ; anal entire4; subcaudals 355 or 44-59.

The above description, being based on literature only, should be re-
ceived with reserve and is subject to correction.

Color. Above, head black with white blotches, one anterior, one
posterior, to the eye, the second blotch larger and extending upwards
to the anterior temporal; a black transverse band, edged anteriorly
with yellow, on nape; back and tail reddish brown, a small brown spot
in the centre of each scale, the spots forming longitudinal lines. Below,
white.

Size. Total length of type, 272 mm. from Gambos, of a 9 , 330
(285 -f- 45) mm. from between Benguela and Bihe.

Remarks. Bocage (1895a), in addition to a true capensis from
Bibala (if the ventral count of 161 is correct, in which case it is the
only example from the West with so low a count), had two snakes with
entire nasals from Gambos and Novo Redondo. Without seeing them,
Boulenger (1895h) made these the cotypes of a new species — bocagii,
of which he had no material in the British Museum. Bocage, however,
had also listed a specimen from Quindumbo with a divided nasal, on
account of this character, Boulenger added its high ventral count
(180) to that of his new species guentheri and added Angola to the
range of guentheri. I regard guentheri as a synonym of c. capensis, a
species which, though normally possessing an entire nasal, produces

1 Not in contact in Sternfeld's two Chifumbazi specimens of lunulalus.

2 In type of lubberti, and others.

3 Said to be 161 in a snake from Bibala, fide Bocage.

* Bocage has corrected his (1895a) misstatement about divided anals and paired subcaudals.
6 In the Benguela to Bihe specimen which I suggest has a mutilated tail.

204 bulletin: museum of comparative zoology

individuals here and there in the east in which it is divided. In this
connection it is interesting to note that such cases appear rare indeed
in the southeast for FitzSimons writes me that the nasal is entire in all
32 examples of c. capensis and 7 (of what I should call) bocagii in the
collection of the Transvaal Museum.

Since its description in 1895, only one specimen of bocagii has been
recorded (Boulenger, 1905c) as such, for I reject Sternf eld's (1908b,
p. 219) record of its occurrence in Togo, a statement apparently based
on a misidentified specimen of the recently-described liddiardae.

Later, however, Sternfeld (1910c) described lubberti, differing only
from bocagii in having a divided nasal and six upper labials, of which
the second and third (left), or third (right) only enter the orbit, conse-
quently the fourth (instead of the fifth) upper labial is largest and in
contact with the parietal. This reduction in labials is a variation com-
mon to four species of the genus at least.

I am less confident in suggesting that Sternfeld 's (1908c) lunulatus
material with 168-172 ventrals is referable to bocagii, much turns on
the stability of the character involving the forming of a suture by the
first pair of lower labials. The snake requires reexamination.

Witte (1933m) has recorded a snake from the southern Belgian
Congo under the name of punctatolineatus, but gives no scale counts.
On geographical grounds I tentatively refer it to bocagii for, if its
second and third labials enter the orbit, as one may justifiably assume,
then it would naturally run down to punctatolineatus in Boulenger's
key.

The snake that Mertens (1937b) refers to nigriceps has a similar
labial condition but its high ventral count (178) precludes its reference
to nigriceps (108-123) or capensis (131-166) though the record involves
extension of the range of bocagii south to the Transvaal. On appealing
to Mr. V. FitzSimons for light on this point he, with customary kind-
ness, furnished me with scale counts of all "capensis" in the Transvaal
Museum. These show that what I regard as bocagii, i.e. snakes with
169-180 ventrals, occurs sparsely throughout the Transvaal, both
bocagii and c. capensis occurring together at Lydenburg, Pretoria, and
Johannesburg. Were we to assume that snakes from these three places
were all of one form we would be confronted with the absurdly large
ventral range of 138-180 for so small a snake. Personally I feel that
bocagii is little more than a race of c. capensis, such treatment, how-
ever, would involve regarding almost the entire Transvaal as an area
of intergrades, at the present stage of our knowledge, therefore, it
seems advisable to treat them as subspecies.

lovekidge: African snakes 205

Localities. ?Belgian Congo: Elisabeth ville. Angola: Benguela
to Bihe; Bibala1; Bingondo; Gambos; Novo Redondo; Quindumbo.
South West Africa: Okanjande to Omaruru. Transvaal:
Gravelotte; Johannesburg2; Lydenburg2; Malelane Camp, Croco-
dile River, Kruger National Park; Pretoria2; Vaalwater. Mozam-
bique: Chifumbazi.

Range. Angola and South West Africa, east through southeastern
Belgian Congo to extreme western Mozambique, south through the
Transvaal where it occurs alongside c. capensis in several localities.

Aparallactus capensis capensis Smith

1849. Aparallactus capensis A. Smith, 111. Zool. S. Africa, 3, App., p. 16:

Kaffirland to the eastward of Cape Colony.

1895h. Boulenger, p. 173.

1896d. Boulenger, p. 259.

1896. Tornier, p. 79.

1898. Sclater, p. 100.

1898. Werner, 1896-7, p. 146.

1902b. Boulenger, p. 18.

1907J. Boulenger, p. 487.

1907a. Roux, p. 81.

1908b. Boulenger, p. 230.

1908. Gough, p. 33.

1908. Odhner, p. 5 (? part, nigriceps).

1909a. Chubb, p. 596.

1909b. Chubb, p. 36.

1910b. Boulenger, p. 516.

1910a. Stemfeld, p. 36.

1912. FitzSimons, F. W., p. 128.

1913. Hewitt & Power, p. 164.
1915a. Boulenger, p. 216.
1915c. Boulenger, p. 634.
1923e. Loveridge, p. 889.
1924b. Loveridge, p. 7.

1925. Werner, 1924, p. 158.

1934. Pitman, p. 298.

1935b. FitzSimons, V., p. 323.

1 936 j . Loveridge, p. 268.

1937a. FitzSimons, V., p. 263.

1937f. Loveridge, p. 496.

1 But said to have 161 ventrals.

2 Where it occurs alongside c. capensis.

206 bulletin: museum of comparative zoology

1937a. Parker, p. 630.

1937e. Hewitt, p. 64, pi. xviii, fig. 1.

1937. Uthmoller, p. 123.

1939b. FitzSimons, V., p. 24.

1940. Bogert, p. 43.

1941. Moreau & Pakenham, p. 109.

1849. Elapomorphus capensis A. Smith, 111. Zool. S. Africa, 3, App., p. 16:

Kaffirland to the eastward of Cape Colony.

1863a. Cercocalamus collaris Gunther, Ann. Mag. Nat. Hist. (3), 11, p. 21,

pi. hi, fig. A: "Central America" (in error).

1895. Gunther, Biol. Centrali Americana, Rept., p. 157.

1865a. Uriechis capensis Gunther, p. 89.

1866. Jan, livr. 15, pi. i, fig. 5.

1882a. Bocage, p. 288.

1882a. Peters, p. 112.

1884a. Rochebrune, p. 154 (ignored).

1887h. Boulenger, p. 175.

1888b. Gunther, p. 324.

1889. Boettger, p. 293.

1891a. Boulenger, p. 308.

1891a. Matschie, p. 609.

1892. Muller, p. 207.

1893. Gunther, 1892, p. 555.
1895a. Bocage, p. 127.
1896a. Bocage, p. 94.

1898. Johnston, p. 361a.

1895h. Aparallactus guentheri Boulenger (part), Ann. Mag. Nat. Hist. (6),

16, p. 172: East and Central Africa (exclude Angola).

1896d. Boulenger (part), p. 259, pi. xi, fig. 2.

1902b. Boulenger, p. 18.

1910b. Boulenger, p. 516.

1910a. Sternfeld, p. 36.

1912. FitzSimons, F. W. (part), p. 128.

1915a. Boulenger (part), p. 216.

1915c. Boulenger (part), p. 634.

1924b. Loveridge, p. 7.

1925. Werner (part), 1924, p. 158.

1934. Pitman, p. 298.

1937f. Loveridge, p. 496.

1937b. Monard (part), p. 129.

1941. Moreau & Pakenham, p. 109.

1895h. Aparallactus punctatolineatus Boulenger, Ann. Mag. Nat. Hist. (6),

16, p. 173: Angola.

1896d. Boulenger, p. 261.

1897a. Bocage, p. 201.

loveridge: African snakes 207

1915a. Boulenger, p. 217.

1915c. Boulenger, p. 634.

1925. Werner, 1924, p. 158.

1934. Pitman, p. 298.

1895h. Aparallactus nigriceps Boulenger (part, not Peters), p. 173.

1896d. Boulenger (part), p. 260.

1896. Tornier, p. 79.

1897. Tornier, p. 65.
1910a. Sternfeld, p. 36.

1915c. Boulenger (part), p. 634.

1924b. Loveridge, p. 7.

1924. Werner (part), 1924, p. 158.

1897a. Uriechis punctatolineaUis Bocage, p. 201.

Further citations of 'capensis', 'guentheri', and ' punctatolineatus' will
be found under c. bocagii.

Names. Cape Centipede-eater or Black-headed Snake (English);
yamitera (Makonde: Tanganyika).

Description. Rostral nearly twice as broad as deep, the portion vis-
ible from above one third to two thirds as long as its distance from the
frontal ; nostril in an entire, semidivided, or divided nasal ; internasals
shorter than the prefrontals ; frontal once and a half to once and two
thirds as long as broad (in the middle), much longer than its distance
from the end of the snout, as long as, or slightly shorter than, the
parietals, once and two thirds to twice as broad as a supraocular; no
loreal; preocular 1, in contact with the nasal; eye small, its diameter
equal to, or greater than, its distance from the mouth; postocular 1,
not, or but very rarely1, in contact with the anterior temporal; tempo-
rals 1 + 1 ; upper labials 7, sometimes 6, the third and fourth, or some-
times the second and third, entering the orbit, fifth largest and in
contact with, very rarely separated from2, a parietal ; first lower labial
not in contact (unless lunulatus is a synonym) with its fellow behind the
mental; 3 lower labials in contact with the anterior sublinguals, which
are as long as, or slightly longer than, the posterior. Midbody scales in
15 rows, smooth; ventrals 131-167; anal entire; subcaudals 34-593.

Based on original descriptions, literature, data of those in Transvaal
Museum, and many Kenya, Tanganyika, Mozambique and Transvaal
specimens in the M.C.Z.

1 On right side only of a Lumbo snake (M.C.Z. 16364).

2 On right side only of a Lumbo snake (M.C.Z. 16364).

3 Or 63 if one includes a snake from Ingwavuma, Zululand (T.M. 15315) with 154 ventrals,
these counts checked by FitzSimons who has another Zululand specimen (from Ntambanana)
with 133 ventrals and 50 subcaudals.

208 bulletin: museum of comparative zoology

Color. Two principal color phases exist, one, which is intermediate
between the uniformly plumbeous, sylvicoline c. ulugurensis and the
pale brown, savanna and coastlands c. capensis, occurs in deforested
or adjacent areas and is likely to present so many stages that it is
unworthy of recognition by name. The following description is based
on a pair, which also happen to be the most northerly examples of c.
capensis known.

<? 9 . Mt. Mbololo. Above, head black, a five-scale wide transverse
black band edged before and behind by ill-defined scale-wide light
bands (which broaden on the sides) on nape; back and tail uniformly
iridescent plumbeous or steely blue. Below, throat white, body also
but so thoroughly infuscated with gray as to appear dusky.

cf. Mlalo, western Usambara Mtns. Above, as last, but the light
bands on nape are broader, sharply distinct, and white (in alcohol) ;
also there is a light spot present at the base of each dorsal scale. Below,
similar to the Mbololo snakes.

cf. Lydenburg, Transvaal. Above, head black, a four-scale wide
transverse black band edged before by a scale-wide buff band on nape;
back and tail pinkish brown or pale brown with a fine brown vertebral
line. Below, white. The width of the nuchal band varies from 3 to 6
rows in South African specimens in which the vertebral line may be
black, if present, or absent altogether.

Size. Total length of ?d\ 290 (239 + 51) mm. from Lumbo; total
length of ? 9 , 410 (335 + 75) mm. from Cape Province (Werner, 1898,
sex not stated. This is 100 mm. above any other record and may be a
misprint). #

Remarks. It would appear that after sending the description of
Elapomorphus capensis to press, Sir A. Smith, realizing the name for
his new genus was preoccupied, forwarded the manuscript of Aparal-
lactus capensis to his publishers for substitution, for it is inserted under
the caption of Sauria ! Fortunately it has paragraph priority over the
other description.

FitzSimons (1937a), after reexamination, doubts that either of the
specimens in the British Museum represent the type, though consid-
ered so by Boulenger (1896d).

A. guentheri of Boulenger was a composite of several specimens of
capensis with divided nasals, which he had, and an example of bocagii
with a divided nasal mentioned by Bocage, which Boulenger had not
seen.

A. punctatolineatus Boulenger was based on another capensis men-
tioned by Bocage which had only 6 upper labials, the second and third

loveridge: African snakes 209

entering the orbit, a condition occurring spasmodically throughout
the range of capensis. Later Boulenger (1896d) referred a Nyasaland
snake to this species.

A. nigriceps of Boulenger was a composite of a specimen of capensis
with 6 upper labials, etc., and the data derived from Peters' original
description of nigriceps, which is distinguished from all other Aparal-
lactus by its low ventral count (108-123).

The name capensis has not been applied to any other species except
bocagii, which is distinguished by its much higher ventral count (168—
191).

Dentition. Bogert (1940) after examination of three snakes from
Nyasaland and Transvaal, found that the maxillary teeth ranged from
5 to 7 followed, after a diastema, by 2 enlarged grooved fangs.

Hemipenis. This is discussed at length by Bogert (1940).

Sexual dimorphism. There appears to be a slight overlap in the
ventral count of this species for a Lydenburg c? in the Museum of
Comparative Zoology has 148, while Boulenger (1896d) has recorded
a 9 from Zanzibar with that number.

Breeding. On April 27, at Mbanja, a 9 held 2 eggs measuring 31 x
4 mm.

Diet. At Lumbo, one morning, a Leptotyphlops longicauda was cap-
tured together with several Lycopkidion semiannulis and Aparallactus
c. capensis. They were all put into a cigarette tin until evening when I
should have time to attend to them. On opening the tin about 6 p.m.
I was disagreeably surprised to find the valuable worm snake missing.
As escape was out of the question I held up the other snakes, one by
one, against a strong acetylene lamp until I found the worm snake
doubled up in the stomach of a Cape black-headed snake.

So run my typescript notes of 25..viii.l918, but in recent years I
have often wondered whether I did not make a mistake for a worm
snake would be natural prey for a wolf snake, whereas, except for a
snail, there is no other record of an Aparallactus eating anything else
but a centipede. On the other hand see note below.

Temperament. At Lumbo, where I took sixteen of these little snakes,
I observed that they bit quite fiercely at times but their tiny teeth
failed to break through the skin. A nine-inch female attacked a five
and a half inch snake, the latter seized its aggressor so that together
they formed a struggling circle. If I had left them alone it seems
possible that the larger would have dined off the lesser.

Habitat. This Lumbo series were mostly dug from about the roots
of shrubs and grass though some were found on the surface of the

210

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sandy soil in the early morning. Hewitt states that they are to be
found in termitaria or beneath stones in open country, while V. Fitz-
Simons has taken them beneath stones on granite hills. My Mbololo
specimens were taken in a somewhat similar situation on the mountain-
side just below the forest edge.

_ Localities. Kenya Colony: Mount Mbololo. Tanganyika Ter-
ritory: Marangu, Kilimanjaro; Mbanja; Mlalo, w. Usambara;
Mohorro; Sanya, Kilimanjaro; Tanga; Ujiji. Zanzibar. Mafia
Id. Mozambique: Angoche;Lumbo;Rikatla;Tete. Nyasaland:
Chiradzulu; Lake Nyasa; Mlanje; Shire Highlands; Zomba.
Southern Rhodesia: Bulawayo; Gwelo; Salisbury district; Vic-
toria Falls; Vumba Mountain. Bechuanaland : Serowe. Zulu-
land: Entendweni; Ingwavuma (T.M.); Kosi Bay; Ntambanana
(T.M.); Umfolosi Rivers junction. Transvaal: Blaauwberg
(T.M.); De Kaap Goldfield; Delmas Road near Pretoria; Irene;
Johannesburg (T.M.); Klein Letaba (T.M.); Kraalkop; Krabbe-
fontein; Legogot; Lydenburg district; Modderfontein; Mphome;
Nelspruit (T.M.); Punda Maria (T.M.); Rustenberg (T.M.);
Selati; Shilowane; Woodbsuh (T.M.); Zeekoegat. Orange Free
State. Cape Province: Burghersdorp; East London; Pondoland;
Zingqolo. Angola: Quindumbo.

Range. Southeast Kenya Colony south to Cape Province.

Aparallactus capensis uluguruensis Barbour & Loveridge

1928c. Aparallactus uluguruensis Barbour & Loveridge, Mem. Mus. Comp.

Zool., 50, p. 132: Nyange, Uluguru Mountains, Tanganyika

Territory.
1936j. Loveridge, p. 270.
1937f. Loveridge, pp. 493, 502.
1942e Aparallactus capensis uluguruensis Loveridge, p. 301.

Name. Uluguru Centipede-eater (English); penge (Pokomo).

Description. Rostral nearly twice, or twice, as broad as deep, the
portion visible from above one third to two thirds as long as its dis-
tance from the frontal; nostril in an entire, semidivided, or divided
nasal ; internasals shorter than, or much shorter than, the prefrontals ;
frontal once and a half to once and two thirds as long as broad (in the
middle), much longer than its distance from the end of the snout,
shorter than the parietals, once and two thirds to twice as broad as a
supraocular; no loreal; preocular 1, in contact with the nasal; eye

loveridge: African snakes 211

small, its diameter equal to, or greater than, its distance from the
mouth; postocular 1, very rarely 21, not in contact with the anterior
temporal; temporals 1 + 1, rarely 1 + 22; upper labials 7, rarely 63,
the third and fourth, rarely second and third3, entering the orbit, fifth
largest and in contact with a parietal; first lower labial not in contact
with its fellow behind the mental; 3 lower labials in contact with the
anterior sublinguals, which are as long as, or slightly longer than, the
posterior. Midbody scales in 15 rows, smooth; ventrals 137-159; anal
entire; subcaudals 43-534.

Based on the fourteen known specimens.

Color. Above, uniformly plumbeous or iridescent black. Below, as
above or somewhat paler, more particularly on throat and tail which
may be almost white. Young of 182 mm. are colored precisely like the
adults.

Size. Total length of paratype & (M.C.Z. 23366), 400 (320 4- 80)
mm. from Amani; total length of paratype 9 (M.C.Z. 23364), 387
(333 + 54) mm. from Nyange.

Sexual dimorphism. cTcf have 137-144 ventrals, and 43-52 sub-
caudals. 9 9 have 158-159 ventrals, and 44-51 subcaudals.

Breeding. At Ngatana, on June 17, a 9 held 2 eggs, each measur-
ing 12x5 mm. Four adult 9 9 taken in the Uluguru and Usambara
Mountains in October, November and December, were not breeding.

Diet. Six centipedes (Alipes grandidieri and another species) were
recovered from the type series. A snail as well as two centipedes were
present in the stomach of a Nyange snake. A seventh centipede in a
Magrotto specimen.

Habitat. Beneath logs in montane rain forest and gallery forest.

Localities. Kenya Colony : Ngatana, Tana River. Tanganyika
Territory: Amani, Usambara Mountains; Magrotto Estate,
Magrotto Mountain; Nyange, Uluguru Mountains.

Range. Virgin forest in coastal belt of Kenya Colony and Tangan-
yika Territory.

i On left side only of M.C.Z. 23369.

2 On left side only of an Amani paratype, right side of a Nyange paratype.

3 On right side only of an Amani paratype.
* 58 was a misprint.

212 bulletin: museum of comparative zoology

Aparallactus nigriceps (Peters)

•

1854. Uriechis nigriceps Peters, Monatsb. Akad. Wiss. Berlin, p. 623:

Tete, Mozambique.

1855. Peters, p. 52.
1859?. Bianconi, p. 385.

1882a. Peters, p. Ill, pi. xviii, figs. 1-le.

1884a. Rochebrune, p. 154 (ignored).

1896a. Bocage, p. 100.

1857. Eucritus atrocephalus Jan, 1857, Cenni Museo Civico Milano, p. 44:

Inhambane, Mozambique (non vidi).

1862. Uriechis atriceps Jan, p. 49 (lapsus for nigriceps).
1866. Jan, livr. 15, pi. i, fig. 4.

1895h. Aparallactus nigriceps Boulenger (part), p. 173.
1896d. Boulenger (part), p. 260 (no material).

Further citation of 'nigricepis' will be found under c. bocagii.

Description. Rostral nearly twice as broad as deep, the portion vis-
ible from above one third to two thirds as long as its distance from the
frontal; nostril in an entire nasal; internasals shorter than the pre-
frontals; frontal once and a half as long as broad (in the middle), much
longer than its distance from the end of the snout, as long as, or
slightly shorter than, the parietals; twice as broad as a supraocular;
no loreal; preocular 1, in contact with, rarely separated from, the
nasal; eye small, its diameter equal to, or greater than, its distance
from the mouth; postocular 1, not in contact with the anterior tem-
poral ; temporals 1 + 1 ; upper labials 6, the second and third entering
the orbit, fourth largest, fourth, or fourth and fifth, in contact with a
parietal; first lower labial not in contact with its fellow behind the
mental; 3 lower labials in contact with the anterior sublinguals, which
are as long as, or slightly longer than, the posterior. Midbody scales in
15 rows, smooth; ventrals 108-123; anal entire; subcaudals 20-35.

Based on original descriptions and figures.

Color. Above, head black with white blotches, one anterior, one
posterior, to the eye, black of crown continues on to nape where it is
edged posteriorly with a yellow band ; back and tail uniformly reddish
olive brown except for the trace of a fine black vertebral line. Below,
pale yellow.

Size. Total length of type, 225 (18.0 + 45) mm.

Sexual dimorphism. ?

Localities. Mozambique: Inhambane; Tete.

loveridge: African snakes 213

Range. Mozambique (known only from the type, and Bianconi's
specimens in Milan Museum).

Genus Elapsoidea

1866a. Elapsoidea Bocage, Jorn. Sci. Lisboa, 1, pp. 50, 70 (type guntherii

Bocage).
1896d. Elapechis Boulenger, Cat. Snakes Brit. Mus., 3, p. 358, footnote

(substitute name).

Maxillary bone extending forwards as far as the palatine, with a
pair of large grooved poison-fangs followed by 2-4 small teeth ; anterior
mandibular teeth longest. Head moderate, not or but slightly distinct
from neck; eye small, with round pupil; nasal divided; loreal absent1;
a single preocular. Body cylindrical; scales oblique, smooth, without
pits, in 132 rows; yentrals rounded. Tail very short; subcaudals all or
most in two rows.

Range. Africa south of 15° N., i.e. Senegal; Kordofan; Ethiopia.

Remarks. The substitute name Elapechis was proposed by Boulenger
on the grounds that the correct form of Elapsoidea should be Elapoidea,
which he considered preoccupied by Elapoides Boie, 1827, of Java.

In 1936 I invited attention to the fact that guntherii and nigra from
the equatorial region, exhibit a higher ventral count than was the case
with Angolan snakes being referred to guntherii. As no one investigated
the subject I have attempted to straighten out the somewhat involved
situation myself, though lacking material of the form laticincta occur-
ring to the north of the thousand-mile-wide equatorial belt. All the
remaining five forms are represented in the collections of the Museum
of Comparative Zoology, which has 73 snakes of this genus, from which
I exclude midtifasciata and its synonym duttoni.

Thus we find ourselves left with a monotypic genus, and it has been
no easy task to find characters wherewith to separate the various
forms and, indeed, I have not attempted to do so in the case of lati-
cincta (as opposed to decosteri) on account of the lack of material.

To judge by our single example of typical sundevallii, that race has
an obtusely pointed snout. In this connection, however, it is impor-
tant to note that Boulenger (1896d) had no material of typical sunde-
vallii when he published his key to the 'species' of the genus. Later

1 A small loreal-like scale appears in 1 of the 77 snakes examined.

2 Boettper's statement that boulengeri had 15 was incorrect; multifasciata with 15-17 is not
an Elapsoidea but a cobra.

214 bulletin: museum of comparative zoology

(1897d) he obtained "several" and in his revised (1910b) key he
dropped this character of snout shape which formerly had been
treated as if of major importance. Was this because it broke down in
series of even sundemlliil The snout is most definitely pointed in an
adult 9 decosteri (from Lumbo), less noticeably so in a young 9 of the
same form from Pretoria, or an adult 9 guntherii from northern
Kenya (Guaso Nyiro). The thought occurred that it might be a
sexual difference, but it was found to be rounded in adult and juvenile
9 9 of guntherii from western Kenya (Kaimosi and Loita Plains) just
as much as in adult and juvenile cf cf of semiannulata from Angola or
juvenile decosteri from Northern Rhodesia. I am inclined to the con-
clusion that it is an adaptation to environment, being rounded in speci-
mens of guntherii and nigra living in the equatorial rain-forests where
they have to burrow only in soft leaf mould, more often pointed in
those snakes {decosteri, fitzsimonsi and guntherii) which live under
savanna conditions. In practice one finds so many intermediate condi-
tions between "obtusely pointed" and "rounded" that I could not
utilize it as a key character.

The portion of the rostral visible from above is certainly greater in
typical sundevallii and fitzsimonsi than in other forms with the excep-
tion of decosteri which would appear to link it with the more northern
races.

The length of the internasals in relation to that of the prefrontals
is a useless character, this also applies to frontal length in relation to its
width, or to the length of the parietals, or to its distance from the
end of the snout. I imagine that the relative lengths of the anterior and
posterior sublinguals will probably be found to vary to much the same
extent in all the races as soon as adequate series, such as have been
available to me in the case of guntherii and nigra, have been studied.

Nevertheless, despite the fact that I regard almost all the matter
furnished under the head of Description as practically valueless, I
have included it as showing the carefully ascertained range of variation
within the limits of each race as here defined. Such data having been
obtained from the literature listed in the synonymy, together with
the results of an examination of all the material at my disposal.

Under the heading Localities, the letter T after a locality signifies
that scale counts of one or more specimens in the Transvaal Museum
have been supplied me; the letters M.C.Z. in parentheses are placed
only after such localities as have not yet appeared in the literature,
and are to indicate that the specimen on which the record is based is
in the collection of the Museum of Comparative Zoology.

loveridge: African snakes 215

Key to the Races1

1. Ventrals less than 169 2

Ventrals 162-184 7

2. Ventrals 138-169; subcaudals 13-27 3

Ventrals 151-166; subcaudals 13-30 6

3. Young with light crossbars about half the width of, or at least much nar-

rower than, the interspaces between them; adults uniformly black 4

Young with light crossbars subequal in width to the interspaces between
them; adults black or brown with 14-22 pairs of well-defined white
crossbars (formed by division of the solid bars of the young) 5

4. Range: Southern Ethiopia and Kordofan west through extreme northern

Belgian Congo (Uelle) to northern Cameroons s. laticincta

(p. 216)
Range: Mozambique, Nyasaland, and Northern Rhodesia south through

Transvaal to Zululand and Orange Free State s. decosteri

(p. 217)

5. Range: Southern Belgian Congo and Angola s. semiannulata

(pi 220)

6. Subcaudals in males 23-30, in females 18-21; range: Kenya and northern

Tanganyika Territory (Oldeani; Kilimanjaro) west through Uganda and
Belgian Congo to Senegal s. guntherii

(p. 222)

Subcaudals in males 18-24, in females 13-17; range: eastern Tanganyika

Territory (Magrotto, Usambara, and Uluguru Mountains) s. nigra

(p. 225)

7. Adults and young slaty gray tinged with purplish brown with numerous

white- or yellow-edged, black crossbars; range: Natal s. sundevallii

(p. 228)

Adults uniformly purplish brown above, young barred as in the typical

form; range: eastern Cape Province (Kimberly) west through Bechuana-

land to South West Africa s. fitzsimonsi

(p. 229)

1 Obviously unsatisfactory in some respects due to the lack of material representing laticincta
and paucity for the two following forms; under the circumstances it has seemed best to give
only the range rather than cite differences which may not prove to be constant.

216

bulletin: museum of comparative zoology

Statistical Synopsis of Variation in the Races of Elapsoidea

Maximum

a 5>

length

Race

"ffl

s 0

c3.s

r Labials i
act with
rior sublin

cd
3

"a
u
o

rior nasal
ntact (c)
3ular

Head
and

Tail

cu *^ aj

1)55

3
CD

0. C

£•£■

a:
O

S
H

Post
in c
pre

Body

s. lalicincla

143-150

13-24

7 (3-4)*

3-3

1+2

430

s. decosleri

138-169

13-26

7 (3-4)

3-4

1+2*

650

s. semiannulata

139-153

15-27

7 (3-4)

3-4

1+2

438

s. gunlherii

151-166

18-30

7 (3-4)

3-4

1+2

581

s. nigra

151-162

13-24

7 (3-4)*

3-4

1+2*

548

s. sundevallii

163-184

19-27

7 (3-4)

3-4

1+2*

825

s. fitzsimonsi

162-181

17-23

7 (3-4)

3-4

1+2

766

* See text for rare variation involving one side of one snake.

Elapsoidea sundevalli laticincta (Werner)

1917. Elapechis guentheri Sternfeld (not Bocage), p. 481.

1939c. Scortecci, p. 183, figs. 101-102.

1940a. Scortecci, p. 136, figs. 1-2.

1919. Elapechis laticinctus Werner, Denks. Akad. Wiss. Wien, 96, p. 507,

fig. 8: Kadugli, Kordofan, Anglo-Egyptian Sudan.
1923a. Werner, p. 179.

Description. Snout obtusely pointed ; portion of rostral visible from
above measuring half its distance from the frontal ; internasals half the
length of the prefrontals ; frontal once and a third to once and a half
as long as broad, as long as its distance from the end of the snout, two-
thirds the length of the parietals ; upper labials rarely 6, the second and
third entering orbit (on right side of Filtu snake) ; anterior sublinguals
slightly longer than the posterior.

For character common to all forms see p. 213; for scale counts see
statistical table above.

Color. Young. Above, a goblet-shaped (Uelle specimen) prolonga-
tion of, or a black line from, the black nuchal crossbar extends over
parietal suture, otherwise head grayish ; body barred alternately black
(about 8-10 scales wide) and gray (about 5 scales wide), the dark bars
being about twice as broad as the light interspaces, which apparently
number 14 on body, 2 on tail. Below, whitish.

loveridge: African snakes 217

Adult from Poli. Above, head and body uniformly black. Below,
whitish.

Size. Total length of a supposed (^(with 24 subcaudals), 228 (203
+ 25) mm. from Filtu (Scortecci), or a supposed 9 (with 17 sub-
caudals), 458 (430 -f- 28) mm. from Poli (Loveridge), of juvenile type,
supposedly a 9 (with 13 subcaudals), 237 (220 +17) mm.

Remarks. The above description, as well as data furnished in statis-
tical table and key, is based on the information contained in the above
citations together with the data derived from a large example from
Poli, Garua, northern British Cameroons, submitted to me many years
ago by the Vienna Museum. This is the only member of the northern
form which I have seen and whether I am correct in lumping all to-
gether under the name laticincta remains to be seen. With inadequate
material I have been unable to separate it from the southeastern race
decosteri, to which Werner himself stated that it was related, allegedly
differing from gimtherii by its obtusely pointed snout. Remarks on the
latter character will be found under the genus.

Diet. A skink (Mabuya perroteti mongallensis) in stomach of type.

Localities. Ethiopia: Filtu. Anglo-Egyptian Sudan: Ka-
dugli. Belgian Congo: Angu, Uelle. British Cameroons: Poli
near Garua. (Possibly some other northern Congo references to
gimtherii for which no scale-counts are given, and almost certainly the
northern Nigerian record of Boulenger, repeated by Angel, are refer-
able to laticincta).

Range. Southern Ethiopia and Kordofan west through northern
Belgian Congo (Uelle) to northern Cameroons (and possibly northern
Nigeria).

Elapsoidea sundevallii decosteri Boulenger

1888d. Elapsoidea Decosteri Boulenger, Ann. Mag. Nat. Hist. (6), 2, p. 141 :
Delagoa Bay, Mozambique.

1895. Elapsoidea Boulengeri Boettger, Zoo. Anz., 18, p. 62: Boroma, Zam-
besi, Mozambique.

1922a. Mertens, p. 182.

1937b. Mertens, p. 14.

1895. Elapsoidea Guentheri Giinther (part, not of Bocage), p. 525 (Nyasa-

land only).
1934. Pitman, p. 298.

1937. Pitman (part), p. 346 (N. Rhodesia references).

1896. Elapechis sundevali Peracca (not Smith), p. 4.

1896d. Elapechis guentheri Boulenger (part), p. 359 (Nyasaland only).

218 bulletin: museum of comparative zoologt

1896. Peracca, p. 4.

1908. Gough, p. 33.

1909a. Chubb, p. 596.

1910b. Boulenger (part), p. 519 (Transvaal).

1912. FitzSimons, F. W. (part), pp. 166, 167.

1913. Hewitt & Power, p. 165.
1921a. Angel, p. 44.

1896d Elapechis decosteri Boulenger, p. 360.

1898. Sclater, p. 101.

1908b. Boulenger, p. 230.

1910b. Boulenger, p. 519.

1912. FitzSimons, F. W., pp. 166, 168.

1923a. Werner, p. 179.

1896d. Elapechis boulengeri Boulenger, p. 361.

1898. Boettger, p. 119.

1915c. Boulenger, p. 635.

1923a. Werner, p. 179.

1907a. Elapechis niger Boulenger (not Giinther), p. 12.

1908c. Sternfeld, p. 247.

1912. Peracca, p. 6.

1923e. Loveridge (part), p. 890 (Lumbo only).

1934. Elapsoidea (Elapechis) niger Pitman, p. 298.

1908. Elapechis sundevallii Gough (part, not Smith), p. 34 (Orange River
Colony).

Sternfeld's (1910b, 1910c) reference to decosteri is tentatively re-
ferred to s. fitzsimonsi subsp. nov.

Native name. Kouseband slang (Afrikaans: F. W. FitzSimons).

Description. Snout rounded or obtusely pointed; portion of rostral
visible from above measuring half to three-quarters its distance from
the frontal; half to three-quarters the length of the prefrontals ; frontal
once and a third to once and a half as long as broad, as long as, or
longer than, its distance from the end of the snout, much shorter than,
or two-thirds the length of the parietals; posterior nasal rarely sepa-
rated from the single preocular, temporals only rarely 2 + 2 ; anterior
sublinguals slightly shorter than, subequal to, or slightly longer than
the posterior.

For characters common to all forms see p. 213; for scale counts see
statistical table on p. 216.

Color. Young. Above, a goblet-shaped prolongation of the dark
nuchal crossbar extends over parietal suture on to frontal, sides of
head with dusky markings, otherwise head white or grayish; body
barred alternately with chocolate brown (or black) and pale buff (or

loveridge: African snakes 219

white), the dark bars being much broader than the light interspaces
which number 10-25 (normally 15-20) on body, 2-3 (rarely 4) on tail.

Adult. Above, head and body glossy black (fading to purplish
brown in alcohol), or dark gray with each scale edged with black,
uniform, or the outer row of scales whitish and sometimes with a trace
of narrow white crossbars. Below, whitish or grayish.

Size. Total length of an unsexed specimen (Transvaal Mus. 16800),
715 (650 + 65) mm., from Manabajof a gravid 9 (M.C.Z. 18233), 500
(468 + 32) mm. from Lumbo; type of decosteri (S. Air. Mus.), 450
(400 + 50) mm.; of juvenile type of boulengeri (Senck. Mus.), 170
(156 + 14) mm.

Remarks. E. decosteri was based on an unsexed snake which, to judge
by its 25 subcaudals, is probably a male, for the same reason the smaller
snake with 26 subcaudals allegedly a 9 (Boulenger, 1896d) is likely to
be a male also. E. boulengeri, based on a juvenile with consequently
different color pattern to the old decosteri, was said to differ by the
possession of 15 midbody scale-rows, however Mertens (1937b), on
reexamination of the type found that there are but 13. I therefore
refer it to the synonymy of decosteri.

Peracca's (1896) records of both sundevalli and gunthcriihom Kazun-
gula, S. R., here referred to decosteri may possibly be fitzsimonsi.

Breeding. On January 19, at Broken Hill, a 9 held 10 large and
elongate eggs (Pitman). On July 11, at Lumbo, a 9 held 4 eggs, the
largest measuring 17 x 7 mm.

Habitat. Two were dug from a termite hill in dry savanna bush at
Lumbo, altitude circa 100 feet (A. L.).

Localities. Mozambique: Boroma; Zambesi; Chimbo (Tschim-
bo); Delagoa Bay; Lourenco MarquesT; Lumbo. Nyasaland:
Shire Highlands. Northern Rhodesia: Broken Hill to Bwana
Mkubwa Mine; Lealui; Mbala, east Loangwa District. Southern
Rhodesia: Bulawayo; DekaT, 50 miles south of Victoria Falls;
Eldorado; GatoomaT; Kazungula; Wankie. Transvaal: ErmeloT;
Lydenburg District1, MessinaT; MiddleburgT; Piet RetiefT; Pil-
grims' RestT; Pretoria (M.C.Z.) ; SabieT; Swaziland1"; WaterbergT;
WaterpoortT; White RiverT. Zululand: Kosi Bay; ManabaT;
MaputaT. Orange Free State. (Transvaal Mus. specimen listed
by Gough as sundemllii, but has only 152 ventrals).

Range. Zululand and Transvaal north to Mozambique and the
Rhodesias. It is possible that this race extends north of the Rovuma

T Transvaal Museum specimens, not seen, but the data for which has been kindly supplied
by Mr. F. W. FitzSimons.

220 bulletin: museum of comparative zoology

River into southeastern Tanganyika Territory for Mr. R. de la B.
Barker writes me (l.vi.41) that at Lindi he had found a snake, seven
inches in length, with 14 white crossbars. On the other hand it may be
a young nigra of brighter coloration at sea level than is usual in its
forested montane habitat.

Elapsoidea sundevallii semiannulata Bocage

1873b. Elapsoidea Giintherii Bocage (not Bocage of 1866), p. 224.

1888a. Boettger (part), p. 82 (omit range).

1895a. Bocage (part), p. 129, pi. xiv, figs. 3a-c.

1896a. Bocage (part), p. 79 (Angola record only).

1897a. Bocage (part), p. 202 (Angola records only).

1933. Schmidt, p. 14.

1936h. Loveridge (part), p. 41 (Caconda only).

1937b. Mertens, p. 14.

1938e. Mertens, p. 442.

1940. Bogert, p. 86.

1 882b. Elapsoidea semi-annulata Bocage, Jorn. Sci. Lisboa, 8, p. 303 : Caconda,

Angola.

1887c. Elapsoidea Hessei Boettger, Zool. Anz , 10, p. 650: Povo Netonna,

Banana, Belgian Congo.

1888a. Boettger, p. 83, pi. ii, figs. 6a-e.

1922a. Mertens, p. 182.

1896d. Elapechis hessii Boulenger, p. 360.

1915a. Boulenger, p. 218.

1898. Boettger, p. 118.

1920a. Witte, p. 62.

1920b. Witte, p. 275.

1923a. Werner, p. 179.

1898. Elapechis guentheri Boettger (not Bocage), p. 118.

1937b. Monard, pp. 136, 137 (but omit range).

1900. Elapsoidea guntheri var semiannulata Ferreira, p. 52.

Description. Snout rounded or obtusely pointed; portion of rostral
visible from above measuring a third or half its distance from the
frontal ; internasals two-thirds or three-quarters the length of the pre-
frontals ; frontal once and a half as long as broad, longer than its dis-
tance from the end of the snout, slightly shorter than the parietals;
rarely a small azygous scale between frontal and parietals1; anterior
sublinguals separated from the mental except in the aberrant type of

1 In a specimen from Povo Nemlao (Boettger).

loveridge: African snakes 221

hessei in which they were extensively in contact with the mental, sub-
equal to, or slightly longer than, the posterior.

For characters common to all forms see p. 213; for scale counts see
statistical table on p. 216.

Color. Young Topotype. Above, the apex of a A-shaped prolonga-
tion of the black nuchal crossbar extends over parietal suture to the
frontal; a black circumorbital ring present or absent; loreal region and
sides of head with dusky markings, otherwise head white; body barred
alternately chocolate brown and white, the dark bars being subequal
in width to the light interspaces, whose edges are even whiter, 15
(equal to 15 pairs in adult) white bars on body, 3 on tail. Below,
throat white, rest whitish with dusky mottling especially along the
edges of the scales.

Adult. Above, snout and sides of head olive, but lower portion of
upper labials white, otherwise head and body black, grayish black, or
light brown, the edges of some scales tipped with pure white to form
narrow crossbars, of which there are from 14-22 pairs on body, 2-3 on
taih Below, whitish or yellowish, uniform, or with dusky mottlings
accentuated on the edges of the scales.

Size. Total length of c? (A.M.N.H. 51837), 475 (438 + 37) mm.
from ? Hanha, of 9 (CM. 5914), 337 (296 + 41) mm. from Chitau.

Anatomy. Bogert (1940) has described the hemipenis of an Angolan
semiannulata as follows : "Extends to the eleventh caudal, bifurcating
at the ninth. Sulcus bifurcates at the seventh caudal. Basal portion
with small spines, a row of enlarged spines increasing in size distally on
either side of the sulcus with smaller spines in the intervening space.
On the side opposite the sulcus, a pair of enlarged spines is present in
the region of the sixth caudal. Each fork is armed with small spines in
longitudinal rows."

Diet. Two hymenopterous larvae in a Katanga snake (Mertens).

Localities. Angola: Caconda; Cazengo; Chitau; Cubal; Dondi,
near Bella Vista (M.C.Z.); Galangue (Galanga); Gambos, Mossa-
medes; Kalukembe; Kampulu; Maconjo. Belgian Congo: (south-
ern frontier): Dilolo to Muciacia; Povo Nemlao; Povo Netonna.
Range. Angola and southern Belgian Congo.

222 bulletin: museum of comparative zoology

Elapsoidea sundevallii guntherii Bocage

1866a. Elapsoidea Guntherii Bocage, Jorn. Sci. Lisboa, 1, pp. 50, 70, pi. i,
figs. 3-3b: Cabinda, Portuguese Congo and Bissao, Portuguese
Guinea.

1866b. Bocage, p. 70.

1884a. Rochebrune, p. 192 (ignored).

1884b. Sauvage, p. 201.

1895a. Bocage (part), p. 129 (exclude Angola).

1895. Giinther (part), p. 525 (exclude Nyasaland).

1896. Tornier (part), p. 84 (exclude eastern Tanganyika).
1936h. Loveridge (part), p. 41 (exclude Angola).

1936j. Loveridge, p. 271.

1937c. Loveridge, p. 278.

1937d. Mertens, p. 9.

1937. Pitman (part), p. 346, pi. xiii, fig. 1, pi. N, fig. 3.

1937. Uthmoller, p. 123.
1938a. Pitman, pp. 218, 233.

1938b. Pitman, pp. 40, 81, 199, 257, 316, 331 (reprint with plates).

1938. Uthmoller, p. 46.

1942e Loveridge (part), p. 302 (exclude Magrotto).

1896d. Elapechis guentheri Boulenger (part), p. 359 (exclude Shire).

1896b. Mocquard, p. 45.

1897b. Boulenger, p. 280.

1897. Tornier, p. 65.
1900b. Boulenger, p. 455.
1902a. Boulenger, p. 447.
1906L Boulenger, p. 215.
1907. Lonnberg, p. 16.
1908b. Sternfeld, pp. 220, 234.
1909a. Sternfeld, p. 23.

1910b. Boulenger (part), p. 519 (exclude Transvaal).

1910. Meek, p. 405.

1910a. Sternfeld (part), p. 37 (exclude eastern Tanganyika).

1911c. Boulenger, p. 167.

1915a. Boulenger (part), p. 218 (exclude Angola & Nyasaland).

1915c. Boulenger (part), p. 635 (exclude Angola & Nyasaland).

1916a. Loveridge, p. 86.

1916b. Loveridge, pp. 117, 120.

1917. Chabanaud, p. 381.

1917b. Chabanaud, p. 13.

1918a. Loveridge, p. 324.

1919b. Boulenger (part), p. 294 (exclude Angola & Nyasaland)

1920a. Witte, p. 62.

1920b. Witte (part), p. 274 (exclude Angola & Nyasaland).

loveridge: African snakes 223

1922c. Angel, p. 357.

1923e. Loveridge, p. 889.

1923a. Werner (part), p. 179 (exclude Angola & Nyasaland).

1924b. Loveridge, p. 7.

1933f. Angel (part), p. 185, figs. 69-69c (exclude Angola etc.).

1933m. Witte, p. 96.

1897b. Elapechis moebiusi Werner, Verh. Zool.-Bot. Ges. Wien, 47, p. 400:

Kete, Togoland.

1910a. Elapechis niger Sternfeld (part, not Giinther), p. 37.

1920a. Witte, p. 62.

1920b. Witte, p. 275.

1923e. Loveridge (part), p. 890 (exclude Lumbo, Mozambique).

1934a. Elapechis sp. Schwetz, p. 381.

Among the above citations will be found those with or without the
final 'i', others with the 'ii' or 'ue' rendering, it has not been thought
necessary to indicate such minor deviations. Further citations of
'guntherii' will be found under s. nigra, s. semiannulata, s. decosteri and
s. laticincta.

Native name. Mugoya (Gishu), a name which is applied to any
Typhlops or Lcptotyphlops in Ganda or Soga, fide Pitman.

Description. Snout rounded; portion of rostral visible from above
measuring a third to half its distance from the frontal; internasals
two-thirds to three-quarters the length of the prefrontals ; frontal once
and a quarter to once and a half as long as broad, as long as, or longer
than, its distance from the end of the snout, slightly shorter than, rarely
equal to, the length of the parietals ; posterior nasal rarely separated
from the single preocular; anterior sublinguals slightly shorter than
the posterior.

For characters common to all forms see p. 213; for scale counts see
statistical table on p. 216.

Color. Young, ex. Kenya. Above, head as well as body, dark, the
white crossbars or bands broader than in adults, where each white bar
has split to form a pair.

Adult. Above, head and body plumbeous gray, brown, or black,
uniform, or the edges of some scales tipped with white to form trans-
verse series of white dots arranged in broad or narrow crossbars of
which there are some 15 to 33 pairs on body and 3 to 5 on tail. Below,
yellowish or whitish, each scale lightly edged with dusky, or grayish,
brownish, or black, the gular region usually lighter.

Whether snakes from Kenya (east of Kaimosi, Elgon and the Burnt
Forest) are separable by color remains to be demonstrated. The half-

224 bulletin: museum of comparative zoology

dozen Nairobi specimens which I captured had coral pink or red
centres to the white crossbars separating the black interspaces.
Whether such is the case with snakes from the Guaso Nyiro, Njoro,
and Loita Plains is not known. Pitman records the eyes of two Buka-
lasa snakes as being black with invisible pupil, and translucent with
silver pupil respectively.

Size. Total length of tf1 (M.C.Z. 40721), 630 (581 + 49) mm., and
of 9 (Mich. Mus.), 627 (583 + 44) mm., both from Kaimosi. The
former was erroneously cited as a 9 in a previous paper.

Remarks. The type of moebiusi, measuring 442 (412 + 30) mm.,
was dark brown above and light yellow below. Its author acquiesced
in Boulenger's action in referring it to the synonymy of guntherii.

Snakes from northern Tanganyika Territory recorded by Mertens
and Uthmoller are somewhat intermediate, agreeing more closely with
guntherii than with nigra, however.

Angel (1922c) records an aberrant individual from Clermont House,
Nairobi, in which the rostral, internasals, first and second upper
labials are irregularly broken up into small scales; the posterior nasal
is in contact with the preocular on one side, separated on the other;
the supraocular is divided on the right side only.

Loveridge (1936J) was in error in giving temporals as 1 + 3 and
ventrals 167, the former was a misprint for 1 + 2, the latter a mis-
count for 166.

Breeding. On September 10, at Kaiso, small eggs present in ovary
(Pitman).

Diet. Four lizard eggs, each measuring 8x4 mm., were found in
the stomach of a Nairobi snake.

Defence. Black specimens, apparently uniformly so, by sudden
inflation of the lungs bring into prominence the previously concealed
white-tipped bases of certain scales, thus producing an annulate effect
which is quite startling.

Temperament . Naturally peaceable and inoffensive, biting only upon
real provocation. In Parklands Forest Reserve I nearly trod on one
which was sluggishly making its way through the sparse herbage which
had sprung up with the advent of the rains.

Habitat. Sea level to 7000 feet. Pitman found a specimen in a
recently-planted cotton patch on the Kaiso plain, usually, however,
this somewhat secretive species is encountered on, or at the edge of,
forest. The Burnt Forest record is based on a snake which was cross-
ing the road at dusk, I held it down with my cycle pump, examined,
then released it.

loveridge: African snakes 225

Localities. Uganda: Budongo Forest; Bukalasa, 30 miles north
of Kampala; Bussu; Kabanda; Kagera River mouth; Kaiso-Tonya
Plain on east shore of Lake Albert; Kaliro in Busoga (A.M.N.H.);
Mabira Forest; Ruwenzori - — foot of; Serere, Teso; Sese Ids.;
Sipi, Mt. Elgon. Kenya Colony: Burnt Forest; Guaso Nyiro
(M.C.Z.); Kaimosi; Kajiado; Kakamega; Kijabe; Loita Plains
(M.C.Z.); Nairobi; Njoro; Parklands, Nairobi. Tanganyika
Territory (northern and western): Kagera (as Kagehi); Kibonoto;
Oldeani; Sanya. Portuguese Congo: Cabinda. Belgian Congo :
Dramba; Kunungu; Mahagi Port; Stanley Pool; Stanleyville;
Yakoma. French Equatorial Africa: Fort Sibut; Kuango
River. Nigeria. Dahomey: Agouagou. Gold Coast. Togo:
Kete; Mangu; Misahohe. Portuguese Guinea: Bissau; Rio Cas-
sine. Senegambia: Guidimaka.

Range. Northern Tanganyika Territory and Kenya Colony west to
Senegal.

Elapsoidea sundevallii nigra Giinther

1888b. Elapsoidea nigra Giinther, Ann. Mag. Nat. Hist. (6), 1, p. 322:

"Ushambola" i.e. Usambara Mountains, Tanganyika Territory.
1895b. Werner, p. 193.

1896. Tornier, p. 84.
1928e. Loveridge, p. 117.

1896d. Elapechis niger Boulenger, p. 359, pi. xx, fig. 1.

1897. Tornier, p. 65.

1910a. Sternfeld (part), p. 37, fig. 43 (Nguelo and Tanga only).

1915a. Boulenger (part), p. 218 (omit Congo and Northern Rhodesia).

1915c. Boulenger (part), p. 635 (omit Congo and Northern Rhodesia).

1920. Aders, p. 338.

1923a. Werner (part), p. 179 (omit Congo and Northern Rhodesia).

1924b. Loveridge (part), p. 7 (omit Kenya Colony and Zanzibar).

1940. Parker, Moreau & Pakenham, pp. 310, 313.

1896. Elapsoidea guntherii Tornier (part), p. 84 (Bulwa, Magila, Tanga).

1928c. Barbour & Loveridge, p. 134.

1937f. Loveridge, p. 502.

1942e. Loveridge (part), p. 302 (Magrotto only).

1910a. Elapechis Guentheri Sternfeld (part), p. 37 (Bulwa, Magila, Tanga).

1911b. Nieden, p. 442.

Further citations of "nigra" or "niger" will be found under s. gun-
therii and s. decosteri.

Native name. Kifutu (Sambara).

226 bulletin: museum of comparative zoology

Description. Snout rounded; portion of rostral visible from above
measuring a quarter to a third its distance from the frontal; internasals
rarely half, usually two-thirds to three quarters, rarely equal to, the
length of the prefrontals ; frontal once and an eighth to once and a half
as long as broad, rarely as long as, more usually longer than, its dis-
tance from the end of the snout, slightly shorter than, or two thirds the
length of the parietals ; postoculars only rarely 1 ; temporals only rarely
1 + 1 ; anterior sublinguals subequal to, or slightly longer than, the
posterior.

For characters common to all forms see p. 213; for scale counts see
statistical table on p. 216.

Other aberrations noted are : a small azygous scale between pre-
frontals and frontal in M.C.Z. 23411, and topotypes 23424, 23435; a
minute loreal present in topotype M.C.Z. 23427; the specimen (M.C.Z.
23413) with a single postocular has it on the left side only; a small
azygous scale is present between lower postocular and temporal on
both sides of M.C.Z. 23424, 23427, 23435, and on right side only of
23441.

Color. Young. Above, the apex of a A-shaped prolongation of the
nuchal crossbar extends over parietal suture, otherwise head white or
brownish above; body gray with about a score of black crossbars
narrowly edged with white, the dark bars being subequal in width to
the gray interspaces ; tail uniformly black. It will be observed that the
light transverse bars of the young progressively disappear with age,
from the tail forwards, to produce the uniformly black adults. Below,
whitish anteriorly, grayish posteriorly.

Adult. Above, head and body glossy iridescent black, uniform or
the edges of some scales tipped with white to form transverse series
of white dots or narrow crossbars, of which there may be some 18-24
pairs on the body. Below, usually uniformly black, paler or even white
upon the throat, very rarely a few white patches in the middle of the
ventrals.

Size. Total length of c? (M.C.Z. 48448), 585 (548 4- 37) mm. from
Magrotto, of 9 (M.C.Z. 23455), 494 (460 + 34) mm. from Bumbuli,
of type 9 (Brit. Mus.), 420 (390 4- 30) mm. A 569 mm. specimen,
listed by me as 9 , proves on reexamination to be a cf .

Remarks. Sternfeld's (1910a) statement that the anal is divided was
just a slip.

Breeding. On October 1, at Nyange, a 9 held 2 eggs measuring 40 x
10 mm. On November 29-30, at Amani, four 9 9 held 2, 3, 4, and 4
eggs respectively, of which the largest measured 27 x 10 mm. On

loveridge: African snakes 227

December 16, at Bumbuli, two 9 9 held 4 and 5 eggs respectively, the
latter measuring 37 x 12 mm.

Diet. Caecilians (Boulengerula boulengeri) of which 7 were recovered
from the stomachs of 6 snakes, for details see Barbour and Loveridge
(1928).

Parasites. Nematodes (Kalieephalvs sp.) and their cysts taken from
Amani snakes, cestodes (Proteocephalus or Ophiotaenia elapsoidea, and
0. sp. near monnigi) taken from Amani and Nyange reptiles, mites
beneath ventrals of Amani specimen.

Temperament. If gently handled this montane snake makes no
attempt to bite so cannot be considered a vicious species. Apparently
wholly black examples, their small heads scarcely larger than their
bodies, are sufficiently like burrowing vipers (Atractaspis rostrata) to
be mistaken, others which at first sight appear entirely black, will, if
molested, inflate their lungs and thus bring into prominence the series
of transverse bars, causing them to look like Crotaphopeltis h. hotam-
boeia. Two Magrotto snakes had lost the ends of their tails, perhaps
through fighting; one had a very truncated stump.

Habitat. Vicinity of montane rain forest between 2000 and 3000
feet, in heaps of debris within or without the forest. In the Usambara
forests several were encountered on paths between noon and 3 p.m.,
but at Magrotto I captured four wandering on paths about sunset,
their empty stomachs suggesting the reason for their being abroad.

Localities. Tanganyika Territory : Magrotto Mountain ; Tanga ;
Uluguru Mountains — Nyange; Usambara Mountains — Amani;
Bulwa (Buloa); Bumbuli; near Magila; Nguelo.

Because Sir John Kirk, H. M. Consul at Zanzibar, sent home the
type from Ushambola, Boulenger concluded that Ushambola was
in Zanzibar. Finding the species abundant in the Usambara
Mountains in 1926, I surmised that Ushambola was an archaic
spelling of Usambara for Sir John's duties took him up the Pangani
River in what was once called the Usambara district. Moreover
Aders (1920) remarks that he had never come across the species
on the island, but for further confirmation I approached the Survey
Department in Zanzibar who replied that they had no knowledge
of such a locality as "Ushambola" on the island. Zanzibar,
therefore, must be excluded from the range.

The three records of the occurrence of nigra in Mozambique, Nyasa-
land, and Northern Rhodesia, are referable to the race s. decosteri
whose adults are just as black as nigra but have a much lower ventral
count.

228 bulletin: museum of comparative zoology

Range. Eastern Tanganyika Territory (Usambara, Magrotto, and
Uluguru Mountains).

Elapsoidea sundevallii sundevallii (Smith)

1848. Elaps sunderwallii (sic) A. Smith, 111. Zool. S. Africa, Rept., p. Ixvi:

"South Africa to east of Cape Colony."

1880c. Peters, p. 797, pl. -, fig. 2 (sundevallii, emend).

1887h. Boulenger, p. 180.

1896d. Elapechis sundevallii Boulenger, p. 360.

1897d. Boulenger, p. 375.

1898. Sclater, p. 101.

1898. Werner, 1896-7, p. 147 (see Remarks below).

1908b. Boulenger, p. 230.

1908. Gough (part), p. 34 (omit Orange River Colony).

1910b. Boulenger, p. 519.

1912. FitzSimons, F. W., pp. 166, 168(omit Griqualand and Basutoland).

1923a. Werner, p. 179 (but restrict range to Natal).

1937e. Elapsoidea sundevallii Hewitt, p. 78.

1937a. FitzSimons, V., p. 263.

Another citation of 'simdevalli' will be found under s. decosteri.
From Werner onwards several authors have dropped the final '[' from
sundevallii; it has not been thought necessary to indicate such trifling
deviations.

Description. Snout obtusely pointed ; portion of rostral visible from
above two-thirds of, equal to, or greater than, its distance from the
frontal; internasals half to two-thirds the length of the prefrontals;
frontal once and a third to once and a half as long as broad, slightly
shorter than, as long as, or longer than, its distance from the end of
the snout, slightly shorter than, or two-thirds the length of the parie-
tals; posterior nasal rarely separated from the single preocular; tem-
porals only rarely 1 -f- 1 ; anterior sublinguals subequal to, or slightly
longer than, the posterior.

For characters common to all forms, see p. 213; for scale counts see
statistical table on p. 216.

Color. Young. Above, the apex of a A-shaped prolongation of the
black nuchal crossbar extends over parietal suture forwards to pre-
frontals ; body barred alternately with black and white, the bars being
of equal width, edges of the pale bars lighter and doubtless persisting
to form the narrow crossbars of the adult. Below, yellowish.

Adult. Above, slaty gray with a reddish or purplish brown tinge, or

lovekidge: African snakes 229

rusty brown, at least part of labials and lower temporal region yellow-
ish, body and tail with 14 -+- 31 to 34 + 4 white- or yellow-edged black
crossbars, which are as broad as, or much broader than, the inter-
spaces between them, though often narrowing on the sides. Below,
usually including lower lateral scale-row, yellowish or whitish, with or
without dusky mottling or brownish marbling.

Size. Total length of an unsexed snake (Brit. Mus.), 880 (825 -f-
55) mm., and of a 9 (Werner, 1898), 625 (590 4- 35) mm.

Remarks. Smith (1848) gives a detailed account of the type which
is worth consulting, its accuracy is vouched for by Peters (1880c) who
borrowed the type and a juvenile example from the Royal Swedish
Museum before transferring the species to Elapsoidea of Bocage, which
genus he considers to be closely related to Hemibungarus Peters
(i.e. Calliophis Giinther) of southeast Asia.

Peracca's (1896) identifications of "sundevali" and "guentheri" as
occurring at Kazungula on the south bank of the Zambesi, Southern
Rhodesia, are both referred to s. decosteri though there is a remote
possibility that one might be the young of fitzsimonsi.

Werner's (1898) record of the 'Cape' must be considered erroneous
pending confirmation. His paper deals with three collections, whose
source he indicates by putting a 'K' (Kap), 'N' (Natal), or T' (Trans-
vaal) after the species. I suggest that the *K' which appears after his
undoubtedly typical sundevallii, may well have come from a carelessly
written manuscript 'N'.

Gough (1908) mentions several abnormalities, but his "O.R.C" (i.e.
Orange River Colony, now Orange Free State) record really refers to a
decosteri, as does a Pretoria specimen (M.C.Z. 14194) labeled "sunde-
vallii" when received from the Transvaal Museum in 1920.

Localities. See three preceding paragraphs. Natal: De Deur
near Evanton; Durban; Estcourt; Newcastle.

Range. Natal.

Elapsoidea sundevallii fitzsimonsi subsp. nov.

1910b. Elapechis Decosteri Sternfeld (not Boulenger), p. 31.

1910c. Sternfeld, p. 57.

1913. Elapechis sundevalli Hewitt & Power (not Smith), p. 165.

1935b. FitzSimons, V., p. 326.

1937b. Elapsoidea sundevallii Mertens (not Smith), p. 15.

1 The lower number apparently results from loss of the white edging and consequent fusion
of two dark crossbars, which results in the dark bars being much narrower than the interspaces.

230 bulletin: museum of comparative zoology

Type. Chicago Natural History Museum, No. 17667, an adult d1
from Gomodimo Pan, Kalahari Desert, Bechuanaland Protectorate,
collected by Herbert Lang, April, 1930.

Paratypes. C.N.H.M. No. 17668, a cf with same data as type;
C.N.H.M. 17666, a ? taken between Gomodimo and Kuke; also
Transvaal Museum (not seen, but data supplied by V. FitzSimons)
from Okwa River and Damara Pan respectively, all collected by the
Vernay-Lang Kalahari Expedition in April, 1930.

Diagnosis. Differs from typical sundevallii, with which it agrees in
other respects, in being uniformly purplish brown when adult.

Description. Nasal and preocular in contact ; preocular 1 ; postocular
2; temporals 1 + 2; upper labials 7, the third and fourth entering
the orbit; lower labials 6-7 of which the first 3 (3-4 in paratypes)
are in contact with an anterior sublingual; ventrals 181 (162-177,
not 164-178); subcaudals 21 (17-23).

Snout obtusely pointed; portion of rostral visible from above two-
thirds of, or equal to, its distance from the frontal; internasals half to
two-thirds the length of the prefrontals; frontal once and a third to
once and a half as long as broad, slightly shorter than, as long as, or
longer than, its distance from the end of the snout, slightly shorter
than, or three-quarters, the length of the parietals ; anterior sublinguals
subequal to the posterior. (This paragraph is based only on the three
specimens examined.)

Color. Young. Above, the apex of a A-shaped prolongation of the
black nuchal crossbar extends over parietal suture forwards to pre-
frontals; a dark spot behind eye; sides of head yellowish; body barred
alternately with dark slaty brown or black, and pale yellowish or white,
the bars being of equal width, edges of the pale scales usually dark-
edged. Below, yellowish.

"Adult and halfgrown. Above, uniform dark slaty gray, with a
reddish or purplish brown tinge, paler posteriorly and on sides; upper
lip, outer row of scales and lower half of adjacent row, creamy white.
Below, uniform creamy white, snout and sides of tail tinged with
pinkish."

Size. Total length of type <? (C.N.H.M. 17667), 714 (671 + 43)
mm., surpassed by a cf (T.M. 887), 766 (716 + 50) mm.; paratype 9
(C.N.H.M. 17666), 606 (570 + 36) mm.

Remarks. Named for Mr. Vivian FitzSimons of the Transvaal
Museum, who supplied me with numerous scale-counts, and who was
the first to invite attention to this form, and whose admirable account
of the coloration I have given above.

loveridge: African snakes 231

FitzSimons (1935b) suggests, and probably correctly, that Stern-
feld's (1910b) record of decosteri from South West Africa is the same as
these Bechuanaland specimens. If correct, then Sternfeld's figureof 157
ventrals requires checking as it is slightly lower than the accepted range
(162-181).

Whether Peracca's (1896) record of sundevallii from Kazungula on
south bank of Zambesi, Southern Rhodesia, is referable to this form
or to s. guntherii where I have put it, is open to question.

Habits. These active snakes were usually found in the vicinity of
pans, where their coloration rendered them conspicuous on the light
colored sand, in early evening following rain (FitzSimons).

Localities. See remarks above. Bechuanaland : Damara Pan;
Gomodimo Pan; Gomodimo to Kuke; Okwa River. Cape Prov-
ince: Kimberly; Kimberly Mine. South West Africa: Gobabis
(as decosteri); Okahandja (Transv. Mus.); Okanjande near
Otjiwarongo (Mertens).

Range. Eastern Cape Province (Kimberly) west through Bechuana-
land Protectorate to South West Africa.

Genus Paranaja gen. nov.
Genotype. Naja midtifasciata anomala Sternfeld (M.C.Z. 22380).

Maxillary bone extending forwards as far as the palatine, with a pair
of large grooved poison-fangs followed by 2 small teeth ; anterior mandi-
bular teeth longest. Head moderate, slightly distinct from neck; eye
moderate, with round pupil; nostril between two nasals and an inter-
nasal; loreal absent; a single preocular. Body cylindrical; scales
oblique, smooth, without pits, in 15-17 rows; ventrals rounded. Tail
short; subcaudals in two rows.

Range. Africa from the Belgian Congo to French Cameroon.

Remarks. Though agreeing with Elapsoidea in the degree of forward
extension of the maxillary bone, the new genus is readily distinguish-
able from it by external characters such as the larger eye, oblique
nature of scalation, and relatively longer tail which in Elapsoidea is
very short.

The interesting species on which this monotypic genus is based, was
first referred to Naia (=Naja) by its discoverer (1902a) but later was
transferred to Elapechis ( = Elapsoidea) with a query by Boulenger
(1915a) who had already described it (1904d) as an Elapechis. Stern-
feld (1917), with only a head, described under the appropriate name
anomala as a full species what appears to be a recognizable race.

232 bulletin: museum of comparative zoology

Bogert, who has recently been studying cobra skulls (1943), and to
whom I am indebted for the following notes based on his dentitional
examination, considers Paranaja most closely related to Pseudohaje
for the palatine in both genera is almost identical in shape, exhibiting
a long internal process above the point where it is connected to the
pre vomer (vomer of most authors), moreover the external process
(processus maxillaris) is elongated as in Pseudohaje in contrast to the
condition obtaining in Naja and Elapsoidea; in the latter the internal
process is greatly reduced, while in Naja it is more or less truncate,
the curved extension at the posterior end of the process being absent in
most species and only feebly developed in Naja haje. In view of the
similarity of the palatine, as well as agreement with Pseudohaje in
general shape of the ectopterygoid, pterygoid, and maxillae, it appears
probable that other skull characters (no skull being available for
examination at the present time) will be found to conform with those
of Pseudohaje.

Though the distal aperture of the fang in atiomala is small, as in
Pseudohaje, Bogert finds the fang length — -1.6 mm. (measured by
Klauber's method) is contained 250 times in total length, and is,
therefore, proportionately about half as large again as in Pseudohaje
and but slightly larger than in most Naja. The further dentition of
atiomala, as shown by M.C.Z. 22380, is : palatine 9-9, pterygoid 19-19,
dentary 17-17. Though the palatine count is higher than encountered
by Bogert in any African species of Naja, it is sometimes exceeded in
the Asiatic N. naja naja.

The scalation is so irregular that any count is apt to be a trifle con-
fused, for on the specimen in question Bogert found a row dropping
out and then reappearing again, the formula being something like
19-17-15-16-15-13, and adds that in all Pseudohaje examined by him
the preanal count was constantly 11.

The diameter of the eye in Pseudohaje is about half its distance from
the mouth, whereas in Paranja it is about equal to the distance; such
comparisons, however, are likely to be affected by the age of the snake.
The ratio of tail into total length is .16, or nearly twice that of Elap-
soidea, though our specimen is a female.

Thus in the number of teeth on the respective bones, in the number
of scales and their imbrication, in size of eye, and in relative tail length,
Paranaja agrees more nearly with Naja than with Pseudohaje.

If I were asked to speculate on lines of descent, I would suggest that
Naja, entering Africa from the northeast, gave rise to both the allegedly
arboreal Pseudohaje and the presumably terrestrial Paranaja, the latter

loveridge: African snakes 233

retaining many Naja characteristics. Both are sylvicoline as is Naja
melanoleuca which likes to bask in trees or is ready to take to them; on
the other hand only once have I found the savanna-haunting N. n.
nigricollis off the ground, and then only in a Bougainvillea.

Key to the Races

Upper labials 7, sixth largest; range: Belgian Congo, i.e. south of the River

Congo m. multifasciata

(p. 233)
Upper labials 6, fifth largest; range: French Cameroons, i.e. north of the River

Congo m. anomala

(p. 234)

Paranaja multifasciata multifasciata (Werner)

1902a. Naia multifasciata Werner, Verh. Zool.-Bot. Ges. Wien, 52, p. 347:

Upper Maringa River, Belgian Congo.

1923a. Werner, p. 183.

1904d. Elapechis Duttoni Boulenger, Ann. Mag. Nat. Hist. (7), 14, p. 15:

Leopoldville, Belgian Congo.

1915a. Boulenger, p. 218.

1920a. Witte, p. 62.

1920b. Witte, p. 275.

1923a. Werner, p. 180.

1933a. Witte, p. 70.

1915a. Elapechis ? multifasciatus Boulenger, p. 219.

1920a. Witte, p. 62.

1920b. Witte, p. 275.

1933a. Witte, p. 70, figs. 1-3.

Description. Snout broadly rounded; rostral broader than deep, the
portion visible from above measuring from a third to two-thirds its
distance from the frontal ; internasals three-quarters the length of the
prefrontals; frontal once and a half as long as broad, slightly shorter
than, as long as, or longer than, its distance from the end of the snout,
two-thirds or four-fifths the length of the parietals ; posterior nasal in
contact with the single preocular; eye as long as, or longer than, its
distance from the nostril; postoculars 2-3; temporals rarely 1 + 2,
usually 1 + 3, rarely 2 + 3; upper labials 7, the third and fourth
entering the orbit, sixth largest; 4 lower labials in contact with the
anterior sublinguals, which are separated from the mental, and slightly
shorter than, or as long as, the posterior. Midbody scales in 15-17

234 bulletin: museum of comparative zoology

rows, 17-19 on neck, 19-21 immediately behind parietals; ventrals
150-174; subcaudals 31-38, all or most in two rows.

Color. Above, snout and sides of head whitish, a dark transverse bar
across internasals, a dark streak over labial suture beneath eye, fol-
lowed by others on labial sutures and temporal region; head (from
prefrontals) and body pale brownish, each scale with a dark spot
posteriorly resulting in the formation of a series of angular crossbands.
Below, whitish.

Size. Total length of multifasciata type (Mus. Roy. Hist. Nat.
Bruxelles, presumably a d" as with 38 subcaudals), 525 (462 + 63)
mm., of duttoni type (Brit. Mus., presumably a 9 as with 31 sub-
caudals, though Boulenger suggested that it might be a cf), 520 (450
+ 70) mm.

Remarks. Werner (1902a) considered that multifasciata was most
nearly related to "Naja goldii" from which it was readily distinguished
by its low subcaudal count. Witte (1933a), who reexamined and
figured the type, states that the two suboculars described by Werner
are nonexistent, and finds 19 instead of 17 scale-rows on the neck,
perhaps the explanation may be found in Witte's count having been
made nearer the head (vide Remarks under genus). Witte also amends
the ventral count from 172 to 174, and that of the subcaudals from 36
to 38, strangely enough he has a specimen from Balombo with pre-
cisely the same counts — 174 and 38.

Boulenger claimed that duttoni was closely related to E. s. nigra,
an entirely erroneous view; indeed it is somewhat surprising that he
ever should have considered placing "duttoni" in the genus "Elapechis"
from which it differs in the head being slightly distinct from the neck,
the moderate eye, the nostril being between two nasals and an inter-
nasal, midbody scales in 15-17 (instead of 13) rows, with 17-19 on
neck, and a tail which is only moderately short.

Localities. Belgian Congo: Besankusu; Bolombo (Bolombe);
Leopold ville; Upper Maringa River.
Range. Belgian Congo.

Paranaja multifasciata anomala (Sternfeld)

1917. Naja anomala Sternfeld, Wiss. Ergeb. Zweit. Deutschen Zent.-Afr.-
Exped. 1910-1911, 1, p. 482, pi. xxiv, fig. 9: Assobam Forest region,
French Cameroons.

1922a. Mertens, p. 182.

1923a. Werner, p. 182.

1924b. Werner, p. 45.

loveridge: African snakes

235

Description. Differs from the typical form only in those characters
mentioned in the key and also, though of no consequence, in the por-
tion of the rostral visible from above measuring from a quarter (type)
to two-thirds (M.C.Z. specimen) its distance from the frontal, and the
anterior sublinguals being slightly longer than the posterior (type and
M.C.Z.) • It may be thought by some that these are rather slender
grounds on which to recognise a race; in view of a definite geographical
barrier, however, the action seems justifiable. As the species is so rare,
I append the scale-counts so far published, those of Werner's type as
amended by Witte.

P. m. mxdtifasciata

Source Locality

Midbody

scales

Ventrals

Sub-
caudals

Upper labials Post-
Center orbit) oculars

Type U. Maringa

174

7 (3rd & 4th) 2

duttoni Leopoldville

150

7 (3rd & 4th) 3

Witte 1 Basankusu

164

? ?

1933a J Bolombo

17
P. m

174

. anomala

? ?

Source Locality

Midbody
scales

Ventrals

Sub-
caudals

Upper labials Post-
Center orbit) oculars

Type Assobam F.

6 (3rd & 4th) 2

M.C.Z. Bitye, Ja R.

164

6 (3rd & 4th) 2

Werner \ ?

157

6 (3rd & 4th) ?

1924b /

Size. Total length of larger (Werner, 1924b, probably 9 ), 515
(440 + 75) mm., of known 9 (M.C.Z. 22380), 463 (400 + 63) mm.

Remarks. Sternfeld (1917) based his anomala on a head only, while
noting its similarities to multijasciata he was misled by Werner's mis-
statements regarding the presence of suboculars. He believed that
anomala has a deeper rostral and larger eye, and went on to compare
it with Naja melanoleuca of the same size which had an even larger
eye. Werner (1924b) records a second example in the Vienna Museum.

Localities. French Cameroons: Assobam (Assobom); Bitye,
Ja River.

Range. French Cameroons.

236 bulletin: museum of comparative zoology

BIBLIOGRAPHY

of 319 works mentioning the species dealt with in this paper from
1735-1943. No attempt has been made to complete those prior
to 1880, but 1766 contributions to African herpetology published
since 1880 have been searched. Titles of papers are omitted for

the sake of brevity.

Aders, W. M.

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1921a. Bull. Mus. Hist. Nat. Paris, 27, pp. 42-44, figs. 1-3.

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1922. Sierra Leone Studies (Freetown), 5, pp. 7-37.

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1859? "Specimina Zoologica Mossambicana." (Bologna), pp. 1-282, pis.
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1870. "Observations on the Geology and Zoology of Abyssinia, . . ."
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Bocage, J. V. Barboza du

1866a. Jorn. Sci. Lisboa, 1, pp. 37-56.
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loveridge: African snakes 237

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1887a. Jorn. Sci. Lisboa, 11, pp. 177-211.

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BOETTGER, O.

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BOULENGER, G. A.

1887b. Proc. Zool. Soc. London, p. 127.

1887h. Zoologist (3), 11, pp. 171-182.

1888d. Ann. Mag. Nat. Hist. (6), 2, pp. 136-141.

1890b. Ann. Mag. Nat. Hist. (6), 6, pp. 91-93.

1891a. Proc. Zool. Soc. London, pp. 305-309.

1893a. Ann. Mag. Nat. Hist. (6), 12, p. 273.

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238 bulletin: museum of comparative zoology

1898a. Ann. Mus. Civ. Stor. Nat. Genova (2), 18, pp. 715-723, pis. ix-x.

1900b. Proc. Zool. Soc. London, pp. 433-456, figs. 1-2, pis. xxvii-xxxii.

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loveridge: African snakes 239

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1860. Proc. Acad. Nat. Sci. Philadelphia, pp. 241-266, 553-566.

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1928. Proc. Zool. Soc. London, pp. 923-961, pis. i-iv.
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1826. "Neue Classification der Reptilien . . ." (Wien), pp. i-viii + 1-66,
pi. i.
FitzSimons, F. W.

1912. "The Snakes of South Africa; their Venom and the Treatment of
Snake Bite." (Cape Town) 2nd. ed., pp. i-xvi + 1-547, figs. 1-162
+ A-Z, pi. col.
FitzSimons, V.

1935b. Ann. Transvaal Mus., 16, pp. 295-397, figs. 1-30, pis. x-xi.
1937a. Ann. Transvaal Mus., 17, pp. 259-274, pi. x.
1939b. Ann. Transvaal Mus., 20., pp. 17-46.
Gendre, E.

1909. Extr. Comptes Rendus Soc. Linn. Bordeaux, in Actes, 63, pp. cv-
cvi.
Gmelin, J. F.

1788. "Caroli a LinnS . . . Systema Naturae ..." (Lipsiae), ed. 13, 1,
part 3, pp. 1033-1516.

240 bulletin: museum of comparative zoology

Gough, L. H. •

1903. Zool. Jahrb. Syst., 17, pp. 457-468.
1908. Ann. Transvaal Mus., 1, pp. 17-45.

GUNTHER, A.

1858c. "Catalogue of the Colubrine Snakes in the Collection of the British

Museum." (London), pp. i-xvi + 1-281.
1859a. Ann. Mag. Nat. Hist. (3), 4, pp. 161-174, pi. iv.
1862b. Ann. Mag. Nat. Hist. (3), 9, pp. 52-59, 124-132, pis. ix-x.
1863a. Ann. Mag. Nat. Hist. (3), 11, pp. 20-25, pi. iii.
1864b. Proc. Zool. Soc. London, pp. 303-314, pis. xxvi-xxvii.
1865a. Ann. Mag. Nat. Hist. (3), 15, pp. 89-98, pis. ii-iii.
1865b. Zool. Rec, 2, pp. 139-162.

1866a. Ann. Mag. Nat. Hist. (3), 18, pp. 24-31, pis. vi-vii.
1872a. Ann. Mag. Nat. Hist. (4), 9, pp. 13-37, pis. iii-vii.
1881b. in Oates, F., "Matabele Land and the Victoria Falls." App., pp.

229-330, pis. C-D.
1888b. Ann. Mag. Nat. Hist. (6), 1, pp. 322-335, pis. xviii-xix.
1889a. in Oates, F., "Matabele Land and the Victoria Falls." ed. 2, App.,

pp. 229-330, pis. C-D.
1893. Proc. Zool. Soc. London, 1892, pp. 555-558, pis. xxxiii-xxxy.
1894a. Proc. Zool. Soc. London, 1893, pp. 616-628, pis. liii-lvii.
1895. Ann. Mag. Nat. Hist. (6), 15, pp. 523-529, pi. xxi.
1895b. "Biologia Centrali-Americana. Reptilia and Batrachia." 1885-

1902, (London), pp. i-xx + 1-326, pis. i-lxxvi.

Hallowell, E.

1844. Proc. Acad. Nat. Sci. Philadelphia, pp. 58-62.

1852b. Proc. Acad. Nat. Sci. Philadelphia, pp. 203-205.

1854a. Proc. Acad. Nat. Sci. Philadelphia, pp. 98-105.

1857. Proc. Acad. Nat. Sci. Philadelphia, pp. 48-72.
Hermann, J.

1804. "Observationes zoologicae quibus novae complures aliaeque
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Hesse, P.

1889. Zool. Garten (Frankfurt am. Main), 30, pp. 257-267.
Hewitt, J.

1910a. Ann. Transvaal Mus., 2, pp. 50-71.

1912. Rec. Albany Mus., 2, pp. 264-285.
1913e. Ann. Natal Mus., 2, pp. 475-484, pi. xxxix.

1937e. "A Guide to the Vertebrate Fauna of the Eastern Cape Province,
South Africa. Part II. Reptiles, Amphibians and Freshwater
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Hewitt, J. and Power, J. H.

1913. Trans. Roy. Soc. S. Africa, 3, pp. 147-176.

loveridge: African snakes 241

HOBLEY, C. W.

1912. Journ. E. Africa Uganda Nat. Hist. Soc, No. 5, pp. 43-56, figs.
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1857. Cenni Museo Civico Milano (non vidi).

1858. Revue Mag. Zool. (2), 10, pp. 438-449, 514-527.

1859. Revue Mag. Zool. (2), 11, pp. 122-130, 148-157, 505-512, pis. iv,
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1862. Arch. Zool. Anat. Fisiol., 2, pp. 1-76, pis. v, vii, xvii-xviii.

1863. "Elenco sistematico degli ofidi descritti e disegnati per l'lcono-
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1865-. "Iconographie generale des Ophidiens." (Milano), livre.
1869. 13-32, pis. (which are numbered afresh for each part).

Jeude, T. W. van Lidth de

1895. Notes Leyden Mus., 16, pp. 227-230.

1898. "Catalogue" Mus. Hist. Nat. Pays-Bas, 10, part 2, pp. 1-54 (fish)
+ 1-52 (reptiles) + 1-11 (amphibians).
Johnston, H. H.

1898. "British Central Africa." (London), pp. 355-361a.

1906. "Liberia." (London), 2, pp. 521-1183, Mus. maps.
Kuhl, H.

1820. "Beitrage zur Zoologie . . ." etc. (Frankfurt am Main), pp. 1-151.
Lampe, E.

1911. Jahrb. Nassau Ver. Naturk., 64, pp. 137-236.
Lampe, E. and Lindholm, W. A.

1902. Jahrb. Nassau Ver. Naturk., 55, pp. 1-66.

Latreille, P. A. see Sonnini etc.

Lepri, G.

1910. Boll. Soc. Zool. Ital. (2), 11, pp. 317-328.

Linnaeus, Carl von

1758. "Systema Naturae." (Holmiae), ed. 10, 1, pp. 1-824.

1760. "Systema Naturae." (Halae Magdeburgicae), ed. 12, 1, pp. 1-532

Lonnberg, E.

1907. in Sjostedt, Y., "Wissenschaftliche Ergebnisse der Schwedischen
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1911. Svenska Vetensk.-Akad. Handl. (Stockholm), 47, No. 6, pp. 1-42,
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Lonnberg, E. and Andersson, L. G.

1913. Arkiv Zool., 8, No. 20, pp. 1-6.

Loveridge, A.

1916a. Journ. E. Africa Uganda Nat. Hist. Soc, 5, pp. 77-87.
1916b. Journ. E. Africa Uganda Nat. Hist. Soc, 5, pp. 109-214.
1918a. Journ. E. Africa Uganda Nat. Hist. Soc, No. 13, pp. 315-338.

242

bulletin: museum of comparative zoology

1923e. Proc. Zool. Soc. London, pp. 871-897.

1924b. Journ. E. Africa Uganda Nat. Hist. Soc, Spec. Suppl. No. 3, pp

1-16.
1928d. Proc. U. S. Nat. Mus., 73, Art. 17, pp. 1-69, pis. i-iv.
1928e. Bull. Antivenin Inst. America, 1, pp. 106-117.
1928g. Bull. Antivenin Inst. America, 2, pp. 32-41.
1928J. Bull. Antivenin Inst. America, 2, pp. 72-76.
1929h. Bull. U. S. Nat. Mus., No. 151, pp. 1-135, pi. i.
1933h. Bull. Mus. Comp. Zool., 74, pp. 197-416, pis. i-iii.
1935c. Bull. Mus. Comp. Zool., 79, pp. 1-19.
1936f. Proc. Biol. Soc. Washington, 49, pp. 59-62.
1936h. Zool. Ser. Field Mus. Nat. Hist., 22, pp. 1-111.
1936j. Bull. Mus. Comp. Zool., 79, pp. 209-337, pis. i-ix.
1937c. Proc. Acad. Nat. Sci. Philadelphia, 89, pp. 265-296.
1937f. Bull. Mus. Comp. Zool., 79, pp. 481-541, pis. i-iv.
19S8d. Proc. New England Zool. Club, 17, pp. 49-74.
1941e. Proc. U. S. Nat. Mus., 91, pp. 113-140.
1942e. Bull. Mus. Comp. Zool., 91, pp. 237-373, pis. i-vi.
Martinez y Saez, Don F. de P.

1886. Ann. Soc. Esp. Hist. Nat. Madrid, 15, p. 339.
Matschie, P.

1891a. Zool. Jahrb. Syst., 6, pp. 605-611.
1891b. Zool. Jahrb. Syst., 5, pp. 612-618.
1892. Sitzb. Ges. Naturf. Freunde Berlin, pp. 101-110.
1893c. Mitt. Fors. Gel. Deutschen Schutzgeb., 6, pp. 207-215.
Meek, S. E.

1910. Zool. Series Field Mus. Nat. Hist., 7, pp. 403-414.
Merrem, B.

1790. "Beitrage zur Naturgeschichte." (Duisburg & Lemgo), pp. i-ii +
1-47, col. pis. i-xii.
Mertens, R.

1922a. Senckenbergiana, 4, pp. 162-183.

1937b. Abh. Senckenberg. Naturf. Ges., No. 435, pp. 1-23.

1937d. Veroftenlich. Deutschen Kolon.-Ubersee-Museum Bremen, 2, pp.

1-9.
1938b. Abhand. Senckenberg. Naturf. Ges., No. 442, pp. 1-52, pis. i-x.
1938e. Senckenbergiana, 20, pp. 425^42, figs. 1-6.
1940a. Zool. Anz., 131, pp. 239-250, fig.

MOCQUARD, F.

1887b. Bull. Soc. Philom. Paris (7), 11, pp. 62-92, pis. i-ii.
1889. Bull. Soc. Philom. Paris (8), 1, pp. 145-148.
1896b. Compte-Rendu Soc. Philom. Paris, pp. 44-45.
1897b. Bull. Soc. Philom. Paris (8), 9, pp. 5-20.
1897c. Bull. Mus. Hist. Nat. Paris, 3, pp. 122-123.

loveridge: African snakes 243

1899a. Bull. Mus. Hist. Nat. Paris, 5, pp. 218-219.
1902b. Bull. Mus. Hist. Nat. Paris, 8, pp. 404-417.
1905a. Bull. Mus. Hist. Nat. Paris, 11, pp. 76-79.
1905b. BuU. Mus. Hist. Nat. Paris, 11, pp. 285-290.
1908a. Bull. Mus. Hist. Nat. Paris, 14, pp. 259-262.
1908b. in Foa, E., "Resultats scientifiques des Voyages en Afrique
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Monard, A.

1931. Bull. Soc. Neuchatel. Sci. Nat., 55, pp. 89-111, figs. 1-5.
1937b. Arqu. Museu Bocage (Lisboa), 11, pp. 1-154, figs. 1-3.

Muller, F.

1885a. Verh. Naturf. Ges. Basel, 7, pp. 120-165, pis. ix-xi.
1885d. Verh. Naturf. Ges. Basel, 7, pp. 668-717, pis. ix-xi.
1890b. Verh. Naturf. Ges. Basel, 8, pp. 685-705, pi. x.
1892. Verh. Naturf. Ges. Basel, 10, pp. 195-215, pis. iii-iv.

MfJLLER, L.

1910. Abhand. K. Bayer. Akad. Wiss. (Miinchen), Kl. 2, 24, pp. 543-
626, pi. -.

NlEDEN, F.

1911b. Sitzb. Ges. Naturf. Freunde Berlin, pp. 441-^52.

Noble, G. K.

1921b. Nat. Hist. (New York), 21, pp. 166-171.

Odhner, T.

1908. Arkiv Zool., 4, No. 18, pp. 1-7.

Parker, H. W.

1933. Ann. Mag. Nat. Hist. (10), 12, pp. 544-548, figs. 1-2.

1936c. Novit. Zool. (Tring), 40, pp. 115-146, figs. 1-2.

1936e. Ann. Mag. Nat. Hist. (10), 18, pp. 594-609, figs. 1-13.

1937a. Ann. Mag. Nat. Hist. (10), 20, pp. 629-632.

1940a. Ann. Mag. Nat. Hist. (11), 5, pp. 257-274, figs. 1-3.
Parker, H. W. ; Moreatt, R. E. and Pakenham, R. H. W.

1940. Ann. Mag. Nat. Hist. (11), 5, pp. 309-314.
Peracca, M. G.

1896. Boll. Mus. Zool. Torino, 11, No. 255, pp. 1-4, figs. -

1909. in Abruzzi, Due D', "II Ruwenzori. Parte Scientifica." (Milano),
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1910. Boll. Mus. Zool. Torino, 25, No. 624, pp. 1-6.
1912. Museo Zool. Univ. Ann. Napoli, 3, No. 25, pp. 1-8.

Peters, W. C. H.

1854. Monatsb. Akad. Wiss. Berlin, pp. 614-628.

1855. Arch. Naturg., 21, 1, pp. 43-58 (reprint of 1854).
1867b. Monatsb. Akad. Wiss. Berlin, pp. 234-237, pi. -
1870c. Monatsb. Akad. Wiss. Berlin, pp. 641-652, pis. i-ii.

244 bulletin: museum of comparative zoology

1875a. Monatsb. Akad. Wiss. Berlin, pp. 196-212, pis. i-iii.

1876a. Monatsb. Akad. Wiss. Berlin, pp. 117-123, pi. i.

1877c. Monatsb. Akad. Wiss. Berlin, pp. 611-621, pi. -

1880c. Monatsb. Akad. Wiss. Berlin, pp. 795-798, pi. -

1881d. Sitzb. Ges. Naturf. Freunde Berlin, pp. 147-150.

1882a. "Naturwissenschaftliche Reise nach Mossambique . . . Zoologie,

3, Amphibien." (Berlin), pp. i-xv + 1-191, pis. i-xxx.
1882d. Sitzb. Ges. Naturf. Freunde Berlin, pp. 127-129.

Pfeffer, G.

1893. Jahrb. Hamburg. Wiss. Anst., 10, pp. 71-105, pis. i-ii.

Pitman, C. R. S.

1934. "A Check List of Reptilia and Amphibia occurring and believed to
occur in Northern Rhodesia." in Pitman, "Report on a Faunal
Survey of Northern Rhodesia." (Livingstone), pp. i-xii + 1-500
+ Index i-xxxii, maps. A-K.

1936. Uganda Journal (Kampala), 4, pp. 41-61, pis. vi-vii 4- E-G.

1937. Uganda Journal (Kampala), 4, pp. 220-246, 319-349, pis. xi-xiii,
+ H-M.

1938a. Uganda Journal (Kampala), 5, pp. 160-244, pis. xv-xviii -f-R-W.
1938b. "A Guide to the Snakes of Uganda." (Kampala), pp. i-xxi +

1-382, pis. i-xxviii, col. pis. A-Q, diagrams I-II, maps. (Being a

reprint, repaged, of foregoing articles).

Power, J. H.

1927c. Trans. Roy. Soc. S. Africa, 14, pp. 405-422, fig. 1, pis. xviii-xxii,

map.
1931. Trans. Roy. Soc. S. Africa, 20, pp. 39-49, figs. 1-2.

1935. Proc. Zool. Soc. London, pp. 333-346, figs. 1-10, pi. i.

Prato, A. del

1893. "Le Raccolte zoologiche fatte nel Congo dal Cav. Giuseppe
Corona." (Parma; privately printed pamphlet), pp. 1-14.

Reichenow, A.

1874. Arch. Naturg., 40, 1, pp. 287-298, pi. ix.

Reinhardt, J. T.

1843. K. Danske Vidensk. Selsk. Afhandl., 10, pp. 233-277, pis. i-iii.
1860. Vidensk. Meddel. Naturhist. Foren Kjobenhavn, pp. 209-250,
pis. iii-iv.

ROCHEBRUNE, A. T. DE

1884a. "Faune de la Senegambie. Reptiles." (Paris), pp. 1-221, pis. i-xx
(Mostly ignored as utterly unreliable).

Rose, W.

1929. "Veld and Vlei." (Cape Town), pp. i-xxiii + 1-240, figs. -, photos.
1-125.

loveridge: African snakes 245

Roux, J.

1907a. Revue Suisse Zool. Geneve, 15, pp. 75-86.
1907c. Zool. Jahrb. Syst., 25, pp. 732-742, pi. xxvii.
1936. in Jeannel, R., "Mission Scientifique de l'Omo." 3, pp. 157-190,
figs. 1-4.

Sauvage, H. E.

1884b. Bull. Soc. Zool. France, 9, pp. 199-208, pi. vi.

SCHLEGEL, H.

1837. "Essai sur la Physionomie des Serpens." (Leide), 2, pp. i-xvi +
1-606, and Atlas with pis. i-xxi, maps 1-3.

Schmidt, K. P.

1923. Bull. Am. Mus. Nat. Hist., 49, pp. 1-148, figs. 1-15, pis. i-xxii,

maps 1-19.
1933. Ann. Carnegie Mus., 22, pp. 1-15, figs. 1-2, pis. i-ii.

Schneider, J. G.

1801. "Historiae Amphibiorum naturalis et literariae." (Jena), 2, pp.
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SCHOTJTEDEN, H.

1933. Revue Zool. Bot. Afr., 23, pp. 233-238.

SCHWETZ, J.

1934a. Revue Zool. Bot. Afr., 25, pp. 379-384.
1934b. Revue Zool. Bot. Afr., 26, Suppl., pp. 23-25.

SCLATER, W. L.

1898. Ann. S. African Mus., 1, pp. 95-108, pi. v.

SCORTECCI, G.

1928b. Atti. Soc. Ital. Sci. Nat., 67, pp. 290-339, figs. 1-8, pis. vii-ix.

1931c. Atti. Soc. Ital. Sci. Nat., 70, pp. 203-215.

1934c. Rassegna Economica delle Colonie, 12, Nos. 1-6, pp. 1-83, figs.

1-36.
1939a. Ann. Mus. Civ. Stor. Nat. Genova, 63, pp. 263-291.
1939c. "Gli Ofidi Velenosi dell' Africa." (Milano), pp. i-xvi -f 1-292, figs.

1-151.
1940a. in "Missione Biologica nel paese dei Borana Raccolte zoologiche."

(Roma), 2, pp. 125-150 ( = 5-30 in separates).
Seba, A.

1735. "Locupletissimi Rerum Naturalium Thesauri . . ." etc. (Amstel-

aedami), 2, pp. i-xxx + 1-154, pis. i-cxiv.

Sjostedt, Y.

1897. Kongl. Svenska Vet.-Akad. Handl., 23, Part 4, No. 2, pp. 1-36,
pis. i-iii.
Smith, A.

1838-. "Illustrations of the Zoology of South Africa, Reptiles."

1849. (London), pis. i-lxxviii, App., pp. 1-28.

246 bulletin: museum of comparative zoology

Sonnini de Mannoncourt, C. N. S. and Latreille, P. A.

1802. "Histoire naturelle des Reptiles." (Paris), 4, pp. 1-410.

Steindachner, F.

1867a. in "Reise der Osterreichischen Fregatte Novara . . . Zool." 1, pp.
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Stejneger, L.

1893b. Proc. U. S. Nat. Mus., 16, pp. 711-741.

Sternfeld, R.

1908. Sitzb. Ges. Naturf. Freunde Berlin, pp. 92-95.

1908a. Mitt. Zool. Mus. Berlin, 3, pp. 397-432.

1908b. Mitt. Zool. Mus. Berlin, 4, pp. 207-336.

1908c. Mitt. Zool. Mus. Berlin, 4, pp. 237-247.

1909a. "Die Schlangen Togos," in "Die Fauna der deutschen Kolonien,"
(Berlin), Ser. 2, part 1, pp. i-iv + 1-29, figs. 1-42, map.

1909b. "Die Schlangen Kameruns," in "Die Fauna der deutschen Kolo-
nien," (Berlin), Ser. 1, part 1, pp. i-iv + 1-28, figs. 1-34, map.

1910a. "Die Schlangen Deutsch-Ostafrikas," in "Die Fauna der deut-
schen Kolonien," (Berlin), Ser. 3, part 2, pp. i-iv + 1-47, figs.
1-54, map.

1910b. "Die Schlangen Deutsch-Sudwestafrikas," in "Die Fauna der
duetschen Kolonien." (Berlin), 4, part 1, pp. i-iv -f- 1-45, figs. 1-50,
map.

1910c. Mitt. Zool. Mus. Berlin, 5, pp. 51-56.

1911a. Sitzb. Ges. Naturf. Freunde Berlin, pp. 245-251.

1912b. Sitzb. Ges. Naturf. Freunde Berlin, pp. 384-388.

1912c. "Reptilia," in "Wiss. Ergeb. der Deutschen Zentral-Afrika-
Expedition 1907-1908." (Leipzig), 1913, 4, pp. 197-279, figs. 1-4,
pis. vi-ix.

1917. "Reptilia und Amphibia," in "Wiss. Ergeb. der Zweiter Deutschen
Zentral-Afrika-Expedition 1910-1911." (Leipzig, 1917), 1, pp.
407-510, pis. xxii-xxiv.

Theobald, W.

1868. "Catalogue of Reptiles in the Museum of the Asiatic Society of
Bengal." (Calcutta), pp. 1-88, App., i-iii, four pis.

Tornier, G.

1896. "Die Kriechthiere Deutsch-Ost-Afrikas . . ." (Berlin, reprinted
1897), pp. i-xiii + 1-164, figs. 1-11, pis. i-v.

1897. Arch. Naturg., 63, 1, pp. 63-66.
1901a. Zool. Jahrb. Syst., 14, pp. 85-86.
1901b. Zool. Anz., 24, pp. 61-66.

Uthmoller, W.

1937. Temminckia (Leiden), 2, pp. 97-134, maps.

1938. Zool. Anz., 124, pp. 41-48.

loveridge: African snakes 247

Wagler, J. G.

1830. "Naturliches System der Amphibien, . . ." (Monachii), pp. 1-354,
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Werner, F.

1895b. Verh. Zool-Bot. Ges. Wien, 45, pp. 190-194, pi. v.

1896. Verh. Zool.-Bot. Ges. Wien, 46, 361-364, pi. vi.

1897b. Verh. Zool.-Bot. Ges. Wien, 47, pp. 395-408.

1898. Jahrb. Abhand. Natur. Magdeburg, 1896-1897, pp. 139-148.

1898a. Verh. Zool.-Bot. Ges. Wien, 48, pp. 191-213. figs., -, pi. ii.

1899a. Verh. Zool.-Bot. Ges. Wien, 49, pp. 132-157.

1902a. Verh. Zool.-Bot. Ges. Wien, 52, pp. 332-348.

1913a. "Die Lurche und Kriechtiere," in Brehm, A. "Tierleben." (Leip-
zig), ed. 4, 5, pp. i-xvi + 1-598, figs., pis., maps.

1915c. "Reptilia und Amphibia," in Michaelsen, W. "Beitrage zur
Kenntnis der Land- und Susswasserfauna Deutsch-Siidwest-
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1919. Denks. Akad: Wiss. Wien, 96, pp. 437-509, pis. i-ii, map.

1923a. Arch. Naturg., 89, Abt. A, pp. 138-199, figs. 1-9.

1924b. Sitz. Akad. Wiss. Wien, 133, 1, pp. 29-56, figs. 1-9.

1925. Arch. Naturg., 1924, 90, Abt. A, pp. 108-166, figs. 1-17.

1926. Sitz. Akad. Wiss. Wien, 134, 1, pp. 243-257.
1929a. Zool. Jahrb. Syst., 57, pp. 1-481, figs. 1-48.

WlTTE, G. F. DE

1920a. Revue Zool. Afr., 8, pp. 58-66, figs.

1920b. Revue Zool. Afr., 8, pp. 273-282.

1927d. Revue Zool. Afr., 15, pp. 320-332.

1932b. Revue Zool. Bot. Afr., 22, Suppl., p. 14.

1933a. Ann. Soc. Sci. Bruxelles (B), 53, pp. 70-73, figs. 1-3.

1933J. Revue Zool. Bot. Afr., 24, Suppl., pp. 111-123, figs-

1933m. Ann. Mus. Congo Beige, Zool (1), 3, pp. 153-188, pis. v-xi.

Wyllie, E.

1927. S. African Journ. Nat. Hist., 6, p. 129.

Zenker, G.

1892. Mitt. Fors. Gel. Deutschen Schutzgeb., 5, pp. 181-184.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCV, No. 3

A REVISIONAL STUDY OF THE
FOSSIL SYKPHIDAE

By Frank Montgomery Hull
University of Mississippi

With Thirteen Plates

CAMBRIDGE, MASS., U.S.A.

PRINTED FOR THE MUSEUM

April, 1945

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIV, and Vol. XCV, No. 1, 2 and 3
have appeared and of the Memoirs, Vol. I to LV.

Publication of Memoirs ceased with Vol. LV.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCV, No. 3

A REVISIONAL STUDY OF THE
FOSSIL SYRPHIDAE

By Frank Montgomery Hull
University of Mississippi

With Thirteen Plates

CAMBRIDGE, MASS., U.SA.

PRINTED FOR THE MUSEUM

April, 1945

ZookiVjy

APK 25 1945

t/BRAHt-

No. 3. — A Revisional Study of the Fossil Syrphidae
By Frank Montgomery Hull

TABLE OF CONTENTS BW1f

Introduction and Acknowledgments 251

History of Previous Work 252

Sources of Present Material 253

Geological Distribution of the Syrphidae . . . . . 254

Systematic Descriptions 257

Subfamily Syrphinae 257

Subfamily Cheilosinae 273

Subfamily Chrysotoxinae 326

Subfamily Eumerinae 327

Subfamily Microdontinae 330

Subfamily Eristalinae 331

Subfamily Volucellinae 335

Subfamily Xylotinae 338

Syrphidae incertae sedis 342

Insects misidentified as Syrphidae 344

Conclusions 345

Bibliography 351

Index of genera and species 354

INTRODUCTION AND ACKNOWLEDGEMENTS

Although many fossil Syrphidae have been described during the
past century, no one has attempted a comprehensive treatment of
them. The specific details within this family are often obscure, and a
study of this kind is needed in order to relate the fossil species to the
Recent genera. This effort toward a revision of the fossil Syrphids
began with a study of the excellent collection in the Museum of Com-
parative Zoology at Harvard LTniversity. I am indebted to Dr. F. M.
Carpenter and Dr. C. T. Brues for the suggestion that a study of the
fossil Syrphids might be replete with interest, and furnish some insight
into the phylogenetic history of the group. I wish further to thank
Dr. Carpenter for neverfailing advice of many kinds, for much dis-
cussion and encouragement. Besides allowing me to examine the very
unusual collections of the Museum of Comparative Zoology, he has
secured the loan of a number of types, all of which made this study
possible. In addition, I am deeply indebted to Dr. C. T. Brues for
many kindnesses and much advice. To Professor Nathan Banks I owe
much thanks for helpful suggestions and for permitting me access to

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important collections in which I found representatives of fully half of
the living genera.

My thanks are due to Dr. E. A. Chapin of the U. S. National
Museum and the late Dr. F. E. Lutz of the American Museum, not
only for the loan of material but for placing facilities for study before
me in visits to their institutions; to Professor T. D. A. Cockerell of
the University of Colorado for the loan of types from the university
collections; and to Dr. A. L. Melander of the College of the City of
New York for the loan of material.

Early in this investigation it became apparent that it would be
advisable to study the amber Syrphidae in the university museum at
Konigsberg. This was made possible by a grant from the Penrose Fund
of the American Philosophical Society. Important collections con-
taining rare living genera were studied in Vienna, Berlin, Amsterdam
and London. Additional amber material was obtained in two institu-
tions in Berlin, and amber and other material studied from the British
Museum. I wish especially to thank Dr. K. Andree, Director of the
Geologisch-palaontologisches Institut und Bernsteinsammlung der
Albertus-Universittit of Konigsberg and his associate Dr. Otto Pratje
for kindnesses to me during my visit there and for loan of the material.
I wish also to thank the following individuals who materially aided
this study by the loan of specimens : Dr. W. Janensch of the Geologisch-
Palaontologisches Institut and Museum der Universitiit Berlin; Dr.
J. V. Wumdorf, Preussische Geologische Landesanstalt in Berlin; Mr.
T. H. Withers, Department of Geology, British Museum of Natural
History; and Dr. G. D. H. Carpenter, University Museum of Oxford
University, England.

PREVIOUS WORK ON FOSSIL SYRPHIDAE

The earliest reference to a fossil Syrphid fly in the literature is of
an unidentified species of Microdon listed by Serres in 1829 from the
beds of Aix in Provence, France. In 1837 Germar studied the "car-
bonum fossils" from the Oligocene of Bonn, including one poorly pre-
served Syrphid, which he called Hemophilus? primatius. Brodie
(1845-47) is erroneously credited by compilers with having figured
a Syrphid, which Giebel later (1856) described under the family
Muscidae. Weyenberg (1869) described a Syrphid (Cheilosia dubia)
which must unquestionably be deleted from the lists, since it is totally
unrecognizable.

In the middle of the last century three workers devoted much time

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to a serious study of fossil insects. The first of these was Hope, who
in 1845-47 listed many interesting dipterous genera from Aix in
Provence in France, including a specimen of Rhingia which has re-
posed in the Hope collection unnoticed for ninety years. This was
recently loaned to me by Dr. G. D. H. Carpenter of the University
Museum of Oxford. Oswald Heer, who published in 1849 his "Die
Insektenfauna der Tertiargebilde von Oeningen and Radoboj in
Croatiens," was the second student of the century to give much at-
tention to fossil insects. Herman Loew in 1850 was the first distin-
guished dipterist to become interested in fossil Syrphidae. He described
no species, but mentioned seven genera from Baltic amber. I was
fortunate in finding the material Loew studied in the British Museum,
though the Volucella, which Loew mentioned and which was the most
interesting genus among his material, was missing. It is possible that
he had the species I describe below as Ptilocephala volucelloides (sub-
family Volucellinae) from the collection of the University of Konigs-
berg. Carl von Heyden (1867) named several species of Syrphids which
were taken from the beds of Rott (Siebengebirge in Rheinland). I
found one of his types at the British Museum.

In 1877 Samuel Scudder began his notable investigations of the
fossil insects of the Green River Shales of Wyoming in this country.
These studies appeared at irregular intervals, until as late as 1890,
after which Scudder mentions no more Syrphids. Dr. Samuel Willis-
ton, about this time, examined certain other specimens of Scudder's
and tentatively assigned them to genera. I have confirmed his con-
clusions in all cases where he mentioned catalogue numbers that have
made possible the tracing of the material he handled.

Finally, mention should be made of the work of Professor T. D. A.
Cockerell, pioneer enthusiast in the modern study of fossil insects,
who has added more species to the list of fossil Syrphidae than any
previous writer.

SOURCES OF PRESENT MATERIAL

The specimens upon which this study of fossil Syrphid flies is based
are from ten different deposits, including Baltic amber. The Syrphidae
preserved in rock are from several collections, the most important of
which is in the Museum of Comparative Zoology, containing the
Scudder collection and the specimens mentioned and discussed by
Williston in his 'Synopsis' (1886). All but one of Scudder's five species
have been studied. His specimens were from the Green River shales

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and excepting one species in the National Museum, they are contained
in the collections of the Museum of Comparative Zoology. Specimens
in the American Museum have been examined and I have likewise
been fortunate in studying the types of Dr. Cockerell's fossil Syrphidae,
loaned by the University of Colorado Museum. It has therefore been
possible for me to see most of the Syrphids from North America. Only
one of the fifteen genera and only four of the twenty-five species have
not been seen at the time of writing. There are almost twice as many
species known from the North American beds as from all of the re-
maining beds of Europe, not including the specimens from amber.

I have been able to locate but few of the types of fossil Syrphidae
from European beds ; fortunately, most of these species were described
by two men, Heer and Heyden, and they have been carefully illus-
trated. These illustrations are not of the desirable type, but it is clear
that the fossils belong to the subfamily Syrphinae and to the group
Syrphus in the broad sense. For phylogenetic analysis this can serve
my needs. I have studied the types of Merodon germari Heyden from
Rott (British Museum of Natural History); Platycheirus infumatus
Heer from Radoboj (British Museum of Natural History); and
Rhingia Hope from Aix (Hope Museum, Oxford University).

Of the nine Baltic amber genera mentioned in literature, only seven
received specific names and therefore numbers, which made them
traceable. Of these seven genera I have been able to study five. One
of the two named specimens missing is believed to have been mislaid
at the University of Konigsberg with the Hymenoptera, having been
returned with this family by Dr. Cockerell. The fullest facilities were
extended to me at Konigsberg and I was allowed to examine their
collection of amber Diptera, which included some ten or twelve
thousand specimens and which yielded a total of one hundred and
nineteen specimens of Syrphids. Several other collections of amber
have yielded new species or old types, chiefly those of the Geologisch-
Palaontologisches Institut und Museum der Universitat Berlin, the
Preussische Geologische Landesanstalt Museum Berlin, and the British
Museum of Natural History. All of these amber collections have
furnished a total of twenty new species, making a total of nineteen
genera and subgenera and twenty-nine species known from amber.

THE GEOLOGICAL DISTRIBUTION OF THE SYRPHIDAE

Eleven formations, including Baltic amber, have yielded Syrphids;
seven of these are in Europe and three are in the United States.

hull: revisional study of fossil syrphidae 255

(1) Florissant Shales, Colorado (Miocene)

These shales have yielded an enormous number of beautiful fossils,
among which are species of Syrphidae. The fossiliferous character
of these shales was discovered by A. C. Peale in 1876. Carpenter (1930)
states that only the upper part of the formation bears fossils, and that
the preservation was due to the entombing action of volcanic silt and
ash. The flora associated with these insects was composed largely of
deciduous shrubs and trees and was similar in many ways to that of
the upper Cretaceous, with the addition of such genera as Alnus, Aster,
Fraxinm, Populvs, Rosa, Rhus, Salix and Ulmus. These are all modern
genera of plants and a certain number of well developed flowers are
present. Rosa, Rhus, and Salix present today quite an attraction to
Syrphid flies. The climate must have been warm, for magnolias were
present. Previous writers have noted the presence of tropical elements
in the Florissant fauna, such as Glossina.

(2) Creede Shales, Colorado (Miocene)

This rock seems to be a mixture of very fine sand and clay; it had
the same origin as the Florissant shales. Only one Syrphid has been
taken from this bed thus far. The flora has not yet been fully studied.

(3) Green River Shales, Colorado, Wyoming, Utah (Eocene)

This is the only Eocene bed in this country which has yielded
Syrphid flies. It extends over portions of Colorado, Wyoming, and
Utah, and has yielded nine species of Syrphidae. The biota includes
a very considerable flora and a large number of insects, as well as fish.
Although originally regarded as a gigantic lake bed covering thou-
sands of square miles, the deposit is now thought to have been formed
by series of numerous small lakes, which were at times saline rather
than fresh (Cockerell 1926, Carpenter 1930). As far as can be ascer-
tained, the climate was hot and probably humid. Palms are common
in the shales, as well as Fulgorids, which are not abundant in tem-
perate regions.

(4) Oeningen Shales, Germany (Miocene)

This is an old and famous collecting ground for fossil insects, con-
sisting of mixed marls and limestone, mostly thinly laminated; and
considered to have been spring-fed lake deposits. Two quarries have

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yielded most of the specimens, which include two Syrphid flies. Heer
(1849) made a very careful study of the Oeningen fossils.

(5) Marls of Radoboj, Croatia (Miocene)

These deposits are just east of the top of the Adriatic Sea. Unger
was the first to give serious attention to their fossils. His work was
followed by that of von Charpentier and by that of Heer in 1865. A
few Syrphids are included among the three hundred insects described.

/ (6) Lignite Beds of Rott (Siebengebirge at Bonn, Germany),

(Oligocene)

Three species of Syrphidae have been described from these beds.
One of these belongs to Merodon, a genus which prefers warm climates
and whose home is now upon the Mediterranean shores; it confirms
the conclusions drawn respecting the climate in Miocene times, which
were based upon other species from these beds (Goss, 1878). The
authority upon the insects of this region was Carl von Hey den.

(7) Beds of Aix in Provence, France (Oligocene)

This formation, once considered Eocene but now known to be upper
Oligocene, is stated by Goss (1878) to include the richest beds of its
period on the continent. Marcel de Serres (1829), who was the first
to discover insects here, found some eight genera, including two
Syrphids, both of great interest. Among later students were Murchison
and Lyell (1829); Bonn (1851-6) and the Reverend F. W. Hope
(1845-47). In studying the climate in which the biota existed, Saporta
(1872) found the flora to be even more southerly than the fauna, a
peculiarity which we have noticed for other geological formations in
which insects occur. Microdon which occurs in these beds is a char-
acteristically tropical genus.

(8) Baltic Amber, Germany (Oligocene)

The Baltic amber is one of our richest sources of fossil insects, and
I have been able to study more than a hundred and fifty Syrphid in-
clusions. Several writers have discussed the environment in which the
insects originally lived; the abundance of Psychodidae, Tipulidae,
Mycetophilidae, Empididae, and bark-loving Dolichopodids indicate
a densely shaded environment.

hull: revisional study of fossil syrphidae 257

(9-11) Miscellaneous Occurrences of Fossil Syrphids

One species of a Syrphid was described by Foerster (1891) from
Brunstaat in Alsace (middle Oligocene), and another by Stackelberg
(1925) from the Miocene of Caucasia. In addition, a Syrphid has been
mentioned by Theobald (1934) from Camoins: (Oligocene).

SYSTEMATIC DESCRIPTIONS
Subfamily SYRPHINAE

Syrphus is geologically the oldest Syrphid genus, as far as known.
This early occurrence of Syrphus agrees with the present concept as to
the more generalized of the fourteen subfamilies, which places Syrph-
inae at the bottom. Three species of the genus are known from the
Eocene and all of them certainly appear to belong properly within
that genus, at least in the wide sense. It is impossible to determine
into which of the existing subgenera they should go, if indeed to any
of them.

Pongracz re-allocated two species formerly described in Syrphus to
the genus Platycheirus. He does so on the basis of the wings, but I
believe that this must remain a guess on his part, however shrewd,
as does my own somewhat conjectural assignment of one species (per-
sistens) to Platycheirus, until the fore legs are better known from other
specimens. Platycheirus rests solely on the dilation of the fore tibiae
and tarsi in the male. I assign my species (persistent) to Platycheirus
on the form of the tarsi and also upon the basis of the abdominal pat-
tern, which is not always reliable, but sometimes a good indication.
Its tarsi appear to be modified. The abdominal pattern is very con-
stant in many Syrphid genera, and in persistens it is certainly more
like that of Platycheirus than of Syrphus. The species (quadrata)
Scudder described as a Milesia is obviously out of place and I assign
it provisionally to Asarcin'a, an existing African genus. It surely be-
longs in the Syrphinae, and the pattern is very much like that of
Asarcina today.

These are the only genera in this subfamily represented by fossils.
It is therefore possible that much the greater part of the Recent
genera in Syrphinae are merely subgenera and have evolved in some-
what more recent times.

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Syrphus aphidopsidis Cockerell
Plate 5, fig. 25
Ann. Ent. Soc. Amer., 2: 253, fig. 4. (1909)

Female. Length 14.0 mm.; length of wing 9.2 mm. Head: is broadly
rounded, and the curvature behind suggests that the head was nar-
rowly concave in posterior vertical profile. The head is unusually well
preserved, each facet visible, though parts of the surface are cracked
away. Front broad, about two-fifths of the head width from the ver-
tical aspect. Thorax: and scutellum quite dark and black, the rim of
the latter appears more or less truncate apically, but was probably
semicircular instead, since the pigment of the abdomen was as dense
and dark as that of the scutellum; the demarcation is indistinct.
Abdomen: of this species is elongate and there is a pattern of transverse
bands completely interrupted in the middle but least interrupted on
the third segment; all of them are, therefore, broken up into spots.
The bands are basal in position on the segments, and in extent along
the lateral margin occupy about two-fifths of the length of the seg-
ments. Those on the fifth segment very much smaller. These spots
are roughly triangular in shape, with their posteromedial borders some-
what convex. Abdominal pile setaceous. Legs: very little of the legs
shows in the specimen; the hind femora were short and a little thick-
ened, and their base apparently pale in color. Wings: are noteworthy
for the heavy vena spuria and the exceedingly sigmoid apical cross
vein, joining the third vein barely at right angles, some little way from
the tip of the wing. Third vein ending with the costa quite at the tip
of the wing.

The following description was taken from the obverse, which is in
the British Museum of Natural History :

Head: eyes dichoptic with posterior margins broadly rounded, the
facets not enlarged above. Front long, with a dark colored lunule
above the antennal base; first and second antennal joints short, sub-
equal, the third about as long as second, but larger and deeper; round
in shape. The arista slender, basally thickened, about as long as the
antennae. The ocelli do not show. Thorax: is confused by the presence
of legs, the specimen being preserved by a ventral posterior. Details
cannot be satisfactorily ascertained. Abdomen: the abdominal sternites
appear as dark patches on the third, fourth and fifth segments, with
bristle pairs short and sharp, all directed posteriorly. These begin as
a narrow strip on the second segment, widened to a truncated wedge
on third and becomes a broad trapezoidal area on the fourth segment.

hull: revisional study of fossil syrphidae 259

Legs: hind femora rather thick, but uniformly so; thick for a member
of the genus Syrphus ; covered on their basal and lateral surfaces for
the whole length with short sharp bristles, which are not setigerous.
Hind tibiae slightly curved and also rather thickly covered with sim-
ilar, short, thick bristles. Wings: only the base of the costa of the wing
with spinules or short bristles. The remainder are too minute to be
readily observed. Costa ending at tip of wing just past the union with
the third longitudinal vein. Third longitudinal vein straight.

Locality: Florissant, Colorado. Horizon: Miocene.

Type: reverse in University of Colorado Museum (No. 8566); ob-
verse in British Museum.

Syrphus willistoni Cockerell
Plate 3, fig. A, C; Plate 4, fig. 6, 15
Bull. Amer. Mus. Nat. Hist., 26: 9. PI. I. (1909)

Male and female. Length 11.3-6.4 mm.; abdomen and scutellum
6.8-3.8 mm.; wing length 8.2-5.3 mm.; abdominal width 3.2-2.0 mm.;
these measurements are taken from a series of fourteen specimens, the
numbers of which are noted below.

The agreement of the respective parts is remarkably good. It is not
unusual to find a hundred percent size variation or over in Syrphids.

Head: little detail shows, although on occasional patches facets can
be seen. I am able to determine that the supraocular facets were en-
larged. Antennae short. The third joint appears to be about one and
one-half times as long as wide, to be markedly rounded off dorsally and
distally so as to come to an obtuse point ventrally. The arista is
slightly thickened and held erect. It is but little longer than the third
joint. I can make out setae upon the end of the first joint, but not
upon the second, though they were undoubtedly present. In the
female, the eyes are separated at vertex by about .5 mm. and thence
diverge gradually down the front. Thorax: dark in color. The pile is
not apparent; it was presumably delicate. The scutellum is two and
a fifth times as wide as long with even rim and no trace of bristles or
pile. Abdomen: five segments and the hypopygium of the male are
visible. The first is practically covered by the scutellum; the second,
third and fourth are nearly equal in length and not greatly differing
in width. The second and fourth about the same, the third being
slightly wider. The fifth is half the length of the fourth and is narrower.
Second, third and fourth segments marked with a large yellow spot on
each side, reaching the margin in at least two-thirds of its full length,

260 bulletin: museum of comparative zoology

and separated in the middle, by a complete median band, which is
parallel-sided, and progressively wider from fourth to second segment.
The post median corner of each spot is rounded, the anterior median
corners practically square. The fifth segment is clear anteriorly, but
in the type specimen there is abundant indication that the posterior
half was dark. The whole abdomen with the exception of the anterior
part of the second segment, where it thins considerably, is covered
with abundant, beautifully regular, short, but not unusually short,
stiff, sharp tipped hairs. They grow more numerous on the fifth seg-
ment. It might be described as stiff pile. Number 3955 is an exception
in that the first pair of spots do not reach the margin. Numbers 3961,
3968, 3966 and 3963 show an extra wide abdomen with consequent
alteration in shape of the spots. In view of varying pressures, I do not
believe these differences are important specifically. Legs: unfortu-
nately little is shown. The tarsi appear to be enlarged in No. 3954.
The hind femora are slightly thickened. Wings: of the typical Syrphus
type. The third longitudinal vein is ever so barely sinuous. The first
longitudinal vein again approaches the costa insensibly, but there is
no question of the submarginal cell; it is open. The spurious vein is
quite distinct and runs nearly to the union of fourth longitudinal vein
and postical cross vein. The second longitudinal vein joins costa at an
angle of about fifty-five degrees. The costa and third longitudinal
vein end together at tip of wing.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: Amer. Mus. Nat. Hist. Allotype: No. 3954 M.C.Z.
Additional specimens: Numbers 3955, 3956, 3957, 3958, 3959, 3960,
3961, 3963, 3964, 3965, 3966, 3967, 3968; all Museum of Comparative
Zoology.

Handsome flies with large clear fenestra, or spots upon the abdomen,
which were doubtless yellow in life. These spots are almost rectangular.
The species belongs to that large class of fossil Syrphus with the bands
medially divided.

Syrphus hendersoni James

Canad. Ent., 64: 265 fig. (1932)

This species is unfortunately based merely upon an abdomen. The
pattern of this abdomen is striking, with four segments and perhaps
a fifth visible. The outline of the abdomen is broad and the segments
are somewhat detached from one another. The length is about half
again as long as the width. The spots of the first segment are small,

hull: revisional study of fossil syrphidae 261

oval, well-removed from any of the borders of the segment, widely
separated in the middle and somewhat pointed towards the antero-
lateral corner. Those of the third segment represent narrow lateral
bands continuous basally only for a short distance beginning at the
margin, and strongly bulging posteriorly in somewhat abrupt fashion
after having run with somewhat uniform width for almost half their
total width. The medial portion of these spots might be looked upon
as separate, large, rounded medio-basally squared spots which are
confluent with the narrow basal and lateral spots. Spots of the fourth
segment similar except that the medial rounded portion is smaller,
more widely separated and the neck representing the confluence is
narrower. The pile of the abdomen is short and delicate, but is a little
stiffer in each posterior lateral corner along the posterior border for a
short distance.

Locality : Green River, Colorado, Station 25 Kimball Creek. Hori-
zon: Eocene.

Holotype: No. 15652. University of Colorado Museum.

Contrary to the remarks of James, the pattern of this species is not
greatly different from that of certain species of western Syrphus, such
as amalopis O. S.

Syrphus platychiralis spec. nov.

Sex indeterminate. Length for the two specimens 7.0 mm. and 8.0
mm.; length of scutellum and abdomen 4.4 mm. and 5.0 mm.; for
wing 5.6 mm. and 5.8 mm.; width of abdomen 1.3 mm. and 1.6 mm.
Head: very little detail shows. The antennae were short, with large
third joint and a moderately short arista, remarkably thickened, grad-
ually and evenly, on the basal half. The third appears to have been
obtusely pointed ventro-distally. Thorax: dark in color. No detail of
pile apparent. Scutellum large, one and one-fifth times as wide as
long, and somewhat squared in outline. Margin simple. Abdomen:
slender in form, pale in color with dark posterior cross bands on the
segments and each pale band interrupted or subinterrupted medially.
The second, third, and fourth segments are nearly equal in length.
The fifth is nearly as long as the fourth, but narrower. Hypopygium
quite visible and rounded. The median band is widest and complete
on the second segment, on the third it is quite narrow and appears to
dwindle so it barely reaches the anterior border. The same is true of
the fourth segment. The following segment clear. Pile stiff, bristle-
like, much as in Syrphus wiUistoni Cockerell, but on the posterior seg-

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merits it fails to become condensed and superabundant. Legs: hind
femora decidedly strengthened without being greatly enlarged; about
one and a half times or more as thick as the hind tibiae. Both are
everywhere covered with appressed, close-set, short bristle-like pile,
running in the appropriate direction. The varying vortices of the pile
are easily discernible and it is remarkable that it is continuously of
the short stiff type. Hind femora and tibiae dark; the former appear
to have been pale basally. Wifigs: typically Syrphus-like. I am unable
to detect significant differences from those of Syrphus willistoni
Cockerell, although the marginal cell appears to be more distinctively
open.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: No. 3926. M.C.Z. (Scudder collection). Paratype: No.
3927. M.C.Z. (Scudder collection).

This species, while close to Syrphus willistoni, appears distinct in its
more slender abdomen, semi-interrupted bands of the abdomen, and
differences in pile.

Syrphus carpenteri spec. nov.

Plate 1, fig. B; Text-figure 1.

Female. Length 5.7 mm. (6.2 mm. including antennae); head 1.5
mm.; abdomen and scutellum 2.8 mm.; wing 4.6 mm.; width of wing
1.8 mm. Head: presented from the dorsal aspect, the eyes are widely
separated, almost every facet standing out clearly. The ocelli are very
clear, forming an obtuse triangle, the posterior pair being one and a
half times farther apart than each of them from the anterior one.
Upper anterior facets slightly enlarged. The antennae are longer than
broad and appear slender, but since they are not large this does not
make them appear long. Third joint about equal to first and second
in length; perhaps a little shorter. Thorax: dark in color — no pile
shown. Scutellum short, about two and fifth times as broad as long;
evenly rounded with simple margin and no trace of bristles. Abdomen:
four segments and a trace of fifth visible. First segment dark in color.
Remainder of the abdomen with large, quadrate, pale spots, about
one fifth of each segment being devoted to dark posterior segmental
border, and a progressively narrower median dark stripe, narrower
from fourth to second, where it is quite narrow and is about half as
wide as the posterior border of that segment. The median band of the
fourth segment is three times as wide as the median band of the second
segment. The median stripe reaches the anterior border of each seg-

hull: revisioistal study of fossil syrphidae

263

merit and the lateral borders of each segment are dark, though very
narrowly so in the middle of the second segment. The basi-median
corner of the spots is only slightly convex and the anterior median
corners of second and third quite square; on the fourth segment the
median stripe widens as it approaches the anterior border of the seg-
ment. The abdomen of this species is thus seen to be rather wide, with
unusually large subquadrate spots upon it. But the size of the fly is
small. Abdomen covered with short thick bristly hair, but not notice-

Fig. 1

ably thicker on the posterior segments. Legs: but little shows. Hind
femora slightly thickened. Wings: well preserved. Curious, short and
rather brbad. Costa microsetose; spurious vein very distinct, reaching
nearly to fusion of fourth longitudinal vein and postical cross vein.
Third longitudinal vein nearly straight, what curve it has is not a
sinuosity, but a very low gentle arch over its whole length. It joins
costa at tip of wing. Costa stops just a little beyond. Marginal cell
broadly open and second longitudinal vein joins costa at an angle of
about thirty to thirty-five degrees. First longitudinal vein unusually
heavy. Subapical cross vein not sinuous, nor sigmoid, but angulate,
a trace of spur directed inward.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: No. 3928. M.C.Z. (Scudder collection).

I take great pleasure in naming this interesting species for Dr.
Carpenter, to whom I owe much for his encouragement in the study
of these flies.

264 bulletin: museum of comparative zoology

Syrphus eocenicus Cockerell
Proc. U. S. Nat. Mus., 64: 7. PL 2., fig. 2. (1924)

"Robust, 12 mm. long; abdomen 7 mm. long and 4.5 mm. wide;
head and thorax black, the scutellum not pallid as in modern Syrphus,
but appearing a shade lighter than the abdominal bands; abdomen
with very well-defined markings, the three broad light bands all
interrupted by a narrow median band, somewhat broader, but still
very narrow, on first segment; the light bands reach the extreme
margins, and of approximately equal width, the first however con-
spicuously wider mesally than the others, the upper edge of the second
dark band presenting a double curve on each side ; the other dark bands
also have a double curve on each side, but not sufficiently to make the
light bands appear arcuate; the third light band is much wider at the
sides than mesally; there is a fourth light band, broadly interrupted
mesally, its inner ends pointed. Wings not preserved."

Locality: Green River, Wyoming (Station above head of the Ute
trail above Sellers Ranch, Roan Mountains, Colo., July 1922). Hori-
zon: Eocene.

Holotype: 69179 U.S.N.M. (Not seen).

"A beautiful specimen, although lacking the wings. It appears to be
close to S. umbellatorum Schiner, but considerably larger, with dark
scutellum. It is much larger than S. lithaphidis Cockerell, found fossil
in the Eocene of Cathedral Bluffs."

The above description is that of Cockerell. On my visit to the
National Museum the type could not be located. This is a clearly
marked and valid species, which should be easily recognized.

Syrphus lithaphidis Cockerell

Proc. U. S. Nat. Mus., 57: 253. (1921)

"Length, about 8.2 mm.; head and thorax 4 mm.; length of wing
6.8 mm. Head and thorax fuscous (doubtless black in life); wings
hyaline; abdomen pallid with broad dark bands on hand margins of
segments, and a broad dark median band, evanescent on the apical
half. Venation in general as in modern Syrphus, except that the sub-
costal cell is not nearly so slenderly tapering at the apex, the end of the
first vein being somewhat like that figured by Williston for Paragus
tibialis, though this does not agree well with an actual specimen of
P. tibialis before me. There is a distinct, though not dark cloud filling
the apical part of the subcostal cell, as in modern Syrphus. The wing

hull: revisional study of fossil syrphidae 265

measurements are as follows: end of auxiliary vein to end of first about
1. 6 submarginal cell on first basal about .8 mm.; last posterior on
second basal about .32 mm. ; tip of anal to wing margin about .24 mm.
The general form and appearance entirely agree with Syrphus.

Locality: "Cathedral Bluffs South of Little Tommies Draw at a
point where samples were taken. Colorado. (Winchester 17-5)."
Horizon: Eocene.

Holotype: 66585 U.S.N.M. (Not seen).

"In the markings of the abdomen, this closely resembles S. willistoni
Cockerell."

The description of this species is very inadequate and would prob-
ably not enable its recognition, although there is no doubt that it is
related to Syrphus willistoni. The type could not be found.

Syrphus euphemus Hey den
Palaeontographica, 17: 262. PI. 45, fig. 29 (1850)

The description given by the author presents the following points of
interest and value.

Length 7 mm. Head: large, posteriorly truncate, strongly bulging
towards the front. The eyes are holoptic and large covering the entire
upper surface of the head. Thorax: is much longer than broad, and
broadest posteriorly; the scutellum is large, its margin rounded and
entire. The author states that upon head, thorax and scutellum one
can clearly see a light greenish-brassy color. Abdomen: has five seg-
ments; it was somewhat longer than the thorax and broadest in the
middle; the three first segments were about equally long, the fourth
shorter, the fifth smaller and posteriorly rounded. The color of the
abdomen was chiefly yellow; upon the second, third and fourth seg-
ments and perhaps the first there was a rather broad, sharply bordered
black fascia; all the segments have likewise a black, somewhat smaller,
sharply bordered posterior margin. Wings: the first longitudinal vein
was branched. The author states that the third longitudinal vein arose
from the second beneath the branch of the first with a rather strong
deflection near the base of the wing and went without branching to the
tip of the wing. He notes the presence of the usual oblique cross vein
through which the vena spuria runs parallel to the fourth longitudinal
vein. The first posterior marginal cell is closed and moderately stalked.

Locality: Rott, Siebengebirge at Bonn, Germany. Horizon: Oligo-
cene.

266 bulletin: museum of comparative zoology

Type: One male originally in the Krantz collection. I was not able
to locate it.

The species may be known by the narrow, black margins on the
posterior borders of the abdominal segments and by the broad, com-
plete, medial fascia which divides the pale markings in half.

Syrphus freyeri Heer
Ins. Oen., 2: 244. PI. 17. figs. 12, 12a, 12b. (1849)

The description and illustration given by the author present the
following points of interest and value.

Length without head 6.5 mm.; of thorax 2.5 mm.; its breadth
3.6 mm.; of abdomen 4 mm.; its breadth 2 mm.; of wing 7.5 mm.
The head is wanting. The thorax is oval and shining black; the
scutellum similarly colored. The wings project out over the abdomen
and their venation is clear and similar to Syrphus. The abdomen is
elongate oval and shining coal black; the first segment cannot be
discerned; the second is longer than the third, the fourth much shorter,
the fifth still shorter. Each segment excepting the first has on each
side a light colored spot. They approach one another on the dorsum
as far as the black line and thus form an unbroken fascia. On the
second segment are two almost triangular spots whose more acute
angle lies somewhat medialward; upon the second and third segments
the large spots extend to the basal margins of the segments. These
spots widen towards the middle of the abdominal segments and
leave their posterior margins arcuate. The fourth pair of spots is
much smaller, they are oval and lie upon the short fifth segment.
The legs are not sufficiently well preserved to furnish data. The
wings project out over the abdomen and their venation is clear and
similar to Syrphus.

Locality: Radoboj, Hungary. Horizon: Miocene.

Type: Two specimens in the k.k. montan. Museum zu Wien. Heer
says that one specimen is contained upon a slab with other flies such
as Oedipoda medlanosticta, Limnobia vetusta, etc. I was not able to
locate the types.

Heer says that the fly is closest related to Melanostoma mellinum,
which has a widespread range over the northern hemisphere. It agrees
in size, in the glossy, dark colored thorax and spotted maculation of
the abdomen with the male of M. mellinum. It is unlike that species
in the almost triangular spots of the second abdominal segments, in

hull: revisional study of fossil syrphidae 267

the spots of the fifth segment. He compares it to Platycheirus scutatus
Meig., which however has quadrangular, pale abdominal spots. This
species may be easily recognized by the large, round spots on each
side of the third and fourth segments, and the similar pair of sub-
triangular spots on the second segment.

Syrphus geminatus Heer
Ins. Oen., 2: 245. PI. 17, fig. 13. (1849)

Length of abdomen 5 mm. ; breadth 3 mm. Head: and part of the
thorax are destroyed, only the posterior part of the latter is preserved;
it appears to have been black; likewise the scutellum. Abdomen:
elongate, oval; the first segment very short and dark, the following
three more or less the same length; they are entirely light colored and
were in life probably white or light yellow. The anterior border of
each segment has a clearly contrasted, narrow, linear crossband of
black. The posterior border of each segment is light colored, but
immediately before that there is a second linear, black crossband;
this band lies next to the anterior one of the following segment and
borders the yellow posterior border of each segment. Thus there are
two black cross lines that closely approach one another and which
encloses a light one band formed by the posterior margin of the
segment. The fifth segment is distinctly smaller toward the end and
on the posterior margin only a dark spot is visible. The very short
sixth segment appears to have been light colored. The wings are for
the most part destroyed; yet on the left wing enough of the veins is
preserved so that one can recognize its position in the genus Syrphus.

It belongs near the group of Syrphus balteatus F. (Europe), S. nec-
tarinus Wied. (China) and S. alternans Macq. (from Coromandel),
differing, however, from all of these by the fact that the middle of the
first and second segments are not black, and that always each segment
on the posterior border is light ; the anterior border, however, is black,
while in that one the anterior border is yellow, on the posterior border
black. (Rewritten from original.)

This species is easily recognized by the very narrow complete black
bands just before the posterior margin of the second, third and fourth
segments, as well as similar ones on the anterior margins of the third,
fourth and fifth segments, together with the fact that none of these
bands are interrupted.

Locality: Radoboj, Hungary. Horizon: Miocene.

268 bulletin: museum of comparative zoology

Types: two specimens in the K. K. Montan. Museum at Vienna.
In my visit to the Museum, which Heer states is the repository for
these specimens, I was not able to locate the types.

Syrphus reciprocus Foerster
Abh. Geol. Spezialk. Els., 3, no. (5): 486. PI. 14, fig. 28. (1891)

Female. Whole length 6.5 mm. Head: the anterior side of the
head with its proboscis is turned backwards, due to pressure. The
large eyes do not touch, so this specimen is apparently a female.
Only the basal segment of the antennae is preserved. Abdomen: this
shows principally the ventral side with the sockets of the legs; on the
dorsum the large middle segment with the scutellum is seen from the
side. It is 4 mm. long and 2 mm. broad and consists of six segments.
Legs: only a few remnants are preserved. (Rewritten from original.)

Locality: Brunnstaat, Alsace. Horizon: Middle Oligocene.

Type: I have not been able to find the type.

Foerster states that Syrphus ewphemus Heyden from Rott has the
same size, but is more slender. From the figure it seems characterized
by a wholly pale abdomen.

Platycheirus haidingeri Heer

Heer, Ins. Oen., 2: 243 PI. 17, fig. 11 (1849). Pongracz, Ann. Mus. Nat-
Hungarici, 25: 188 (1928)

Male. Length 13 mm.; length of wings 12 mm.; length of abdomen
8 mm.; width of abdomen 4 mm. Head: largely destroyed, but more
or less hemispherical. Eye facets plainly distinct, somewhat enlarged
above. Head appears to have been a little wider than humeri. An-
tennae do not show. Thorax: dark in color, with evidence of some long
hairs on the side. Scutellum small, margin evenly rounded, about one
and one-half times as wide as long. Abdomen: broad, quite flat, with
distinctly emarginate sides and sparse, quite fine setigerous or bristle-
like pile, that reaches over the segmental margins, but which is not
concentrated along the posterior margins. Pile appressed. Abdomen
dark in color. Pattern apparently consisting of narrow lunules, meet-
ing or practically meeting the mid-line but not reaching the lateral
margin. Present on second and third segments. Hypopygium present.
Legs: details of the legs are not plain, except that on what appears to
have been the mid-femora, the bristles are sparse, long, slender.
Wings: well preserved, on both sides the basal one-half with very

hull: revisional study of fossil syrphidae 269

sharp outwardly depressed short bristles. Third longitudinal vein
turning down slightly opposite the middle of the subapical cross vein.
Thus very slightly convex instead of concave. Submarginal cell open,
color of veins brown, pubescence of wings thick, and well preserved.
Spurious vein conspicuous, very apparently not heavily chitinized.
Posterior margin of wing with distinct micro-nodules on margin of
wing.

Locality: Oeningen, Germany. Horizon: Miocene.

Type: I have no information as to the place of deposit of the type.
I studied a specimen in the British Museum which may have been the
type and on which my description is based.

Platycheirus persistens Hull
Plate 4, fig. 4-5
Psyche, 45: 116. Fig. 3 (1938)

Male. Length 10 mm.; of abdomen and scutellum 6 mm.; of wing
7.2 mm.; second specimen, length 10; thorax and abdomen 6.2 mm.;
of wing 3.8 mm. Head: hemispherical, obviously narrower than thorax.
Eyes narrowly dichoptic. Face dark in color. No details of antennae
visible. Thorax: dark, though very little pigment is preserved, and no
details of pile can be seen. Scutellum semicircular, the margin evenly
convex, the width about one and three fourths greater than the length.
Abdomen: slender, the sides not quite parallel, but slightly convex,
leaving the middle segments barely wider. The first segment juts
beyond the rim of the scutellum by a fifth the scutellum's length.
Second and third segments of nearly equal length, the former longer.
Fourth segment slightly shorter than third. Fifth segment two-fifths
as long as the preceding one. Hypopygium prominent and smoothly
rounded. The segments are marked with brown. The posterior two-
fifths of the second segment with a median wedge, pointing to and
reaching the anterior border, and similar pattern on the two succeeding
segments, the brown of the posterior border on the fourth segment
occupying nearly the whole of the posterior half. The fifth segment is
clear. Legs: slender. For the most part, they are not well preserved,
but one pair of tarsi, apparently the left hind tarsi, are well preserved
and shows decided expansion and thickening of the joints. Wings:
poorly preserved.

Female. Specimen No. 3950 (3951) is without head. The wings
are a little better preserved and show the third longitudinal vein and

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costa ending quite beyond the tip of wing, though not nearly as much
as in Rhingia. The abdominal pattern is quite similar; beyond the
fact that the segments are slightly wider, I am unable to detect dif-
ferences of importance. One whole hind leg (right) is preserved. The
femora were slightly thickened one and two-fifths the width of tibiae
and the tarsi were not dilated. The obverse is fragmentary and poor.
Perhaps a trace of antennae appears upon it. The opposite hind tarsus
is shown, the maculation is deceptive and the abdomen also appears
disproportionately short and wide.

Locality: Creede, Colorado. Horizon: Miocene.

Holotype: No. 3949M.C.Z. Allotype: No. 3950. M.C.Z. (obverse).
Paratype: No. 3951. M.C.Z. (reverse). All are poorly preserved.

Platycheirus infumatus Heer

Heer, Ins. Oen. 2: 246. PI. 17, fig. 14. (1849). Pongracz, Ann. Mus. Nat.
Hungarici, 25: 190, fig. 53 (1928)

I give below an analysis of this fly drawn from the original descrip-
tion and the illustration.

Length: without head 7.5 mm.; of wings 7.5 mm. Head: in the
illustration the head was missing. Thorax: largely destroyed, the
scutellum unrecognizable. Abdomen: well preserved but he finds that
the abdomen though slender and cylindrical does not have the nec-
essary basal contraction which though slight is characteristic of the
recent genus Doros. The second and third segments were of equal
width; the former segment pale in color with a suggestion of a black
middle line at the base. The third segment was quadrangular, black
in color with in the middle a narrow, light colored cross band; the
margins of this fascia were quite sharp and clear. The fourth segment
is of the same size as the third, is light colored in the specimen with
small black, posterior band. The fifth segment is much shorter and
light and beyond there an indistinct, small, contrasting terminal
segment that appears to have been darkly spotted upon the margin.
The entire abdomen densely set with very fine pile. Wings: well
preserved. Heer compares the species to Syrphus and Doros. He notes
the presence of an anterior dark border upon the wing from vena
scapularis to the wing tip, in which respect it is very suggestive of
Doros. Heer found the venation to be typical of Syrphus.

This species should be recognizable upon the basis of the parallel-
sided abdomen together with the brown wing border and the abdominal
pattern. The posterior half of the fourth segment entirely black. It

hull: revisional study of fossil syrphidae 271

is possible that this fly was an early type related to Doros which,
though a small genus, seems to be at home in Europe.

Locality: Radoboj, Croatia. Horizon: Miocene.

Types : Heer states that there are two specimens, one of these upon
the same piece of rock as Formica occulata and Myrmica minutula. I
was not able to examine the types, but Pongracz redescribed this insect
from a specimen in the Geologischen Bundesanstalt (Vienna).

Leucozona nigra spec, now

Plate 2, fig. B

Female. Length 9 mm. ; length of wing 6.2 mm. Head: large, about
as wide as thorax, hemispherical in shape, the antennae short, the
third joint of only moderate size, short, a little longer than broad,
the arista rather longer than the antennae and basally thickened. No
details of either head or thoracic pile can be seen. Thorax: short and
broad, about as broad as long, black in color. Scutellum semi-circular,
at least twice as wide as long; dark. Abdomen: robust, flattened, and
relatively broad. It is one and one-half times as long as wide and five
segments are visible. It appears to have been almost uniformly black,
but there is considerable evidence that the posterior half of the second
segment bore a wide pale medially interrupted transverse band. There
is a bare possibility that there were similarly light colored spots on the
remaining segment but here the specimen has more of the appearance
which might be expected if portions of the chitin had peeled away.
There is an odd shiny luster or vitreus appearance upon the abdomen,
thorax, and scutellum of this specimen which I have not noticed on
other fossil flies. The pile of the abdomen, while not excessively long
as in wooly flies, has the appearance of being longer than usual. Legs:
no details of the legs are apparent. Wings: very little longer than the
abdomen, beautifully preserved. Both the apical cross vein is confluent
with the third vein some way before the tip at a right angle, and the
preceding section of the third vein is slightly curved upward through-
out its whole length. Vena spuria quite distinct. Small cross vein
strongly oblique before the vena spuria and enters the discal cell a
little past that point which corresponds to a third of the way from the
base. The outward portion of the fourth vein which bends down to
join the lower marginal cross vein is very strongly bent indeed, and
this appears to be one of the principal characteristics of the wing. The
marginal cell is open and there is a large quadrate spot in the middle
of the anterior part of the wing.

272 bulletin: museum of comparative zoology

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: one female, coll. Am. Mus. Nat. Hist., No. 157a.

I think that there is no question of this fly being a member of
Leucozona, though the abdomen is not quite so long as in modern
species. The well preserved venation and the characteristic wing spot
make this an especially interesting fly.

Asarcina quadrata Scudder

Plate 2, fig. D

Scudder, Bull. U. S. Geol. Surv. Terr., 4: 752 (1878). Scudder, Tert. Ins.,
557. PL 9, fig. 13 (Milesia) (1890). Williston, Syn. N. A. Syrph., 281
(1886)

Sex indeterminate. Head: large, nearly as broad as the thorax, the
eyes large, the front quite large, prominent, half as broad as the head
and about half as long as broad. The head and thorax while present
show few details ; they were black in color. Thorax: barely longer than
wide and widest in the middle. Abdomen: well preserved and shows a
characteristic pattern. It was oval, of about the same width as the
thorax and its ratio of width to length is 9:13. The ratio of abdominal
length to thoracic length (including scutellum) is 14:10. The abdomen
was thin and flat, with the first segment black or brown and the re-
maining segments chiefly pale in color margined with narrow fascia
and vittae of black or brown as follows : second segment barely over
twice as wide as long, with a narrow, posterior, marginal black fascia ;
the segment is divided in the middle with a narrow black vitta that is
barely wider at its apical end and connection with the posterior fascia.
Anteriorly it expands just before it reaches the anterior border and
sends a slender, bordering fascia part of the way towards each side of
the base of the segment. The third segment has posterior black fascia
of about the same width as the previous one; it is barely wider in the
middle and there is no median vitta whatever; the remainder of the
segment is pale. The fourth segment is quite similar in pattern to the
third, the posterior fascia attenuated just before it reaches the poste-
rior lateral corners. Fifth segment with only a faint trace of a narrow
posterior fascia. The entire abdomen was thickly covered with short
setaceus pile. Wings: while present show very few details. They were
longer than the abdomen in a ratio of 14:20. Scudder states that the
third longitudinal vein originates from the second in the middle of the
wing, is very gently arcuate in its outer half and appears to terminate
just above the tip of the wing. He states further that the fourth longi-

hull: revisional study of fossil syrphidae 273

tudinal vein is united by an oblique cross-vein to the third very near
the origin of the latter, the marginal vein simple with the fourth longi-
tudinal vein bending down at its tip to meet it. I am not able to verify
these details because of past efforts to clear the wing.

There is no question of the subfamily relationships of this species.
It could not be a Milesia and the abdominal pattern is extremely close
to that typical for the large Ethiopian and Asiatic group of flies known
as Asarcina, which actually range into southern Europe. I think the
narrow fascia and broad flat abdomen place it with reasonable cer-
tainty in that genus. '

Locality: Green River, Wyoming. Horizon: Eocene.

Type (male) : no. 14691 in the U. S. N. Museum. Examined by me
in the original and from a photograph kindly furnished by Dr. E. A.
Chapin.

The Subfamily CHEILOSINAE

There are sixteen genera of this subfamily occurring as fossils. Ten
are extinct, and the majority of these are in the Baltic amber fauna
only. This is, I believe, explained if one constructs a map to show the
distribution of all Recent Cheilosia, for certainly in point of density of
species in terms of land area scarcely any Syrphid genus exceeds
Cheilosia. The genus is very abundant today in central Europe.
There are only thirty-six Recent genera and subgenera known today in
the Cheilosinae. Sixteen is also exactly the number of Recent Cheilo-
sene genera and subgenera known today from Europe, and there are
only nineteen from the United States, fifteen of which are the same as
European ones. I cannot avoid the conclusion that the Eristalinae and
the Volucellinae are now dominant groups, whereas formerly the
Cheilosinae were better developed.

It is appropriate to mention here that the Baltic amber Cheilosinae
are not quite like modern ones, though in a somewhat indefinite
fashion, and that the Myioleptas (valida excepted) are certainly not
strictly like the genotype (a European species).

Of these sixteen Cheilosine genera, possibly Eoxylota should go to
the Xylotinae, though how we would enlarge the concept of the latter
to hold it without danger to the former is not clear. I think we must
regard Eoxylota and Hemixylota (Chilean Recent relative) together
with the Myioleptas as definitely transitional between these two sub-
families. Since Myiolepta-\ike forms were so common in the Oligocene
and since there are three such genera from the shales of the Eocene and

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Miocene (Archalia, Cacogaster, Xylotosyrphus) and since one definite
Oligocene Xylotinae genus is found (Megaxylota) it seems reasonable to
conclude that we are witnessing in the array of Oligocene Syrphids
the origin of the Xylotine subfamily, both in point of facial type and
acquisition of megamorphic femora, femoral armament, and migration
of the small cross vein of the wing. Early in these studies I was aston-
ished to find many flies strongly favoring Xylotinae, but precluded
from there by the basal position of the small cross vein. I was there-
fore particularly pleased to discover Megaxylota in the amber, a true
member of the Xylotinae. A study of Xylotosyrphus will show that,
though the wing has altered, the typical Xylotine pattern of the abdo-
men has remained practically unchanged for millions of years. These
beautiful and exquisite tetramaculate species (Xylota, Planes) form
some of the most interesting components of our present Syrphid
fauna. Finally, there is a possibility that some present-day species of
Cheilosia can satisfactorily be allocated to Protorhingia. Possibly I am
wrong in assigning Doliomyia and Palaeopipiza to the Eumerinae as
early foreshadowing types of the modern genus Eumerus. If I am,
these two genera would return to the Cheilosinae.

Cheilosia ampla Scudder

Bull. U. S. Geol. Geogr. Surv. Terr., 4: 753 (1878).

Sex indeterminate. Specimen No. 5160: length 5.8 mm.; abdomen
and scutellum 3.2 mm.; of wing 5.8 mm. Specimen No. 5162: total
length 3.9 mm. (minus head); abdomen and scutellum 2.7 mm.; wing
missing. Head: no details. What I believe may be an antennae (third
joint) appears nearly twice as long as broad, evenly rounded at tip.
Arista wanting. Thorax: dark in color, covered with fairly long stiff
bristly pile, but none of the exceedingly long stiff macrochaetae.
Scutellum evenly rounded, its margin simple, with a few bristles; its
width two and a third times as great as its length, the curvature per-
fectly gradual, so that there is no suggestion of squareness. Abdomen:
five visible segments and what appears to have been male genitalia.
The scutellum covers about one-half the length of the first segment.
Abdomen short and broad, about one and a half times as long as
broad. Second, third, and fourth segments about equal in length, the
fourth a little narrower, and the fifth much narrower and short, about
one-third as long as the preceding segment. The abdomen is pale.
If it was once pigmented it has disappeared from all except a narrow
well-marked posterior border on the second, third, and fourth seg-

hull: revisional study of fossil syrphidae 275

ments, which border is about one tenth the length of the segment in
width. Abdomen covered with very fine delicate fairly long pile.
Legs: hind femora slightly thickened; hind tibiae about two-thirds
as thick as hind femora, and noticeably bowed or arched. Hind basi
tarsi long, about two-fifths as long as hind tibiae and at least as long
as remaining tarsal joints. All the legs covered with a conspicuous
double row of stiff, dark spinules irregularly placed. Wings: poorly
preserved. Surface villose; costa microsetose. Third longitudinal vein
nearly straight with a very slight convexity towards the front, cov-
ering its length. Marginal cell widely open. The second longitudinal
vein joining costa quite acutely, perhaps at an angle of twenty degrees.
The anterior cross vein joins the discal cell quite proximally and both
the subapical and discal cells are rather long and slender. (Redescribed
from the holotype specimen.)

Locality: Green River, Wyoming. Horizon: Eocene.

Holotype: No. 5160; paratype No. 5162, Mus. Comp. Zool.

Scudder placed other specimens (5158, 5159, 5161) as Cheilosia
ampla, and he placed number 5162, curiously, as Cheilosia sp. From a
careful examination of this material, I conclude that number 5162 is
Cheilosia ampla Scudder. Thus there are two specimens. No. 5161 I
conclude to be a new species ; its generic affinities will be discussed later.
Number 5158 and 5159 are certainly indeterminate. They may just
as well remain Cheilosia ampla ? Scudder. Both are from type locality.

I also place here provisionally a specimen from Dragon, Utah
(Green River Shales) which has a markedly setigerous scutellum. Its
abdomen corresponds perfectly with that of Scudder's specimen (num-
ber 5160). As that specimen shows traces of bristles on the scutellum,
perhaps the others were lost. The Dragon specimen may remain here
provisionally; I figure its scutellum. (Plate 4, fig. 13).

Cheilosia miocenica Cockerell

Plate 1, fig. C

Bull. Am. Mus. Nat. Hist., 26: 72, fig. 5a, (1909)

Male. Length 10 mm. ; length of wing 9 mm. Head: elongate, eyes
especially elongate, and the posterior part broad right at the base,
leaving (ventrally) the margin concave and fitting tightly over the
thorax. Front convex, rounded, bulging, though nowise protuberant.
Antennae present but concealed. Front and head dark in color.
Thorax: dark; pilar details obscure. Some long delicate hairs at side

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of mesonotum, just before the wing. Scutellum large, broad, more than
twice as wide as long, the rim gently rounded. Squamae and fringe
beautifully preserved; squamae not large, the fringe large, hairs of the
fringe forked once. Abdomen: short, robust, flattened. Hypopygium
large. Pile delicate and slender, not at all long. Color of abdomen dark
brown, though lighter in the fossil than the thorax, due probably to
thinness. Hypopygial and terminal pile not longer than the remaining.
Hind femora short and quite slender, its pile short, no visible spines.
Wings: costal spinules double-rowed, very sharp, rather long, tuber-
culate. Vena spuria present but quite faint. The radial sector vein
arises by offset separation rather than furcation. The wings are con-
siderably longer than the abdomen and the costa like that of Rhingia
or Protorhingia is extensively carried down past the real end of the
wing. In fact, it is quite possible that this species should be put in
the genus Protorhingia (new genus). The marginal angle of the
second posterior cell is a little bit rounded but has a spur and there is
a very short section only to the fourth longitudinal vein before the
origin of the apical cross vein. Both the marginal cross veins are close
to the wing border and quite parallel to it, except as they are slightly
bowed inward. The small cross vein slightly oblique above the vena
spuria, is not quite one-third of the way from the base of the discal
cell. The stigma was definitely darker in color. (Redescribed from
type).

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: No. 4444, University of Colorado Museum.

Cheilosia scudderi Cockerell and LeVeque

Plate I, Fig. A

Scudder, Tert. Ins., 561. PI. 9, fig. 8. (1890) (Cheilosia sp.) Cockerell &
LeVeque. American Naturalist, 45: 357, fig. 4. (1931)

Female. Length 6.3 mm. (6.5 mm. with antennae); leVigth of wing
7 mm. Head: small, much less wide than the thorax, elongate in form.
Front narrowly separated. Occiput apparently well developed though
not tumid. Antennae beautifully preserved, pale in color. The first
joint quite short, third moderate in size, with a fringe of 17 or 18 short
sharp bristles and a larger one on the outside. Third joint very large,
half again as long as wide, evenly rounded at tip. Arista long, two and
one-half times as long as antennae, thickened on basal half, nowhere
sharply. Color of head dark. Thorax: dark, pile short, delicate, a little

hull: revisional study of fossil syrphidae 277

longer on post calli. No macrochaetae. Scutellum dark, broad, short,
rounded, pile as long as that of calli; no macrochaetae. Abdomen:
round, barely longer than wide, widest at end of second segment. Pile
delicate, not long, only moderately abundant, second segment dark,
with a pair of subquadrate basal spots, wider than long, two-thirds as
long as segment, completely interrupted in the middle, not reaching
margin. A similar pair on base of third segment, less rectangular.
Fourth segment apparently all dark. The merest tip of the small
narrow fifth segment shows. Legs: indistinct. Wings: are consider-
ably longer than abdomen, delicate and practically hyaline, the costa
ending practically at tip of the wing and having a double row of sharp-
spined tubercles. Both the marginal angles of the first and second
posterior cell have long spurs from the fourth and fifth longitudinal
vein. The apical cross vein joining the third vein very acutely and not
far from the tip of the wing. The third vein barely curves over its
whole length outwardly. The vena spuria present, small cross vein
quite oblique above the vena spuria, and entering the discal cell just
past the first fourth-way point. Stigma very faint. (Redescribed from
type).

Locality: Green River, Colorado; Dragon, Utah. Horizon: Eocene.

Holotype: No. 15463a in the University of Colorado Museum, from
Green River, Colorado. Also No. 3952 in the Museum of Comparative
Zoology, from Dragon, Utah, belongs to this species.

This species is unique in having light spots on the second and third
segments of the abdomen. The abdomen considered alone is almost
identical in its roundness, flatness, type of pile and pattern with that of
Phalacromyia of the American tropics today. There are no marked
species of Cheilosia living, with one exception (Europe) and this is of a
different sort.

Cheilosia hecate spec. nov.

Plate 3, fig. D; Plate 4, fig. 14

Female: length 8.5 mm. including head; abdomen and scutellum 4.4
mm.; width of abdomen 2.7 mm.; length of wing 6.5 mm.; width of
thorax 2.8 mm. Head: very little detail can be made out of the head
owing to its position. One antennae with its arista is clearly shown.
Color of antennae light reddish brown throughout. The third joint
oval, tapering to an obtuse point. The arista situated at extreme base
of the joint and consisting of an unusually long bristle, evenly and
gradually and conspicuously strengthened on its basal half, and held

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at an angle of forty-five degrees from the joint. The third joint is
about two times as long as the second and the first appears to be ever
so slightly longer than the second. First and second joints minutely
setose. There is no trace of particularly long setae upon their distal
rims, nor is there trace of pubescence on the arista. Thorax: the dor-
sum is dark in color, and covered with fairly long, soft, bristly hairs
but not in any sense spinose. However, there are present the extremely
long setigerous bristles on both calli and scutellar margin which are
the characteristic of many Cheilosi?ii, Volucellini, and of Ferdinandea.
There appear to be four on each side of the scutellum, the median pair
crossed, and some weaker ones at base and to one side of the scutellum.
The scutellar marginal bristles are at least as long as the scutellum.
There are three equally long bristles on the post calli and shorter ones
just before the wing. The humeri appears to be pilose. The scutellum
is wide with evenly convex rim. Abdomen: five segments are visible.
The greater part of the first is covered by the scutellum. The second
is two and one-half times as wide as long; the third is equally long and
practically as wide, whereas the fourth is again two and one-half times
as long as wide but is considerably shortened in length. It is about
three-fifths as long as the second. The fifth segment is inconspicuous.
The most noticeable feature of the abdomen which is unicolorous and
dark, is the middle posterior cone of extremely strong, long setigerous
bristles. These overlap the following segment. Legs: the hind femora
very slightly thickened. The hind tibiae slender, together with the
hind femora covered with thick moderately long bristly pile. Wings:
these are of the typical Cheilosinae type. The costa is microsetose.
The third longitudinal vein is then strongly deflected on its extreme
distal part without even suggesting a kink or bend. As a matter of
fact, the first longitudinal vein approaches the costa so insensibly that
it could be said the marginal cell was just closed. It would be better
to describe it as just open. The second longitudinal vein joins the costa
at an angle of twenty-nine to thirty degrees. The anterior cross vein
is situated well before the middle of the discal cell, and the spurious
vein is quite distinct to within a short distance of the union of lower
marginal cross vein and fourth longitudinal vein. The veins are pale
brown, the wings quite hyaline, with stigma. Under the high power the
small close set villi of the wing are beautifully apparent. These wings
were a fourth longer than the body.

Male. Length of this specimen 9 mm. ; of wing 6.7 mm. ; of abdomen
and scutellum 5.3 mm.

Specimen No. 3953 is a profile of this species. The long bristles of

hull: revisional study of fossil syrphidae 279

the fourth segment shows nicely and venation is quite indistinguishable
from No. 3939. It shows the same subdistal approximation of both of
the veins, second and third longitudinal, to each other one. The details
of the facial profile are of greatest interest. The antennae are hanging
downward and only its narrowest edge or surface shows. It does not
furnish much of interest. Beneath the antennae the face is first gently
concave, then slowly and slightly produced into what might be termed
a gentle tubercle, since it retreats more suddenly below. There is
present what appears to be facial strips. Several things are very
clearly and indisputably shown: the nature of the profile, the quite
convex front, and the degree of production and development of the
face and cheeks below the eyes. On the margin of the eyes above the
front is a trace of what may have been pile. The specimen is obviously
a male. One hind tarsus shows apicotarsal joint, three long bristles,
bicolored cjaw and one pulvillus. The hind femora had long, though
delicate, bristles beneath. The pile of the thorax was erect, fairly stiff,
short, and abundant.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: No. 3929. M.C.Z. Paratype: No. 3953. M.C.Z. (Scud-
der collection).

Cheilosia pratjei spec. nov.

Plate 11, fig. 93, 94; Plate 12, fig. 104; Plate 13, fig. Ill

Male: Length 10 mm.; length of wing 9 mm. Head: very large*
slightly wider than thorax. Eyes enormous and conspicuous, broadly
touching in the male. Upper facets enlarged vertical triangle very
small, not quite restricted to the ocelli. Front not large, convex, with
a median crease on the upper half. Front thickly long pilose, margin
before the antennae lunulate. Antennae located above the middle of
the head in profile. Third joint large and flat, scarcely longer than
wide, broadly rounded apically. Arista very long, strongly but not
sharply thickened basally; about three times as long as the antennae.
Face below the antennae with a broad obtuse tubercle apparently near
the middle. Due to the angle of the specimen it is impossible to deter-
mine whether or not the face is distorted. Face with a single very large
obtuse tubercle below the antennae. Face not greatly produced down-
ward. Cheeks easily visible below the eyes, but not prominent, some
strong stiff bristly hairs on the lower part of face and cheeks. Upper
part of face and region of tubercle appears to have been pubescent
only. Thorax: very broad; as wide as long, somewhat convex, thickly

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and densely pilose. Pile rather long everywhere, pale whitish in color,
the pale white pile on the sides before the wing longer and somewhat
bristly. Scutellum very large, barely twice as wide as long; the disc
and the rim convex, the former covered with thick upright pile, be-
coming long near the rim and on the margin and rim with forty or
fifty long stiff bristly hairs about half of which are set within tubercles,
all of these are pale in color. Dorsum of thorax dark, scutellum dark
but subtranslucent. Abdomen: but little longer than wide, broad and
robust, about as wide as thorax, somewhat convex, the apex very
broad, the color dark, especially on the posterior halves of the seg-
ments. Each segment light brownish or yellowish brown basally. Sur-
face of abdomen thickly long bristly pilose. Hypopygium large and
thick, but quite concealed. Legs: hind femora elongate, stout but not
thickened, with exceedingly numerous, rather long bristly spines which
are not sharp pointed but not thick along the ventral surface. Legs
dark in color, the bases of the tibiae and the tarsi somewhat brownish
yellow. Wings: much longer than abdomen quite pointed apically,
broadest apically, uniformly dark brownish. Stigma a little darker.
Vena spuria very heavy and chitinized. Small cross vein enters the
very long discal cell about three-eighths of the way from the base.
Marginal cross vein extremely long, practically straight, close to wing
margin and parallelling it. The apical cross vein about five-eighths as
long as discal cell. The first posterior cell apically drawn out into a
very sharp acute point owing to the fact that the apical cross vein
joins the third vein practically at the tip of wing. Third vein and
costa end at tip of wing.

Locality: Baltic amber, Germany. Horizon: Lower 01 igocene.

Holotype: in the University Museum of Konigsberg. The specimen
bears no label, but the slide is labeled K; I affix the No. H 103.

I take pleasure in naming this species for Dr. Pratje, who gave much
courteous and helpful assistance in my study of amber Syrpbids.

Cheilosia oligocenica Theobald

Les Insectes fossiles des Terraines oligocenes de France, p. 353, pi. 3, fig. 1.

Length 8.75 mm. Head: transverse, almost as large as the thorax;
the two large eyes are placed upon either side; the front is slightly
prominent. The vertex is elevated and at the front there is a fragment
of the basal part of the antennae with the arista lacking. Thorax:
oval, the scutellum easily distinguished. Abdomen: ovoid, with five
segments visible. Legs: not mentioned. Wings: these are well pre-

hull: revisional study of fossil syrphidae 281

served and it can be clearly seen that it belongs to the Syrphidae; the
longitudinal veins are a little undulated. The general color of the fly
is black.

Apparently the preservation of this fly is sufficient to provide good
specific characters. A more critical analysis of the wing should throw
more light upon the generic and subgeneric relationships of this fly.

Locality: Aix en Provence. Horizon: Oligocene.

Holotype: Am 24, Mus. Nat. Hist. Natur., Paris.

Cheilosia germanica spec. nov.

Plate 11, fig. 83, 95; Plate 12, fig. 99; Plate 13, fig. 112

Male. Length 8 nim.; length of wings 7.5 mm. Head: large, defi-
nitely wider than thorax. Eyes apparently bare, touching in the male
for some distance. Vertical triangle not very small, rather long and
acute, due to the fact that the eyes are not as widely touching as in
some species, with but nine or ten long curved bristles at the top.
Front steep; short; I cannot discern whether it was pubescent but it
appears to have been bare. Antennae situated a little above the middle
of the head in profile. Third joint large, rather deeper than long,
broadly rounded, all the joints and the arista black, the latter a little
over twice as long as the antennae, strongly thickened basally with
two basal joints discernible. Face and cheeks black, the former with a
conspicuous tubercle in the middle. Face beneath the antennae con-
spicuous. The cheeks very little developed and the face at the angle
between cheeks and face produced not more than length of antennae.
Occiput very narrowly visible in profile, with a fringe of short bristly
hairs. Thorax: convex; its pile bristly, moderately long and thick, a
few short macrochaetae on the posterior calli and side of the thorax
above the wing. Scutellum large, convex on disc and rim, about twice
as wide as long, with three or four pairs of long slender bristles on the
rim and a few shorter ones on the sides above. Abdomen: nearly twice
as long as wide, barely wider than the thorax, dark reddish brown in
color, widest in the middle of the second segment, gradually tapering
past that to the broad and rounded apex of the abdomen. Legs: hind
femora very stout but little thickened, with only stiff bristles ventrally.
Hind tibiae practically as long as femora, rather thickened on the outer
two-thirds, the hind basi tarsi as long as the remaining joints and
considerably thickened, very dark, femora and most of the tibiae
blackish; tarsi apparently very dark brown. Wings: uniformly dark
brown, obscured by being closely folded and overlapping the abdomen.

282 bulletin: museum of comparative zoology

Stigma very dark brown. Vena spuria distinct but weak. The small
cross vein enters the discal cell a little less than one-third of the way
from base. Lower marginal cross vein very obliquely directed away
from wing margin. Apical cross vein at first obliquely directed away
for a short distance then bent outwards, joining the third vein not far
from tip of wing. Third vein and costa end at tip of wing. Wing
pointed, apex broad basally, upper squamae with six or seven very
stiff bristles, acute apically, the basal marginal angles of the first and
second posterior cells spurred.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: at the University Museum at Konigsberg. It bears the
number B 16560.

Cheilosia nigrachaeta spec. nov.

Plate 11, fig. 80; Plate 12, fig. 102, 107

Male. Length about 6 mm.; length of wing 5.5 mm. Head: large,
much broader than thorax. Eyes large and conspicuous, broadly
touching in the male, the front not large, steep, very little produced.
Impossible to determine whether pilose, pubescent or bare. The
antennae above the middle of the head in profile short, third joint
large, not as long as wide, broadly rounded apically, the arista half
again as long as antennae, strongly thickened on the basal two-thirds.
The first and second antennal joints dark brown, third joint lighter in
color, especially basally, rather thick on the bottom. Face concave
below the antennae, with a very distinct tubercle. Owing to poor pres-
ervation, it is difficult to determine the nature of the covering. It
appears to have been densely pubescent. Face not produced away
from eyes for more than the length of the antennae. Crease of the side
margins deep. Occiput scarcely visible in profile. From above, the
posterior profile is considerably concave about the vertex and behind
the ocelli. Thorax: quite convex, the dorsum and scutellum very large,
a little over twice as wide as long, the apex a little flattened. The j
margin with a few long slender bristles ; the calli and sides of the thorax i
with strong short macrochaetae. Abdomen: a little longer than broad,
quite robust, barely wider than thorax, much of it obscured by the
wings and preservation. Legs: more slender, barely thickened a little
in the middle with stiff bristles only ventrally. Legs dark brown, tarsi
somewhat yellowish brown. Wings: uniformly pale brown, marginal
angles of the first and second posterior cells with spurs. Lower mar- |
ginal cross vein short, more or less straight, strongly directed away ;

hull: revisional study of fossil syrphidae 283

from wing margin, the last section of the fourth vein before the origin
of the apical cross vein rather long and two-fifths as long as apical
cross vein. Apical cross vein sinuous at origin, joining third vein near
the tip of the wing. Third vein and costa end at tip of wing. Stigma
not darker than remainder of wing. Vena spuria faint. Anterior cross
vein oblique, joining the discal cell a little less than a third of the way
from base.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum at Konigsberg. The specimen
bears the number B 14281.

Cheilosia bruesi spec. nov.

Plate 11, fig. 90; Plate 12, fig. 98; Plate 13, fig. 118

Male. Length 7.5 mm.; length of the wings 6.5 mm. Head: large,
wider than thorax, the eyes apparently bare, anterior facets enlarged,
touching in the male but not for a great distance. Vertical triangle
large, not restricted to the ocelli, the front small, protuberant at the
level of the antennae, subconvex, somewhat steep, apparently without
pile. The presence of pubescence cannot be ascertained. Antennae
situated about the middle of the head in profile, but a little below the
middle of the eyes. Antennae short, third joint very large, suborbicu-
lar, scarcely longer than wide, all of the joints and the arista dark red-
dish brown, the latter but little longer than the antennae, strongly
thickened on the basal half. Micropubescent. Face beneath the anten-
nae quite concave, rising to the low, obtuse, but otherwise well formed
tubercle, which is situated a little above the level of the bottom of the
eyes, then descending gently below the tubercle to the margin of the
epistoma, a distance half as long as the tubercle itself. Face somewhat
produced at the angle between the cheeks and face at least as long as
the antennae. Facial strips wide and conspicuous. Occiput visible
only on the lower third behind, with some strong bristles above and at
the vertex and some long fine hairs at the bottom near the cheeks.
Thorax: dark in color, very convex, the pile sparse, short. Scutellum
quite large, semicircular, convex on disc and rim. Marginal bristles
short, the apical two bristles the longest, not very heavy and scarcely
half as long as the length of the scutellum. Abdomen: twice as long as
wide, wider than thorax and widest at end of second segment from
which it tapers gradually to the broad obtuse apex of the hypopygium.
Four segments, a small portion of another on the left hand posterior
corner and the exceedingly large globose hypopygium are visible. The

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first two segments occupy scarcely more than a third of the length of
abdomen. Abdomen sparse, short bristly, with a little longer bristly
pile at the sides of the second segment at the base. Legs: hind femora
short, somewhat thickened, the thickening spread over the whole
length, the ventral surface of the femora for the greater part of the
length, especially outwardly, with short spinous bristles. Hind tibiae
seven-eighths as long as femora, considerably thickened on the apical
half, blackish in the middle and very dark brown basally and apically.
Femora very dark brown. Wings: considerably longer than the ab-
domen, the third longitudinal vein scarcely bent downward, ending
with costa practically at tip of wing. Vena spuria faint, the last section
of the fourth vein about two-fifths as long as the apical cross vein,
barely sinuous. Basal marginal angle of first and second posterior cells
short spurred, the apical cross vein very long, barely sinuous, joining
the third vein rather close to apex.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum at Konigsberg. The amber
block was marked "Family Leptidae" by someone. I affix the number
5 B 348 on the block.

Cheilosia sepultula Cockerell
Proc. U. S. Nat. Mus. 51: 96 (1917)

This species was described in a comparative way by Cockerell with
reference to C. miocenica Cockerell. This method is not entirely satis-
factory in such a large and difficult genus and an examination of a
type will be necessary to determine its specific nature. I quote below
the measurements taken by the author of the species.

Length about 8.5 mm.; width of thorax about 3.5 mm.; length of
wings 8 mm. The head and thorax are dark. The abdomen is pale,
thinly pilose with black or very dark, narrow, sutural bands but no
longitudinal vittae. The costa is thick with two rows of minute bristles
as in C. miocenica. The apical angle of the first posterior cell is more
acute than in miocenica. In the following measurements those for
miocenica are given in parenthesis.

Width (depth of marginal cell) .3 mm. from end 35 (.27) mm.

Length of first posterior cell from lower basal corner to upper apical
corner 4 (4) mm.

Submarginal cell on first basal (not allowing for curve)

1.2 (1.28) mm.

hull: revisional study of fossil syrphidae 285

First basal on second posterior 83 (1.09) mm.

Second basal on second posterior 27 (.32) mm.

Second basal on third posterior 3 (.25) mm.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: no. 61994 U. S. N. M. I was not able to locate the type.

"The smaller specimen, assigned to C. miocenica1 and collected by
Mr. Rohwer, belongs to this species."

This species is unrecognizable on the basis of its present description.
There are so many species of Cheilosia differing in minute ways that a
comparison from a photograph is needed.

Genus Cheilosialepta genus nov.

Head: very large, widely holoptic in the male. Eyes bare, anterior
facets enlarged, vertical triangle small with protuberant ocelligerous
tubercle and with a few long curved bristles. Front not large, steep,
flat, densely pubescent only, nowhere markedly swollen or convex.
Antennae short, third joint very large, little longer than broad, thin
above, rather thick below. Arista elongate, strongly thickened basally
and pubescent. Face quite narrow, almost carinate, due to the pinched
and thinned central ridge, which is broadly rounded. There is only the
faint indication of a tubercle in the male, and the female with similar
narrow and protuberant face is without tubercle, and very narrowly
concave. Thorax: convex, the margin of the mesonotum, posterior
calli, upper part of mesopleurae and the rim of the scutellum as well
as posterior margin of the mesonotum before the scutellum everywhere
equipped with enormous heavy machrochaetae suggesting Ferdinandea
(Recent genus), but of course the face is very different from Ferdi-
nandea. Scutellum very large, perfectly semicircular, with convex rim
and disc. Abdomen: large and elongate. Hind femora slender without
spines below. The hind tibiae with strong bristles laterally near the
middle. Wings: very much like those of Myiolepta.

Genotype: Cheilosialepta baltica spec. nov.

This genus is distinguished from Myiolepta by the narrow, almost
carinate face, the more broadly holoptic eyes of the male, the heavy
macrochaetae of thorax and scutellum and the semicircular shape of
scutellum and the absence of spines on the hind femora and the pres-
ence of spinous macrochaetae in the middle of the hind tibiae. It

» Bull. Amer. Mus. Nat. Hist. 26:72. (1909).

286 bulletin: museum of comparative zoology

differs from Cheilosia by the non-tuberculate face of either sex and the
Myiolepta-]\ke venation.

Cheilosialepta baltica spec. nov.

Plate 8, fig. 54; Plate 9, fig. 67, 68, 69; Plate 10, fig. 79

Male. Length 7.5 mm. ; length of wing 6 mm. Head: very large, dis-
tinctly wider than thorax. Eyes enormous; anterior facets but little
enlarged; eyes touching broadly in the male. Vertical triangle small,
restricted to the ocelli. Bristles erect, long, and stiff; they consist of a
cluster of fourteen, two of which are placed at the extreme posterior
end of the triangle. Front not large, bare, densely pubescent, with a
lunulate margin above the antennae. Antennae short; the third joint
quite large, half again as long as wide. Arista long, strongly thickened
basally, about twice as long as antennae; all of the joints reddish brown,
not very dark. Arista blackish. Face reddish brown, quite concave,
below the antennae rising to a large conspicuous tubercle and then
abruptly descending the short distance to the epistoma. Face not
deeply produced, cheeks very short in profile. Occiput scarcely visible
at any point due to the prominence and curvature of the eyes and with
a fringe of delicate hairs. Thorax: convex, barely longer than wide; the
dorsum with thick, short bristles. Humeri pilose; mesopleura with
one excessively long heavy bristle. Side of thorax behind the humeri
with three long bristles. Margin of the dorsum just above the base of
the wing with three heavy bristles and the posterior calli with three
heavy still longer bristles. The dorsum of the thorax just before the
scutellum with four heavy long bristles and the margin of the scutel-
lum with four pairs of very long, very heavy bristles longer than the
length of the scutellum. Disc of scutellum somewhat flattened, thick,
short, bristly, in shape twice as wide as long, extreme apical margin
a little subtruncate, the rim broadly rounded. Abdomen: a little more
than half again as long as wide, tapering considerably from the end of
the second segment; the last two segments scarcely longer than the
first two; only four segments visible. Sides of the abdomen slightly
curled over, quite thin. Abdomen quite convex, the posterior segment
sub-cylindrical. Hypopygium large and rounded, quite concealed.
Abdomen short, appressed bristly, some very long stiff bristles at the
base and sides of the second segment, and a strong radially directed
tuft of shorter bristles at the junction of the first and second segment
on sides. Legs: hind femora a little thickened on the basal two-thirds.
Ventrally equipped only with stiff relatively short bristles. These

hull: revisional study of fossil syrphidae 287

bristles might be described as subspinous but they are not true spines.
Wings: considerably longer than abdomen, the very long, almost strong
apical cross vein joins the third vein almost at the tip of wing. Third
vein a little drawn down on the outer third, ending with the costa, but
not prominently. The marginal cross veins practically parallel the
wing margin and both of the basal marginal angles of the first and
second posterior cells are spurred. Vena spuria distinctly and heavily
chitinized. Wings: pale yellowish. Stigma deep brownish-yellow.

Female. This female is more poorly preserved in many respects than
one that I describe as the male. The hind tibiae bear in the middle on
the outside a row of six or seven long stout bristles, the coxae bear a
fan-like row on the outside of four stiff bristles and others at the point
where they touch; the metasternum bears a very few long hairs but
is mostly bare. The color of the legs is dark brown ; bases of the tibiae
and anterior tarsi somewhat lighter. The apex of the middle femora on
the outer lateral side bears a little cluster of three stiff long bristles.

Locality : Baltic amber, Germany. Horizon : Lower Oligocene.

Holotype : in the University Museum at Konigsberg. This specimen
bears the number 11B665 on the block itself; the slide on which it is
mounted bears the number 2665 and also bears the following designa-
tion: VII. 2. 288 (Museum Stantien and Becker).

Paratype: in the University Museum at Konigsberg. The specimen
bears the label B 16787, the slide the number 16787.

Genus Arctolepta gen. no v.

Head: large, much broader than thorax, in profile from above very
concave. Front quite flat, somewhat prominent. Antennae barely
above the middle of the head in profile, short, third joint large, not
longer than broad, very orbicular. Arista elongate, face deeply con-
cave. Epistoma thrust forward. Occiput visible in profile throughout.
Thorax: very little longer than broad, quite convex; scutellum large,
convex on disc and rim with many long bristles on the margin of the
scutellum but none on the mesonotum before it. Mesonotum on sides
and posterior calli with some long bristles. Abdomen: elongate, over
twice as long as wide, rather flattened basally, somewhat convex on the
terminal segment. Pile short, appressed, bristly. Legs: hind femora
stout, but not greatly thickened, slightly arcuate with many stiff
bristly spines below and apically. Hind tibiae practically as long as
femora. Wings: elongate, not as long as abdomen. Apical cross vein
joining third vein some distance before the tip of wing. Vena spuria

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present and weak. Apical cross vein slightly sinuous ; last section of the
fourth vein relatively short.

Genotype : Arctolepta calarnitans spec. nov.

This fly cannot go in Cheilosia for lack of tuberculate face or Myio-
lepta since the scutellum is chaetate and the apical cross vein is conflu-
ent remote from tip of wing. It differs from Sericolepta in the spiny
scutellum, orbicular antennae, and the simple femora and long tibiae,
small size, much shorter apical cross vein, and the wings not being as
long as the abdomen. In Sericolepta in spite of the large abdomen, the
wings are much longer. From C heilosialepta it is at once very distinct
in the remote point of confluence of the apical cross vein.

Arctolepta calamitans spec. nov.
Plate 6, fig. 36; Plate 10, fig. 74-76

Female. Length 10 mm.; length of wing 6.4 mm. Head: very large
Eyes bare, front and vertex and cheeks black in color, the front with
some scattered light erect pile. Face pubescent only. Antennae light
brownish orange, the basal bristles black. Arista black. Occipital
bristles above stiff and black, the pile in the middle and below is pale.
Thorax: dorsum of thorax and scutellum black in color, aeneus. The
pile on the sides pale and erect. The many short bristles that broadly
cover the mesonotum, together with the macrochaetae on the sides and
scutellum, are black. Abdomen: black throughout, the short stiff spin-
ous bristles appressed and black and very sharp and very little longer
even at the extreme tip of the abdomen. Bristles of genitalia pale.
Legs: throughout black in color except that the anterior tarsi are
brownish red. Hind basi tarsi as long as the remaining segments,
somewhat thickened, the bristles below brush-like and thick. The pile
at the base and along the dorsal length of the hind femora, the bristles
and spines below and on the tibiae and the strong spines and the apex
of the hind tibiae as well as the middle tibiae black. Wings: uniformly
brownish. Last section of fourth vein about half the length of the
marginal cross vein, the small cross vein is extremely oblique above the
vena spuria and enters the discal cell almost at the midway point but
is basal. Both the marginal angles of the marginal cells with very
short spurs.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum at Konigsberg, from Baltic
amber, number X444.

hull: revisional study of fossil syrphidae 289

Genus Protorhingia genus nov.

Head: large, the eyes particularly conspicuous and bare. Upper
facets somewhat enlarged, broadly touching in the male, the vertical
triangle in the male quite small, restricted to the ocelli which are
strongly raised above the surface. Front small, rather steep, scarcely
protruding. The antennae located distinctly below the middle of the
head in profile. Antennae short, but the third joint quite enlarged and
about twice as long as wide and suborbicular. Arista very long and
slender, not noticeably thickened at base, bare. Face below the
antennae gently concave and retreating, the epistoma barely produced
forward. Face without tubercle. Face below the eyes but little pro-
duced and in length about equal to length of the antennae. Occiput
narrowly visible on the lower part of the head, not in the slightest dis-
cernible on the upper two-thirds of head. Thorax: broad, about as
wide as head across the wings, dorsum quite convex. Scutellum very
large, hemispherical, convex on the disc and rim. Thorax and ab-
domen short pilose, the former with a few macrochaete. Scutellum
with many long stiff bristles about apex. Abdomen: quite short, broad,
compact, the terminal segments narrow, curling down and thin at the
sides. Legs: hind femora short, very little thickened with only a few
stiff bristles disto-ventrally. Wings: elongate, much longer than ab-
domen, with typical Syrphus venation, except that the costa and with
it the third vein are grossly and conspicuously pulled down beyond and
below the tip of the wing for some distance and are as conspicuously
bent downward as most Volucellas are bent upward. This leaves the
sub-marginal cell grossly widened and flared at apex. The marginal
cross veins are long, due to the length of the wing, close to wing border,
almost parallel to it, and the basal posterior angles of both their
respective cells are spurred. Wings villose throughout.

Genotype: Protorhingia carpenteri spec. nov.

The genus Protorhingia, it seems to me, is a well founded one, for
the wing in its way is extremely unique. A very few of the two hundred
or more living species of Cheilosia show a somewhat similar wing in
regard to the great extension of the costa around the end of the wing.
Since this is a characteristic of Rhi7igia, which is, of course, abundantly
distinct on the basis of its conoidal snout, it seems at once apparent
that the form that I now describe is a true connecting link between
Rhingia and Cheilosia and it is possible that these few living members
of Cheilosia should be put in Protorhingia. There is this difference,
however, between Protorhingia: modern Cheilosia must have the face

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tubereulate in the male. In fact, a large number of Cheilosia-like forms
from the amber, species which in every way suggest our Cheilosia
probably cannot be placed in the genus Cheilosia in the modern sense
purely because one of their sexes is non-tuberculate. It will be remem-
bered among the living genera of the subfamily Cheilosinae that there
are a considerable group of genera sharply separated from Cheilosia
and its congeners on this very character of only males tubereulate.
It seems evident with so many concaved-faced species and genera from
the amber that we are witnessing, when combined with living forms,
the three types of facial combination :

a. Faces of both sexes distinctly non-tuberculate.

b. Faces of one sex tubereulate (always male?).

c. Faces of both sexes distinctly tubereulate.

Protorhingia carpenteri spec. nov.

Plate 10, fig. 77

Female. Length 10 mm.; length of wing 9 mm. Head: front and
vertex not very widely separated, perhaps separated by three-tenths
of a millimeter. Antennae with the third joint apparently quite large
and but little longer than broad, its shape is somewhat obscured owing
to whitish excretion. Face not greatly produced but the cheeks are
certainly deeper than in Protorhingia magnipennis; at the lower part
of the face there is a certain amount of vesicular swelling that incloses
a few bubbles. Thorax: dorsum of thorax with one bristle behind the
humerus and two quite long ones on the posterior calli. The pile of the
dorsum of the thorax is quite short with an occasional longer bristle.
Scutellar disc also very short, bristly pilose with only a few long bristles
on the margin. One or two of these are broken off but there appears
to have not been more than six or eight bristles two-thirds of the
length of the scutellum. Scutellum convex with convex rim. Color of
thorax dark, shining brown with three vittae black in color fused on the
anterior two-thirds and V-shaped posteriorly. It is possible that in life
these were not discernible or at least obscure. Abdomen: a little over
half again as long as wide, tapering gradually from the end of the
second segment. Base a little bit wider than the thorax and narrowly
luteous or subtranslucent in color. The remainder of the abdomen, in-
cluding the last half of the second segment, quite dark. Abdominal
pile short, thick, and bristly. Legs: hind femora short, very little thick-*
ened, with only a few stiff bristles disto-ventrally. Wings: hyaline in
color, the basal and anterior marginal area, including the stigma, ob-

hull: revisional study of fossil syrphidae 291

scured by bad preservation. The last section of the third vein past the
confluence of the apical cross vein is very short and is about one-fourth
the length of the subapical cross vein.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: no. 3969, Mus. Comp. Zoology; consists of a crude speci-
men preserved in an uncut piece of Baltic amber, purchased by the
author from the Bernstein company in Konigsberg.

Protorhingia magnipennis spec, now

Plate 10, fig. 78

Male. Length about 12 mm., the length of the wing 12 mm. Head:
quite large, the posterior occipital fringe very short and consisting of
but a few rows of hairs except in the region of the very short slightly
convex cheeks where it is a little longer. Facial strips narrower than
carpenteri, the antennae dark reddish, lighter above and near the base ;
the apex and lower portion blackish. The arista dark in color, pile of
vertex longer than elsewhere on the head, erect and somewhat stiff.
Front very slightly convex, with scarcely any trace of median impres-
sion and densely pubescent; apparently pale in color. The cheeks,
narrow sides of face and apparently the greater part of the face except
for a somewhat triangular area on the sides pale pubescent or pollinose.
Thorax: densely erect, bristly hairy. The pile becoming somewhat
appressed on the posterior part of the dorsum. Surface pile of scutel-
lum dense, longer than that of dorsum of thorax, the marginal bristles
quite stiff, nearly two-thirds as long as scutellum and composed of
about fifteen or more on the semi-circle. The color of the thorax and
scutellum appears to have been brilliant coppery or violet. Humeri
pilose, two small bristles directly in front of the wing, three others on
a level with or behind the humeri, three on the posterior calli. Apex
of the dorsum before the scutellum without marked bristles. Abdomen
subshining metallic. Pile erect, thick, and bristly and not very long,
appressed only at the extreme margins of the terminal segment. Legs:
hind femora obscured by poor preservation, the anterior femora small,
slightly arcuate, considerably thickened, and with a row of long stout
bristles on the posterior side. Wings: more or less hyaline, especially
apically, the basal point a little yellowish or brownish, the stigmal cell
deep yellowish brown. Vena spuria weak and faint but with a conspicu-
ous node just past the origin of the second and third longitudinal veins.
Alulae quite large and appear to be rather dark; their fringe is long.
The veins of wings are strong and the section of third vein from the

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confluence of the apical cross vein to the tip is not quite one-third of
the length of the apical cross vein.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum at Konigsberg, No. K7559
("collection Klebs).

Rhingia species Hope
Trans. Ent. Soc. London, 4: 252 (1845^7)

Sex indeterminate. Length 11 mm. This specimen is very unsatis-
factory in many ways. While it appears to be a Rhingia, the venation
cannot be worked out with sufficient certainty to determine its nature.
I therefore refrain from assigning a specific name.

Locality: Aix in Provence, France. Horizon: Oligocene.

Holotype : Hope Museum, Oxford, England. The type was studied.

Rhingia zephyrea spec. nov.

Male (apparently). Length 7.8 mm.; of abdomen and scutellum
4.6 mm.; of wing 6.6 mm. Head: preserved from dorsal view. The
epistoma projects conoidally in front of the face. The antennae are
scarcely discernible. No details can be seen. Face dark in color.
Thorax: dark in color. Pile not discernible. Scutellum evenly convex on
margin; about one and three-fourths wider than long, its margin with
one or two long bristles and numerous short, fine hairs. Abdomen:
oblong, apparently more so than in present day species, which have the
abdomen almost round. Only four segments including the first show
with any degree of distinctness. There are faint indications of the out-
line of the fifth. The abdomen was uniformly dark in color with the
possible exception of the basal half of the second segment. Much of the
pigment is gone, but almost exactly the right posterior quarter of that
of the second segment remains, and from the even gradation of this
pigment-area's anterior margin it seems likely that the base of the
segment was pellucid. Legs: wanting. Wings: well preserved in places.
The anterior cross vein is quite before the middle of the discal cell.
The costa ends quite beyond the tip of the wing as it should in the
modern concept of this genus ; the second and third longitudinal veins
are, therefore, decidedly arched and convexed distally. The subapical
cross vein is more or less evenly convex on its outer edge.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype : No. 3946, in the Museum of Comparative Zoology. This
is the specimen referred to by Williston (1886).

hull: revisional study of fossil syrphidae 293

PlPIZA MELANDERI Spec. nOV.

Plate 2, fig. C

Female. Length 13 mm. ; length of abdomen 7.5 mm. ; of wing 7 mm.
Head: quite globular, smaller than the thorax. The antennae set appar-
ently a little below the middle of the head in profile, short, the third
joint of moderate size, scarcely longer than wide. Arista not showing.
Thorax: small, short, and broad and dark in color. The pile delicate
and apparently erect and not very long. The outline of the scutellum,
since the fly is a little bit twisted to one side, appears to have been less
than twice as wide as long with an evenly circular margin. Abdomen:
slender and preserved in such a way that there is a slight appearance
of narrowness at the base. This is probably due to the partial side wise
position of the fly. Abdomen two and one-half times as long as wide,
dark in color with a pair of basal pale-colored spots on both the second
and third segments. These spots are odd in shape and rather widely
separated. There is a trace of such spots on the fourth segment. Pile
of abdomen fairly long and delicate. Legs: the hind femora are visible.
They are slightly thickened and their pile, at least on the ventral part,
is short but stiff. The tibiae practically as long as the femora but
slightly arcuate. Wings: well preserved and with typical Pipiza-\ike
venation. The marginal cross veins are slightly bowed inward near the
middle and there is a suggestion of an inwardly directed spur. There is
a short spur directed upwards towards the end of the vena spuria from
the fourth longitudinal vein just before that vein joins the lower mar-
ginal cross vein. Small cross vein enters the discal cell barely more than
a fourth the distance from the base.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: no. 154, coll. American Museum of Natural History.

Pipiza venilia Heyden
Palaeontographica, 17: 260. PI. 45, fig. 28. (1870)

The description and illustration given by the author present the
following points of interest and value.

Length 7 mm. The specimen lies upon its side so that the wing is
partly obscured by the abdomen. The head is fairly large ; the eyes are
large and holoptic; the front is projecting and rounded. The antennae
are wanting. Thorax: is longer than broad. The scutellum was about
half as long as the dorsum of the thorax with entire unspined margin.

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Abdomen: was elongate, twice as long as broad, and a little broader in
the middle. Five segments are visible, the first four of almost equal
length. There is some black pile posteriorly upon the segments. Legs:
are almost wholly wanting; only traces of the femora are visible.
Wings: the middle of the wing is clearly shown and the vena spuria is
visible and determines definitely its place within the Syrphidae.
Heyden states that the first posterior submarginal cell is prominent
over the discoidal cell and the upper anterior angles of the first posterior
cell is pointed ; the small cross vein lies upon the basal side of the discal
cell; the tip of the wing is not clear so that it cannot be seen if the sub-
marginal cell is open or closed.

Locality: Rott, Germany. Horizon: Oligocene.

Type: One specimen in the Krantz collection. I was not able to
locate the type.

This species is based upon a well preserved specimen, but since the
living flies in the group Pipizini are especially difficult to classify and
recognize, it is obvious that the type should be re-studied.

Genus Pseudopipiza subgenus nov.

Since the species antiqua is not strictly congeneric with modern
Pipizas it seems to me sensible to place it in a separate genus.
It cannot be determined definitely that the eyes are pilose though
apparently they are. If so, this would be like modern Pipiza to this
extent. The face is bulging out definitely about the region of the epis-
toma. The eyes prominent and face and head thickly hairy as in
Pipiza. Thorax: convex, thickly, delicately pilose without macro-
chaetae. The scutellum large, semicircular but with the rim thin and
much flattened in contrast to Palaeopipiza. The abdomen drooping
as in Pipiza or Palaeopipiza, much convex but rather more densely
pilose. The hind femora are a little more thickened and spinose per-
haps, the apical cross vein is quite long and joins the third vein prac-
tically at tip of wing, more closely than in the living species.

Genotype : Pseudopipiza antiqua spec. nov.

Pseudopipiza differs from the present-day Pipiza in the face and in
the confluence point of the apical cross vein being practically at wing
tip. It differs from Palaeopipiza in the different face, very pilose head
and thorax and the much thinned scutellar rim. In the latter genus
the margin of the scutellum is thick and convex.

hull: revisional study of fossil syrphidae 295

PSEUDOPIPIZA ANTIQUA Spec. nOV.

Plate 11, fig. 86; plate 13, fig. 113

Male. Length 4.5 mm.; length of wing 3.3 mm. Head: eyes touch-
ing broadly, apparently pilose. Antennae situated two-fifths of the
distance from the top of the head. First two joints very short ; third
large, slightly subquadrate, barely longer below than wide. Arista
basally thickened, slender on the apical half. Face below antennae
very slightly concave for some distance, bulging out a little near the
epistoma; not tubercular. Face, front, cheeks and occiput dark.
Antennae brown, arista pale. Pile of entire head pale silvery, very
thick on the face and cheeks, present on front. Thorax: and scutellum,
dark in color, the margin of the scutellum very thin and impressed and
the pile of scutellum and thorax everywhere quite long and delicate
and very dense. Abdomen: drooping from the base, very convex and
subcylindrical past the second segment; its pile thick delicate, sub-
appressed, everywhere pale. Hypopygium large, rounded, prominent,
together with the whole abdomen dark in color. Abdomen and thorax
obviously shining or aeneous. Legs: hind femora somewhat thickened,
with thick bushy pile above; a few bristly delicate spines, perhaps only
bristles apically. Wings: with venation like Pipiza. Marginal angles
of first and second posterior cells spurred, lower cross vein straight,
apical one long, curved inward on the first half but is actually almost
imperceptibly curved over its whole length, the curve upward, the last
section of the fourth vein before the origin of the apical cross vein
straight and a little longer even than the lower marginal cross vein.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the Geologisch-Palaontologisches Institut and Mu-
seum der Universitat Berlin. I affix the number H 220 to the specimen.
Allotype: H 221, female; same museum.

Pseudopipiza europa spec. nov.
Plate 11, fig. 96; plate 13, fig. 121

Female. Length 5 mm.; length of wing 4 mm. Head: barely wider
than thorax, hemispherical. The front wide and the eyes broadly
separated even at the vertex. Antennae set below the middle of the
head in profile; short; third joint moderately large, as long as wide and
broadly rounded. The arista is a little longer than the antennae and
is strongly thickened on the basal two-fifths. The face below the an-
tennae is gently concave and has only a slight suggested protuberance
about the oral margin. The face is short and not at all produced. The

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cheeks, while not conspicuous are well developed. The occiput is very
narrowly visible above and below. The posterior occipital margin
viewed from above is shallowly concave. There is some short, sparse
pile on the front and some longer, forward-directed, bristly hairs at
the top of the vertex. The cheeks are short pilose. There seems to be
a few erect hairs on the face, which are very sparse. Thorax: convex,
short, erect pilose. The scutellum is short and small, its margin semi-
circular with some delicate, upturned-bristles on the margin. There
are a few long, slender bristles on the calli and before the wing. Abdo-
men: nearly twice as long as wide and short pilose; the posterior seg-
ments are narrow and quite convex. The abdomen was apparently
aeneous in life. Legs: hind femora short and quite slender with a barest
suggestion of a thickening just past the middle; equipped ventrally and
distally with very fine, sharp bristle-like spines. The hind tibiae are
slightly arcuate and thickened distally and end transversely.

Holotype: a female in the Geologisch-Palaontologisches Institut
und Museum der Universitat Berlin No. H 221. Found in Baltic
Amber (Lower Oligocene).

Chrysogaster antiquaria spec. nov.

Female. Length 5.0 mm.; of wing 3.8 mm.; of abdomen and scutel-
lum 2.4 mm. Head: small dark colored fly, preserved in exact profile.
The farthest projection of the face seems to be about one-third the
height of the head from below, this seems to be sharply but obtusely
angulate rather than tuberculate and below this point the face recedes
away. Above this point the face recedes to the point of antennal
attachment and above the point of antennal attachment the long front
is rather convex but wrinkles cannot be seen. It must be admitted
that in this specimen the point of attachment and characteristics of the
antennae are open to two interpretations, since below the point of
attachment as described, and arising from the angular point of the face,
there is a slender attached body which may consist of two or three
slender antennal joints. If this is true, the face just above the antennae
is concave and the antennae would be situated very low upon the
profile. This seems improbable. At the point of the face described as
the locus of the antennae, the profile is slightly damaged, but a body
does arise there which consists quite probably of the real antennae.
Three joints can perhaps be made out. The first two joints are short
and the third is oval, about two or two and a half times as long as
wide. The fly belongs, therefore, to the subgenus Orthoneura. The eye

hull: revisional study of fossil syrphidae 297

facets are not visible. Thorax: gently convex, a little sloping just in
front of the head. Scutellum small and flat. No pile can be discerned
on the thorax. Abdomen: fat, the tergite and sternite wide apart;
obviously the abdomen was tumid with eggs. Six segments can be
seen and there are others long drawn out and tapering, as characteris-
tically precede the ovipositor of these flies, but they are obscured by
other matter and the exact number is uncertain. No pile is visible.
Legs: small and doubled up. They were mostly pale in color, the distal
halves of the tibiae and the tarsi dark in color, the basal parts of the
legs pale. Some fine hair can be discerned upon them. The hind femora
is slightly thickened; it is about one and a half times, or less, as thick
as its tibiae. Wings: hyaline; if infuscation was present it is not evi-
dent. The last section of the sub-apical cross vein joins the third vein
nearly at right angles. It might be described as slightly recurrent.
The venation is difficult to make out, but from one position, with the
right slant, the more important details can be very well seen.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: No. 12621, in the Museum of Comparative Zoology. This
is the specimen mentioned by Williston (1886).

Genus Cacogaster genus now

Head: large and broad. Front and face quite wide and rounded in
profile from above. Arjsta elongate and slender. The antennae appar-
ently low upon the profile of the head since no trace of them shows.
Thorax: broad and robust. Abdomen: short and wide, being shorter
than in Syrphus but not quite as round and flat as in Rhingia. The
pattern of the abdomen is unique and the many small isolated spots
suggest patterns that are often seen in certain gadflies but not in
Syrphids. Wings: venation Syrphus-Mhe. Marginal cell open. Third
longitudinal vein straight. Anterior cross vein well before the middle
of the discal cell. Spurious vein present and extensive. Costa and third
vein end at the tip of the wing.

Genotype: Cacogaster nooamaculata spec. nov.

The particular characteristics of this form lie in the wide front and
face, its convexity, and the short abdomen with its unique type of
maculation. Better specimens are needed to place it finally in its
phyletic position, but this can be said of many fossils since the key
characters to exact position are usually so minute that they are seldom
preserved except in amber.

298 bulletin: museum of comparative zoology

Cacogaster novamaculata spec. nov.
Plate 6, fig. 37

Sex indeterminate. Length 8.8 mm.; length of body and scutellum
5.2 mm.; length of wing 6.3 mm.; width of abdomen 3.0 mm. Head:
very little detail shows. The eyes seem to be separated, antennae are
not visible, both aristae appear, and are long and thickened; the an-
tennae themselves were certainly quite short. Thorax: short and broad
and dark in color. Very short pile present on thorax. Scutellum about
one and three-eights times as broad as long. Margins simple, evenly
rounded and without pile or bristles. Abdomen: with five segments and
three others visible terminally. First segment dark, nearly covered by
the scutellum. The succeeding three segments marked each with three
subtriangular spots on the posterior border, a median one and one on
each side, in each postero-lateral corner. The median spot on the
second segment is largest and continuous as a band to anterior margin
interrupting the segment; the median ones of third and fourth seg-
ments failing to reach the anterior margin and progressively smaller.
The median spot of third segment almost an equilateral triangle.
Second segment a little longer than third; third and fourth equal.
Third segment two and three quarters to three times as wide as long.
Abdomen covered with short decumbent dark bristly pile, somewhat
thicker on the posterior part of fourth segment. Legs: hind femora
moderately thickened, hind tibiae slightly so. Hind femora appears to
be without spines or setae apically. Hind tibiae slightly thickened and
covered with thick short ventro-decumbent bristly hair. Wings: these
are of the Syrphine type. Marginal cell open. Third longitudinal vein
straight; anterior cross vein well before the middle of the discal cell.
Costa microsetose. Spurious vein evident, reaching almost to fusion of
fourth longitudinal vein and postical cross vein. Third longitudinal
vein practically straight until close to tip, then with a gentle downward
concavity before joining costa at tip of wing, where costa ends. Second
longitudinal vein joins costa at an angle of about twenty-five degrees.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype : no. 3940 in the Museum of Comparative Zoology. (Scud-
der collection.)

This is a very peculiar form which is so aberrant from any Syrphid
type I am acquainted with that I hesitate to place it with Syrjihus.
The body is intermediate in shape between Syrphus and Rhingia; the
maculation reminds one of a large group of Tabanus but of no Syrphids
I have seen.

hull: revisional study of fossil syrphidae 299

Myiolepta valida spec. nov.
Plate 11, fig. 89; Plate 12, fig. 97, 108

Female (apparently). Length 7.5 mm.; length of wing 5.6 mm.
Head: large, wider than thorax; eyes conspicuous, rather widely sepa-
rated, nowhere approximated. Antennae located about the middle of
the head in profile, due to the position of the specimen the exact
position of the antennae cannot be ascertained. Antennae short, third
joint large, perfectly rounded, but little thickened, and flat on the
sides. Arista three times as long as the third joint, strongly thickened
basally and pubescent. Antennae dark reddish brown, face apparently
without tubercle but deeply concave below the antennae and the epis-
toma bluntly protuberant. The antennae reaching almost to the
epistoma and certainly reaching below the middle of the face in profile.
Head very much obscured by whitish exuvia. Thorax: well preserved,
thickly, short appressed setate. The sides of the thorax before the wing
with four large stiff bristles, one such bristle on the mesopleura and
three on the posterior calli ; none on the apex of the dorsum before the
scutellum but the margin of the scutellum has three pairs of exception-
ally stiff heavy long bristles and a few weaker ones basally. Disc of
scutellum with many short bristles. Margin of scutellum convex, form
semicircular. Abdomen: short, broad, and robust, wider than thorax,
together with the legs much obscured by white exuvia. Wings: nearly
hyaline, very pale brownish, thickly long villose, the stigma a little bit
darker. Vena spuria quite faint, small cross vein very little oblique
above the vena spuria, joining the discal cell exactly one third of the
way from base. Posterior basal angle of the lower marginal cell spurred,
lower marginal cross vein straight, the posterior marginal angle of the
first posterior cell without any spur, the subapical cross vein gently
sinuous at first and then proceeding straight and acutely to join third
vein practically at tip of wing.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype : in the University Museum at Konigsberg. The specimen
is not marked; the slide is marked S. I affix the Number H 104 to the
slide. Paratype : no. 22195 in the British Museum of Natural History.

Myiolepta andreei spec. nov.

Plate 11, fig. 88; Plate 12, fig. 103, 105; Plate 13, fig. 114

Female: Length 6.5 mm.; length of wing 4.3 mm. Head: large,
wider than thorax viewed from above. The head is rather thick, eyes

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rather broadly separated, nowhere angularly approximated. Front a
little produced but not conspicuous. Antennae situated in middle of
head in profile, short, third joint large, broader than long, rounded
apically; the arista but little longer than the antennae, strongly thick-
ened on the basal half, pubescent; the entire antennae almost black.
Face below the antennae distinctly concave. Epistoma not greatly
produced. The face at this point being barely as long as third antennal
joint, cheeks in profile narrow. Occiput strongly visible and conspicu-
ous on the lower three-fourths of the eye margin but quite invisible
above this point, its upper margin is set with stout spines, the lower
part short pilose. Thorax: scarcely longer than wide, the posterior
part of the thorax past the wings much narrower. Scutellum of mod-
erate size, a little more than one and one-half times as wide as long,
a little bit flattened, the apex subtruncate, the margin with a few
short stiff bristles. Thorax without the second segment tapering gradu-
ally to the broadly rounded tip. Surface of abdomen short setaceous,
with some longer pile at the sides and base of the second segment.
Terminal segments very convex, color of abdomen light brown; sub-
translucent, but this is probably due to the preservation. Wings:
nearly hyaline. Stigma barely darker. Vena spuria very faint. The
last section of the fourth vein before the origin of the apical cross vein
is long but not quite as long as lower marginal cross vein, the latter
slightly bent inward and its origin spurred from the fifth vein. Apical
cross vein barely bent inward shortly after its origin, the remainder
straight, joining third vein a short distance from tip of wing, but not
as close to it as is usual in Myiolcptas.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum at Konigsberg. The specimen
is B 14586 and the slide is 14586. This species is named for the Director
of the Konigsberg Museum.

Myiolepta germanica spec. nov.

Plate 11, fig. 91, 92; Plate 12, fig. 101, 106; Plate 13, fig. 120

Female. Length 8 mm.; length of wings 6.3 mm. Head: much
broader than thorax, very large. Eyes conspicuous, bare, broadly
separated, nowhere approximated. Front short pilose, pubescent only
on the lower part. Antennae set at about the middle of the head in
profile, short. Third joint large, a little longer than broad. Arista
strongly basally thickened. Micro-pubescent. Face in profile quite
concave, the blunt epistoma projecting and the face opposite the

hull: revisional study of fossil syrphidae 301

epistoma produced about as long as third antennal joint. The face
appears to be pubescent only. Facial strip broad, conspicuous, the
crease deep. The occiput visible throughout in profile, rather strongly
developed posteriorly on upper part of the head, with a few spinous
bristles near the vertex. Thorax: convex, very short, setaceous, the
bristles appressed. Scutellum quite large, a little more than twice as
wide as long, the posterior part subtruncate. The disc a little flattened
and a few quite short bristles on the rim. Abdomen: quite broad and
robust, wider than thorax, a little longer than broad, much concealed
by the wings. The pile short, much appressed and setaceous. A stiff
surface patch of long bristly hairs very dense in the basal corner of
the second segment. Legs: the hind femora slender, a little thickened
in the middle and with only very short numerous spinous bristles ven-
trally. Hind tibia over three-fourths as long as hind femora, some-
what thickened posteriorly the hind basi tarsi about as long as re-
maining joints, and somewhat thickened. It is difficult to ascertain
the correct proportions of the legs and their joints, due to the fact that
the legs are either uniformly swollen in regular fashion or their con-
tents have shrunken, faithfully reproducing them but with greater
thickness. There appears to be a gaseous envelope about these limbs
which, however, bears all the spines and bristles and armature and
pile of each joint. It is therefore difficult to know whether the dark
central contents have shrunken. Nevertheless, the pile and armament
can be faithfully and correctly ascertained. Wings: with the outer
marginal angles of the second posterior cell spurred, its cross vein
straight, strongly directed away from wing margin and only a little
longer than the last section of the fourth vein, the latter with mere
trace of spur. Apical cross vein quite sinuous basally, then sharply
drawn out toward tip of wing to join the third vein a short distance
before the apex. Third vein and costa end at tip of wing. Vena spuria
weak but not faint. Wings light brownish, stigma darker. The anterior
cross vein joining the discal cell one-third of the way from base, the
cross vein strongly oblique above the vena spuria.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype : in the University Museum at Konigsberg. The specimen
is No. B435 II and the slide is number 2435 (XII 2.200).

Myiolepta luhei Cockerell

"Probable length about 11 mm., wing 9 mm., or a fraction less; legs
rather robust, black, with dark hair; a dark cloud traverses the wing

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in the region of the forking of the veins 2 and 3, and above and below
(the same is seen, less developed, in the living M. varipes); and vena-
tion agreeing with M. varipes, Lw., in nearly all respects. The following
table brings out the venational characters:

"Second vein ending much nearer to third than to first

M . bella Williston

1. First posterior cell ending almost on margin of wing; outer side

of discal cell nearly straight; fourth vein more strongly bent near

end of discal cell M. luhei, n. sp.

First posterior cell ending some little distance from margin of
wing; outer side of discal cell bent inwards; fourth vein less

strongly bent near end of discal cell M. varipes, Lw."

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.
Holotype : Dr. Cockerell says "in the University Museum at Konigs-
berg." Since I was not able to locate the type of this species, I quote
the description of Cockerell above. I believe that this species can be
recognized on the basis of the dark cloud upon the wing.

Myiolepta woteni spec. nov.

Female. Length about 6 mm. Head: broad, rather less than hemi-
spherical; upper portion of the front rather broad; the ocelli in an
equilateral triangle ; the pile of front delicate, abundant, pale in color.
The face black without tubercle; the antennae blackish, rather large,
the third joint about one and one-half times as long as wide. Thorax:
black, with delicate, erect pile that is pale in color. There are no stout
bristles on thorax or scutellum. The scutellum is broad, with an un-
usually well developed crimped or emarginate rim and only fine, erect,
delicate pile. Along the margin the hairs are slightly longer. Abdomen:
completely obscured by the preservation and the wings folded over
it. It appears to be robust, about one and one-half times longer than
its width and dark in color. Legs: easily seen from one side; black in
color; the hind femora considerably thickened, at least twice as thick
as the middle of the slightly flattened and slightly arcuate hind tibiae.
This thickening of the hind femora is distributed over most of its
length and it is narrowed shortly before its apex and base. On the
ventral surface of the hind femora there is considerable short, stubby,
slightly appressed, black pile but no spines. Wings: much of the vena-
tion is obscured by the preservation, but the marginal and submar-
ginal, and first posterior cells and lower marginal and subapical cross
veins are visible. The marginal cell is widely opened; the subapical

hull: revisional study of fossil syrphidae 303

cross vein is long, with a gentle curve in its middle, directed proximally
and is confluent with the third vein almost at the apex of the latter at
the costa; the angle thus formed is quite acute. The last section of
the fourth vein from end of lower marginal cross vein to the beginning
of the subapical cross vein is quite long and is as long or longer than
the lower marginal cross vein. I see no trace of a vena spuria in the
lower portion of the first posterior cell but much of this cell is obscured
and there may have been a fold or a trace of one in the proximal
portion.

Locality : Baltic amber, Germany. Horizon : Lower Oligocene.

Holotype: one specimen no. 9080 in the Museum of Comparative
Zoology: Haren collection.

This species appears to be distinct from others on the bases of the
venation and the hind femora.

Genus Sericolepta genus nov.

Head: very large. Eyes strongly separated in the female. Front
prominent, flattened, somewhat concave. Arista short, third joint
large, a little longer than broad. Arista somewhat longer than the
antennae, basally thickened, pubescent. Face deeply concave below
the antennae, but the face itself is little produced. Thorax: longer
than broad, convex, scutellum large, semicircular, its disc and margin
convex, not thinned or impressed. Abdomen: quite robust, wider than
the thorax, widest on the second segment. Legs: hind femora stout and
elongate, a little the widest in the middle but not very narrowed either
basally or apically, the ventral outer half equipped with many spines,
much more than in Myiolepta. Tibiae about four-fifths as long as
femora, stout and ending transversely. Wings: with marginal cross
vein very long, the outer one strongly sinuous, joining the third vein
some distance from the end of the wing. Vena spuria weak, anterior
cross vein approximately one-third of the way from base, but quite
oblique.

Genotype: Sericolepta maculata spec. nov.

This genus is, I believe, a well marked one; though it is like
Myiolepta in many respects, the confluence of the very sigmoidal apical
cross vein some distance from the tip of the wing is quite unlike that
of the modern species of Myiolepta in the strict sense. Moreover, this
is an exceptionally large species, over twice as large as present-day
Myioleptas and without the flattened somewhat impressed scutellum
found in Myiolepta or the appressed setaceous pile. It is worthy of

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note, however, that two of the species that I describe and leave for the
time being under Myiolepta, namely andreei and germanica, are some-
what like the present species in that the apical cross vein does not
joint the wing at the tip and they are unlike both the present form and
Myiolepta in lacking any spines on the femora. Myiolepta valida,
appears to be a true Myiolepta beyond doubt. These other species and
subgenera, it seems to me, we must regard as more highly developed
inasmuch as the apical cross vein confluence point has moved more
basalward.

Sericolepta maculata spec. nov.

Plate 11, fig. 84, 85, 87; Plate 12, fig. 100, 109, 110; plate 13, fig. 119

Female. Length 13 mm.; length of wing 10 mm. Head: very large,
wider than thorax. Eyes conspicuous, thickest somewhat away from
the middle when seen from above. Front wide; eyes widely separated,
nowhere approximated. The front flattened, somewhat concave,
pilose only on the upper half. Somewhat protuberant at the level of
the antennae. Antennae about the middle of the head in profile,
short; third joint large, half again as long as wide, slightly pointed
apically though the end is actually rounded. The arista elongate,
less than half again as long as antennae, basal half strongly thickened
and pubescent. The head from above, behind the eyes quite concave.
The occiput at the top well developed, but not thicker than it is on the
sides, with much long delicate pile and four or five spines that do not
begin until near the middle of the head. Face below the antennae
strongly concave, the epistoma bluntly protuberant. Thorax: very
broad, somewhat longer than wide, thickly covered with very delicate
erect pile. The scutellum large, semicircular, very convex with a few
slender stiff hairs posteriorly, nowhere with macrochaetae. Abdomen:
half again as long as wide, very broad and robust, the second and third
segments black with large quadrate brownish yellow spots in the basal
corners widely separated in the middle. The fourth segment appears
to have been wholly dark, but it is difficult to say for sure. Abdomen
thickly erect, long, delicately pilose at least on the basal half. Legs:
hind femora stout and long, rather thickened especially right in the
middle. On the basal half the hind femora are brownish yellow; on the
outer half black, the outer half ventrally with very numerous short
spines not confined to a single row. Hind tibiae rather stout, less than
four-fifths as long as the hind femora, ending transversely. Wings:
very long, considerably longer than abdomen. The marginal cross vein
quite long, the lower one slightly sinuous, the outer one strongly

hull: revisional study of fossil syrphidae 305

sigmoid joining the third vein some way from the tip though not re-
motely and the third vein and costa ending at tip of wing. The basal
marginal angles of the first and second posterior cells with spurs. The
vena spuria weak, but visible throughout. The small cross vein
oblique above the vena spuria, and entering the discal cell about three-
sevenths of the distance from the base. Wings uniformly brownish.
Stigma darker.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum of Konigsberg. The number
on the specimen is 2B642 and on slide is 2642 and 229.

Genus Archalia genus nov.

Head: large. Eyes small. The antennae are not well preserved;
short and the third joint short and rounded. Face below the eyes
extensive apparently and in profile retreating below the antennae.
There is no evidence of a tubercle. Thorax: short and broad and
robust. Scutellum large, apparently semi-circular. Abdomen: thick,
convex posteriorly, very robust and with the terminal part broadly
rounded in what is probably a male hypopygium. Legs: hind femora
particularly short and massive, much as in the Syritta of today of an
altogether different subfamily. Ventrally before the apex there are
many small setae. Wings: venation well preserved. Small cross vein
quite basal the vena spuria strong and the apical cross vein very sig-
moid, and confluent with the third vein some distance from the tip of
the wing. Third vein and costa join the wing margin at the tip. Mar-
ginal cell broadly open.

Genotype: Archalia femorata spec. nov.

This appears to be an unique fly. In general it is well preserved, the
legs and wing particularly well preserved. The profile and details of the
antennae are poor. This is the specimen which Williston (1886)
referred to as related to Myiolepta. However, this relationship does not
appear to be at all close. The anterior cross vein more basal, the con-
fluence point of apical cross vein quite different and the massive
femora make it unrelated. Its dark coloration and basal cross vein
throw it into the Cheilosinae.

Archalia femorata spec. nov.

Plate 3, fig. B; Plate 4, fig. 10, 11 '

Male (apparently). Length 8.2 mm.; length of wing 5.6 mm. Head:
the specimen is placed laterally and a fairly good profile shows. The

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head is broad and short, and the front is round and bulging or promin-
ent despite the face that it is extremely short and steep. The specimen,
which is almost certainly a male had the eyes holoptic and the upper
facets are enlarged. Below the antennae the face appears to be pro-
duced ever so little as a steep shallow convexity and then to retreat to
the oral margin. The most remarkable peculiarity about the species
seems to be the small eyes, leaving the cheeks and occiput well devel-
oped. The antennae were short, the first and second joints subequal,
together a little longer than the third, and the third joint short and
rounded. There is only an extremely faint indication of an arista.
Thorax: dark in color. No details can be made out. The scutellum is
short, smooth rimmed and quite broad. No pile or bristles appear.
Abdomen: short and broad and dark in color, presumably without
pattern, though this cannot be ascertained with certainty. Five
visible segments and a large hypopygium. On the third segment there
are traces of thick, moderately short hair. Legs: hind femora enor-
mously thickened, about as in Syritta, with the same stoutness and type
of thickening apparent in that genus. There is a double row of short
heavy setigerous spinules along a ventral flange on the posterior two-
fifths of the hind femora. At the point where these begin, the two-
fifths way point, there is the not uncommon obtuse, outward produc-
tion. The hind tibiae were very slender and closely applied to the
femora, and were bent near the middle to correspond with the above
mentioned production of the hind femora. The other femora and
tibiae were slender. The legs appear to have been dark in color.
Wings: characterized by widely open submarginal cell and practically
straight third longitudinal vein with only the merest of sinuosities.
The subapical cross vein is beautifully and strikingly sigmoid. The
veins were quite heavy, and wing clear, and reaching a little beyond
the tip of abdomen. The third vein joins the costa at the tip of the
wing, and the first longitudinal vein joins the costa at an angle of about
seventy-seven degrees.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: No. 3941, in the Museum of Comparative Zoology.
(Scudder Collection).

Genus Palaeoascia Meunier

Ann. Soc. Ent. France for 1893, 259 f. (1893). Ann. Soc. Sci. Bruxelles, 19: 7
(1895)

Head: large, subglobular. Eyes occupying most of the head; broadly
holoptic in the male. Front small, a little bit convex. Antennae short,

hull: eevisional study of fossil syrphidae 307

located in the middle of the head in profile, third joint not large. Arista
a little longer than antennae, basally thickened. Face below antennae
with well developed but small tubercle. Face below the tubercle
descending a short distance to the epistoma, which is not more pro-
duced than the tubercle is. The occiput extensive throughout in profile
in contrast to Palaeosphegi?ia. Thorax: longer than broad, very convex,
with a few short spines on the sides above the wings as in Palaeos-
phegina. Scutellum hemispherical, rather considerably broader than
long, the apex with a single pair of widely spaced very strong tubercu-
lous bristles and occasionally with a smaller pair on the outside.
Abdomen: elongate, about two and one-half times as long as wide,
rather thick, and somewhat convex on the surface, covered with some-
what scattered short appressed bristles. Male hypopygium large and
rounded. Legs: the hind femora elongate, a little thickened especially
on the dorsal surface, somewhat more slender than in Palaeosphegina
in the males, with a double row of very long slender sharp pointed
spines which reach practically to the extreme base of the femora. The
outer lateral surface of the hind femora just past the middle with two
to four similarly long sharp spines. Hind tibiae sometimes with a few
spines near the middle of the outer surface. Absent in the type. Wings:
with the venation very much like that of Palaeosphegina; so much so
that I am at a perfect loss to find any difference not covered by the ex-
treme venation of that genus. However, the wings of Palaeoascia do
not seem to be quite so variable as Palaeosphegina; on the other hand
they appear to be larger for the respective size of the insect.

Genotype : Palaeoascia uniappendiculata Meunier.

The genus constantly differs from Palaeosphegina in the matter of
the swollen and tumid occiput. In addition the hind femora of the
male is spinose on the basal half as well as distal half, the lateral
femoral spines are constant in Palaeoascia but I believe that they some-
times occur in Palaeosphegina but rarely.

Palaeoascia uniappendiculata Meunier

Plate 5, figs. 16-20; Plate 7, figs. 43-48; Plate 8, fig. 51

Jahrb. Preuss. Geol. Landesanst., 24: (2) 201. (1904)

Male. Length 5 mm.; length of wing 4 mm. Head: front, vertex,
face and occiput flat, the front and the upper part of the face quite
thickly pilose and this is in contrast to Palaeosphegina. The antennae
located below the rather convex front, directly in the middle of the

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head, are short, the third joint is about one and one-half times as long
as wide with bluntly rounded apex and the color, including the arista,
light reddish brown. Thorax: dorsum of thorax black. Scutellum
reddish, the pile of the former is sparse but more abundant than that of
Palaeosphegina, is quite erect and light in color. Abdomen: pigment
of abdomen is destroyed in one type and largely obscured in another;
in the third specimen the whole color is somewhat pallid, perhaps
teneral. Nevertheless it is easily possible to see that the abdomen is
banded with dark brown or black on the bases of all of the segments
and a little bit more narrowly on the posterior margins. In one speci-
men, No. K5098, the whole of the terminal segment appears to have
been black. "Legs: hind femora definitely black on the basal half in
specimen No. 2407K, and the base of the femora in this one yellow,
the base of the tibiae, the basal two-thirds of the remaining tibiae and
all of the fore and middle tarsi pale yellow. In the other specimens the
color of the legs is somewhat uncertain and the femora appear to have
been unicolorous.

Of the females it may be said that the front near the vertex is much
narrower, as can be seen from Plate 9, fig. 35.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Types : in the University Museum at Konigsberg, from Baltic amber.

One male, No. K5098, 72247 (K5098-Klebs) ; another male which I
judge is a type as it is labelled in Meunier's handwriting, but the
number is not one of those given by him,— No. 2497k (2407-Klebs) ;
and two females No. Z2086, K2631, (2631-Klebs); and 21692, K263B
(2632-KIebs). The following twenty-six females have been deter-
mined as belonging to this species: No. B14 (14649-Klebs); No.
11B856 (2856, VII. 2.282-Klebs) ; specimen unnumbered (V) ; unnum-
bered (0) ; unnumbered (P) ; unnumbered (Y) ; unnumbered (I) ; No.
3345 (345.VIL 2.217-Klebs); No. XX134676 (24676-Klebs) ; No. B
19985(G); No. B446D (2446-VII. 2. 210-Klebs); No. B433 (433-VII.2.
212-Klebs); No. 5B393 (393 VII.2.209-Klebs); No. IIB546 (2546 VII.
2.202-Klebs) ; No. XB4555 (24555-Klebs) ; No. X127 (unmounted);
No. X151 (unmounted); X144 (unmounted); X337 (unmounted);
K 1930 (unmounted); K1939 (unmounted); K1984 (unmounted);
No. 11 B 369 (2869); No. B 14681 (14681); No. B 4683 (24683) (front
of this specimen very narrow at the top) ; unnumbered (Q) (I find this
an extremely small specimen, which may be a distinct species. Length
4 mm.). The following ten males belong here: No. 3B593 (3995 VII
2.457-Klebs) ; No. 3B404 (3494 VII.2.219-Klebs) ; No. 492 unmounted)
No. IIB363 (H 109); No. 11B785 (12785-Klebs); X4485, XB (4485-

hull: revisional study of fossil syrphidae 309

Klebs); B14678 (14678-Klebs); unnumbered (W); No. 11B899 (2899
VII. 2.220-Klebs); No. 11 B 656 (2656).

In the Geologisch-Palaeontologisches Institut and Museum der
Universitat Berlin are seven females as follows: T. M. B. 3 (H 201);
specimen unnumbered (H 202); T. M. B. 10 (H203); 2 (H 206); 4
P.M.B. (H-208); P. M. B. 5 (H-207); P. M. B. 6 (H-210).

In the British Museum of Natural History are two females : no. 22194
and 22196 from the Loew collection, marked Miocene amber and with
his label of 'Syrphici', and one male No. 22197 with the same designa-
tion, except the accompanying slip marked 'Syrphus'.

Palaeoascia uniappendiculata brachypennis var. nov.

Female. This specimen I designate as a variety of Palaeoascia uniap-
pendiculata on the basis of the broader and shorter wing, the propor-
tions of which differ. The subapical cross vein is more sigmoid and the
wing is a little bit more brownish. The legs are uniformly darker in
color, nowhere light yellowish. Facial tubercle black. Pile of front not
thick.

Holotype: in the University Museum at Konigsberg. No. B395 (395
VII.2. 216-Klebs).

Palaeoascia uniappendiculata obtusa var. nov.

Female. Length 3.5 nim.; length of wing 3.2 mm. In this form the
tubercle of the face, while present, is so extremely blunt and perfectly
straight between tubercle and antennae, suggesting the manner in
which it is formed in Palaeosphegina, that I think it merits varietal
distinction.

Holotype: a specimen in the University Museum at Konigsberg.
The specimen itself is without number; the slide is No. 403 VII. 2. 210
Klebs).

Palaeoascia atrata spec. nov.

Plate 5, fig. 21-24.

Female. Length 5 mm.; length of wing 4.3 mm. Head: this is a
very black species sharply distinguished, in addition to the jet black
coloration of the entire face, tubercle, antennae and head, by the much
more thickly and somewhat long pilose front and vertex. The face
also appears to be more pilose and especially below the tubercle and

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on the sides. The arista is strongly swollen on the basal fourth but
not sharply swollen. The third joint of the antennae is at least half
again as long as wide, perhaps a little more broadly rounded. Thorax:
jet black, scutellum almost as dark. Abdomen: banded but quite dark,
the basal fascia occupying nearly a third of the segment and no apical
fascia on the fourth and fifth segments. Legs: dark brown or black.
Wings: deeply tinged with brown throughout.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum at Konigsberg. The specimen
is labelled B27261 (B27261-Klebs). Paratypes: a specimen unlabelled,
slide No. 'B', Konigsberg; another specimen a male, British Museum
of Natural History. No. XIII B327 (13327-353 In 18666).

I associate the following four specimens from Konigsberg with the
species atrata, basing my conclusion on the similar blackish color, dark
brown wings, long bristly pilose scutellum and thickly pilose front and
blackish antennae. Two of them are males as follows: No. 11B885
(2885 VII. 2.197-Klebs); without number on the specimen (H 110-
Klebs). The two females bear these numbers: the first XIIB788
(XIIB788); the second, lacking number on specimen but with the
affixed letter of X, 2H111).

There is a large male, No. 3B620, which I believe is abundantly dis-
tinct from atrata, but because of the great variability of these flies, I
prefer to leave it under atrata for the present. In the first place the
antennae are much shorter, dark brownish, the third joint scarcely
longer than wide, obtusely rounded, very thick and long pubescent.
The arista sharply swollen on the basal fifth. The face and head
everywhere jet black, the front rather thick, long pilose. The tubercle
prominent, concave, and face above tubercle very hairy. Thorax
and scutellum jet black and the scutellum in particular, in addition
to the strong pair of heavy long spines, has a number of stiff, quite
long black bristles on the disc. Its length is 5.2 mm. ; of wings 4.2 mm.

The full number of this aberrant specimen which is in the University
Museum at Kongisberg is No. 3B620 (3620 VII. 2.456 Klebs).

Palaeoascia nigra spec. nov.

Plate 7, fig. 49

Female. Length 4.8 mm.; length of wings 4.2 mm. Head: this speci-
men is distinguished by the very black coloration, sharp prominent
tubercle on the face, the third antennal joint is one half again as long
as wide, but is light reddish brown. This arista swollen on the basal

hull: re visional study of fossil syrphidae 311

fifth rather sharply. The pile of the front is fairly long but certainly not
as thick as found in atrata and the pile of the face is very sparse indeed.
Thorax: jet black. Scutellum with only two strong spines and no long
stiff bristles on the dorsum. Abdomen: is sharply banded with black
on the third and fourth segment, and on the apex of the second seg-
ment and the base of the fifth segment. Legs: hind femora small,
quite short, evenly thickened above and below but mostly in the
middle, a little spindly at base with only two lateral spines and these
are near the apex and very long and stiff. Only the apical third or
two-fifths at most, of the hind femora are black and only the distal
two-fifths of the hind tibiae. The middle femora pair of legs are en-
tirely light yellowish brown and the anterior pair practically the same.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum at Konigsberg. The specimen
is unnumbered, the slide is No. (N. H108).

Genus Spheginascia Meunier

Allg. Zeitschr. Ent. (B) 6: 71. (1901). Jahrb. Preuss. Geol. Landesanst., 24: (2)
205 (1904)

Small sized flies with a robust abdomen much like Cheilosia.

Head: large, eyes conspicuous, broadly touching in the male. Vertex
and front long bristly, the latter with a deep median crease. Antennae
short, third joint half again as long as wide, broadly rounded, very
short pubescent, the arista quite long, thickened on the basal third
but not abruptly, the apical bristles of the second segment quite long.
Front above the antennae in the male convex. Face below the an-
tennae in the male for a short distance concave, then rising to a strong
and prominent bulge, broadly rounded. This bulge or broad tubercle
descends equally abruptly to the epistoma. Face nowhere deeply pro-
duced. Face in the female below the antennae concave, only without
the tubercle. The epistoma a little bit thrust forward, sharp at the
edge. Occiput visible in profile only in the female. Thorax: scarcely
longer than broad, convex, long sparse pilose. Scutellum at least twice
as wide as long, the margin almost semi-circular, the disc and rim con-
vex, although the scutellum is not thick. Abdomen: twice as long as
wide, widest at end of second segment, barely less than half as wide
at the end of fourth segment. Hypopygium large, broadly rounded
from above. Abdomen rather long pilose, especially on the sides.
Legs: hind femora slender, long bristly pilose on the sides and above
and ventrally with many long stiff bristly spines, sharp pointed but

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nowhere thick, not definitely confined to rows. Wings: characterized
by the marginal cross vein being strongly and obliquely directed away
from wing margin. Apical cross vein a little bit sigmoid joining the
third vein remote from tip; never recurrent. Both the marginal angles
of the first and second posterior cell with long spurs. Vena spuria
present.

Genotype: Spheginascia biappendiculata Meunier.

SpHeginascia biappendiculata Meunier
Plate 7, Fig. 50; Plate 9, Fig. 61, 66, Plate 10, Fig. 70, 71
Jahrb. Preuss. Geol. Landesanst., 24. (2) 205. PL 13, fig. 4 and 5 (1904)

Male. Length 5 mm.; length of wing 4.5 mm. Head: eyes large,
bare. The face, front and vertex dark brownish black. The antennae
and arista very dark brown. The arista is strongly thickened on three-
fourths of its length, more so at base. The pile of the front and face
is black. Thorax: is black with some evidence of having been aeneus
in life. The bristles on the dorsum of the thorax are very long, scat-
tered and delicate on the upper part of the pleura. Before the wing
there are several long stout stiff bristles and a few others on the pos-
terior calli. They are not to be compared to the stiff spinous bristles of
flies in the living genus Ferdinandea. The scutellum black, with two
pairs of long stiff tuberculous bristles near the apex and some long,
very sparse, slender, bristly hairs on the disc. Abdomen: uniformly
dark brown, perhaps a little bit darker on the posterior segments of
the abdomen and a little bit lighter on the dorsal half. Abdomen a
little wider than thorax, the pile quite sparse but conspicuous, long and
bristly. Legs: hind femora dark brown, a little lighter near the apex.
All the tibiae dark apically and light yellowish brown basally. All the
tarsi brown or fuscous.

Female. In the female the eyes are unusually broadly separated and
the face is concave.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Types: in the University Museum at Konigsberg. One male, No.
Z3874 (K2549, Z3874-Klebs). One female No. K4233, Z587 (K4233-
Klebs). Paratypes : there is one male in the Geologisch Landesmuseum,
Berlin from Baltic amber with no number on the specimen, the slide
bearing the number (101).

I have determined the following specimens from the University
Museum of Konigsberg: four males— No. 111B366 (3266 VII.2. 195-

hull: revisional study of fossil syrphidae 313

Klebs) ; one specimen unlabeled, slide bearing the letter M, I affix the
number H 112. No. IIB468 (2469 VII.2.250-Klebs); 11B563 (2563
VII. 2.203-Klebs). Six females— No. SB368 (368 VII.2.208-Klebs);
No. 547 (unmounted); No. XB6335 (26335-Klebs) ; No. XIIIB311
(13331 -(Klebs); No. X60 (unmounted; and one specimen without
number, the slide bearing the number (13548 VI, 22, 7781-Phys. Oek.

Ges.).

In the Geologisch-Palaontologisches Institut und Museum der Uni-
versitat, Berlin there are two females-— No. 9 (H 205); and No.
7 P M B (H 209), and one male No. H 211. The British Museum of
Natural History has three females— No. 22200, 22201 and 22202,
which are from the Loew collection. In the American Museum of
Natural History one Male No. 502-129.

Spheginascia biappendiculata rectinervis var. nov.

Male. Length about 5 mm.; wings 4.5 mm. This variety differs
from the typical form in respect to the venation of the wing and the
details of the scutellum. The scutellum has three pairs of strong, long,
black bristles on the margin and one or two pairs of shorter ones near
the base on each side. In the wing the angle formed by the lower
marginal cross vein with the last portion of the fourth longitudinal
vein is considerably more obtuse, the last section of the fourth longi-
tudinal vein between the two cross veins is straight instead of curved
downward, as is also the lower marginal cross vein.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: one specimen no. 9076, in the Museum of Comparative
Zoology (Haren collection).

Genus Pseudosphegina genus nov.

Head: broadly subglobular in the male with the eyes narrowly sepa-
rated a distance a little more than the width of the third antennal
joint; no where approximated. Anterior facets a little enlarged. Eyes
bare, the antennae set above the middle of the head in profile, small,
the third joint a little longer than wide. Arista short, basally thickened
and bare. Face below the antennae straight for a distance down to
the conspicuous tubercle which lies nearly opposite the level of the
bottom of the eyes. Face past the tubercle abruptly retreating, to the
epistoma. Cheeks quite small. Occiput not visible in profile. The
flanges of the lower part of the posterior surface of the occiput very

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strong and conspicuous. Thorax: a little longer than wide, somewhat
convex, almost bare with a few scattered short bristles. Scutellum
small, wider than long, nearly bare with two strong apical bristles and
a weaker pair on either side of these. Abdomen: elongate, about three
times as long as wide, relatively thin at base and considerably thicker
at apex due to the development of the hypopygium and genitalia.
Legs: hind femora quite simple and slender with five or six well de-
veloped, spines ventrally on the outer half. Wings: rather longer than
abdomen, not wider at base than in the middle. Apex somewhat
rounded, the venation quite like that of modern Sphegina except that
the vena spuria is quite absent.

Genotype: Pseudosphegina dichoptica spec. nov.

The genus Pseudosphegina is certainly of peculiar interest because
it is in every way like modern Sphegina except that the face is tubercu-
late, whereas in all living Sphegina the face is concave and the face
thrust forward. The genus Palaeosphegina of Meunier is somewhat
misnamed for it is even less like modern Sphegina; in addition to having
the face tuberculate the males were strongly holoptic and as is well
known, the true Spheginas have only dichoptic males. Thus, out of
the four possible combinations three of the combinations are known
to have developed. Two of these exist as fossils. The third form
Sphegina is not known positively to exist as a fossil, although I have
placed a specimen in the genus Sphegina from the Florissant, from
which specimen it can not be discerned whether the face is tuberculate
or developed as it is in modern Sphegina. It may well be that the
specimen of the Colorado form that I have placed in Sphegina is a
true member of that genus but we must wait for other specimens to
know what its face is like. The known combinations of face and eyes
are as follows:

a. Face tuberculate; males dichoptic {Pseudosphegina).

b. Face tuberculate; males holoptic {Palaeosphegina) .

c. Face non-tuberculate; males dichoptic {Sphegina).

Pseudosphegina dichoptica spec. nov.

Plate 6, Fig. 27-28; Plate 7, Fig. 38, 39

Male. Length 5.5 mm. ; wings 4 mm. Head: the face, front and ver-
tex black. Antennae and arista black, third joint of antennae a little
pointed at apex, one and one-half times as wide as long. Arista half
again as long as antennae, the pile of the occiput short, confined to a
single row of collar-like bristly hairs. Thorax: black, pleura brownish,

hull: revisional study of fossil syrphidae 315

the dorsum of the thorax exceptionally sparsely pilose, with only a
scattered bristle here and there. Scutellum black, abdomen with the
second, third and base of the fourth segment dark brownish, the
remainder of the abdomen and the prominent rounded hypopygium
shining black. Apical portions of genitalia light brown. Abdomen:
everywhere thickly appressed bristly. Bristles of hypopygium erect
and short, the last sternite thick bristly, the preceding ones with very
few bristles. Legs: hind femora blackish brown on the apical half,
paler brown basally, the outer half with a few stiff appressed bristles
and five or six long stiff sharp pointed bristles ventrally. Wings:
with the marginal angles of the first posterior cell long spurred, the
corresponding angle of the second posterior cell with a merest trace
of a spur. The lower marginal cross vein almost joining fourth vein
at right angles, and last section of fourth vein before the origin of the
apical cross vein quite long, longer than the apical cross vein itself.
Apical cross vein somewhat curved or sigmoid, on its outer half joining
the third vein at right angles, the last section of the third vein about
five-sixths as long as subapical cross vein. Third vein and costa end
at tip of wing. Wings uniformly pale brownish, the stigma darker.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum at Konigsberg. The amber
bears no label. The slide upon which it is mounted bears the label of
'J'. I affix for future designation the number H 101 to the slide.

PSEUDOSPHEGINA WITHERSI Spec. nOV.

Plate 11, fig. 81, 82; Plate 13, fig. 115

Male. Length 6 mm.; length of wing 4.7 mm. Head: large, much
wider than thorax. Eyes extensive, very narrowly separated indeed,
more so than in dichoptica. Front and vertex and occiput, except
narrowly below, dark in color. Face, antennae and cheeks light
yellowish brown to brownish orange, the face a little darker, the arista
quite pale yellowish. Tubercle well developed, pile of head everywhere
pale, including the finely pilose occipital fringe. Thorax: convex,
dark in color apparently aeneous, almost bare. Scutellum with two
very stiff thick apical bristles; smaller bristles apparently absent and
the dorsum almost bare. Abdomen: dark on the first and almost the
whole of the second segment except for the apical band. Anterior half
or more of third segment dark and the whole of the terminal segment.
Hypopygium particularly large and broad and dark. Abdominal pile
pale. Legs: hind femora brown, darker apieally with few black spines,

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four of them exceptionally long. Distal three-fifths of the hind tibiae
dark, the whole of the middle pair of legs and all of the front pair of
legs except the anterior basal two-thirds of the fore femora light yel-
lowish brown. Wings: distorted but the apical cross vein very slightly
curved and the last section of the fourth vein definitely curved out-
ward slightly.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the British Museum of Natural History (Loew col-
lection). The specimen is unnumbered, the box No. (22270).

Genus Palaeosphegina Meunier

Jahrb. Preuss. Geol. Landesanst., 24: (2) 204. (1904)

Head: broadly hemispherical, the eyes extremely large occupying
almost all of the head, broadly touching in the male, rather widely
separated in the female, the width of the front above the antennae
not quite twice as wide as width across ocelli. Ocelli at top of head in
a nearly equilateral triangle. Front with only a very few scattered
hairs and a small rounded convex tubercle above the antennae. In
profile the back of the head from the ocelli down to the vertical
portion of the head is very broadly rounded, the antennae are barely
situated below the junction of the middle and upper thirds of the head.
Antennae short, third joint large. Arista not quite twice as long as
antennae, very slender at tip, strongly thickened at base. Face with a
prominent tubercle that descends below a little bit more abruptly than
the rise of the tubercle above. The tubercle is almost opposite the
lower third of the eyes. Face very little produced below the tubercle,
scarcely as long at the point of junction of cheek and face as the width
of the third antennal joint. The occiput scarcely visible in profile but
somewhat narrowly below, equipped with an occipital collar or fringe
of pile of several rows of hairs. Thorax: barely longer than broad, very
convex; it is practically bare or with scattered short hairs. The scutel-
lum is large, semicircular, convex on disc and rim, with two to six
very strong long bristles on the rim, the outer ones shorter. The
scutellum seems to vary considerably in shape from circular to ob-
tusely pointed. Abdomen: elongate, about three times as long as wide,
widest distally on the fourth segment, the abdomen being spatulate
rather than petiolate in most of the forms, sometimes flattened and
again strongly convex. The pile of the abdomen is appressed, short
bristly; the metanotum is conspicuous. Legs: the hind femora are
always elongate, always a little thickened and this thickening is usually

hull: revisional study of fossil syrphidae 317

past the middle ; occasionally it is in the middle (specimen No. X 153).
Without exception the distal ventral half of the femora is equipped
with a double row of sharp stout long spines, and it should be noted
that these spines never encroach on the basal half except as very
slightly overgrown appressed bristles in contradistinction to Pal-
aeoascia, in the male of which the spines are as strong on the base as
they are apically. This is far from being an important difference, but
it is one distinction between the two genera and I think it shows three

things :

A. That the spinal armament originated from ordinary bristly setae.

B. That it began at the tip of the femora in all cases and moves

basalward in a varying degree.

C. And lastly, that it has enjoyed a much greater impetus of de-

velopment in the male sex.

The hind tibiae end transversely; the middle tibiae end with a series
of blunt stout spines of which one is sometimes much longer and
stouter. Wings : varying from elongate and slender to quite broad and
short. There is always this uniform feature of venation that the mar-
ginal cross veins are strongly obliquely directed away from wing mar-
gin, leaving the last section of the fourth vein before the end of the
wing, varying over one hundred per cent but never recurrent. Some-
times the apical cross vein is perfectly straight; more often sigmoid and
occasionally quite strongly sigmoid. The vena spuria is absent but it
must be remembered that it is quite faint in present day Spheginas.
The males differ in no essential particular from the females, except for
the strong holopticism, the prominent hypopygium and genitalia
resulting in a little more petiolate abdomen.

Genotype : Palaeosphegina elegantula Meunier.

Palaeosphegina elegantula Meunier

Plate 9, fig. 63-65

Jahbr. Preuss. Landesanst. 24: (2) 204, PL 13, fig. 2 and 3. (1904)

Female. Length 8 mm.; length of the wing 6.2 mm. Head: face,
front and cheeks dark coppery brown to black. Antennae and arista
light orange. The facial tubercle is large, evenly rounded, not quite as
much produced as the cheeks. Occipital pile dense and light brownish
in color. The third joint of antennae is one and one-fourth times as
long as wide. Thorax: dark coppery brown or reddish above ; there are
faint indications of a trivittate condition. Scutellum brownish red,

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lighter than the mesonotum, the spines of the thorax above the wing
are quite short, the pleura are lighter in color. Abdomen: is light
reddish or orange brown in color, in the type obviously banded but in
this specimen the wings are matted over the abdomen and largely
obscure it. Data taken from other specimens show that the basal
part of each segment contains a narrow black or dark brown band of
variable length and interrupted in the middle in more or less variable
fashion. Sometimes the posterior portion of the segment is very
narrowly dark fasciate as well but not interrupted. Hind femora dark
reddish brown, nearly black on the outer half, much lighter in color
on the basal two-fifths or half and very narrowly just before the apex.
The hind tibiae a little darker on the distal half. All the tarsi reddish,
the middie tibial spurs contain one very long stout spine. Wings: of
average length in contrast to the extremes. The marginal angles of the
first and second posterior cells both with very short spurs, or the dis-
tance from the confluence point of the apical cross vein to the end of
the wing is about four-fifths as long as the apical cross vein. The last
section of the fourth vein before the origin of the apical cross vein is
longer than the apical cross vein. These remarks are chiefly taken from
the type specimen on which the species is primarily fixed and I have
indicated in the discussion of the genus the principal nature of the
variations which are encountered in the individuals of this genus.
There are probably several species involved in this complex, but in
view of the tremendous variability I have had very little success in
isolating these and in correlating the numerous differences. I prefer
to leave them for the present as a group of variable individuals.

Locality : Baltic amber, Germany. Horizon : Lower Oligocene.

Type: in the Preussische Geologische Landesanstalt in Berlin. The
specimen, labelled G. L. 99, was studied. There is supposed to have
been another type at the University of Konigsberg, which, however,
I was not able to locate; but there certainly were other numerous speci-
mens in their collections, all of which I have before me at time of writ-
ing. These total thirty-three specimens not including one each of two
varieties nor the specimen of uncertain determination.

The specimens from the University Museum at Konigsberg are as
follows :

Females, specimen IIB857 (2857 VII. 2.284-Klebs) ; No. 14708
(14708-Klebs); No. B901 (VII. 2. 217-Klebs); No. B14285 (14285-
Klebs); No. IIB662 (2662.Vii.2.287-Klebs); No. X122 (unmounted);
No. X161 (unmounted); No. X120 (unmounted); No. X164 (un-
mounted); X529 (unmounted); No. X 124 (unmounted); No. K2416-

hull: revisional study of fossil syrphidae 319

343 (unmounted); No. X IIB786 (13786-Klebs); No. 16562 (16562);
No. XB4365 (24365-Klebs) ; No. 14287; No. K 7871; the following
nine specimens have no number of the specimen but the slides are
designated thus: No. (E); (R); (H); (A): (340); (H 106); (T); (Z);
(L). Seven males: No. IIB915 (2915 VII.2.195-Klebs); No. IIB900
(2900 VII.2.922-Klebs) ; No. X153 (unmounted); No. X 331 (un-
mounted); No. X 336 (unmounted); No. X111B228 (13228-Klebs);
no number on the specimen, the slides labeled (U). There is also at the
Geologische-Palaontologisches Institut und Museum der Universitat
Berlin one male No. T. M. B. (H 204), and one female at the British
Museum of Natural History No. XIIIB229 (13229, 351, In 18664),
and one female at the American Museum of Natural History; all of
these have been studied by me.

With regard to the following specimen, No. B14138 (14138-Klebs),
I cannot positively decide whether or not it is a male or a female ; if it is
a female it would go into the genus Paleosphegina since it is definitely
dichoptic. Should it be a male, on the basis of the hypopygium (which
is obscure) it would go into the genus Pseudosphegina. The evidence
which is available from the terminal portion of the abdomen indicates
a female. Specimen at the University Museum, Konigsberg.

Palaeosphegina elegantula tristis new variety

Female. I designate this as a variety based on the extremely narrow
front. The antennae are dark brown and the quite short arista sharply
thickened on the basal third. The front and vertex are almost black;
Bcutellum black; the legs dark brown, with the tarsi somewhat lighter.
The hind tibiae pale on the basal half.

Locality : Baltic amber, Germany. Horizon : Lower Oligocene.

Type: specimen without number in the University Musetfm at
Konigsberg; the slide is numbered F H107.

Palaeosphegina elegantula atrox var. no v.

Male. Size large. Head: face, front and cheeks jet black. First an-
tennal segment black, third very dark brownish. Arista sharply and
heavily thickened on the basal fifth and quite pubescent on the basal
fifth. Abdomen: with first two segments and the base and apex of
third and of all the very long fourth segment and basal part of hypopy-
gium jet black. The third segment has a very brown yellowish band.
Legs: the hind tibiae sharply jet black only on the apical third. Hind

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femora jet black past the middle except narrowly at the apex. Re-
mainder of the legs entirely pale yellowish.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Type: specimen in the University Museum at Konigsberg, number
11B891 (2891 VII.2.215-Klebs).

Palaeosphegina pilosa spec. nov.

Female. Length 5.2 mm. ; length of wing 4.2 mm. Head: very large,
much wider than thorax; everywhere, face antennae and the arista jet
black. The pile of front pale brownish, nowhere thick. Face apparently
with a little short pile above the tubercle. Tubercle rather large,
formed as in elegantula. Thorax: and scutellum shining jet black.
Pile of both of these very sparse, nearly bare, a few long black bristles
before the wing, and scutellum, with two pair of unusually thick stiff
bristles which are long, the inner pair widely spaced, and in closing a
pair of very short bristles. Outer pair shorter. Abdomen: broadly
petiolate, jet black with black appressed short bristles. Legs: every-
where jet black, the bristles on the femora confined to the apical half,
one long and one short bristle on the outer lateral surface of the hind
femora, a short distance from the end. Wings: very brownish. Stigma
extremely dark. Apical cross vein rather sigmoid, both the marginal
angles of the first and second posterior cells spurred.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype : in the University Museum at Konigsberg, number B331
(331 VII 2.206-Klebs).

This species has the occiput and facial tubercle quite as expected
for Palaeosphegina. It is a slightly smaller species, the form a little bit
more suggestive of Palaeoascia and the lateral femoral spines suggest
Palaeoascia. From elegantula Meunier it is at once distinct in the
totally black coloration, besides the smaller size and the arrangement
of the spines on the scutellum.

Palaeosphegina baccha spec. nov.

Female. Length about 4 mm. Head: large, considerably wider than
thorax; hemispherical in shape; the occiput slightly swollen when
viewed laterally and broadly pilose. The vertical posterior ocular
margin straight and not indented. Ocelli set in approximately an
equilateral triangle; the smooth, short pilose front widening gently
down to the antennae which are set above the middle of the head in
profile. There is a small tubercle on the face below the antennae. The

hull: revisional study of fossil syrphidae 321

first and second joints of the antennae are quite short, the third joint
is broadly rounded, a little longer than wide with a slender dorsal
arista that is slightly thickened on the basal third. Thorax: black,
sparsely short pilose, scutellum of the same color, with a pair of rather
widely separated blackish bristles which are about as long as the length
of the scutellum. Abdomen: elongate and rather flat, slightly crumpled.
Five segments are present; the third, fourth and fifth have a dark
colored transverse band or fascia lying across the base of the segment,
its width comprising about two-fifths the length of the segment. Upon
the third segment the lateral width of the fascia appears to occupy a
little more than half the length of the segment and on this same seg-
ment the fascia appears to be interrupted in the middle truncately.
The nature of the first and second segments can not be ascertained.
Legs: light brownish yellow in color, the hind femora are simple and
slender. Wings: the apical third of both wings is destroyed. From
the remainder, the venation can be described as follows: a wing sug-
gesting Palaeosphegina in the absence of the spurious vein and in that
the last section of the fourth vein is quite remote from the subapical
cross vein, in fact, almost recurrent; the lower marginal cross vein is
quite straight, making a right angle with the last section of the fourth
vein and about a 70 degree angle with the fifth vein.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: one specimen no. 9029 in the Museum of Comparative
Zoology (Haren collection).

There can be very little doubt of the family affinity of this specimen,
although the disposition of the subapical cross vein and of the second
and third longitudinal veins can not be determined. The preserved
portion of the right wing is a little greater and a little longer than that
of the left one and is truncated obliquely from along the costa just at
the end of the first longitudinal vein. All of the outer half of the sub-
costal cell dark brown ; the venation of this wing is not greatly different
from that of other specimens of Palaeosphegina except that the discal
cell is very much longer than usual ; the lower marginal cross vein more
recurrent; the fly itself appears to be rnore slender; the head perhaps
larger and the hind femora appear to lack the slender bristly spines
characteristic of Palaeosphegina.

Palaeosphegina fumosa spec. nov.

Male. Length 5 mm.; wing 4.3 mm. Head: hemispherical, entirely
black in color, eyes touching for about half the length from vertex to

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antennae ; the pile of front and upper part of face above tubercle black
in color, rather abundant and dense; the occipital pile is also black.
Tubercle of face small but well developed ; eyes bare ; the antennae dark
brownish black, the third joint about one and one-third times as long
as wide; the black dorsal arista thickened on the basal fifth; slopes of
the face below the tubercle without pile. Thorax: and scutellum black
with sparse delicate pile. The scutellum is rather long, apparently
almost as long as wide, with a pair of rather close-set, long, black
bristles arising from the apical margin. Abdomen: the color can not be
positively ascertained ; the first three segments are almost wholly dark
brown and translucent, as is much of the right half of the fourth seg-
ment, but on the greater part of the fourth segment there is an exten-
sive black pigmented area and there appear to be traces of such pig-
ment on the more obscured portions of the basal segments, hence it is
entirely probable that the abdomen was black in color; it is overlaid
by the smoky right wing. The sternites of the fly are certainly brown-
ish yellow in color. The hypopygium appears to have been large and
well developed with lobes and processes, resembling Recent Sphegina
in this respect ; these structures can not be determined because of over-
lying, glistening air spaces. Legs: dark reddish brown, the hind femora
slender, very little thicker than the other pairs; apical three-fifths
black in color and the lateral, ventral margin equipped with ten widely
spaced, rather long and slender black bristly spines which begin not
far from the base of the femur. Wings: venation beautifully pre-
served ; the entire wing smoky grey ; stigmal cell yellowish brown ; the
subapical cross vein joining the third vein at right angles with a
rounded bend just before it joins the third vein and its initial section
straight. The first section of the fourth vein straight and quite long;
a little longer than the lower marginal cross vein and equal in length
to the last section of the third vein, running from wing tip back to
subapical cross vein.

Locality : Baltic amber, Germany. Horizon : Lower Oligocene.

Holotype: one specimen, no. 9075 in the Museum of Comparative
Zoology (Haren collection).

Sphegina abrasa Theobald

Les insectes fossiles des terrains oligocenes de France, 1937, p. 243; pi. 18,
fig. 12.
Head: The base of the head preserves some traces of black; it is
semi-circular and as large as the thorax. The eyes are bare, large,

hull: revisional study of fossil syrphidae 323

projecting and of an oval shape. The face is flattened or depressed and
the antennae are narrowly visible. Thorax: The body of this insect is
of a reddish yellow coloration. The thorax is elongate, subrectangular,
the corners rounded, the scutellum rounded posteriorly. Abdomen:
clearly demarcated from the thorax, swollen and inflated into a club
which is more extensive in length than the thorax ; four segments are
visible, the fourth of a darker color than the preceding ones. The
body is covered with a fine pubescence. Legs: slender, finely pilose;
the hind femora are swollen and carry two rows of spines. Wings:
the venation is well preserved except towards the apex of the wing.
The subcosta ends toward the middle of the anterior border; the R is
elongate, the Radial sector is bifurcate, joining itself to 'M' through a
transverse vein ; 'M' bifurcate, "une cellule discoidale", the anterior
branch of 'M' joins itself to 'Rs'; 'Cu' bifurcate, joining itself to 'M'
(v. Figure). Dimensions: entire length- 8 mm.; wing- 5 mm.

Relationships: the venation of the wings is that of the Syrphidae.
In the genus Sphegina the third femur is swollen and arme Brun. of
India has the wings of the same color.

The author notes that Foerster has described Syrphus reciprocity in
slabs of Brunnstatt. This species lacks the wing in his single indi-
vidual specimen. It measures 6.5 mm.; perhaps it is identical with
Sphegina abrasa.

Locality: Kleinkembs, France. Horizon: Oligocene.

Holotype: R 186. Coll. Mieg. Mus. Ball. I have not seen the type;
I give above restated and rearranged the important particulars from
the author's description.

Sphegina obscura spec. nov.

Plate 4, fig. 1-3

Female (apparently). Length of the abdomen 7 mm. Head: and
front half of thorax missing. This is an obscure specimen, but the
shape of the abdomen and details of the femora can be made out fairly
well. Thorax: scutellum large, with strongly and evenly convex mar-
gin, almost hemicircular, one and one-half times wider than the second
segment at its smallest width. Abdomen: considerably constricted and
petiolate. The last two segments flared and expanded, as is typical in
the genus. The figure gives some idea of the relative shape and pro-
portion of the segments. There are traces of spots in the third and
fourth segments. It will be seen that the shape of the abdomen is not
greatly different from Sphegina infuscata Loew. Legs: hind femora

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thickened, but not spindle-shaped, the thickening begins quickly and
ends more or less abruptly, although the taper, as in many species, on
the distal end of the femora is gentle. There is a series of strong spines
arranged on the distal ventral part of the hind femora. The distal
third appears to have been dark in color. Wings: not easily discernible.
The fore part of the wing is best preserved and the convexity of costa,
and of second and third longitudinal veins is a matter of interest.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: No. 3947 in the Museum of Comparative Zoology. This
is the specimen seen by Williston (1886).

Genus Eoxylota genus nov.

This form differs from Hemixylota Shannon in the slightly tubercu-
late face and in the presence of the very numerous short spines on the
ventral part of the hind femora. The apical cross vein is less sigmoid
in Eoxylota and the margin of the scutellum in Hemixylota is distinctly
impressed. From true Xylota, Eoxylota is at once distinguished by the
cross vein being basal and by the small tubercle.

Genotype : Xylota pulchra Meunier.

Meunier described it m the genus Xylota. Because of the slightly
tuberculate face and the strongly basal cross vein, I am obliged to
remove it from Xylota and I place it in the genus Eoxylota. In many
respects it resembles the Chilean genus recently described by Shannon,
Hemixylota, which is like "the present form, with the basal cross vein
throwing it out of Xylotinae and into Cheilosinae, in the modern sense.
It is very odd that this genus should find its nearest living relative in
Chile. I have recently seen a specimen of Cyrtid fly from the Bern-
stein fauna which also finds its nearest living relative in Chile and
South Africa, bearing a mute witness to the flight of time necessary for
such present isolated distribution. Eoxylota strictly speaking differs
from Hemixylota only in the somewhat tuberculate face, slight differ-
ences in wing and facies. I have compared it with a specimen of the
genotype of Shannon's genus kindly sent to me by Dr. Edwards.

Eoxylota pulchra (Meunier)

Plate 6, fig. 29-30; Plate 10, fig. 72-73

Jahrb. Preuss. Geol. Landesanst., 24, (2): 207. PI. 13, fig. 6 and 7 (1904)

Male. Length 11 mm.; length of wing 8 mm. Head: broadly oval,
eyes very large, touching in the male but not for a great distance. The

hull: revisional study of fossil syrphidae 325

front small, black, flattened, somewhat protuberant at the level of the
antennae. Antennae located above the middle of the head in profile,
first two joints short, third not very large, half again as long as wide,
rounded apically, the arista short, thickened throughout, strongly on
the basal half. Face deeply concave below the antennae, apparently
with a small tubercle lying a short distance above the epistoma. The
junction of the cheeks and face is quite angular and obtusely ridged,
but scarcely more produced than the face at epistoma. Thorax: is
broad, longer than wide, considerably flattened, with a few stiff bristles
on the sides at base of wing. The mesonotum with thick delicate quite
erect pile, in color uniformly black. Scutellum large, rather elongate,
not quite twice as wide as long, the disc considerably flattened, erect
pilose, the margin with about six pairs of long stiff tuberculous bristles;
the apex seems to be rather thin, its color uniformly black. Abdomen:
elongate, two and one-half times as long as wide, widest at the end of
the third segment. Second, third and fourth segments subequal in
length, the latter rather broad at the apex, but little narrowed and
convex only on the sides. The hypopygium wide, broadly rounded and
not greatly visible from above. The surface of the abdomen as a whole
quite flattened, uniformly dark in color, appressed setaceous in the
middle, erect pilose on the sides. Legs: the hind femora elongate, a
little bit stout, not thickened, the whole ventral length with numerous
short sharp spinous bristles, the femora black, narrowly reddish api-
cally, the tibiae almost entirely light brownish red, a little bit darker,
narrowly, in the middle. Tarsi light reddish brown, the hind tibiae
much widened apically, pinched in before the apex, with many stout
short terminal bristles or setae. Wings: elongate, longer than the ab-
domen, uniformly tinged with brown. Stigma darker. Vena spuria
very faint; small cross vein a little oblique, long, entering the discal
cell about three-eighths of the way from the base, the marginal cross
vein long, almost paralleling the wing margin. Cross vein long, almost
paralleling the wing margin, the outer one joining the third vein some
distance but not remotely from tip. The basal marginal angles of the
first and second posterior cells short, spurred ; the last section of the
fourth vein before the origin of the apical cross vein somewhat long,
but only a third or less than the length of the apical cross vein.

Locality : Baltic amber, Germany. Horizon : Lower Oligocene.

Type: No. 8 in the University Museum at Konigsberg.

The above description as well as the illustration I give are taken
from the type specimen. In describing this species, he makes much of
the curious, irregular, jagged-edged spots on thorax and abdomen.

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Unfortunately he mistook for a definite pattern what is only the frag-
mented condition of the surface pigment. This species is a uniformly
blackish fly without pattern or marks.

The Subfamily CHRYSOTOXINAE

This small subfamily has at most but two living genera, but the
genus Chrysotoxum itself, is a world wide aggregate of many species,
all of them with a characteristic appearance. The fly described below
is the only fossil insect that has yet appeared that seems to show any
relationship to Chrysotoxum.

Genus Protochrysotoxum genus nov.

Large flies with prominent robust, very convex abdomens, much as
in present day species of Chrysotoxum. Apparently there are narrow,
basal, light colored fascia on the segments. The small cross vein is
located well before the middle of the discal cell. The third longitudinal
vein is straight whereas it is curved in Chrysotoxum. The head is
wanting in the specimen and cannot be described. It is sufficiently
well preserved, however, so that additional specimens, if they are ever
found can be recognized.

Genotype: Protochrysotoxum sphinx spec. nov.

Protochrysotoxum sphinx spec. nov.

Plate 2, fig. A

Male. Length 12.5 mm. Head: large, rounded, more or less sub-
globular, not quite as wide as thorax; no details of antennae visible.
Thorax: dark, broad; scutellum obscure, apparently quite large with
a circular rim. Abdomen: very large, evidently quite convex with
emarginate rim. Four segments and the tip of a fifth visible. The ab-
domen is dark, short, stiff pilose and on the base of the fourth and fifth
segment, particularly on the sides, there appears to be a pair of spots
wedged-shaped, with a straight posterior edge, the acute portion of the
wedge turned inward. Legs: absent. Wings: preserved in frag-
mentary fashion. The vena spuria is well developed, the whole of the
discal cell is visible and the small cross vein enters this cell practically
at the middle, its upper portion is strongly oblique. The lower marginal
cross vein is upright on the basal portion and then is acutely drawn out

hull: revisional study of fossil syrphidae 327

to join the fourth vein, the last portion of which is drawn down to
meet it. The outer cross vein, while not erect on the basal half ap-
proaches this condition and gives off an inward spur and then is drawn
out acutely but its confluence point and the tip of the wing are not
visible.

Locality: Florissant, Colorado. Horizon: Miocene.

Holotype: No. 159, coll. Amer. Mus. Nat. Hist.

The Subfamily EUMERINAE

I place here Meunier's genus Palaeopipiza, which bears very little re-
lation to Ascia (now Neoascia) and also my own genus Doliomyia. It
seems to me the wings of these two flies suggest Eumerus in venation,
as does the large, slightly quadrate third antennal joint, low set an-
tennae, and much thickened arista, and straight and subconcave face
types. Nothing is lacking but slight further changes in the wing and
the development of megamorphic femora.

Genus Palaeopipiza Meunier

Ann. Soc. Sci. Bruxelles, 26: 103, fig. 6. (1902); Jahrb. Preuss. Geol. Lande-
sanst., 24:209 (1904)

Head: eyes extensive. Condition in the male unknown. The back
of the head in profile very rounded and convex. Antennae set about
the middle of the head in profile, possibly a little below. The antennae
short, the first and second joints particularly so, the third joint very
large, flat, a little longer than broad, slightly more developed on the
ventral surface leaving the joint somewhat asymmetrical but not ex-
actly quadrate. Arista set in the middle of the upper surface midway
between the base and end, very short, two basal segments visible and
grossly thickened throughout although the tip is slender; surface
pubescent. Face below antennae distinctly concave, flat to the level
of the eyes, the epistoma a very little produced and rounded. Front
apparently bare. Face with some sparse pile. Occiput well developed
behind the ocelli and for a short distance along the eyes then scarcely
visible. Thorax: quite short, as broad as wide, at least convex; short
pilose without macroehaetae. Scutellum quite large, perfectly semi-
circular. Margin convex, with a few slender bristles, only the apical
two of any length. Abdomen: not quite twice as long as wide, broad
basally. Scarcely less broad at the beginning of the fourth segment at

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which becoming very convex, it tapers to the narrowly rounded tip
of the fifth segment. Abdomen everywhere short pilose, except at
basal corners. Posterior pile appressed. Legs: hind femora short and
slender. I cannot discern if spines are present. Hind tibiae as long as
femora, considerably thickened on the outer three-fourths. Wings:
venation as Palaeoascia. The apical cross vein much longer, paral-
lelling wing margin, joining third vein not far from tip. Vena spuria
appears to be faintly present; this is uncertain.
Genotype : Palaeopipiza xenos Meunier.

Palaeopipiza xenos Meunier

Plate 8, fig. 55-57; plate 13 fig. 117

Ann. Soc. Sci. Bruxelles, 26: 103; fig. 6. (1902). Jahrb. Preuss. Geol. Lande-
sanst., 24: (2) 209 (1904)

Female. Length 6 mm.; length of wings 4.7 mm. Head: the eyes
apparently dark in color. Antennae dark brown, the third joint orange
brown in parts. Arista brown, the pile of the face white in color. The
front at the region of the antennae is very little prominent. Concavity
of the face not very deep. There are a few long pale bristles about the
ocelli and some still longer dark bristles on the vertex behind the ocelli.
Thorax: and scutellum dark in color, apparently aeneus, the pile
everywhere pale and the bristles of the scutellum pale. Ahdomen: con-
colorous with the thorax throughout. The sides of the segments very
narrowly yellow, the pile of the abdomen without exception everywhere
pale. Legs: femora black, possibly aeneus; the bases of the tibiae to
a varying extent, on the hind tibiae almost the whole, dark brown,
remainder brownish yellow. Wings: somewhat brownish, the stigma
a little darker.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Type: in the University Museum at Konigsberg. No. K4304, 2492
(K4304-Klebs).

Genus Doliomyia genus no v.

Small short robust species. Head: large, barely wider than thorax,
not very elongate nor yet is it flattened, the males probably holoptic,
the front long, very convex in profile. The antennae distinctly set
above the middle of the head in profile, quite short but the third joint
very large, suborbicular. The arista somewhat longer than antennae
basally, thickened but not conspicuously, bare and slender pointed.

hull: revisional study of fossil syrphidae 329

Face below antennae perfectly straight, gently retreating. Cheeks
practically nonexistent, so that at the epistoma the face is only half
as long as the third antennal joint. Occiput tumid for a short distance
above and close to the vertex. Thorax: about as long as wide, the
scutellum short and quite wide, a little over twice as wide as long, the
disc convex and the rim narrowly emarginate and impressed, without
strong bristles. Dorsum of thorax and scutellum exceedingly short
microsetate. Abdomen: short and broad, barely wider basally than
the thorax, convex and very short microsetaceous. Legs: femora
short, slender, not thickened and with short setaceous bristles instead
of spines ventrally. Wings: a little longer than abdomen, broad basally
and broadly rounded at apex. The third longitudinal vein ending with
costa a little beyond the tip of the wing. The lower marginal cross
vein straight, directed considerably away from the margin of the wing
and thus oblique rather than parallel. The last section of the fourth
vein before the origin of the apical cross vein quite long, the apical
cross vein angulate a little before the middle, the angle directed in-
ward but without spur. The confluence of the apical cross vein with
the third vein is not very far from the tip, but the remaining distance
of the third vein to the tip of the wing is about one-fourth of the length
of the apical cross vein. Both the basal angles of the first and second
posterior cells spurred. Wings villose throughout. Vena spuria very
faint if not actually absent.

Genotype: Doliomyia ckalybea spec, no v.

This genus differs from Meunier's Palaeopipiza in the fact that the
antennae are definitely located above the middle of the head in profile.
The arista of Palaeopipiza in contrast to Doliomyia is very short,
strongly thickened basally, pubescent and not very slender on the
apical half. The third joint of Palaeopipiza is somewhat truncate on
the dorsal area and the face somewhat concave in profile. The abdomen
of Palaeoascia is elongate, very convex, strongly drooping from the
base and is suggestive of Paragus (Recent genus). The subapical cross
vein is not strongly angulated; both of these genera remind one very
closely of the genus Eumerus among Recent Syrphids. The wings of
Doliomyia, the large third antennal joint, the shape of the face all
suggest Eumerus. They are sharply distinct from Eumerus through
the absence of a grossly thickened, swollen hind femur, with its den-
ticulate, ventral apex and from Citibaena, a close relative of Eumerus
with slender femora, again with the absence of spines on the femora
and the quite bare eyes. In Citibaena the eyes are densely long pilose.

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DOLIOMYIA CHALYBEA Spec. nOV.

Plate 8, fig. 58-59; Plate 9, fig. 62; plate 13, fig. 116

Female. Length 6 mm. ; length of wing 5 mm.

Head: broad. Face, front and vertex black in color, the antennae
very dark brown to black. Arista black. Pile of front very short, be-
coming still shorter before the antennae, but if bare at any point it
would be very narrowly before the antennae. Pile of vertex about the
ocelli a little bit longer, the occiput on the sides very short pilose and
short pilose below. I cannot distinguish any pile on the face. The face
is well preserved and so it is quite possible the pile was absent or if
present was only in the form of micropubescence. Thorax: and scu-
tellum jet black, the pile exceptionally short without macrochaetae
anywhere. There is a narrow, convex, impressed margin on the ex-
treme end of the dorsum of the thorax just before the scutellum. One
specimen suggests that the thorax was dark shining bluish. Abdomen:
dark blackish in color and all of the legs dark black. Legs: the tibiae a
little bit. lighter perhaps and the apical joints of the tarsi light brown.
Wings: strongly infuscated and smoky throughout, the stigma darker.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: specimen No. B16788 (Slide no. 16788) in the University
Museum at Konigsberg.

Paratype: a specimen labelled B438 (slide numbered 438). Both
are from the collection of Dr. Klebs.

The Subfamily MICRODONTINAE

Serres assigned a species to this subfamily in 1829, which he neither
described nor illustrated ; since he did not give a type number, I have
been unable to trace the specimen. I am inclined to believe he was
right in his placement, for Microdon is so characteristic, that it is diffi-
cult to see how he could have been mistaken. Perhaps no single fossil
would be as interesting as this one, for the subfamily Microdontinae
is very peculiar, consisting of hundreds of species of very unstable
nature.

Genus Microdon species Serres

Geognosie des Terrains Tertiaires, p. 253, (1829).

Serres stated that this fly closely approached the species Aphritis
auropubescens described by Latreille.

Locality: Aix, Provence, France. Horizon: Oligocene.
Type: I was not able to locate the place of deposition.

hull: revisional study of fossil syrphidae 331

The Subfamily ERISTALINAE

Four genera are assigned here. At least two, possibly three or even
all of the genera are very questionable.

a. Helophilus primarius Germar. This record should be deleted, in
my opinion, from the family. I can not see that there is the slightest
reason for placing this fly in Helophilus. I did not locate the type.

b. Merodon germari Heyden. I located this type in the British
Museum. It is extremely interesting and the body and legs beyond
doubt suggest Eristalis, but even when discovered the wing was in-
sufficiently preserved to say that it should definitely go into the
Eristalinae. To definitely place a fly in the subfamily Eristalinae, we
must rely on the third vein being kinked, because if it were not (and
it fortunately is in over seven hundred species which are in about
sixty genera and subgenera) we should have to fall back, in present
knowledge, on the basal femoral patches of setae, a character that
varies to some extent. Besides the kinked vein, it is true that the Eris-
talinae almost always possess very robust abdomens and often large
squamae, but we can hardly define the subfamily in these terms. The
ancestry of Eristalinae is too poorly known for us to conjecture much
upon its origin. Their face is nearly always tuberculate, only three of
many genera excepted, and the origin of both of these, as well as the
Sericomyinae is obscure. I believe that through Mallota, Brachypalpus
and Criorrhina and even better through Sericomyia, we find a certain
amount of closeness which is more than resemblance. This points
vaguely to a relationship with Xylotinae but at an extremely early
period, because most true Xylotinae have the opposite type of face.
It is significant that plumose aristae are restricted (one exception in
Cheilosiinae) to Eristalinae (a few genera), Volucellinae (all genera)
and in Sericomyinae (all genera). We have seen that the Volucella-
form existed complete, except for the wing, in the lower Oligocene.

c. Scudder described Eristalis lapideus from Green River, Wyoming,
but Williston, who saw the specimen, did not believe it was an Eris-
talis. Since the type is now lost, we will have to wait until it is again
found to know its real affinities. Williston was almost certainly right
in assigning it elsewhere.

d. Palaeoeristalis tessellatus, new genus and species (described be-
low). I place this species in an Eristalinae genus on the basis of the
short rotund abdomen and the slight curvature of the third vein.
Since it is poorly preserved, this is all that I can discover of significance.
There are other genera (non-Eristalinae) with the third vein shallowly

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dipped. This is quite possibly an early type of Eristalis in which the
third vein has not yet become deeply and definitely kinked.

Helophilus villeneuvi Theobald

Les insectes fossiles des terraines oligocines de France, 1937, p. 351 ; pi. 7, fig.
12.

Length 12 mm. Head: large, as wide as the thorax, semicircular in
shape. The eyes do not quite touch. The vertical triangle is large,
the front short, the posterior border of the head almost straight. The
ocelli and antennae not visible. Thorax: black, oval and strongly
pubescent. Abdomen: wider and longer than the thorax, compressed
and ovoid; black in color with clear spots on the borders and clear
transverse bands. Legs: wanting. Wings: long and over-reaching the
extremity of the abdomen. The venation is obscure but the costa,
subcosta and radial sector veins can be distinguished. The radial
sector is bifurcated. The longitudinal fold of the vena spuria can be
discerned.

The author places this insect in Helophilus largely upon the basis of
size, compact shape and an abdominal pattern somewhat similar to
that of Helophilus quadrivittatus Wied. He states further that the first,
second, third and fourth segments of villcneuvei carry the clear lateral
spots; the fifth carries besides a transverse clear band.

The close approximation of the eyes suggests the genus Mesembrius
instead of Helophilus; the marked pubescence of the thorax might in-
dicate Mallota or some subgenus of this group. There are considerable
number of closely related subgenera in the Helophilini.

Locality: Aix-en-Provence, France. Horizon: Oligocene.

Type: L'Ecole des Mines de Paris.

Helophilus miocaenicus (Stackelberg)

Tubifera mioeaenica Stackelberg, Revue Russe d'Ent., 19, p. 89-90, Taf. II
(1925).

This species was described by Stackelberg from the Middle Miocene
of Voroshilovsk, Caucasia. His description is accompanied by an ex-
cellent photograph which shows clearly much of the venation of the
wing and shows the pattern of the abdomen well. This pattern with
its large, paired, pale spots, almost touching, and black posterior
borders is not greatly dissimilar to that of pendulus Linnaeus, a Recent
species. The character of the venation clearly places the fly in the

hull: revisional study of fossil syrphidae 333

subfamily Eristalinae. The loop of the third vein is deep and evident.
The marginal cell appears to be open as is required in the genus
Helophilus. The description is not quoted here in as much as it will be
necessary for the student to refer to the original illustration.

Merodon germari Heyden
Plate 5, fig. 26; Plate 8, fig. 60
Palaeontographica, 78: 10; fig. 5 (1862)

Size quite large. Head: very large and not so wide as humeri, some-
what shallow, but it is not long. Eyes touching, mid-line facets not
enlarged, occipital eye margin seen from above laterally sharp and
with two little borders as far as can be ascertained. Thorax: broad,
robust, black in color. Scutellum cannot be made out clearly, but
appears to have been about one and three-fourths times as broad as
long. Thorax short, pilose. Abdomen: broad, slightly emarginate, with
two distinct black patterns with markings sharp and not cloudy on
edges. Posterior margin overrun with post bristles, long and sharp.
Pattern as follows : a black posterior border on second segment, widen-
ing toward sides and produced as a median stripe, until it reaches the
margin of the first segment, where it spreads out narrowly. Second
segment with much wider posterior margin, only very narrowly wider
laterally, with similar median stripe reaching second segment and not
spreading out as it does so. Third and fourth segment with a similar
band. Fifth appears to have been entirely dark. Hypopygium small.
Legs: hind femora very thick, but not greatly thickened out of propor-
tion to the size. There is some evidence of a few setigerous bristles
apicoventrally on the hind femora. Hind femora thickly covered with
short sharp, slender bristles. Hind tibiae similarly covered. Wings:
poorly preserved. Base with a few short sharp spinules. Size and pa-
tern resembling that of a species of Milesia.

Locality: Rott, Germany. Horizon: Upper Oligocene.

Holotype: British Museum of Natural History; the type was
studied.

Genus Palaeoeristalis genus nov.

Head: hemispherical. Thorax: short and broad. Scutellum one
and three fourths broader than long. Abdomen: quite robust and short.
Five segments can be seen. The abdomen is broadly rounded past the
second segment and evidently quite convex. Legs: hind femora very
much thickened, the thickening spread more or less over the whole

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length but a little greater basally. Ventro-distally the femora is
equipped with many small setae rather than spines. Wings: with the
third longitudinal vein gently curved down into the first posterior cell.
The apical cross vein joins the third vein not far from the tip of the
wing; it is sinuous. The lower cross vein is near to the margin of the
wing and more or less parallels it. Third vein ending near tip of wing.
Genotype : Palaeoeristalis tesselatus spec. nov.

Palaeoeristalis tesselatus spec. nov.
Plate 4, Fig. 7-9

Head: hemispherical with detail apparent. Thorax: short and
broad. No pile can be seen, nor any bristles. Scutellum about one and
three-fourths times as broad as long, rather square in outlines, with
simple margin. Abdomen: five visible segments; form broad and
rounded. Second segment two and one-half times as broad as long,
third twice as broad as long, and the remaining two, short, evenly
rounded off. Abdomen indistinctly marked but the second and third
segments have each a wide posterior border dark in color, which is
slightly and gradually widened as it approaches the lateral margin,
and which medially interrupts the anterior pale band in a rather broad
fashion. The margin of the spots are vague and not sharply delineated.
The color is darkest along the median line. The fourth and fifth seg-
ments more or less wholly dark. Posterior halves of second and third
segments, especially the dark areas, and the terminal segments
covered with short, quite thickly set, bristle-like hair, which in the
middle is directed posteriorwards and on either side obliquely towards
the lateral margins. Legs: one hind femora shows as a pale enormously
thickened structure with many sharp black setae on the ventral
distal two-thirds. The hind tibiae are long. Wings: poorly pre-
served. Marginal cell open. Anterior cross vein joining discal cell
way beyond middle, somewhat drawn out in oblique fashion but not
markedly so. Subapical cross vein apparently angulated. Costa
microsetose. Third longitudinal vein with a decided flexure without
in the least being kinked.

Locality: Dragon, Utah. Horizon: Eocene.

Holotype: no. 3948 in the Museum of Comparative Zoology, (F. M.
Carpenter, collector).

This is a remarkably interesting form. With its small size, broad
round abdomen, thickened femora, it suggests chiefly an Eristalis.
Present day Eristalis must have a kinked third longitudinal vein.

hull: revisional study of fossil syrphidae 335

Eristalis paucisinuatus Theobald

Les insectes familes des terraines oligocenes de France, 1937, p. 352; pi. 23;
fig. 12.

Length 11.5 mm. Head: large and transverse; the facettes of the
two large eyes are visible, they appear to touch on the front. The
anteriorly produced antennae have a pointed arista which is not
plumose. Thorax: large, black, finely hairy with numerous bristle.
Scutellum large and rounded. Abdomen: oval, rounded at the ex-
tremity with four segments visible. The pattern of the abdomen sug-
gests slightly that of Eristalis solitus Walker; the pattern of the second
and third segments especially differing. Legs: at least the hind femora
swollen and enlarged and hairy. Wings: the wings lie folded upon the
abdomen ; almost all the venation is visible. The vena spuria is clearly
visible. There is a curve in the third longitudinal vein which is not so
deep as that in E. solitus Walker.

This fly would appear to be a satisfactory example of the Eristalinae.
The more gently curved third longitudinal vein is a matter of special
interest as in all Recent members of this subfamily this vein is rather
deeply kinked.

Locality: Aix en Provence, France. Horizon: Oligocene.

Type: No. 1005, Theobald collection.

The Subfamily VOLUCELLINAE

The Volucellinae is one of the dominant Recent types of Syrphidae;
its species are preeminently characteristic of the Neotropical Region.
Very few fossil flies have been associated with this subfamily. Loew
mentioned an undetermined specimen from the Baltic amber which he
assigned to Volucella. The only other instance is the unique new genus
Ptilocephala from the amber. It is clearly Volucella-\\ke in eight
particulars.

Genus Ptilocephala genus nov.

Head: small. Eyes large and flattened, the front especially steep,
oblique, flattened and concave, but a little bit convex before the an-
tennae. Antennae situated three-fourths of the way up from the
bottom of the head and at the junction of the middle and upper thirds
of the eyes. First two joints short, third joint elongate, three or four
times longer than wide. The arista a little longer than antennae,
plumose with twenty rays above and with many shorter ones below.

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The head below the antennae very flat, not produced near the bottom
of the eyes, the face is abruptly and conspicuously produced into a
rounded and protuberant knob or tubercle which is beset with long
bristles. Below the tubercle the face recedes gently and then drops
vertically for a considerably greater distance. Thorax: with stiff ma-
crochaetae along the margin and on the margin of the scutellum.
Scutellum with an oval concave depression on the disc. Abdomen:
short, appressed setose. Legs: hind femora short, somewhat thickened
in the middle, and spread out over the whole length. Tibiae very short
and thickened on the distal two-thirds and about three-fourths as long
as femora. Details of the ventral surface of femora cannot be ascer-
tained. Wings: large, but short and broad, a little longer than the
abdomen, broadest basally, and but little less broad toward the apex.
The vena spuria faint, but present. Alulae well developed. Stigma
pale. Costal cell very wide, the costa with a double row of unusually
heavy, long, erect, sharp-pointed microspinulae; the lower marginal
cross vein straight, directed away from wing margin. Spurs of fourth
and fifth longitudinal vein very short or absent, the apical cross vein
sinuous, joining the third vein at right angles but not recurrent some
distances from tip of wing but not remotely.

Genotype: Ptilocej)hala volucelloides spec. nov.

This genus is unique in many ways, and is the only certain fossil
member of the subfamily V olucellinae . Loew mentioned having seen
a species of the Volucella from the amber, but he did not describe it
or assign any number to it. I have studied all the Syrphids mentioned
by him except the Volucella, which I did not find at the British
Museum with his other specimens. The present species forms a true
connecting link between the Cheilosinae and Volucellinae because it
has the venation not greatly different from Cheilosia and the tubercu-
late face, and the chaetae of Cheilosia and has the plumose arista of
Endoiasiamyia (Hiatomyia). Indeed, when I first thoroughly examined
the specimen I considered it to belong to this genus. However, the
knobbed, bristly, tuberculous, deep conical face, the macrochaetae are
all characters which are strongly Volucella-\\ke and the facies of the
insect is more like that of Volucella than it is of Cheilosia. Lastly, were
the apical cross vein recurrent and did the third vein join costa far
above tip of wing one would have practically transformed it into a
modern Volucella-Wke type. The depression on the scutellum reminds
one of Phalacromyia but it is much more like that of Graptomyza and
the faint vena spuria and the concave flattened front also suggests
this genus. It is a peculiar and interesting fly, which is clearly Volu-

hull: revisional study of fossil syrphidae 337

cella-like in eight particulars; the wing however is but little suggestive
of Volucella. Has body-type here, as in Xylotosyrphus been laid down
before wing shift? Or is this but an aberrant off-shoot? I do not see
how the former conclusion is to be avoided. Ptilocephala combines
some of the characters of the most specialized and of the most simple
genera of the Volucellinae.

Ptilocephala volucelloides spec. nov.

Plate 6, fig. 31-35

Female. Length 4.8 mm.; length of wings 4.6 mm. Head: broader
than thorax. Eyes large, flattened, widely separated in the female, the
top of the head obliquely flattened as in certain species of Volucella
or Graptomyza. The front between the ocelli and the slightly convex
portion just above the antennae is shallow, concave, and flat and erect
sparse pilose. Antennae situated about three-fourths of the way up
from the bottom of the head or nearly at the junction of the upper and
middle third of the eyes. Antennae elongate, the first two joints short,
the third joint three and one-half to four times as long as wide, dark
brownish black in color. The arista barely longer than antennae,
strongly thickened basally and long stiffly plumose above and below,
the upper basal rays the longest. About twenty rays on each side.
Face below the antennae in profile apparently flat with the level of the
eyes, then a short distance from the level of the bottom of the eyes
strongly and abruptly produced out as a rounded blackish bristle-beset
knob, which is very protuberant, rounded, and below which the face
continues to drop, after a brief recession, for some distance. Cheeks
not very conspicuous, the actual epistomal-cheek-occipital profile is
obscured by exuvia. Presumably the face is sharply conical and the
oral profile concave. Thorax: convex, with two stiff bristles on the
side before the wing, two above the wing, two on the mesopleura,
three on the posterior calli, one bristle on each side before the scutellum
and a little bit to the outside. Scutellar margin with three pairs of
bristles, all of which are exceedingly stout and thick. Pronotum of
thorax with a collar of short stiff bristles. Scutellum with the disc
broadly and symmetrically flattened and concave, the margin of the
concavity lined with bristles and a very few bristles on the concavity.
The scutellum suggests that of Graptomyza flavorhjncha Hull in the
matter of this oval concavity. Abdomen: short, broad, a little longer
than wide, black in color; as far as can be discerned, with thick short
appressed bristles. Much of the abdomen is hidden and obscured by

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the wings. Legs: hind femora a little bit thickened in the middle, short,
the color obscure but mostly dark. Wings: light brownish, stigma a
little darker.

Locality: Baltic amber, Germany. Horizon: Lower Oligocene.

Holotype: in the University Museum of Konigsberg (No. X 118).

Volucella species Loew

Ueber den Bernstein und die Bernsteinfauna Meseritz Progr. k. real schule
p. 1-48. (1850)

The dipterist Loew mentioned having seen a specimen of this genus
preserved in amber but he gave no name to it. The specimen should
be in the British Museum, but I was not able to locate it among the
other material examined there.

The Subfamily XYLOTINAE

Two genera have been assigned here by name without numerical
or species designation. They are Criorrhina Giebel and Tropidia
Handlirsch. I was privileged to search through the collection of the
late Dr. Handlirsch in Vienna, but without finding the Tropidia. I
have been unable to trace Giebel's collection. Tropidia, to occur in the
modern sense, must have a megamorphic femora, a femoral plate and
carinate face.

Xylotosyrphus is to be doubtfully placed here. It has all that
Xylotinae should have except a definitely post basal small cross vein.
It is an early type, connecting through the Myioleptini, the Cheilo-
sinae and Xylotinae.

Megaxylota is a large and beautiful fly from the amber. It is a true
Xylotinae in every respect, and is not unlike, in some ways, Brachypal-
pus or Calliprobola.

Genus Criorrhina species Giebel

Zietschr. Geol. Nat., 2: 87 (1870)

There is considerable uncertainty whether this genus occurs in the
Baltic amber. Whether Giebel was dealing with such a species or not
is uncertain, because in his brief remarks he compares the fly to a
Dolichopodid, and of course no Criorrhina could be anything like this.
It will be necessary to wait until the type is located to determine the
matter.

hull: revisional study of fossil syrphidae 339

Genus Xylotosyrphus genus nov.

Head: more or less destroyed, about as wide as thorax. Thorax.
long and broad, somewhat convex, dark in color, without pattern.
The scutellum one and one-half times as wide as long, with simple
semi-circular rim. Abdomen: two to two and a half times as long as
wide with prominent rounded hypopygium and the sides of the ab-
domen nearly parallel. Abdomen marked with two pairs of prominent,
subquadrate, pale spots on second and third segments. Legs: hind
femora considerably thickened. Wings: with venation more or less
like that of Xylota but the small cross vein definitely basal in position
and entering the discal cell less than half of the way from the base.

Genotype: Xylotosyrphus pulchrafenestra spec. nov.

This genus seems to be in every respect like certain species of present
day Xylota, except that the small cross vein is definitely well before the
middle of the discal cell and in our modern scheme of things this would
preclude placing it in the Xylotinae. Consequently many things such
as general body form and pattern have a more ancient history than the
relative position of the small cross vein, long relied upon as an im-
portant index of relationship. It may be that more fully preserved
specimens will later show other differences between Xylotosyrphus and
present day Xylotas.

Xylotosyrphus pulchrafenestra spec. nov.
Plate 4, fig. 12

Male. Length 10 mm. Width of abdomen 2.7 mm. Length of wing
indeterminate (about 6.8 mm.).

Head: the head shows little detail. Part of the head past the occi-
put being destroyed. Thorax: stout, broad, and relatively short in
profile with a gently forward downward slope from the middle. Scutel-
lum rather small, evenly rounded. Abdomen: the sides of the abdomen
are parallel, with a beautifully rounded (hemicircular) tip, suggestive
of the hypopygium of the male. Five segments visible. Abdomen
marked with four pale quadrate windows, a pair each on the second
and third segments, each of these segments is divided by a median
black stripe and the anterior median corners of the second pair of spots
and the post median corners of the first pair of spots are markedly
rounded and convex, leaving the black concave. Remaining segments
and hypopygium dark in color. Abdomen largely covered with ex-
tremely short, thick, decumbent, black, bristly pile. Legs: hind femora

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considerably thickened, greatest in the middle or just past the middle;
in length about three times as long as greatest width. Remaining
femora slightly thickened. Hind femora with numerous stiff black
spinules ventrodistally. The hind tibiae appears to have been spinu-
lose basi-ventrally. No further details of the legs can be seen. Wings:
very little shows. In one specimen it can be ascertained that the
anterior cross vein is beyond the middle of the discal cell and is pulled
down distally to an oblique angle. Morover, the terminal section of the
fourth longitudinal vein as it joins the postical cross vein, is strongly
curved downward or deflected. It joins at an angle of some sixty or
seventy degrees. Marginal cell open.

Two beautiful specimens which show, as far as can be ascertained,
all the characteristics of our modern Xylota. It is impossible to as-
certain whether they would come in the neotropical Planes, which is
a Xylota with carinate face.

Locality: Dragon, Utah. Horizon: Eocene

Holotype: No. 3942 in the Museum of Comparative Zoology (F. M.
Carpenter, Collector.)

Para type: one specimen, with the obverse and reverse, Nos. 3943
and 3944, in the Museum of Comparative Zoology.

Genus Megaxylota genus nov.

Head: large and broader than thorax, rather wide and not particu-
larly elongate. The females are broadly dichoptic. The front convex,
somewhat protuberant, the antennae situated a little below the middle
of the head in profile; short; third joint large, a little longer than broad;
rounded. The arista quite long and slender. Face below antennae
deeply concave; the epistoma thrust forward at face itself, short, it is
but little more produced at junction of face and cheek and not angu-
larly ridged. Occiput scarcely visible in profile from above, the back
of the head is gently convex and the flanges on the lower part of the
occiput behind head are conspicuous. Thorax: half again longer than
wide, quite convex, short, dense, erect pilose. Scutellum not large,
convex on the rim, flattened on the disc, the apex subtruncate, the
margin with a few long slender bristles. Abdomen: a little over twice
as long as wide and a little bit convex, especially terminally, broadest
in the middle, not greatly tapering towards the end. Legs: hind
femora very massive, especially on the basal three-fourths, rather
narrowed apically, the ventral part of the hind femora with a slight
bulge shortly before the end and beset with many rather short stiff

hull: revisional study of fossil syrphidae 341

bristles, but no spines. Hind tibiae very short, only three-fifths as
long as femora, thickened throughout, very arcuate, ending trans-
versely. Wings: elongate, pointed apically, broadest basally. The
small cross vein is very oblique; it joins the discal cell about three-
fifths of the way from base. Wings Xylota-Wke in venation. Marginal
angles of the marginal cells spurred, apical cross vein sigmoid, joining
the third vein not very far from the end of the wing. Vena spuria
prominent. Alulae present.

Genotype: Megaxylota magnifemur spec. nov.

These are large flies. This genus differs principally from Xylota in
the grossly thickened femora, which unlike Xylota lacks completely
all spines on the femora. The very short, quite arcuate transverse-
ending tibia is characteristic.

Megaxylota magnifemur spec. nov.
Plate 7, fig. 40-42; Plate 8, fig. 52-53

Female. Length 16 mm. ; length of wing, 10 mm. Head: broad, eyes
apparently bare, the ocelli but little protuberant. The pile of the front
very short but dense and apparently restricted to the upper half, indi-
cating that the front of the male is pubescent only. Face and front as
nearly as can be ascertained under the whitish exuvia, dark in color.
Face pubescent only, the facial strips fairly wide and conspicuous and
with a fringe of hairs. Cheeks near the occiput with some long hairs
and the pile of the occiput throughout is thick, though not very long.
No spines near the top. Occiput broadly rounded near the margin.
Humeri and pleura densely pilose. Thorax: the pile of thorax and
scutellum seems to be pale in color throughout. Scutellum black, the
posterior calli brownish orange and due to the exuvia it is impossible
to tell the color of the mesonotum or of the abdomen. Legs: the femora
seem to have been orange brown, somewhat lighter apically, and the
tibiae and tarsi were beyond question light in color. Wings: are some-
what matted to the abdomen and also coated and while the venation
shows well, the abdomen is somewhat obscured. Wings were pale
yellowish brown in color.

Locality : Baltic amber, Germany. Horizon : Lower Oligocene.

Holotype: in the University Museum at Konigsberg. This specimen
bears no number; the slide is numbered 'D'. I affix the number H 105.

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Genus Tropidia species

Handlirsch, Handbuch Palaeont. und Zool. 4 (2): 1024 (1908)

The genus Tropidia is a fairly well marked and characteristic genus,
but I cannot say definitely whether or not Handlirsch was sufficiently
familiar with the genus to recognize it. Consequently it must await
further study. I was unable to locate the specimen in Handlirsch's
collection in Vienna. He gave no specific name to the specimen.

INCERTAE SEDIS
Syrphus curvipetiolatus Meunier
Jahrb. Preuss. Landesanst. 24: 210, pi. 13, fig. 8, 9 (1904)

Male. Length of body: 7.5 mm.; length of wings 4.5 ? mm. Head:
oral margin is conspicuous. Facial tubercle not distinguishable.
Antennae small. Eyes touching on the front. The first two segments
on the antennae short, the third rounded off, the antennal arista
moderately short and thick. Thorax: scutellum black, faint, hairy.
Abdomen: transparent. It is impossible to discern whether it has
bands or stripes as in Recent forms. Legs: claws large, bent. Pulvilla
strongly developed. Wings: this form is characterized by a very short
appendicular vein proceeding to the posterior cell, which is always
clearly evident in Recent species of Syrphus. (Translated and re-
arranged from the original).

Locality: Baltic, Germany. Horizon: Oligocene.

Type: No. 4445 from the Kgl. Bernstein collection at Konigsberg.
(not seen).

This is the only species described by Meunier of which I was not
able to find the type or a specimen. This fly apparently does not belong
to the genus Syrphus. The type must be examined before it can be
definitely determined to what genus it belongs.

Syrphus bremii Heer
Die Urwelt der Schweiz, fig. 314 (1865)

The note below is the only comment that Heer makes about this
species and Syrphus shellenbergi, which he figures.

"The hoverflies (Syrphiden), however, are in two forms, which are
marked as in the living species with their pretty banded abdomens
(Fig. 314 and 315)."

This species may be recognized by the narrow black bands on the

hull: revisional study of fossil syrphidae 343

base of the third and fourth segments, each broadly interrupted in the
middle, and the large flat triangle in the middle of the second segment.
This maculation makes it unique and easily recognizable, but as it is
characteristic for certain Stratiomyids and as the wings in this fossil
fly seem to have been very imperfectly preserved, there is a consider-
able possibility that Syrphus bremii is a Stratiomyid.

Locality: Oeningen, Germany. Horizon: Miocene.

Type : Not located.

Syrphus schellenbergi Heer
Die Urwelt der Schweiz, fig. 315 (1865)

See the comments under Syrphus bremii. Characterized by the fact
that all of the second, third, fourth and fifth segments are solid black
on all the central portions; narrowly margined everywhere with pale
color.

Locality: Oeningen, Germany. Horizon: Miocene.

Type : Not located.

Helophilus? primarius Germar

Insekt. Protogae. Spec, Fasc. 19: 25, pi. 25 (1837). Giebel. Fauna der Vorwelt.
Die Insecten und spinnen der Vorwelt, 2: 201 (1856)

This species is unrecognizable from the description and figures.
Locality: Bonn, Germany. Horizon: Oligocene.
Type : Not located.

Eristalis lapideus Scudder

Bull. U. S. Geol. Geogr. Surv. Terr., 3 (4): 756 (1877). Tert. Ins. sp. 558 pi.
5. fig. 48. (1890). Williston, Synopsis N. A. Syrph., p. 281-283 (1886)

The following comments based upon Scudder's description are
pertinent.

Length of thorax 3.5 mm. ; breadth of thorax 3.5 mm. The specimen
was poorly preserved but fixed in dorsal aspect with the wings partly
expanded. The head was almost wanting and the thorax without
markings. The abdomen was long, broadest in the middle of the basal
half and posteriorly tapering considerably; the tip of abdomen was
rounded. The apical half of the first segment was black and formed a
distinct transverse fascia. There appeared ?to be five segments. The
venation of the wings shows only upon the basal half of the wing and

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there poorly ; the alula was distinct and quite large with oblique, dark,
transverse ridges indicative Scudder says of Volucella or Oestrus.

Locality: Chagrin Valley, White River, Colorado (W. Denton).
Horizon: Eocene.

Type: one specimen. I was not able to locate the type.

It is rather doubtful if this fly belonged to Eristalis but a better
specimen is required for a decision as to its affinities. Williston, who
also doubted its affinity with Eristalis, stated, "Of the thirty or more
specie's which Mr. Scudder had separated out, I was first struck with
the fact that probably all belong to the first division of the family with
a basal cross-vein, a conclusion at which Mr. Scudder had already, in-
dependently, arrived. There are two possible exceptions to this rule,
but both of them are doubtful, in view of the general relationships of
the other material studied. The first possible exception is the specimen
which Mr. Scudder had doubtfully referred to Eristalis."

Insects Misidentified as SYRPHIDAE

Several names have in the past been associated with the family
Syrphidae, but do not belong there. These are briefly mentioned
below.

Cheilosia dubia Weyenbergh

Archives du Musee Teyler, 2: 259 (1869)

Cheilosia dubia Weyenbergh should be deleted from the list of fossil
Syrphidae. I quote from a recent letter of Dr. E. Dubois of the Teyler
Museum, who had a photograph made of it recently:

"The imprint is very ill-defined. In my opinion and that of one of
my palaeontological pupils, specialized as an entomologist, it only
shows to be of an insect. We can understand that Handlirsch says
of it, '1st sicher kein Cheilosia und keine Syrphide und vermutlich
uberhangs kein Dipteren'."

Psilota tabidosa Scudder
Tert. Ins., 13: 561 pi. 9, fig. 9 (1890)

Length 5.0 mm.; wing 3.9 mm.; abdomen 3.8 mm.

The type of this species has been examined (Mus. Comp. Zool.).
It consists of a body, one wing and a very imperfectly preserved head
and thorax that must necessarily be completely disregarded. The
venation is obscure. Nevertheless, certain portions show tolerably

hull: revisional study of fossil syrphidae 345

clearly. Everything considered, the size, shape of body and wing, and
the apparently six or seven segmented body and venation remove this
fly from any possible location in the Syrphidae. It has in a strong
degree the habitus of a Scenopinid or window fly. It must be noted
that, while the apparent segmental lines of the abdomen may not all
be true segmental divisions, still there are no Syrphids known with
such additional creases midway between segments. If the speci-
men is viewed as a Syrphid the rounded nodose termination of
the abdomen would be equivalent to a hypopygium; thus the specimen
would be a male and limited to five segments. I believe, the species
must be removed from the Syrphidae, perhaps placed in the Sceno-
pinidae.

Remalia sphinx Brodie
Hist, fossil insects second, rocks England. 1845

This species was wrongly included in the Syrphidae. A glance at the
original pagination and subtitles will show that it was originally
described by Giebel (1856) under the family Muscidae, Brodie (1845)
having listed it only.

Syrphopsis globosiceps Zeuner

Fortschr. Geol. Paleont. 11 (28): 316 (1931)

This species is based on a very insufficient specimen. I can make
nothing of it from my examination of the type specimen, and see no
basis for including it among the Syrphidae.

CONCLUSIONS

This study of fossil Syrphid flies warrants consideration along three
different lines:

1 . A general consideration of the relationship of fossil genera, species,
and individuals to the Recent fauna and the origin of the family.

2. Special evidence afforded as to morphological changes.

3. Evidence afforded as to variation of species.

A table is presented below, showing the species and genera of fossil
Syrphids placed in their respective horizons. It shows that the largest
number of species and genera, so far as yet discovered, occur in the
Oligocene. This is certainly due to the abundance of this family in the
Baltic amber (Lower Oligocene), for there are fourteen extinct genera

346

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in this period and only five Recent ones, as compared with three
extinct (possibly only two) genera and eight Recent genera in the Mio-
cene. These figures point to a rapid process of elimination of generic
types in the Lower Oligocene and a consequent period of marked
transition of extinct genera to more Recent ones. Wheeler (1914)
found a somewhat similar situation, when summarizing his conclusions
from a study of the ants of the Baltic amber; no data are afforded as
to the number of extinct genera of Miocene ants, but he notes that
forty-four percent of the Lower Oligocene ant types do not appear in
the Recent fauna. Wheeler further notes that Ulmer, in his study of
amber Trichoptera, finds forty-six and four-tenths per cent extinct
genera. Carpenter (1930) finds that forty percent of the Miocene
(Florissant) genera of ants are extinct types. The percentages for
Lower and Upper Oligocene and for Miocene are contrasted in the
table below.

It will be seen that there is a general elimination of generic types in
several groups of insects throughout this period although the extent
of this elimination varies somewhat.

Lower Oligocene
Amber Syrphidae

19 genera
14 extinct

74% "

Lower Oligocene

Amber Ants

(Wheeler)

43 genera
19 extinct

44% "

Lower Oligocene
Amber Tipulidae

38 genera
9 extinct

24.% "

Upper Oligocene
Syrphidae

6 genera
0 extinct

M iocene
Syrphidae

11 genera
3 extinct

27% "

I am strongly inclined to believe that when Syrphus cnrvi petiolatus
Meunier, the Criorrhina of Giebel, and the Tropidia of Handlirsch, the
only modern genera remaining in the amber list except Chcilosia and
Myiolepta, are located and studied, they will also prove to belong at
least to extinct subgenera. Should this prove correct, the amber
Syrphid fauna would consist of over ninety percent extinct forms, and
we would be able to say that their extinction proceeded almost twice
as fast as that of ants and Trichoptera, as far as this period is con-
cerned.

We can hardly say definitely whether a total of one hundred and
fifty to two hundred specimens of amber Syrphids, out of a total
number of amber specimens of insects exceeding one hundred thousand
and perhaps as much as one hundred and fifty thousand, is an indica-
tion of individual abundance or paucity. We know that in some com-
munities today the Syrphid fauna is particularly rich, but almost no

hull: revisional study of fossil syrphidae 347

satisfactory logical work has been done upon individual and species
population. Mr. Charles Johnson reported finding as many as one
hundred and eighteen species in one spot in New England, but this is
certainly exceptional. Williston reported taking in a few days from one
group of elderberry bushes as many as forty-seven species. The highest
list of species from a state is perhaps that of New Jersey with about
two hundred species, to which others have been added since.

As for the records of fossil species, we are safe in saying that con-
siderable morphogenesis was taking place among Syrphids as far back
as the Lower Oligocene, and almost surely extending back well into the
Eocene, since at least six of the seven principal subfamilies had already
differentiated by this time. A study of Recent types would indicate
that the Syrphinae and Cheilosinae are the most generalized groups
among the fourteen subfamilies. Obviously, the fossil record supports
this conclusion. In a certain sense the Syrphinae represented the off-
shoot from which most of the plain and patterned Syrphids arose, and
the Cheilosiinae, with their characteristic melanic coloring, are cer-
tainly the ancestors of the Xylotinae and the Callicerinae. In these
last two subfamilies the dark or aeneous coloration becomes the com-
monplace, and so perhaps they are ancestral to all of these melanoid
and aeneous types. Many Syrphid genera secondarily acquire a paler
appearance owing to the strong development of light yellow pile and
pale, even silvery pubescence. One is tempted to believe that the
possible desirability of pale coloration provides an explanation for the
abundance of the sheen-like pubescence and pollen of many forms;
nevertheless, this is conjecture.

It seems fairly certain, in view of the decided differentiation of the
two main subfamilies in the Eocene, as well as the beginnings surely of
two others, that the Syrphidae originated sometime in the late Cretace-
ous. The flora of this period, containing as it did Platanus, Fagus,
Quercus, Viburnum, Liriodendron, Acer, Liquidamber, etc., in short
many recent deciduous types of shrubs and trees, might easily have
furnished food for Syrphids, for these flowers are highly polliniferous,
and they are the genera today upon which we often find Syrphid flies,
albeit there is now a distinct preference for definitely white rather than
colored blossoms. It is not wholly improbable that the origin of the
Syrphidae goes even farther back in time, for positively identified
Tipulidae and even some higher Diptera have been found in the
Jurassic.

Lastly, I think it is of interest to note that at least one very prom-
inent Oligocene branch was highly developed in a structural sense, and

348

bulletin: museum of comparative zoology

leaves not even a modified descendant today. This is the group of
genera and species centering around Palaeosphegina and including
Palaeoascia and Pseudosphegina.

B3 ^fe-:

1
A

i '

-»

Fig. 2

2. There are two great fossil face types (middle face only con-
sidered), the tuberculate-faced flies and the epistomally produced
flies. We have the choice of regarding one or the other as the more
generalized. The question has particular significance if we wish prop-

hull: revisional study of fossil syrphidae 349

erly to evaluate forms in Syrphid flies, and quite important in other
families of Diptera. It seems to me that the evidence is in favor of the
idea that the epistomal face is more specialized. The text figures given
below illustrate the principal fossil types of face in Syrphid flies.

The following points can be urged as evidence in favor of regarding
the epistomal face as more specialized:

(a) In that subfamily (Xylotinae) to which, excepting the Cerioid-
inae, we accord much the highest specialization, the definitely concave,
"epistomal-thrust-face" is the predominant type; facial tubercles are
quite scarce. Moreover, that subfamily contains very many forms
with the produced front, which as an antennal prominence, necessarily
borrows from the face, and which according to the concept outlined
in the text figure must be derived from this same concave type. Also,
the Cerioidinae, the last and most specialized subfamily, is wholly made
up of such forms ; the upper part of their faces is well thrust forward,
although the oral margin may be slightly retreating. Since these two
very specialized subfamilies contain so many derivatives of the con-
cave face, it seems to indicate that such a face is more specialized than
the convex, tubercle type. Doubtless both concave and convex face
types have arisen from a more or less straight type of face as shown in
text figure 2.

(b). The two lowest subfamilies on the basis of wing development,
usually have a recessive epistomal (convexo-tuberculate) face.

The Oligocene types then, were more specialized in eyes and more
generalized in face. The specialized acquisition of holoptic eyes has
occurred many times in many families, and this acquisition should not
have been difficult. At least eighty percent of Syrphid genera are
holoptic. The well-known dipterist, Osten Sacken, has adduced some
interesting correlations between pedestrian habits and dichopticism,
and the aerial habit with its associated holopticism. In the main his
views seem to be well warranted, but the exceptions of which he
speaks are still largely exceptions. He is wrong in classifying Sphegina
and Neoascia with the aerial flies.

(c). A very few genera have carried epistomal development to an
extreme (Rhingia, Rhinobaccha, Graptomyza, Lycastris, Lycastrir-
rhyncha). I do not see how we are to avoid the contention that these
are among the most specialized members of the family. Since this is a
further specialization of the epistoma, it seems to me to indicate that
the concave face is more specialized.

3. Concerning individual variation in fossil Syrphids, I would like to
point out that Palaeosphegina is of peculiar interest because it displays

350 bulletin: museum of comparative zoology

such a medley of variations. Out of the total of thirty -three specimens
(almost twenty -five per cent of all of the amber material studied) this
genus shows wide variation in many particulars, varying as much or
more than one hundred per cent; it is much more variable than the
closely related Palaeoascia. This latter genus comprises an even
greater percentage of my material. The material on hand in Palaeo-
sphegina, may possibly represent variations produced over quite a long
period of time. This may account for such a striking lot of variations
in a small number of specimens, but regardless of how we account for
it, the genetic pattern was evidently a highly unstable one. This
genus barely differs from modern Sphegina, except in two points, the
tuberculate face and holoptic males. That means that its legs, wings,
abdomen and thorax are practically what we find in modern Sphegina.
Palaeosphegina however, cannot bear the slightest connection with
Sphegina because of its specialization in the matter of the eyes. The
American species of Sphegina are sharply and definitely distinct, the
individuals of any one species highly stable, showing as a rule, only
one or two percent variation in any particular character.

hull: revisional study of fossil syrphidae 351

BIBLIOGRAPHY

Brodie, P. B.

1845. A history of the fossil insects in the secondary rocks of England.
London. 130 pp., pi. 4, fig. 4.

Bronn, Henrich G.

1851-56. Lethaea Geognostica oder Abbildung und Beschreibung der
fur die Gebirges-Formationen. Bezeichnendsten. Versteinerunge.
Teil I, p. 44.

Carpenter, F. M.

1930. The Fossil Ants of North America. Bull. Mus. Comp. Zool., 70,
no. 1, pp. 1-66, pi. 1-11.

1935. Tertiary Insects of the Family Chrysopidae. Jour. Palaeontology,
vol. 9:259-271.

Carpenter, F. M., Snyder, T. E., Alexander, C. P., James, M.,Hull,F. M.
1938. Fossil Insects from the Creede Formation, Colo. Psyche, 45: 105-
120. fig. 3.

COCKERELL, T. D. A.

1909. Fossil insects from Florissant, Colorado. Bull. Am. Mus. Nat.
Hist., vol. 26: 67-76.

1909. New fossil insects from Florissant, Colorado. Annals Ent. Soc.
Am., 2:251-256.

1910. Some insects in Baltic amber. Entomologist. 43:153-155.
1919. Some American fossil insects. Proc. U. S. Nat. Mus., 51 : 96.
1921. Eocene insects from the Rocky mountains. Proc. U. S. Nat. Mus.,

57:232-260.
1924. Fossil insects in the United States National Museum. Pt. II.

Fossil insects and an Arachnid from the Eocene shales of Colorado.

Proc. U. S. Nat. Mus., 64 (13): 1-15; pi. 2, fig. 2.
1926: A fossil alga from the Eocene of Colorado. Torreya, 27: 111-112.

COCKERELL, T. D. A., AND C. LeVeQUE

1931. The antiquity of insect structures. Am. Natural., 45: 351-359.

Foerster, B.

1891. Die Insekten des Plattigen Steinmergels von Brunstatt. Abhand.
Zur Geol., Special Karte B. 3 (5): 335-593.

Germar, E. F.

1837. Insectorum Protogaeae Specimen sistens; Insecta Carbonum Fos-
silium. Fauna Insectorum Europae. Fasc. 19: 25; pi. 25.

Giebel, C. G.

1852. Deutschlands Petrefacten. Der Deutschen Geol. Gesellschaft,

Leipsig. p. 642.

352 bulletin: museum of comparative zoology

1856. Insecten und Spinnen. Fauna d. Vorwelt mit stetter Berruck-
sichtigung der lebenden Thiere. Monographisch dargestellt.
F. U. BrockhaUs, Leipzig. II. pp. 200-202.

Goss, H.

1878. The insect Fauna of the Recent and Tertiary Periods and the
British and foreign formations of those periods in which insect
remains have been detected. Proc. Geol. Association, 5: 282-343.

Handlirsch, A.

1908. Die Fossilen Insekten und der Phylogenie der Resenten Formen.

pp. 102-1025.
1913-1921. in Schroder, Handbuch der Entomologie, 3: 117-306. ill.

Heer, Oswald

1849 Die Insektenfauna der Tertiargebilde von Oeningen und Radoboj

inCroatien. 2:243-246.
1865. Die Urwelt der Schweiz. Figs. 314-315.

Heyden, Carl von

1867-70. Fossil Dipteren aus der Braunkohle von Rott im Siebengebirge.
Palaeontographica. 17: 260-263; pi. 44, 45.

Hope, F. W.

1845-47. Observations on the fossil insects of Aix in Provence. Trans.
Ent. Soc. London, 4: 250-255.

James, Maurice T.

1932. A new Eocene Syrphid from Colorado (Diptera). Canad. Ent.,
64:264.

Loew, Hermann

1850. Ueber den Bernstein und die Bernsteinfauna. Meseritz Progr. k.
realschule. p. 1-48.

Meunier, M. F.

1893. Une Note sur les Syrphidae fossiles del, Ambre tertiaire (Dipteres).

Ann. de la Soc. Entomologique de France, 259.
1895. Observations sur quelques diptgres Tertiares et Catalogue Biblio-

graphique complete sur les insectes fossiles.de cet ordre. Ann.

Soc. Scient. de Bruxelles. 19: 1-16.
1901. Ueber die Syrphiden des Bernstein. Allgemeine Zeitschrift Ent.,

6:70-72.

1903. Description de quelques dipteres de l'ambre. Ann. Soc. Sci. de
Bruxelles. 26:96-104.

1904. Beitrag ziir Syrphiden-Fauna des Bernsteins. Jahrb. Preuss.
Geologisch. ges., 24: 201-201, pi. 13.

Murchison and Lyell

1829. Edinburgh's New Philosophical Journal, Oct. pp. 287-297.

hull: revisional study of fossil syrphidae 353

Pongracz, Alexander

1928. Die Fossilen Insekten von Ungarn, Mit besondere berucksichti-
gung der Entwickelung der Europaischen Insektenfauna. Annales
Musei Nationalis Hungarici. 25: 188-191.

Saporta, Count

1872. Etudes sur la vegetation du sudest de la France a l'epoque
Tertiaire.

Scudder, Samuel H.

1878. Fossil insects of the Green River shales. U. S. Geol. Surv. Terr.

Bull. 4: 747-776.
1890. The Tertiary insects of North America. U. S. Geol. Surv. Terr.

Bull. 13: 557-562.
1890. A classed and annotated Bibliography of Fossil Insects. Bull.

U. S. Geol. Surv. 69: 1-101.
1895. The Miocene Insect fauna of Oeningen, Baden. Geolog. Mag., 4:

116-172; pi. 4.

Serres, Marcel de

1829. Geognosie des Terrains tertiaires du Midi de la France, pp. 254-
258.

Stackelberg, A. v.

1925. Eine neuer fossiler Vertreter der Gattung Tubifera Mgn. (Diptera,
Syrphidae). Revue Russe d' Entom., vol. 19, p. 89-90.

Theobald, Nicholas

1937. Les Insectes fossiles des terrains Oligocenes de France, pp. 1-473.
Nancy.

Unger, Grans.

1839. Fossile Insekten Radoboj. Verhandl. Kais. Leop, Akad. der
Naturf. 19: 412-428.

Weyenbergh, H.

1869. Sur les insectes fossiles du Calcaire Lithographique de la Baviere,
que se trouvent au Musee Teyler. Archives du Musee Teyler,
2:247-294, pi. 34.

WlLLISTON, S. W.

1886. Synopsis of the North American Syrphidae. Bull. U. S. N. M.,
31: 1-335; pi. 1-12.

Zeuner, Friedrich

1931. Die Insektenfauna des Battiger Marmors. Fortschritte der Geol.

und Palaeont., 11 (28): 316.

354

bulletin: museum of comparative zoology

INDEX OF GENERA AND SPECIES

Archalia 305

femorata Hull 305

Arctolepta 287

calamitans Hull 288

Asarcina 272

quadrata Scudder 272

Cacogaster 297

novamaculata Hull 298

Cheilosia 274

ampla Scudder 274

bruesi Hull 283

dubia Weyenbergh 344

germanica Hull ' 281

hecate Hull 277

miocenica Cock 275

nigrachaeta Hull 282

oligocenica Theobald 280

pratjei Hull 279

scudderi Cock, and Le Veque 276
eepultula Cock 284

Cheilosialepta (e) 285

baltica Hull 286

Chrysogaster . 296

antiquaria Hull 296

Criorrhina 338

sp 338

Doliomyia 328

chalybea Hull 330

Eoxylota 324

pulchra Meunier 324

Eristalis 335

lapideus Scudder 343

paucisinuatus Theobold .... 335

Helophilus 332

miocaenicus 332

primarius Germar 343

villeneuvi Theobald 332

Leucozona 271

nigra Hull 271

Megaxylota 340

magnifemur Hull 341

Merodon 333

germari Heyden 333

Microdon 330

sp 330

Milesia 333

Myiolepta 299

andreeiHull 299

germanica Hull 300

luhei Cock 301

valida Hull 299

woteni Hull 302

Palaeoascia 306

atrata Hull 309

nigra Hull 310

uniappendiculata Meunier . . 307
uniappendiculata brachypen-

nis Hull 309

uniappendiculataobtusaHull 309

Palaeoeristalis 333

tessellatus Hull 334

Palaeopipiza 327

xenos Meunier 328

Palaeosphegina 316

baccha Hull 320

elegantula Meunier 317

elegantula atrox Hull 319

elegantula tristis Hull 319

fumosa Hull 321

pilosa Hull 320

Pipiza 293

melanderi Hull 293

venilia Heyden 293

Platycheirus 268

haidingeri Heer 268

infumatus Heer 270

persistens Hull 269

Protochrysotoxum 326

sphinx Hull 326

Protorhingia 289

carpenteri Hull 290

magnipennis Hull 291

Pseudopipiza 294

antiqua Hull 295

europa Hull 295

Pseudosphegina 313

hull: revisional study of fossil syrphidae

355

dichoptica Hull 314

withersi Hull 315

Psilota 344

tabidosa Scudder 344

Ptilocephala 335

volucelloides Hull 337

Remalia 345

sphinx Brodie 345

Rhingia 292

sp 292

zephyrea Hull 292

Sericolepta 303

maculata Hull 304

Sphegina 322

abrasa Theobald 322

obscura Hull 323

Spheginascia 311

biappendiculata Meunier . . . 312

biappendiculata rectinervis

Hull 313

Syrphopsis 345

globosicepsZeuner 345

Syrphus 258

aphidopsidis Cock 258

bremii Heer 342

carpenteri Hull 262

curvipetiolatus Meunier .... 342

eocenicus Cock 264

euphemus Heyden 265

freyeri Heer 266

geminatus Heer 267

hendersoni James 260

bthaphidis Cock 264

platychiralis Hull 261

reciprocus Foerster 268

schellenbergi Heer 343

willistoni Cock 259

Tropidia 342

sp 342

Volucella 338

sp 338

Xylotosyrphus 339

pulchrafenestra Hull 339

PLATES

PLATE 1

Hull — Fossil Syrphidae

PLATE 1

A. Cheilosia scudderi Cockerell & Leveque; (holotype).

B. Syrphus carpenteri Hull; (holotype).

C. Cheilosia miocenica Cockerell; (holotype).

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 1

■ \ <

PLATE 2

Hull — Fossil Syrphidae

PLATE 2

A. Protochrysotoxum sphinx Hull; holotype.

B. Leucozona nigra Hull; holotype.

C. Pipiza melanderi Hull; holotype.

D. Asarcina quadrata Scudder; holotype.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 2

PLATE 3

Hull — Fossil Syrphidae

PLATE 3

A. Syrphus willistoni Cockerell; 9 . (specimen no. 3960, M.C.Z.)

B. Archalia femorata Hull; holotype.

C. Syrphus willistoni Cockerell; allotype.

D. Cheilosia hecate Hull; holotype.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 3

v ''X

PLATE 4

Hull — Fossil Syrphidae

PLATE 4

Fig. 1. Sphegina obscura Hull; wing of female (type).

Fig. 2. Sphegina obscura Hull; abdomen of female (type).

Fig. 3. Sphegina obscura Hull; hind femur of female (type).

Fig. 4. Platycheirus persistens Hull; abdomen of male (type).

Fig. 5. Platycheirus persistens Hull; tibia and tarsi of male (type).

Fig. 6. Syrphus willistoni Cockerell ; antenna with arista.

Fig. 7. Palaeoeristalis tessellatus Hull; hind femur and tibia (type).

Fig. 8. Palaeoeristalis tessellatus Hull; abdomen (type).

Fig. 9. Palaeoeristalis tessellatus Hull; wing (type).

Fig. 10. Archalia femorata Hull; profile of face (type).

Fig. 11. Archalia femorata Hull; hind femur (type).

Fig. 12. X ylotosyrphus pulchrafenestra Hull; hind femur (type).

Fig. 13. Cheilosia sp. ; scutellum.

Fig. 14. Cheilosia hecate Hull; profile of head (paratype).

Fig. 15. Syrphus willistoni Cockerell; antenna with arista.

3ULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 4

<g&2>~

Hull — Fossil Syrphidae

PLATE 4

Fig. 1. Sphegina obscura Hull; wing of female (type).

Fig. 2. Sphegina obscura Hull; abdomen of female (type).

Fig. 3. Sphegina obscura Hull; hind femur of female (type).

Fig. 4. Platycheirus persistens Hull; abdomen of male (type).

Fig. 5. Platycheirus persistens Hull; tibia and tarsi of male (type).

Fig. 6. Syrphus willistoni Cockerell ; antenna with arista.

Fig. 7. Palaeoeristalis tessellatus Hull; hind femur and tibia (type).

Fig. 8. Palaeoeristalis tessellatus Hull; abdomen (type).

Fig. 9. Palaeoeristalis tessellatus Hull; wing (type).

Fig. 10. Archalia femorata Hull; profile of face (type).

Fig. 11. Archalia femorata Hull; hind femur (type).

Fig. 12. Xylotosyrphus pulchrafenestra Hull; hind femur (type).

Fig. 13. Cheilosia sp.; scutellum.

Fig. 14. Cheilosia hecate Hull; profile of head (paratype).

Fig. 15. Syrphus willistoni Cockerell; antenna with arista.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 4

PLATE 5

Hull — Fossil Syrphidae

PLATE 5

Fig. 16. Palaeoascia uniappendiculata Meunier; hind femur (male).

Fig. 17. Palaeoascia uniappendiculata Meunier; face from in front.

Fig. 18. Palaeoascia uniappendiculata Meunier; antenna with arista (type).

Fig. 19. Palaeoascia uniappendiculata Meunier; abdomen of male.

Fig. 20. Palaeoascia uniappendiculata Meunier; abdomen of female.

Fig. 21. Palaeoascia atrata Hull; wing tip.

Fig. 22. Palaeoascia atrata Hull; wing tip.

Fig. 23. Palaeoascia atrata Hull; abdominal segment and genitalia of male

Fig. 24. Palaeoascia atrata Hull; antenna with arista.

Fig. 25. Syrphus aphidopsidis Cockerell; antenna (obverse of type).

Fig. 26. Merodon germari Heyden; abdomen and hind legs (type).

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 5

PLATE 6

Hctll — -Fossil Syrphidae

PLATE 6

Pseudosphegina dichoptica Hull; head from above (type).
Pseudosphegina dichoptica Hull; hind femur and tibia (type).
Eoxylota pulchra (Meunier) ; hind femur, tibia and basi tarsi (type).
Eoxylota pulchra (Meunier); profile of head (type).
Ptilocephala volucelloides Hull; profile of head (type).
Ptilocephala volucelloides Hull; wing (type).
Ptilocephala volucelloides Hull; hind femur and tibia (type).
Ptilocephala volucelloides Hull; thora and scutellum (type).
Ptilocephala volucelloides Hull; antenna with arista (type).
Arctolepta calamitans Hull; scutellum (type).
Fig. 37. Cacogaster novamaculala Hull; head from above (type).

Fig.

27.

Fig.

28.

Fig.

29.

Fig.

30.

Fig.

31.

Fig.

32.

Fig.

33.

Fig.

34.

Fig.

35.

Fig.

36.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 6

PLATE 7

Hull — Fossil Syrphidae

PLATE 7

Fig. 38. Pseudosphegina dichoptica Hull; wing (type).

Fig. 39. Pseudosphegina dichoptica Hull; abdomen and hypopygium (type)

Fig. 40. Megaxylota magnifemur Hull; abdomen of female (type).

Fig. 41. Megaxylota magnifemur Hull; wing (type).

Fig. 42. Megaxylota magnifemur Hull; head from above.

Fig. 43. Palaeoascia uniappendiculata Meunier; hind femur of male.

Fig. 44. Palaeoascia uniappendiculata Meunier, var. brachypennis Hull;

hind femur.

Fig. 45. Palaeoascia uniappendiculata Meunier; head of female from above.

Fig. 46. Palaeoascia uniappendiculata Meunier, var. brachypennis Hull;

wing.

Fig. 47. Palaeoascia uniappendiculata Meunier; hind femur.

Fig. 48. Palaeoascia uniappendiculata Meunier; head of male from above.

Fig. 49. Palaeoascia nigra Hull; head in profile (type).

Fig. 50. Spheginascia biappendiculata Meunier; wing (type).

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 7

PLATE 8

Hull — Fossil Syrphidae

PLATE 8

Fig. 51. Palaeoascia uniappendiculata Meunier; lateral view (type).

Fig. 52. Megaxylola magnifemur Hull; profile of head (type).

Fig. 53. Megaxylola magnifemur Hull; hind femur and tibia (type).

Fig. 54. Cheilosialepta baltica Hull; antenna and arista.

Fig. 55. Palaeopipiza xenos Meunier; antenna and arista (type).

Fig. 56. Palaeopipiza xenos Meunier; profile of head (type).

Fig. 57. Palaeopipiza xenos Meunier; scutellum (partial reconstruction

from type).

Fig. 58. Doliomyia chalybea Hull; scutellum (type).

Fig. 59. Doliomyia chalybea Hull; head from above (type female).

Fig. 60. Merodon germari Heyden; head from above (type).

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 8

PLATE 9

Hull — Fossil Syrphidae

PLATE 9

Fig. 61. Spheginascia biappendiculata Meunier; wing (type).

Fig. 62. Doliomyia chalybea Hull; profile of head (type).

Fig. 63. Palaeosphegina elegantula Meunier; hind femur and tibia (type).

Fig. 64. Palaeosphegina elegantula Meunier; tibia and basitarsus (type).

Fig. 65. Palaeosphegina elegantula Meunier; profile of head (type).

Fig. 66. Spheginascia biappendiculata Meunier; profile of head (type).

Fig. 67. Cheilosialepta baltica Hull; head from above, male.

Fig. 68. Cheilosialepta baltica Hull; head from oblique view.

Fig. 69. Cheilosialepta baltica Hull; scutellum.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 9

-^g*»^>>^

PLATE 10

Hull — Fossil Syrphidae

PLATE 10

Fig. 70. Spheginascia biappendiculata Meunier; head from above (type).

Fig. 71. Spheginascia biappendiculata Meunier; profile of head (type).

Fig. 72. Eoxylota pulchra (Meunier); abdomen (type).

Fig. 73. Eoxylota pulchra (Meunier); wing (type).

Fig. 74. Arctolepta calamitans Hull; head from below (type).

Fig. 75. Arctolepta calamitans Hull; scutellum (type).

Fig. 76. Arctolepta calamitans Hull; wing (type).

Fig. 77. Protorhingia carpenteri Hull; wing (type).

Fig. 78. Protorhingia magnipennis Hull; wing (type).

Fig. 79. Cheilosialepta baltica Hull; wing.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 10

PLATE 11

Hull — Fossil Syrphidae

PLATE 11

Fig. 80. Cheilosia nigrachaeta Hull ; profile of face.

Fig. 81. Pseudosphegina withersi Hull; face from above.

Fig. 82. Pseudosphegina withersi Hull; profile of face.

Fig. 83. Cheilosia germanica Hull; profile of face.

Fig. 84. Sericolepta maculata Hull; face from above.

Fig. 85. Sericolepta maculata Hull; antenna from outside.

Fig. 86. Pseudopipiza antiqua Hull ; profile of face.

Fig. 87. Sericolepta maculata Hull; profile of face.

Fig. 88. Myiolepta andreei Hull; profile of face.

Fig. 89. Myiolepta valida Hull ; face from above.

Fig. 90. Cheilosia bruesi Hull ; face from above.

Fig. 91. Myiolepta germanica Hull; profile of face.

Fig. 92. Myiolepta germanica Hull; face from above.

Fig. 93. Cheilosia pratjei Hull; face from above.

Fig. 94. Cheilosia pratjei Hull; hind femur.

Fig. 95. Cheilosia germanica Hull; hind femur.

Fig. 96. Pseudopipiza europa Hull; vertex, occiput and ocelli.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 11

PLATE 12

Hull — Fossil Syrphidae

Fig
Fig
Fig
Fig
Fig
Fig
Fig
Fig
Fig
Fig
Fig
Fig
Fig
Fig

PLATE 12

97. Myiolepta valida Hull; scutellum.

98. Cheilosia bruesi Hull; scutellum.

99. Cheilosia germanica Hull; scutellum.

100. Sericolepta maculata Hull; scutellum.

101. Myiolepta germanica Hull; scutellum.

102. Cheilosia nigrachaeta Hull; scutellum.

103. Myiolepta andreei Hull; scutellum.

104. Cheilosia pratjei Hull; scutellum.

105. Myiolepta andreei Hull; hind femur and tibia.

106. Myiolepta germanica Hull; hind femur and tibia.

107. Cheilosia nigrachaeta Hull: wing.

108. Myiolepta valida Hull; wing.

109. Sericolepta maculata Hull; abdomen.

110. Sericolepta maculata Hull; hind femur and tibia.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Pi_ate 12

PLATE 13

Hull — Fossil Syrphidae

PLATE 13

Fig.

111.

Fig.

112.

Fig.

113.

Fig.

114.

Fig.

115.

Fig.

116.

Fig.

117.

Fig.

118.

Fig.

119.

Fig.

120.

Fig.

121.

Cheilosia pratjei Hull; wing.

Cheilosia germanica Hull ; wing.

Pseudopipiza antiqua Hull; wing.

Myiolepta andreei Hull; wing.

Pseudosphegina wither si Hull; wing.

Doliomyia chalybea Hull ; wing.

Palaeopipiza xenos Meunier; wing. From type.

Cheilosia bruesi Hull; wing.

Sericolepta maculata Hull; wing.

Myiolepta germanica Hull; wing.

Pseudopipiza europa Hull, wing.

BULL. MUS. COMP. ZOOL.

Hull. Fossil Syrphidae. Plate 13

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. XCV, No. 4

THE ARGIOPIDAE OF HISPANIOLA

By Elizabeth B. Bryant

With Four Plates

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

May, 1945

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIV, and Vol. XCV, No. 1, 2, 3 and
4 have appeared and of the Memoirs, Vol. I to LV.

Publication of Memoirs ceased with Vol. LV.

Bulletin of the Museum of Comparative Zoology
. AT HARVARD COLLEGE
Vol. XCV, No. 4

THE ARGIOPIDAE OF HISPANIOLA

By Elizabeth B. Bryant

With Four Plates

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

May, 1945

ogy <rf.

MAY 15 1945

No. 4. — The Argiopidae of Hispaniola1
By Elizabeth B. Bryant

Many of the Argiopidae are large and showy spiders, but though
calculated to attract the attention of collectors, only one species was
first described from Hispaniola. In 1895, E. Simon founded the genus
Alcimosphenus on the species licinus, from S. Domingo and Jamaica.
It has since been found on several of the islands of the Caribbean,
but the male is still unknown.

Mr. Banks (1903) published a brief paper entitled "A List of
Arachnida from Hayti, with descriptions of New Species." This
paper noted in all sixty-three species, seventeen of which belong to
the family Argiopidae, though today several are regarded as synonyms.
Eustala prompta (Hentz) = anastera Walck., has not been found again
on the island, though it is common on many of the Antilles ; Wagneriana
unidcm-tuberculata (Keys.) has been found on several of the islands
of the Caribbean and may be rediscovered on Hispaniola when
collections are made at different seasons. Shiga crewi Banks, described
as new from a female, has recently been rediscovered with the male
at Port-au-Prince, the type locality.

The classification of the Argiopidae is probably as satisfactory as
that of any other large and widely distributed group, where the male
and female differ greatly in size, color and secondary characters.
While it is often difficult to place the females in the various sub-
families, males are distinguished by one or more striking characters.
F.O.P.-Cambridge in his "Arachnida of Central-America", estab-
lished a number of new genera, but even in so restricted an area, he
was obliged to relegate several species, known from both sexes, to
the genus Aranea, for lack of definite characters, though he stated
that no Central American species was congeneric with the European
genotype. The same is true of the fauna of Hispaniola. Many of
the Central American genera of F.O.P.-Cambridge have been found
on that island, but no species of Aranea, as restricted today.

In all, the writer has seen fifty -nine species of the Argiopidae.
Some of these are widely distributed, like Nephila clavipcs (Linn.),
found everywhere in the tropics of the New World; Cyclosa oculata
(Walck.), first reported from the vicinity of Paris, an undoubted
horticultural importation, and known today from northern South
America and most of the islands of the Caribbean; Wendilgarda
theridionina Simon, described first from Venezuela and since found
to be widely distributed; Acacesia hamata (Hentz), a common species

1 Published with the aid of a special gift from Mr. George R. Agassiz.

360 bulletin: museum of comparative zoology

in the southern part of the United States, is also found on many of
the islands ; Parawixia carnbridgei Bryant, so far known only from the
type locality, Pico Turquino, Eastern Cuba. Two genera are definitely
of South American origin, Agriognaiha and Ogulnius. The former
is based on a species from Peru, known by both male and female.
Three species have been found in Central America, one is reported
from St. Vincent and three additional species have been added from
Hispaniola. The genus Ogulnius was based on probably the smallest
spider known (0.5 mm. long) from the Amazon. Two more species
of this genus have been reported from St. Catharina, Brazil, one from
the island of St. Vincent, and another from the foot-hills of the
Cordillera Central, south of Santiago, Hispaniola.

Of the fifty-nine species seen, twenty-seven are described as new.
Many of these are small, less than 2.0 mm. in length, and were taken
by Dr. P. J. Darlington on isolated peaks of the Hispaniola mountains,
indicating a restricted distribution.

I gladly express my appreciation of the skill and understanding of
Dr. Darlington, who has collected so much of the material upon
which this paper is based. My sincere thanks are also due to Dr.
Alexander Petrunkevitch, who so kindly compared specimens of
Shiga crewi Banks with his type of Larinia coamensis Petr. from
Puerto Rico; to Dr. W. M. Mann, who collected in Haiti during the
winter of 1912-1913, taking several species not found by other col-
lectors; to Dr. Andre Audant, I am indebted for several interesting
species; to Donald Hurst for material from Puerto Plata. I am
especially grateful to Mr. Nathan Banks for never failing interest and
encouragement.

Family ARGIOPIDAE

Key to Subfamilies of Argiopidac found in Hispaniola

1. Small spiders; eyes heterogeneous; central spinners with posterior pair form

a straight line Thcridiosomatinae

Larger spiders; eyes homogeneous; spinners closely grouped 2

2. Mandibles usually without a boss, or boss rudimentary; genital fold with-

out a furrow; legs with hairs or bristles or few spines 3

Mandibles with a boss; genital fold with a furrow; legs with true spines . . 4

3. Boss absent; paracymbium of male palpus long, narrow and parallel to

cymbium Tetragnathinae

Boss rudimentary; paracymbium short, chitinized and at right angles to
cymbium Metinae

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 361

4. Legs very long, metatarsi and tarsi longer than tibia and patella 5

Legs relatively shorter, metatarsi and tarsi shorter than patella and tibia .6

5. Vulva of female simple ; posterior eye row straight ; carapace convex ; labium

longer than broad Nephilinae

Vulva of female developed; posterior eye row strongly procurved; carapace
flat, labium broader than long Argiopinae

6. Spinners surrounded by a corneous ring; abdomen hard. .Gasteracanthinae
Spinners not surrounded by a corneous ring; abdomen soft Araneinae

Subfamily ARGIOPINAE
Key to species

1. Eyes of anterior row equidistant Gea heptagon

A.M.E. further from laterals than from each other 2

Females

2. Abdomen lobate on sides Argiope argentata

Abdomen evenly rounded on sides Argiope trifasciata

Males

Embolus with lateral barbs near tip Argiope argentata

Embolus smooth Argiope trifasciata

Genus Argiope Audouin 1826

Argiope argentata (Fabr.)

Aranea argentata Fabricius, 1775, p. 433. [ 9 ] "in Indiis"

3 cf 9 Dom. Rep.; Puerto Plata, April-May 1941, (Hurst)
cf 9 Haiti; Diquini, November 1912, (Mann)

9 Haiti; Port-au-Prince, 2 October 1934, (Darlington)
9 Haiti; Kenskoff, 4,300 feet, May 1935, (Roys)

Argiope trifasciata (Forskal)
Aranea trifasciata Forskal, 1775, p. 86. [ 9 ] "Kahirae"

9 pullus Haiti; Kenskoff, 4,300 feet, 2 May 1935, (Roys)

Genus Gea C. Koch 1843
Gea heptagon (Hentz)

Epeira heptagon Hentz, 1850, p. 20, pi. 3, figs. 5, 6; reprint, 1875, p. 122, pi. 14,

figs. 5, 6, 'V 9 North Carolina; Alabama"

9 Haiti; Ennery, 10 September 1934, (Darlington)
9 Dom. Rep.; Puerto Plata, April-May 1941, (Hurst)

362 bulletin: museum of comparative zoology

Subfamily ARANEINAE
Key to Genera

Males

1 . Patella of palpus with 2 spines at apex 2

Patella of palpus with 1 spine at apex 5

2. First coxa without a hook Metepeira

First coxa with a hook 3

3. Lateral eyes on a prominent tubercle projecting forward in advance of

anterior margin of mandible; palpus without a large paracymbium. . . .

Eriophora
Lateral eyes not on a prominent tubercle, palpus with a large para-
cymbium : 4

4. Clavis of genital bulb with a characteristic lateral spur, but not distinctly

bifid Neoscona

Clavis of bulb distinctly bifid Neosconella

5. Third tibia with tuft of cilia on anterior basal side M angora

Third tibia with no tuft of cilia 6

6. First coxa without a hook, anterior row of eyes strongly procurved

Marxia
First coxa with a hook, anterior row of eyes straight or slightly recurved .7

7. Second tibia with a stout apophysis about middle 8

Second tibia with no apophysis about middle : 9

8. Apophysis with two stout spines at apex Verrucosa

Apophysis more slender with a single, long stout apical spine. . . .Edricvs

9. Quadrangle of eyes not narrowed behind, p.m.e. two diameters apart and

larger than a.m.e Wixia

Quadrangle of eyes narrowed behind, p.m.e. one diameter apart, smaller
than a.m.e 10

10. Eye groups very prominent, lateral eyes on a tubercle that projects

beyond anterior margins of mandibles 11

Eye groups not prominent, lateral eyes not projecting forward 12

11. Second tibia incrassate, with a prolateral double row of 5-6 stout spines,

5-6 long ventral spines, third and fourth femora with 5-6 ventral

spines Parawixia

Second tibia scarcely incrassate, with a single prolateral series of three
fine spines, third and fourth femora without spines, or one or two on
third femur Cyclosa

12. Abdomen with shoulder, lateral and apical cusps Wagneriana

Abdomen without cusps 13

13. Second tibia with specialized holding spines, joint somewhat incrassate .14
Second tibia not incrassate, nor with any specialized holding spines .... 15

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 363

14. Specialized spines not arranged in a series but with two or more spines

towards base, one or two very long curved prolateral spines, one or two

shorter, stouter curved ventral spines Acacesia

Specialized spines arranged in a more or less definite series Eustala

15. Posterior femora with ventral spines, fourth coxa not modified

Metazygia

Posterior femora with no ventral spines, fourth coxa with a short black

spine on posterior margin Aranea(?)

Females

1 . Abdomen with cusps or tubercles 2

Abdomen with no cusps or tubercles 8

2. A dorsal basal tubercle as well as lateral tubercles Marxia

No dorsal basal tubercle 3

3. Femora with ventral spines 4

Femora with no ventral spines . . 5

4. Metatarsi with no ventral spines, scapus very long Verrucosa

Metatarsi with few ventral spines, scapus short Parawixia

5. P.M.E. larger than A.M.E Wixia

P.M.E. smaller than A.M.E 6

6. Cephalic portion not separated from thoracic by a semi-circular furrow. .

Wagneriana
Cephalic portion separated from thoracic by a deep semi-circular furrow .7

7. Median quadrangle of eyes narrower behind Cyclosv

Median quadrangle of eyes not narrowed behind Edricus

8. Third tibia with tuft of curved cilia on anterior basal side Mangora

Third tibia with no tuft of curved cilia 9

9. Femora with ventral spines 10

Femora with no ventral spines 12

10. First metatarsus + tarsus longer than first tibia + patella, scapus

recurved Metepeirs

First metatarsus -\- tarsus shorter than first tibia + patella la

1 1 . Scapus very long Eriophorl

Scapus very short . Neoscona

12. First and second tibiae with ventral spines la

First and second tibiae with no ventral spines NeosconellS

13. Anterior margin of cephalothorax very broad Metazygia

Anterior margin of cephalothorax narrow 14

14. Scapus directed forward Eustala

Scapus directed backward 15

15. Scapus short and broad, spiders small Aranea(?)

Scapus protruding from plane of abdomen, spiders larger Acacesia

364 bulletin: museum of comparative zoology

Genus Acacesia Simon 1875

ACACESIA HAMATA (Hentz)

Epeira ? hamata Hentz, 1847, p. 474, pi. 31, fig. 10; reprint, 1875, p. 114,

pi. 13, fig. 10. "c? Alabama, August"
Epeira foliata id., ibid., p. 475, pi. 31, fig. 14; reprint, p. 116, pi. 13, fig. 14.

" 9 Alabama, June, July"
Epeira folif era Marx, 1889, p. 545
Araneus hallucinor Petrunkevitch, 1911, p. 296.

3 cf Haiti; Port-au-Prince, (Crew), Banks Coll.

Genus Aranea Linnaeus 1756

Aranea (?) creavi (Banks)

Figures 1, 3

Singa crewi Banks, 1903, p. 342, pi. 15, fig. 8. " 9 Hayti"
Larinia coamensis Petrunkevitch, 1930, p. 335, figs. 221-224. " 9 Coamo
Springs, Porto Rico"

Male. Length, 4.0 mm., ceph. 2.0 mm. long, 1.6 mm. wide, abd.
2.6 mm.

Cephalothorax olivaceous, margins and median line clouded with
black, eye area dark, quite low and flat, highest anterior to groove
and gibbose each side of groove, sides gently rounded, anterior margin
abruptly narrowed, a long bristle posterior to lateral eyes, thoracic
groove long and distinct, no median row of short bristles posterior
to second eye row as in female; eyes cover entire anterior margin,
anterior row strongly recurved, eyes equidistant, a.m.e. largest of
the eight, carried forward, separated by less than a diameter, a. I.e.
about a radius of a.m.e., posterior row slightly recurved, lateral eyes
subequal and almost touching, p.m.e. little larger than p. I.e., separated
by a line; quadrangle wider in front, longer than wide; clypeus almost
wanting below a.m.e.; mandibles vertical, long, weak, basal half
swollen; labium pale, fused to sternum, tip strongly rebordered;
maxillae pale, more than twice as long as labium, inner margins
parallel, tips widened; sternum about two-thirds as wide as long,
black, with a pale median stripe from labium to beyond the middle,
carried between II and III coxae, IV coxae almost touching, with a
basal tubercle and a short black spine on posterior margin, I coxae
with a hook at retrolateral angle, and a large basal tubercle ; abdomen
pale, black at base and sides, oval, flattened with a pair of faint gray
lines that form an indistinct folium, two pairs of muscle spots, venter

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 365

dark, with a pale median spot; legs much broken, 1-2-4-3, femora
pale, with distal dark bands, tibiae with dark basal and distal dark
bands, spines, I pair, femur, ventral, 3 short spines about middle,
prolateral, 3, dorsal, 4 median, patella, 1 long spine at tip, 1 at base,
retrolateral, 2, prolateral, 0, tibia, scattered spines, II pair not enlarged,
with no specialized spines, femur, ventral, 2 short spines about middle,
patella same as I pair, tibia, spines heavier than I pair, III pair,
femur, ventral, 2 short spines about middle, patella, 1 at tip and
1 at base, lateral, 1-1, tibia, scattered, IV pair, coxa, 1 long sharp
spine on posterior margin, femur, ventral, 3 very long spines near
base, patella, 1 at tip, 1 at base, lateral, 1-1; palpus shorter than
cephalothorax, patella as long as wide, 1 very long bristle at tip, tibia
shorter than patella, with 1 very long bristle, palpal organ large, cym-
bium with a deep notch on retrolateral side with a slender lobe that
ends in a long white bristle, ending near tip of cymbium, embolus
near tip, a long, slender black tube beneath a broad, thin sheath.
Allotype cf Dom. Rep.; Puerto Plata, July 1940, (Hurst)
cF 9 Dom. Rep.; Puerto Plata, July, August 1940, (Hurst)
cf1 9 s Haiti Port-au-Prince, July 1941, (Audant)
Shiga crewi Banks was based on a female, probably collected near
Port-au-Prince. The description of the color markings is quite com-
plete, but no mention is made of the eye arrangement, spines on the
legs or the epigynum, a figure of which is not very clear. Petrunkevitch
who redescribed this species as Larinia coamensis, also from a female,
collected near Coamo Springs, Puerto Rico, figures the entire spider,
eyes and epigynum. His figure shows the cephalothorax much more
slender than in the Hispaniola specimens, but Dr. Petrunkevitch has
kindly compared a pair of the Port-au-Prince material with his type
and reports them to be the same.

Larinia silvestris Bryant, known only from the type, a male, from
Puerto Rico, is congeneric with Sitiga crewi Banks. Both have similar
palpi, with the long embolus at the tip of the bulb, protected by a
sheath-like conductor, and both have the posterior coxae of the male
modified. Larinia silvestris has two small black cusps on the pro-
lateral margin of the third and fourth coxae, while Singa crewi has
a sharp black spine on the posterior margin of the fourth coxa.

These two species undoubtedly belong near Aranea incerta (O.P.-
Cambr.), found in Mexico, Guatemala and Costa Rica. This species
has great variation in size and color markings. The male has a small
black cusp at the upper prolateral angle of the fourth coxae, which
was not noted by either O.P.-Cambridge or F.O.P.-Cambridge.

366 bulletin: museum of comparative zoology

These three species cannot be placed in either the genus Singa or
Larinia, as the males of both of these genera have two stout bristles
at the tip of the patella of the male palpus, a character that is appar-
ently trivial, but is constant as far as known in all Argiopid genera.
F.O.P.-Cambridge left the species incerta in the genus Aranea but
states that it is not typical of the genus.

In the Hispaniola specimens of crewi, there is some variation in
color markings in the females, but little variation in size. A few have
no dark median line on the cephalothorax, or dark bands on the legs,
and the dorsum is pale with dark sides that extend to the base.

Aranea(?) hispaniola spec. nov.
Figure 2

Female. Length, 4.0 mm., ceph. 1.8 mm., abd. 2.6 mm. long,
2.0 mm. wide.

Cephalothorax yellow, with a gray triangle from eyes to near posterior
margin, a median row of five long, overlapping bristles to posterior
eye row, anterior margin little over half the greatest width, groove a
shallow depression, cephalic portion high; eyes, anterior row recurved,
eyes equidistant, a.m.e. largest of the eight, separated by a diameter,
posterior row very slightly recurved, p.m.e. two-thirds diameter of
a.m.e. and separated by less than a diameter, lateral eyes contiguous,
subequal and on a low tubercle; quadrangle wider in front than behind
and wider than high; clypeus less than a diameter of a.m.e.; mandibles
pale, shaded with gray, no boss, three teeth on superior margin, and
three teeth on inferior margin; labium dark, wider than long, tip
rebordered; maxillae more than twice as long as labium, outer half
dark, sides parallel; sternum four-fifths as wide as long, heart-shaped,
not extending between the IV coxae, black with a wide median stripe
of yellow that extends past the middle, IV coxae separated by half a
diameter; abdomen oval, anterior end extending in a slight point over
cephalothorax, dorsum with a wide median dark stripe that almost
covers width of abdomen, lateral margins irregular and darker, sides
pale, venter with a black rectangle from fold to spinnerets containing
a median pale spot below the epigynum; legs, 1-2-4-3, pale, with
many black spots, no dark rings, spines, no ventral femoral spines,
I pair, femur, prolateral, 2, each from a black spot, retrolateral, 0,
dorsal, 2 on basal half, patella, prolateral, 2, retrolateral, 1, dorsal,
2, apical and basal, tibia, prolateral, 3, retrolateral, 4, dorsal, 2,
ventral, 2-2, metatarsus, dorsal, 1 at base, prolateral, 2, retrolateral,

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 367

2, ventral, 2-2; posterior pairs with dorsal basal spine on tibiae;

epigynum area wider than long, parts very pale, a broad, very short

median scape with sides near tip rebordered, lateral margins of area

with a broad transparent chitinized cover, the depressed area each

side of scape with little structure.

Holotype 9 Haiti; Kenskoff, 4,300 feet, 1 May 1935, (Roys)
Paratype 9 Dom. Rep.; San Jose de las Matas, 1,500 feet+, June

1938, (Darlington)

Unfortunately the anterior legs in both type and paratype are

broken, so that the relative length of the joints cannot be noted.

Genus Cyclosa Menge 1866
Cyclosa bifurca (McCook)

Cyrtophora bifurca McCook, 1887, p. 342. " 9 Florida; Merrit's Island,

Indian River"
Cyclosa fissicauda O.P.-Cambridge, 1889, p. 49, pi. 9, fig. 7. " 9 Guatemala,

between Dolores and Chacallal."

9 s Dom. Rep.; Puerto Plata, April-August 1941, (Hurst)
c? Haiti; Kenskoff, 4,300 feet, 3 May 1935, (Roys)

Cyclosa oculata (Walckenaer)

Epeira oculata Walckenaer, 1802, p. 421. V 9 Paris"
Turkheimia walckenaeri O.P.-Cambridge, 1889, p. 47, pi. 8, fig. 6. " 9 Guat-
emala, Volcan de Fuego and Chiacan."

c? 9 Haiti; Kenskoff, 4,200 feet, 2 May 1935, (Roys)

9 Haiti; La Visite, 6,000-7,000 feet, 16-23 September 1934,

(Darlington)
9 Haiti; Furcy, Mt. Cabaio, 7,000 feet, 1940, (Folk)
9 Dom. Rep.; Puerto Plata, April-May 1941, (Hurst)

Genus Drexelia McCook 1892
Drexelia minor spec. nov.
Figure 10

Female. Length, 5.0 mm., ceph. 2.0 mm., abd. 3.4 mm. long, 1.6
mm. wide.

Cephalothorax pale, with quite a wide dark line from p.m.e. to
near posterior margin, a broken marginal dark line, cephalothorax

368 bulletin: museum of comparative zoology

low and flat; eyes cover anterior margin, anterior row recurved,
a.m.e. largest of the eight, separated by more than a diameter and
from a.l.e. by fully two diameters, posterior row very slightly pro-
curved, lateral eyes subequal and touching, p.m.e. separated by less
than a radius and only slightly larger than p. I.e.; quadrangle broader
than long, about half as wide behind as in front; clypeus less than
a diameter of a.m.e.; mandibles pale, vertical; labium pale, wider
than long, tip rebordered; maxillae pale, twice as long as labium and
parallel; sternum with a wide black margin and a pale center, two-
thirds as wide as long, ending in a point in front of IV coxae; abdomen
more than twice as long as wide, extending in a point over cephalo-
thorax, tip rounded, with five narrow dark stripes, separated by
narrower pale stripes, no indication of the paired dark spots found in
D. directa (Hentz), sides mottled, venter with a long central pale spot
surrounded by a broad black frame; legs, 1-2-4-3, brown, pale with
parallel ventral and lateral rows of black dots on femora, no ventral
femoral spines, spines broken; epigynum a broad median scape, with
tip rebordered and a large circular depression each side.
Holotype 9 Haiti; Port-au-Prince, July 1941, (Audant)
Drexelia minor is very closely related to D. directa (Hentz), the
genotype, but it is smaller and more delicate. An immature male,
also from Port-au-Prince, has the same markings. The epigynum is
much broader in proportion than in directa and the openings are
larger.

Genus Edricus O.P.-Cambridge 1890

Edricus crassicauda (Keyserling)

Epeira crassicauda Keyserling, 1865, p. 806, pi. 18, figs. 3, 4. " 9 N. Grenada,
Keyserling Collection."

9 pullus Dom. Rep.; Puerto Plata, July-August 1941, (Hurst)
3 9 Dom. Rep. ; foot hills of Cordillera Central, south of Santiago,
1,000-3,000 feet, June 1938, (Darlington)

Genus Eriophora Simon 1864

Eriophora minax (O.P.-Cambridge)

Epeira minax O.P.-Cambridge, 1893, p. 112, pi. 15, fig. 1. "9 Mexico;
Acaguizotla in Guerrero, 3,500 feet."

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 369

3 9 Haiti; Diquini, November 1912, (Mann)
9 Haiti; Port-au-Prince, July 1941, (Audant)
9 Dom. Rep.; Puerto Plata, April-May 1941, (Hurst)
9 pullus Haiti; Port-au-Prince, (Crew), Banks Coll.

Genus Eustala Simon 1895

This genus was based by Simon on the species anastera of Walck-
enaer, which was described from an unpublished colored drawing by
Abbot. The species varies greatly, both in size and color pattern
and as it was Abbot's practice to name each spider delineated, the
same species was illustrated under many names. In using the Abbot
drawings, Walckenaer accepted most of these names. Hentz, whose
material was from the same general region, did not have access to the
Abbot figures and he added four names to the list. Later Keyserling
added one more.

Many years later, McCook saw the Abbot drawings in the British
Museum and he placed eleven of the Walckenaer species based on
Abbot names and several of his varieties in synonymy. The result
of all of this is, that we have a plethora of names for a single species.
Simon and F.O.P.-Cambridge have accepted McCooks identification
of this species. It has a widely known distribution from New England,
south to Florida and west to Utah. F.O.P.-Cambridge reports it
from Mexico, Guatemala and Costa Rica and quotes from the collector,
H. H. Smith, its abundance, especially in the cacao plantations.

Cambridge states in his definition of the genus, in the Biol. Central
America, vol. 2, p. 503, that the fourth femur of the male has no
ventral spines. This apparently is a very variable character, as in all
the specimens of E. anastera examined, from Massachusetts and
North Carolina, the fourth femur has two converging ventral rows
of spines; in four of the five species of the genus found in Hispaniola,
the males have ventral spines as well as retrolateral and prolateral
spines on the fourth femur. Eustala fusco-vittata (Keyserling) has
no ventral femoral spines and it also varies from others in the genus,
by the extension of the abdomen beyond the spinnerets.

In both Simon's and Cambridge's definition of the genus, the
a.m.e. are described as larger than the p.m.e., with the difference
greater in the male than in the female. But in Eustala bisctosa spec,
nov. from Hispaniola, the a.m.e. are smaller than the p.m.e., in
both male and female.

The palpi of all species of Eustala are remarkably similar, and the

370 bulletin: museum of comparative zoology

epigynums in all but one species has the same general structure. Eustala
perdita spec. nov. has a vulva that suggests Metazygia or Wagneriana.
The male and female both have the same pattern of dark spots on
the cephalothorax and the folium is alike in both sexes.

It is greatly regretted that the specimens of Eustala prompta (Hentz),
(which is considered a synonym of anastera) identified by Banks from
the vicinity of Port-au-Prince are not in .the collection as no other
specimen of that species has been seen from Hispaniola.

Eustala bisetosa spec. nov.
Figures 5, 6

Male. Length, 4.0 mm., ceph. 1.7 mm., abd. 2.4 mm.

Cephalothorax pale, no dark spots, cephalic portion shaded with
gray, eye area carried forward and narrowed, almost as wide as long,
thoracic groove long; eyes, both rows recurved, lateral eyes small and
contiguous, a.m.e. larger than a. I.e., separated by more than a diam-
eter, p.m.e. largest of the eight, separated by little over a radius,
a long, dark bristle between a.m.e. and p.m.e.; quadrangle of median
eyes wider in front and not as high as wide; clypeus about a radius
of a.m.e. ; mandibles small and vertical ; labium as wide as long, tip
pale and pointed; maxillae twice as long as labium, pale, inner margins
parallel; sternum grayish, little longer than wide, ending in a slender
point between fourth coxae; abdomen triangular, with a dark folium
on distal three-quarters, a median basal spear mark extending to
folium, margin of folium undulating and dark, scattered long hairs
on dorsum, sides with dark flecks, venter pale; legs much broken,
first pair very long, first coxae with hook, others not modified, femora
pale at base, other joints with alternate pale and dark bands, most
distinct on posterior pairs, spines, I pair, femur ventral, 2 median
spines, prolateral, 2 very long spines, median and distal, and several
short and weak dorsal and prolateral spines, patella, prolateral,
2, retrolateral, 1, tibia and metatarsus, spines slender; II pair, femur,
ventral, 2 distal, patella, lateral, 2-2, tibia not enlarged, but all spines
either heavy or modified, prolateral, 3, distal and median very heavy
and from a raised base, below a ventral row of 4 shorter spines and a
dorsal row of 3 spines, retrolateral, 2 very short spines; III pair, femur
ventral, 2 short stout median spines from a raised base, IV pair,
femur, ventral, 2 short heavy spines from a raised tubercle near base
and 3 longer retrolateral distal spines; palpus, patella with 1 long

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 371

bristle, palpal organ with a long white clavis, embolus rather short
and heavy, uncus broad with lower corner produced in a point half
hidden by a large white plate.

Female. Length, 4.6 mm., ceph. 1.5 mm., abd. 3.5 mm.

Cephalothorax brown, shaded with gray, no spots, margins dark,
many long white hairs, thoracic groove very short, carapace mod-
erately convex, but not gibbous as in typical species, anterior margin
not as much narrowed as in the male; eyes not carried forward as far
as in male, both rows recurved, a.m.e. separated by more than a
diameter, lateral eyes on a common tubercle, p.m.e. larger than a.m.e.
but not as large as in the male, separated by more than a diameter;
quadrangle wider in front and as high as wide; clypeus equals radius
of a.m.e.; mandibles vertical, superior margin with three teeth, middle
one largest, inferior margin with three subequal teeth ; labium, maxillae
and sternum as in male; abdomen triangular, widest at base, pointed
above spinnerets with a slight tubercle, dorsum dark, with a pale
margin, folium impossible to see, venter pale; legs, I pair much the
longest, femora much darker at distal end, tibiae and metatarsi
with wide dark bands, spines, no ventral spines on femora, I pair,
femur, prolateral, 3, retrolateral, 2 distal, dorsal, 0, patella, lateral,
2-2, tibia, prolateral, 4, retrolateral, 3, ventral, 0, dorsal, 1 at tip,
metatarsus, irregular, II pair, femur, dorsal, 2 distal, prolateral,
0, retrolateral, 0, tibia, prolateral, 3, retrolateral, 2, ventral, 0, dorsal,
1 at tip, posterior femora with no trace of the retrolateral spines found
on male; epigynum projecting forward, basal half free, tip pale and
very short, usual wrinkled area very small.

Holotype cT Dom. Rep.; foot hills of Cordillera Central, south of
Santiago, 1,000-3,000 feet, June 1938, (Darlington)

Allotype 9 Haiti; Kenskoff, 4,500-5,500 feet, 2 September 1934,
(Darlington)

Paratype & Haiti; Kenskoff, 4,500-5,500 feet, 2 September 1934,
(Darlington)

Paratypes 5 9 s Haiti; Kenskoff, 3,500, feet, 3 May 1935, (Roys)

Eustala bisetosa differs in both male and female from the typical
species of the genus, as the p.m.e. are larger than the a.m.e. and by
a lower cephalothorax, which is not gibbous each side of the groove.
The male has ventral spines on the posterior femora, a character
found in a few species of the genus. However, in general features,
both the palpus and the epigynum agree with the typical forms.

372 bulletin: museum of comparative zoology

EUSTALA DELASMATA spec. nOV.

Figure 11

Male. Length, 4.2 mm., ceph. 2.0 mm., abd. 2.5 mm.

Cephalothoraz pale brown, middle area shaded with gray, four
darker spots can be faintly seen near posterior end and a pair anterior
to the thoracic groove, thoracic groove deep and long; eyes, anterior
row recurved, a.m.e. largest of the eight, separated by more than a
diameter, lateral eyes on a tubercle, subequal, separated by a diameter,
p.m.e. separated by a diameter; quadrangle wider in front, and as
high as wide; clypeus less than a radius of a.m.e.; mandibles small
and vertical; sternum gray about margins, with a wide median pale
stripe; abdomen oval, a dark folium with wavy margins, does not
reach base and connects with a short median dark stripe that extends
to base, venter dark with a median pale spot; legs, I pair longest,
I coxa with hook, anterior femora dark, posterior femora banded,
spines, I pair, femur, prolateral, 4, basal spine quite short, dorsal,
3, retrolateral, 2 distal, patella, lateral, 2-2, tibia and metatarsus
with many unpaired spines; II pair, femur, prolateral, 2, dorsal,
3, retrolateral, 2, ventral, 2 distal, 3 median, patella, lateral, 2-2,
tibia, not incrassate, prolateral, 4, dorsal, 2 very long, retrolateral,
3 very small, ventral, distal, 2, metatarsus, with median and basal
whorls of spines; III pair, femur, prolateral, 4, dorsal, 3, retrolateral,
3, ventral, 2; IV pair, femur, prolateral, 3 near base, dorsal, 3, retro-
lateral, 5, all femoral spines on IV pair from a raised base; palpus,
patella with one long bristle, palpal organ similar to E. anastera with
the conductor very much shorter and smaller and the embolus longer,
with clavis smaller and more slender.

Holotype c? Dom. Rep. ; San Jose de las Matas, 1 ,500+ feet, June
1939, (Darlington)

Paratypes 2 d71 Dom. Rep.; San Jose de las Matas, 1,500+ feet,
June 1938, (Darlington)

Paratype d71 Dom. Rep. ; Sanchez, July 1938, (Darlington)

Eustala fusco-vittata (Keyserling)

Epcira fusco-vittala Keyserling, 1864, p. 129, pi .6, figs. 7, 8. " 9 N. Granada,
St. Fc de Bogota."

9 Haiti; hills near Port-au-Prince, 2,000 feet, October 1934,
(Darlington)

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 373

9 Haiti; foot hills north-east of la Hotte, 3,000-4,000 feet,
3 October 1934, (Darlington)
c? 9 Haiti; Diquini, November 1912, (Mann)

9 Dom. Rep.; Puerto Plata, August 1941, (Hurst)

EUSTALA PERDITA spec. nOV.

Figures 4, 9

Male. Length, 3.0 mm., ceph. 2.0 mm., abd. 2.0 mm.

Cephalothorax greenish-yellow, darker about eyes, with a transverse
row of two pairs of small dark spots at groove and a widely separated
pair of larger dots about the middle, anterior margin much narrowed,
thoracic groove faint; eyes cover anterior margin, anterior row strongly
recurved, a.m.e. largest of the eight, convex, separated by a diameter
and carried forward, a. I.e. less than a radius of a.m.e. and separated
from them by a diameter of a. I.e., posterior row straight, p.m.e. little
more than a radius of a.m.e., separated by a radius, lateral eyes
touching and subequal; quadrangle longer than wide and much
narrowed behind; elypeus below a.m.e. about a radius of a.m.e.;
mandibles vertical, weak, boss, small and distinct, fang groove short,
horizontal, superior margin with three small teeth, inferior margin
with two teeth; labium dark, wider than long, tip rebordered; maxillae,
more than twice as long as labium ; sternum triangular, as wide as
long, IV coxa separated by half a diameter; abdomen oval, widest at
base which extends over cephalothorax, generally dark with a pair of
transverse pale spots at base, and an undulate pale stripe each side
on posterior half that marks an indistinct folium, venter pale, shaded
with dark gray; legs, 1-2-4-3, I right missing, basal half of femora
pale, tibiae and metatarsi with dark bands about middle, wider on
anterior pairs, ventral side of femora spotted, I coxa with a small
hook, II pair with no specialized spines, I pair, femur, dorsal, 2 small
on basal half, prolateral, 3 about middle, ventral, 3? median, most
of spines broken off and those remaining very short, patella, 0, tibia,
dorsal, 0, prolateral, 3, retrolateral, broken, metatarsus, 0; II pair,
not modified, femur, dorsal, 2 on basal half, ventral, median row of 3,
prolateral, 0, retrolateral, 1, patella, 0, tibia, dorsal, 3, prolateral, 3,
retrolateral, 3; III pair, femur, ventral, 2 median basal, IV femur,
2 median about middle; palpus, shorter than cephalothorax, patella
and tibia of equal length, tibia with a ventral wing-like projection
the entire length of the joint, paracymbium small, embolus slender

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and almost straight, half hidden beneath a semi-transparent cap at
tip, unca dark with a broad obtuse tip, clavis long with distal third
narrowed.

Female. Length, 5.0 mm., ceph. 2.1 mm., abd. 3.0 mm. long,
3.0 mm. wide.

Cephalothorax like male, but darker about eyes, not much swollen
each side of groove but with the same dark spots; eyes, a.m.e. not
as large as in male, and not carried forward, p.m.e. separated by
almost a diameter; quadrangle and clypeus same as in male; abdomen
oval, widest at base, dark brown with many minute spots, no pair
of transverse spots as in male and lateral pale margins of folium
very indistinct, venter pale with a faint median dark spot ; legs 1-2-4-3,
slender, same as in male but dark rings wider and more distinct and
dark spots on femora more conspicuous, no ventral spines on femora,
spines, I pair, femur, dorsal, 2 small, prolateral, 3, tibia, prolateral,
3, retrolateral, 3, ventral, 0, patella, 0, tibia, dorsal, 0, prolateral, 2,
retrolateral, 2, ventral, 2-2; epigynum, a convex chitinized plate, wider
than long, darker about margins with a short, narrow finger at anterior
end directed forward.

Holotype d71 Haiti; Port-au-Prince, July 1941, (Audant)
Allotype 9 Haiti; Port-au-Prince, July 1941, (Audant)
Paratypes 9 Haiti; Port-au-Prince, July 1941, (Audant)
Paratype 9 Haiti; Kenskoff, 5,000-7,000 feet, September 1934,
(Darlington)

The generic position of this species is confusing. The palpus of
the male is very similar to the typical forms of Eustala but the spe-
cialized spines of the second leg are missing, the vulva is totally unlike
that of any in the genus but is similar to some of the Central American
species that have been placed in the genus Aranea. The male and
female are alike in having the same distinctive pattern on both
cephalothorax and abdomen.

Eustala vegeta (Keyserling)
Figure 7

Epeira vegeta Keyserling, 1865, p. 819, pi. 19, figs. 31-34. " d1 9 N. Granada,
Keyserling Coll."

9 s Haiti; Diquini, November 1912, (Mann)

cT Haiti; Port-au-Prince, July 1941, (Audant)
cf 9 Dom. Rep.; Puerto Plata, April-August 1941, (Hurst)

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 375

Genus Mangora O.P.-Cambridge 1889
Mangora striatipes spec. nov.
Figures 17, 18

Male. Length, 2.0 mm., ceph. 1.0 mm., abd. 1.1 mm.

Cephalothorax pale, anterior margin much narrowed, two-thirds as
wide as long, thoracic groove very long and deep, covered by a black
line, thoracic portion much higher than cephalic, highest near end
of groove; eyes cover anterior margin, anterior row strongly recurved,
a.m.e. largest of the eight, separated by a diameter and from a. I.e.
by a little more, posterior row straight, p.m.e. heavily ringed with
black so they appear larger than a.m.e., separated by about half a
radius, lateral eyes touching; quadrangle wider in front and higher
than wide; clypeus less than a radius of a.m.e., retreating; mandibles
vertical, small, weak, no boss; labium as wide as long, tip narrower
than base; maxillae twice as long as labium; sternum pale, margins
with a narrow black line, triangular, little longer than broad, tip
carried between IV coxae in a lobe; abdomen oval, narrow base extends
over cephalothorax, dull gray, covered with irregular pale blotches
and black dots, one small median black dot at base, 'followed by a
pair that are widely separated, on posterior half, two parallel rows
of five black spots with area between a dull gray, venter a dull yellow
with an irregular dark stripe each side; legs, 4-1-2-3, pale, anterior
pairs with a black ventral line on femora ending in a dot, spines long,
black, fragile, I coxae with a large hook, I femur with 2 long prolateral
spines from a raised base, II tibia with 3 heavy, short ventral spines
near base, with a longer retrolateral basal spine and a heavy retro-
lateral spine about the middle, III tibia with 2 prolateral diagonal
rows of plumose cilia near base, each row of 5 slender cilia from a
raised base, often the cilia are broken but the scar can be easily seen,
no ventral trichobothria on posterior femora; palpus shorter than
cephalothorax, patella and tibia of equal length, patella with one
long bristle at tip, embolus long and ending with an abrupt turn,
free portion half covered with a semi-transparent leaf-like lobe, clavis
at base with two short black points, much smaller than Mangora
mobilis O.P.-Cambridge, from Central America.

Female. Length, 3.0 mm., ceph. 1.7 mm., abd. 2.7 mm.

Cephalothorax and eyes as in male, but the black rings about p.m.e.
much wider, so that eyes appear larger than a.m.e.; mouth parts same
as in male; sternum dark with a pale spot below labium; abdomen
as in male but base much narrower and black spots on posterior half

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fused into two black lines, venter with a median dark spot and smaller
dark spots each side; legs, 4-1-2-3, pale, spines from a black base,
so that legs appear spotted, anterior femora with a ventral black line
with a basal dot as in male, no short ventral spines on II tibia, III tibia
with two parallel diagonal rows of cilia as in male, a long median
ventral trichobothria on IV femur near base and several scars that
may be from trichobothria, IV tibia, prolateral, 2 cilia very near base,
IV patella, a long, retrolateral spine, tibia with a very long median
basal spine; epigynum, a pale narrow median scape with a recurved
tip protruding from the plane of the venter, as in Mangora placida
(Hentz), each side a clear oval space.

Holotype d1 Haiti; Diquini, November 1912, (Mann)
Allotype 9 Dom. Rep.; Puerto Plata, May 1941, (Hurst)
Paratype 2 cf Haiti; Diquini, November 1912, (Mann)
Paratypes 6 9 Haiti; hills near Port-au-Prince, 2,000 feet, 2 October
1934, (Darlington)
Paratypes 2 9 Haiti; Kenskoff, 4,300 feet, 1 May 1935, (Roys)
Paratypes 9 s Dom. Rep.; Puerto Plata, July 1941, (Hurst)
Mangora striatipes probably belongs to Simon's second group of
the genus with M . fornicata (Keys.) as the type, although the a.m.e.
are larger than the p.m.e. It must be a common spider on the island
from the number of specimens collected. The males have no ventral
trichobothria on the posterior femora, but the females have one long
one and several scars. It is difficult to count or describe the spines,
for in most cases, they have been broken and only the scars remain.
The spines are black, long and slender. In the female the median
dorsal basal spine on the fourth tibia is very conspicuous.

Mangora picta O. P. -Cambridge, the genotype, has three short
ventral spines at the base of the II tibia. In neither volume of the
Biol. Centr. Amer., is the number or arrangement of the cilia on the
third tibia noted. This character, while apparently trivial, evidently
is constant in both male and female of a species. Chamberlin and
Ivie, in the descriptions of three new species from Panama, also fail
to mention the cilia.

Genus Marxia McCook 1895
Marxia stellata (Walckenaer)

Epeira stellata Walckenaer, 1805, p. 65, fig. 54. "Bosc, manuscrit sur les
araignees de la Caroline, pi. 1, fig. 1."

c? 9 pullus Haiti; Trou Caiman, 4 September 1934, (Bates)

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 377

Genus Metazygia F.O.P.-Cambridge 1904

Metazygia manni spec. nov.

Figures 12, 13, 23

Male. Length, 5.6 mm., ceph. 3.0 mm., abd. 3.4 mm.Jong, 2.5 mm.
wide.

Cephalothorax chestnut-brown, smooth and shining, rather flat,
cephalic portion higher, anterior margin not much narrowed, thoracic
groove deep, a long slender bristle posterior to lateral eyes and a long
bristle between p.m.e. and p. I.e., median row of small bristles from
groove to posterior eyes; eyes cover anterior margin, anterior row
recurved, a.m.e. largest of the eight, carried forward, separated by
less than a diameter, a. I.e. little over a radius of a.m.e., posterior row
slightly recurved, lateral eyes touching and subequal, p.m.e. slightly
larger than p. I.e., almost touching; quadrangle wider in front than
behind, and wider than high; clypeus retreating, less than a diameter
of a.m.e.; mandibles dark brown, vertical, long, boss small, median
margin on distal half deeply excavate, with a broad flat lobe ending
in an incurved hook above the fang groove, fang groove horizontal,
fang with a very thick base, distal half of fang short, compressed and
sinuous; labium wider than long, tip rounded and rebordered, sides
parallel; maxillae twice as long as labium; sternum three-fifths as
wide as long, emarginate, pointed between IV coxae, I coxae with a
small hook, and a basal lobe, IV coxae swollen; abdomen elliptical,
dorsum slightly convex, with numerous short hairs and long bristles,
pale brown, folium outlined by black emarginate stripes, each section
with a small dark spot, venter pale; legs, 1-2-4-3, pale brown, tibiae
darker, spines, I pair, femur, ventral, 0, dorsal, 4, prolateral, 2,
slender, retrolateral, 2, patella, slender bristle at tip, prolateral,
2, retrolateral, 1, tibia, dorsal, 1, prolateral, 2, retrolateral, 2, ventral,
2-2-2-2, metatarsus, prolateral, 2, retrolateral, 2, ventral, 2-2,
II pair, femur, dorsal, 4, prolateral, 2, retrolateral, 2, ventral, 0, patella,
slender spine at tip, tibia, dorsal, 1, prolateral, 2, large and heavy,
retrolateral, 2, ventral, 2-2-2-2-2, basal pair small, 2p and 3p above
base large and heavy, metatarsus, dorsal, 2, prolateral, 2, retrolateral,
2, III pair, femur, ventral, 2 small stout spines, widely separated,
patella, slender spine at tip, lateral, 1-1, IV pair, femur, ventral,
oblique row of 3 short stout spines at tip, followed by a short space
and a row of 6 spines ending at base, patella, slender bristle at tip,
lateral, 1-1; palpus large, shorter than cephalothorax, patella little

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longer than wide, 1 bristle at tip, tibia shorter than patella with a
large ventral lobe, equal in size to joint, with many long bristles,
paraeymbium small and dark, clavis the most prominent piece, sides
almost parallel, protrudes with tip recurved, embolus a slender
straight tube with tip resting against clavis.

Female. Length, 8.5 mm., ceph. 3.5 mm., abd. 6.1 mm.
Cephalothorax bright brown, cephalic portion much darker, anterior
margin little narrowed, thoracic groove a deep pit rather than a
groove, no bristles posterior to lateral eyes as in male; eyes as in male;
mandibles mahogany brown, geniculate, boss distinct, median margin
not excavate, superior margin of fang groove with two large teeth,
inferior margin with two small teeth; labium, maxillae and sternum
as in male; abdomen elliptical, dorsum slightly convex, yellowish-gray,
with folium outlined with quite wide emarginate black stripes, from
base to near spinnerets, each section with a dark spot, venter dull
brown; legs, 1-2-4-3, brown, anterior patellae spines as in male, no
ventral spines on femora as in male; epigynum, a heavily chitinized
plate, wider than long, with a small anterior turned scape, each side
depressed ovals that probably are the openings.

Holotype cf Haiti: Cap Haitien, January 1913, (Mann)
Allotype 9 Haiti; Cap Haitien, January 1913, (Mann)
Paratypes 4 9 Haiti; Cap Haitien, January 1913, (Mann)
The generic position of this species is very uncertain. The male
agrees with Parawixia in the general structure of the palpus and the
row of ventral spines on all femora. The female however, has the
elliptical abdomen without tubercles and a folium and epigynum
very similar to Metazygia wittfeldae (McCook), the genotype. Male
and female undoubtedly belong together as they were collected at
the same time and have the same abdominal markings. Typical forms
of both Metazygia and Paraivixia, have the second tibia of the male
modified. M. manni however, has no specialized spines on that joint.
The mandibles of the male are modified, which is quite characteristic
of Hispaniola species.

Genus Metepeira F.O.P.-Cambridge 1903

Metepeira inerma spec. nov.

Figure 20

Female. Length, 4.1 mm., ceph. 1.6 mm., abd. 2.6 mm. long, 2.1
mm. wide.

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 379

Cephalothorax yellowish-brown, paler about eyes and on posterior
portion, anterior margin about half the greatest width, head little
raised, no thoracic groove; eyes cover anterior margin, anterior row
recurved, eyes equidistant, a.m.e. separated by fully a diameter and
a half, posterior row very slightly recurved, p.m.e. smaller than
a.m.e., separated by a diameter, lateral eyes touching and subequal;
quadrangle narrower behind than in front and wider than high;
clypeus equals more than a radius of a.m.e.; mandibles vertical, pale
brown, boss very small, fang groove with three teeth on each margin;
labium wider than long, tip rebordered and pointed; maxillae twice
as long as labium; sternum brown, almost as wide as long, (2.5 : 3.0),
sides emarginate; abdomen rather faded, basal half with a pair of
transverse pale spots near middle, posterior half with a pair of con-
verging scalloped dark lines with area between dark, entire abdomen
in life probably covered with long dark hairs from pits, as many
remain on sides, venter with a wide dark stripe from pedicel to spinner-
ets with a central pale spot, each side of spinnerets a small pale spot;
legs, 1-2-4-3, pale, with tips of joints darker, I pair quite long, meta-
tarsus plus tarsus equals tibia plus patella, spines few and weak, all
patellae with a long distal and basal spines, III and IV tibiae with
long dorsal basal spines, I pair, femur, ventral, 0, dorsal, 0, prolateral,
1, retrolateral, 0, tibia, dorsal, 2, distal and basal, prolateral, 1, retro-
lateral, 1, prolateral and retrolateral rows of very long bristles, ventral,
0, metatarsus, ventral, 1, basal; epigynum small for the size of the
spider, median scape colorless, slender with a recurved tip, each side
are chitinized lobes.

Holotype 9 Haiti; Cap Haitien, March 1934, (Utawana Expedition)

Para types 3 9 s Haiti; Cap Haitien, March 1934, (Utawana
Expedition)

Metepeira inerma belongs to the group of which labyrinthea (Hentz)
is the most widely distributed and best known. It is separated from
that species by the smaller size, fewer spines on the legs and by the
epigynum, where the scape is almost colorless, slender and rather short
and the openings are widely separated. It does not agree fully with
the original definition of the genus by F.O.P.-Cambridge, as the
lateral eyes are almost touching, and the first tibia plus the patella
equals the first metatarsus plus the tarsus.

380 bulletin: museum of comparative zoology

Genus Neoscona Simon 1864

Neoscona oaxacensis (Keyserling)

Epeira oaxacensis Keyserling, 1864, p. 121, pi. 5, figs. 12-16. " 9 Oaxaca,

(Mexico)"
Epeira theisii Banks, 1903, nee Walckenaer, 1837

9 Haiti; hills near Port-au-Prince, 2,000 feet, 2 October 1934,

(Darlington)
cf 9 s Haiti; Kenskoff, 4,200 feet, 2 May 1935, (Roys)

Neoscona vulgaris (Hentz)

Epeira vulgaris Hentz, 1847, p. 469, pi. 30, fig. 6; reprint, p. 108, pi. 12, fig. 6.

"9 South Carolina"
Epeira volucripes Keyserling, 1884, p. 528, pi. 13, fig. 27. " 9 Central America,

Panama and Hayti, Universitat Cambridge."
nee Epeira nautica Simon, 1897, Petrunkevitch, 1930, Gertsch.

The synonymy of this species has been confused for many years.
Emerton thought he recognized this species as a northern spider and
added several northern localities as foot notes in the reprint of Hentz
Spiders of the United States, 1875, and in 1884, he placed E. vulgaris
as a synonym of the European spider sclopetaria of Clerck. The two
species belong to different sections of the genus but the error has been
continued for many years. Banks recognized the Hentz species and
in 1900, placed volucripes of Keyserling as a synonym in a short paper
on Alabama spiders but this synonymy has been overlooked by later
workers.

In 1897, Simon identified Epeira nautica L. Koch in a collection of
spiders from St. Vincent. Epeira nautica was described by Koch
in 1875 from Egypt and Syria. In the Museum collection are two
females from Arabia, received from the Paris Museum. Both are
pale and are much larger than any specimens of volucripes or vulgaris
in the collection and while size and color are uncertain characters
to use, the epigynum show decided differences. The scape of nautica
is much longer in proportion to the width and when turned back,
as so often is the case, the darkened areas are connected and the
depressed spots are not the same.

Males of vulgaris and volucripes (type material of the latter), have
been carefully compared, the palpi are identical in every part, but
the number of short stout spines on the second tibia varies. In a

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 381

specimen from Santiago de las Vegas, Cuba, there is a row of ten
short black spines; in the type of volucripes from Haiti, there are but
five or six spines; in a specimen from Shreveport, Louisiana, there
are five spines irregularly placed. This difference may be only indi-
vidual, but it is hardly enough to separate into two species.
cT 9 type Haiti; Epeira volucripes Keys.
3 9 Haiti; 1858, Dr. Weinland.

2 9 Haiti; Cap Haitien, March 1934, (Utowana Expedition)
9 Haiti; Cap Haitien, January 1913, (Mann)
9 Haiti; Dame-Marie, 1941, (Audant)

Genus Neosconella F.O.P.-Cambridge 1904

Neosconella parva spec. nov.

Figures 19, 24

Female. Length, 3.8 mm., ceph. 1.7 mm., abd. 2.G mm. long,
2.1 mm. wide.

Cephalothorax dull yellow, lateral margins and cervical grooves
darker, cephalic portion not elevated behind posterior row of eyes,
no thoracic groove; eyes cover anterior margin, anterior row of eyes
recurved, eyes small and subequal, a.m.e. separated by more than a
diameter with a pair of bristles between, posterior row straight, p.m.e.
largest of the eight, surrounded by wide dark rings that touch, lateral
eyes touching, p. I.e. slightly larger than a. I.e.; quadrangle much
narrower behind than in front and as high as wide in front; clypeus
less than a diameter of a.m.e.; mandibles yellow, vertical, boss small,
fang groove horizontal, superior margin of groove with three teeth,
inferior margin with three smaller teeth; labium brown, wider than
long, tip rebordered ; maxillae brown, more than twice as long as labium,
sides parallel ; sternum brown, three-quarters as wide as long, convex,
pointed in front of IV coxae; abdomen oval, no shoulder tubercles,
basal half high, a pale diamond on basal half with heavy brown marks
each side, on sides irregular dark spots, venter with a broad dark
stripe from groove to spinnerets with a pale spot in center; legs rela-
tively short, yellow, with no darker markings, varying little in length,
but anterior pairs longer, very few spines, femora with no ventral
spines, anterior tibiae with very few ventral spines, posterior tibiae
with a very slender dorsal basal spine; epigynum large for the size of
the spider and protruding from the venter, median scape short, broad,

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with median area depressed, each side of scape are large openings with
heavily chitinized margins.

Holotype 9 Dom. Rep.; foot hills of Cordillera Central, south of
Santiago, 2,000-5,000 feet, June 1938, (Darlington)

Paratypes 9 Dom. Rep.; Loma Rucilla Mountains, north of
Cordillera Central, 5,000-8,000 feet, June 1938, (Darlington)

Neosconella parva does not agree in all points with the description
of the genus given by F.O.P. -Cambridge in the Biol. Centr. Amer.,
1904, 2, p. 474. The p.m.e. are the largest of the eight and the posterior
row of eyes is straight, rather than recurved. Also, there are three
teeth on the upper margin of the fang groove instead of four. How-
ever, the spines on the legs are very few, the legs vary little in length,
and the epigynum is of the same type.

Genus Parawixia F.O.P.-Cambridge 1904

Parawixia cambridgei Bryant

Parawixia cambridgei Bryant, 1940, p. 342, figs. 104-106. "a71 9 Cuba;
Oriente, coast below Pico Turquino, June 1936"

9 Haiti; Diquini, November 1912, (Mann)

Parawixia darlingtoni spec. nov.
Figures 14, 16, 21

Male. Length, 4.1 mm., ceph. 2.1 mm., long, 1.9 mm. wide, abd.
2.5 mm. long, 2.0 mm. wide.

Cephalothorax low and flat, cephalic portion a bright yellow triangle
to groove, remainder a chestnut-brown, both cephalic and thoracic
portions with many fine white hairs, a crest of white hairs posterior to
lateral eyes, anterior margin half the greatest width, thoracic groove
long and distinct; eyes cover anterior margin, anterior row strongly
recurved, a.m.e. largest of the eight, on a small lobe that is carried
forward, eyes separated by less than a diameter, posterior row not as
strongly recurved, p.m.e. little more than a radius of a.m.e. and sepa-
rated by more than a diameter, lateral eyes touching and on a distinct
tubercle, subequal, smaller than p.m.e., a. I.e. and p.m.e. form a straight
line; quadrangle wider in front than behind and wider than high;
clypeus below a.m.e. less than a diameter of a.m.e.; mandibles dark
brown, vertical, basal half granular, fang groove horizontal, and fang

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 383

short; labium wider than long, tip rebordered; maxillae twice as long
as labium; sternum triangular, not quite as wide as long, pale, shaded
with gray about margins, pointed in front of IV coxae, I coxae with
a small hook and a large basal lobe, IV coxae with a strong, dark spur,
IV trochanter with a dark spine; abdomen pale, with no folium, almost
as wide as long, (7:8), a pair of small tubercles between shoulder
angles and not on lateral margins, basal third with many short bristles
directed upward, each from a dark base, venter gray with a pair of
conspicuous white spots anterior to spinnerets; legs, 1-2-4-3, rather
short, not differing greatly in length, pale with a wide dark band on
femora, distal and median bands on tibiae and metatarsi, spines,
I pair, femur, dorsal, 4 small spines in a median row, prolateral, 4,
second from distal end very long, retrolateral, 3, all small spines,
ventral, a retrolateral row of 6 short spines on distal half, a slender
prolateral spine opposite the retrolateral row, patella, dorsal, a slender
spine at tip, a small spine at base, prolateral, 2, retrolateral, 1, tibia,
dorsal, a small spine near base and a long spine near tip, prolateral,
3 long spines on distal half, retrolateral, 2, ventral, a prolateral row
of 5 long spines, retrolateral, 0, metatarsus, 0, II pair, slightly larger
than I pair, femur, dorsal, median row of 3 small spines, prolateral, 2,
retrolateral, 2, ventral, a strong median basal spine longer than the
diameter of the joint from a raised base, a retrolateral row on distal
half of 6 short, strong spines, patella, same as I pair, except that the
basal spine is missing, tibia, dorsal, median row of 3 slender spines,
prolateral, 5, retrolateral, 0, ventral, a prolateral row of 4 graduated
spines more than diameter of joint, a retrolateral row of 3 on basal
third, all from a raised base, basal spine almost twice the diameter of
the joint, metatarsus, sinuous, with 1 dorsal median spineaboutmiddle,
III pair, femur, ventral, 0, patella, dorsal, 1 slender spine at tip, lateral,
1-1, IV pair, femur, dorsal, median row of 3 slender spines, ventral,
a retrolateral row of 8 graduated spines, 2 distal longest, patella,
dorsal, 1 slender spine at tip, 1 near base, lateral, 1-1, tibia, dorsal,
1 median spine at base, ventral, 3 median; palpus, shorter than
cephalothorax, patella as long as wide, with 1 long bristle at tip,
paracymbium large and dark, tip widened and turned back, but not
divided, clavis very large, tip ending in two points.

Female. Length, 4.1 mm., ceph. 1.9 mm., abd. 3.0 mm. long,
3.0 mm. wide.

Cephalothorax chestnut-brown, longer than wide, cephalic portion
with many silvery hairs, slightly convex, anterior margin narrowed,
no thoracic groove; eyes cover anterior portion of head, median eyes

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carried forward, anterior row recurved, a.m.e. only slightly larger
than a. I.e., separated by little more than a diameter, lateral eyes
on a low tubercle and touching, posterior row recurved, p.m.e. little
larger than a.m.e., separated by fully a diameter; quadrangle only
slightly wider in front and as high as wide; clypeus below a.m.e.
equals a diameter of a.m.e. ; mandibles, small, brown, vertical, superior
margin of fang groove with three teeth, inferior margin with two
teeth; sternum brown, little longer than wide, pointed in front of
IV coxae, IV coxae touching; abdomen as wide as long, widest near
base, a pair of widely separated small tubercles between shoulder
angles, slightly larger than in male, a pair of dark basal spots extending
to tubercles and vague paired dark marks on posterior two-thirds,
that suggest a folium, on a cream color ground, many dark bristles,
venter dark with a pair of cream-colored spots anterior to spinnerets ;
legs, 1-2-4-3, rather short, dark brown, tibiae with median pale
band, very few spines, femora, 0, anterior patellae, dorsal, 1 long
spine at tip, 1 short spine at base, prolateral, 2, retrolateral, 1 ; epigynum
rather small, area wider than long, a slender wrinkled white scape at
anterior end, with tip much narrowed and abruptly turned out, best
seen in a lateral view, above the fold are two widely separated black
spots and slightly above them, smaller circles that are probably the
openings.

Holotype c? Dom. Rep.; Villa Altagracia, July 1938, (Darlington)
Allotype 9 Haiti; La Visite, 6,000-7,000 feet, 16-23 September
1934, (Darlington)

The male and female of this species were not found together, but
they have similar markings and are about the same size. They are
the only species seen from the island with tubercles between the
shoulder angles. The genus was erected for Central American species,
all much larger than darlrngtoni, with small tubercles at the tip of
the abdomen. The presence or absence of spines on the posterior
trochanters is a variable character in the genus. The arrangement
of spines on the second tibia of the male is very similar to that found
in Wixia.

Genus Verrucosa McCook 1888
Verrucosa arenata (Walckenaer)

Epeira arenata Walckenaer, 1837, 2, p. 133. "d" 9 La Georgie"

9 Haiti; Grande Riviere, January 1913, (Mann)

9 Dom. Rep.; Puerto Plata, 30 August 1938, (Darlington)

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 385

Genus Wagneriana F.O.P.-Cambridge 1904
Wagneriana tauricornis (O.P.-Cambridge)

Epeira tauricornis O.P.-Cambridge, 1889, p. 44, pi. 6, figs. 2, 3. 'V 9
Guatemala Panama"

9 Haiti; Port-au-Prince, (Crew), Banks Coll.
9 Haiti; Port-au-Prince, July 1941, (Audant)

c? Haiti; hills near Port-au-Prince, 2,000 feet, 2 October 1934,
(Darlington)

Subfamily METINAE

Key to Genera

1. Fourth femur with a prolateral row of curved cilia 2

Fourth femur with no prolateral row of cilia 4

2. Curved cilia on basal half of femur, abdomen with posterior truncate

tubercle Alcimosphenus

Curved cilia on entire length of femur, abdomen with no posterior
tubercle 3

3. First tibia and metatarsus with rows of triangular cuspules .... Plesiometa
First tibia and metatarsus with no rows of cuspules Leucauge

4. Lateral eyes widely separated on separate tubercles Azilia

Lateral eyes touching on a common tubercle 5

5. Mandibles with group of black nodules on inner margin Metargyra

No black nodules on mandibles 6

6. Third femur with one or two rows of long straight colorless trichobothria

(both cf and 9 ), paracymbium of male palpus much divided

Pseudometa
Third femur with no rows of colorless trichobothria 7

7. P.M.E. larger than A.M.E ' Meta (?)

A.M.E. larger than P.M.E Meta (?)

Genus Alcimosphenus Simon 1895
Alcimosphenus licinus Simon

Alcimosphenus licinus Simon, 1895, p. 930. " 9 Jamaica, S. Domingo."

Simon based this genus on females of this species from Jamaica and
San Domingo. It probably is found on all of the islands of the West
Indies, but as far as known, it has never been reported from either
Central or South America. The male is still unknown. Because of
the rows of curved cilia on the fourth femur, it should be placed with
Leucauge in the Metinae.

386 bulletin: museum of comparative zoology

9 Haiti; Trouin, 26 June 1934, (Loomis)

9 s Dom. Rep.; Puerto Plata, July-August 1941, (Hurst)

Genus Azilia Keyserling 1881
Azilia sp.

9 pullus Haiti; La Visite, 6,000-7,000 feet, 16-23 September 1934,
(Darlington)

This specimen is probably one moult from maturity and it may be
Azilia montana Bryant, known only from the holotype, a female
found on the south side of Pico Turquino, Cuba, at an elevation of
from 3,000-5,000 feet.

Genus Leucauge White 1841
Leucauge regnyi (Simon)

Argyroepeira regnyi Simon, 1897, p. 871. " d1 9 St. Vincent"
Meta bigibbosa Banks, 1903, p. 341 nee Keyserling, 1863

9 Haiti; Grande Riviere, -January 1913, (Mann)
9 Haiti; Kenskoff, 4,500-5,500 feet, 2 September 1934, (Dar-
lington)
9 Haiti; Trou Caiman, 4 September 1934, (Bates)
9 Haiti; hills near Port-au-Prince, 2,000 feet, October 1934,
(Darlington)
& 9 Haiti; Kenskoff, 4,300 feet, May 1935, (Roys)
<? 9 Dom. Rep.; Puerto Plata, April-May 1941, (Hurst)

Genus Meta C. Koch 1836

Meta (?) blanda spec. nov.

Figure 22

Female. Length, 2.6 mm., ceph. 1.0 mm., abd. 1.5 mm.

Cephalothorax pale, with a wide dark line above coxae, moderately
convex, anterior margin narrowed, no thoracic groove, a long bristle
posterior to lateral eyes; eyes cover anterior margin, anterior row
recurved, eyes equidistant, a.m.e. slightly larger than a. I.e., posterior
row only slightly recurved, lateral eyes touching on a common tubercle,
p.m.e. heavily ringed with black, separated by about a radius and
from p.l.e. by more than two diameters, p.m.e. slightly larger than

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 387

a.m.e.; quadrangle narrower behind and higher than wide; clypeus
less than a diameter of a.m.e.; mandibles pale, vertical, no boss,
fang groove oblique, superior margin with three small teeth; labium
wider than long, tip rebordered; maxillae more than twice as long
as labium, margins parallel; sternum brown, slightly convex, trian-
gular, almost as wide as long, ending in a blunt point in front of
IV coxae; abdomen oval, white with an indistinct folium on basal
third, and a heavier dark folium on distal third, venter dark with an
irregular white spot in middle and curved white lines each side, that
do not reach spinnerets j legs, 1-2-4-3, pale with a few hairs and
not many true spines, all patellae with a long apical spine, spines
I pair, femur, 1 prolateral about middle, patella, 1 dorsal near base,
1 apical, tibia, dorsal, 2, prolateral, 2, retrolateral, 1, ventral, 0,
metatarsus, no true spines but a prolateral row of stiff bristles;
epigynum, area wider than long, a very broad short median scape that
protrudes outward, openings on extreme lateral margins, large,
surrounded by a wide heavily chitinized ring, dark tubes below the
sldn lead to small black dots below tip of scape.

Holotype 9 Haiti; Kenskoff, 4,300 feet, 1 May 1935, (Roys)

Paratypes 9s Dom. Rep.; Loma Rucilla Mountains, Cordillera
Central, 5,000-8,000 feet, June 1938, (Darlington)

Meta (?) blanda is congeneric with Meta adjuntaensis Petrunkevitch,
from Puerto Rico, also described from a female. Dr. Petrunkevitch
describes another species of Meta from Puerto Rico in the same paper
and states that they do not conform strictly to the type of the genus.

The genus Meta is based on menardi, a common species of Europe
and North America. It is large, with an oval abdomen, rounded and
very high at the base, quite unlike the species reported here. In
1899, O.P.-Cambridge erected the genus Metabus for the female
gravidus found in Guatemala. The diagnosis differs little from Meta
and F.O.P.-Cambridge in 1903, (Biol. Centr. Amer., 2 : 446) places
Metabus gravidus in the genus Meta but states that it is possible that
with material of both sexes, the genus Metabus could be separated
from Meta. It is not impossible, that blanda and adjuntaensis Petr.
belong to this genus, for they certainly are not congeneric with Meta.

Meta (?) hotteiensis spec. nov.

Figure 25

Female. Length, 4.0 mm., ceph. 1.5 mm., abd. 3.0 mm. long,
2.2 mm. wide.

388 bulletin: museum of comparative zoology

Cephalothorax pale yellow, slightly darker in eye area, with a few
long bristles on cephalic portion, anterior margin less than half the
greatest width, cephalic portion not high, and not separated from
thoracic, thoracic groove a small round depression ; eyes cover entire
anterior margin, anterior row strongly recurved, a.m.e. largest of the
eight, separated by a diameter and from a. I.e. by less, a. I.e. little more
than a radius of a.m.e., posterior row almost straight, lateral eyes
touching and subequal, p.m.e. more than a radius of a.m.e. and
separated by a radius ; quadrangle wider in front and higher than wide ;
a long bristle between a.m.e. ; clypeus below a.m.e. about a radius
of a.m.e. ; mandibles pale, geniculate, vertical, no boss, superior margin
of fang groove with three teeth, inferior margin with three smaller
teeth; labium, dark, wider than long, with a rebordered tip; maxillae,
margins dark, twice as long as labium; sternum pale, with a few hairs
about margins, sides almost parallel, pointed in front of IV coxae;
abdomen oval, grayish-brown, no pattern but scattered silvery dots,
venter with a dark rectangular spot from fold to spinnerets, pale each
side; legs, 1-2-4-3, anterior femora dark, posterior femora pale with
dark ring at tip, tibiae with broad middle and distal dark rings, most
distinct on ventral side, spines, femora, no ventral spines, spines few
and weak on tibiae, no spines on metatarsi; epigynum, a broad, short
median scape, with lateral margins wrinkled and median area de-
pressed, tip abruptly narrowed and turned out, best seen in a lateral
view, a pair of very long divergent bristles at base of scape, on each
side are broad ear-shaped openings with heavily chitinized margins;
a pair of small black dots at tip of scape.

Holotype 9 Haiti; foot hills northeast of Massif de la Hotte,
3,000-4,000 feet, October 1934, (Darlington)

This species has an epigynum that is similar to Aranea detrimentosa
(Cambr.) from Guatemala. But in this species, the tip of the scape is
rounded, the openings on each side are smaller and the pair of long
bristles at the base of the scape are lacking.

Genus Metargyra F.O.P. Cambridge 1903

Metargyra maculata spec. nov.

Figure 15

Female. Length, 6.0 mm., ceph. 2.2 mm., abd. 4.0 mm., long
3.6 mm. wide.

Cephalothorax pale, cephalic portion high, covered with short hairs

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 389

and a few long bristles in the middle area anterior to the depression,
no thoracic groove; eyes do not cover entire anterior margin, anterior
row recurved, a.m.e. largest of the eight, separated by a radius, and
from a. I.e. by a diameter, posterior row straight, lateral eyes touching
and subequal, p.m.e. separated by less than a diameter and from
p. I.e. by more than a diameter; quadrangle slightly wider in front and
higher than wide; elypeus below a.m.e. less than a radius of a.m.e.;
mandibles pale, vertical, geniculate, no boss, superior margin of fang
groove with three teeth, and a row of six stiff bristles above groove,
each bristle from a distinct nodule, and on inner margin of mandibles,
above the groove a group of black nodules, inferior margin of groove
with three teeth; labium wider than long, heavily rebordered; maxillae
twice as long as labium, sides parallel ; sternum pale, with many long
bristles, convex, three-quarters as wide as long, ending in a truncate
lobe in front of IV coxae; abdomen oval, almost as wide as long, dull
gray, with small pale spots and irregular darker spots, venter gray,
with a median square of pale dots with a darker triangle in the middle;
legs, 1-2-4-3, anterior pairs very long, I femur as long as cephalo-
thorax, legs pale, femora of first and second pairs with dark ventral
dots, no specialized cilia on III and IV tibiae, spines, femora with no
ventral spines, I pair, femur, prolateral, 3, retrolateral, 1 small spine
near tip, dorsal, 1 small spine at base, patella, dorsal, 1 slender spine
at tip, 1 smaller spine at base, lateral, 0, tibia, dorsal, 2, prolateral,
2, retrolateral, 1, ventral, 0, metatarsus, 0, but many hairs and
bristles, II pair, femur, dorsal, 1 small spine at base, lateral, 1-1,
patella, same as I pair, tibia, dorsal, 1 at tip, 1 at base, prolateral,
1, ventral, 0, metatarsus, 0, III pair, femur, ventral, 2 parallel rows
of long bristles, posterior pairs with very few spines; epigynum, area
wider than long, a broad median septum with ear-shaped openings
each side.

Holotype 9 Haiti; Ennery, 7 September 1934, (Darlington)
Paratype 9 Dom. Rep!; Cordillera Central, 2,000-4,000 feet,
Constanza to Jarabacoa, August 1938, (Darlington)

The generic position of this species is doubtful. In the key for
genera, given by F.O.P.-Cambridge, in the Biol. Centr. Amer., 1902,
2 : 439, for the genera of Metinae, the first femur is three times as
long as the carapace for the genus Metargyra. This character may be
confined to males, but not females. The row of bristles from nodules
above the margin of the fang groove is very unusual.

390 bulletin: museum of comparative zoology

Genus Plesiometa F.O.P.-Cambridge 1903

Plesiometa argyra (Walckenaer)

Tetragnatha argyra Walckenaer, 1837, 2, p. 219, pi. 19, fig.l . " d1 9

Antilles, La Guadeloupe"

Haiti; Port-au-Prince, (Crew), Banks Coll.
9 Haiti; Diquini, November 1912, (Mann)
9 Haiti; Grande Riviere, January 1913, (Mann)
cf Haiti; Trou Caiman, 4 September 1934, (Bates)
cf Dom. Rep.; Sanchez, July 1938, (Darlington)
cf 9 Dom. Rep.; Puerto Plata, April-August 1941, (Hurst)
9 Haiti; Dame-Marie, 1941, (Audant)

Genus Pseudometa F.O.P.-Cambridge 1903

The genus Pseudometa was based on the species /lava of O.P.-
Cambridge from Tabasco, Mexico, known only from the male. Unfor-
tunately, no specimen of this species is in the museum collection.
The definition of the genus is short, and emphasis is laid on how it
differs from related genera. It is separated from Leucauge and
Mecynomcta by the lack of curved cilia on the fourth femur and in
the male by the fine silky hairs on the legs. The male palpus, however,
is distinct, as the embolus and conductor are in a flat coil at the apex
of the bulb, and the paracymbium is broken into several irregular
apophyses. F.O.P.-Cambridge places in the genus six species, three
known only from males and three from females. The figures of the
epigynums of the three females are of two distinct types and there is a
possibility that all are not adult.

The museum collection has males and females of alboguttata (O.P.-
Cambr.) from Barro Colorado, C. Z., a male of bella Banks from
Costa Rica, and the following from the West Indies, distincta Bryant,
cf , 9 , Cuba, linguiformis (Frang.) 9 , Cuba, hamata Bryant, cf , 9 ,
Puerto Rico, and four species from Hispaniola, cornuta Bryant,
cf, 9, conspersa Bryant;, 9, linguiformis (Frang.), 9, and obscura
Bryant, 9 .

A careful examination of the females of all these species, show on the
third femur, a retrolateral row of long colorless bristles. In some
species, there are but four bristles near the base, again, the row ex-
tends the length of the joint and in linguiformis (Frang.), there are two
parallel ventral rows. In the males examined, the bristles are some-

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 391

times absent. In cornuta spec, nov., there are retrolateral rows on the
third and fourth femora, and in hamata Bryant, the row extends the
entire length of the femur, as in the female, but distincta Bryant, male
has no bristles. So probably these bristles have a generic value only
in the female and can not be used, as are the short curved cilia found
on the fourth femur of Leucauge.

PSEUDOMETA CONSPERSA Spec. nOV.

Figure 29

Female. Length, 5.0 mm., ceph. 2.1 mm., abd. 3.1 mm.

Cephalothorax pale, darker from eyes to groove, cephalic portion
high, rising behind posterior eyes and falling gradually to groove, a
few bristles above groove; eyes about cover anterior margin, anterior
row recurved, a.m.e. separated by less than a diameter and from a. I.e.
by more than a diameter, posterior row only slightly recurved, same
length as anterior row, lateral eyes subequal on a common tubercle and
separated by a line, p.m.e. largest of the eight, separated by less than
a diameter and from p. I.e. by more than a diameter; qvadrangle about
square; clypeus less than a radius of a.m.e. ; mandibles brown, vertical,
swollen at base, boss faint, superior margin of fang groove with three
teeth, one farthest from base of fang largest, above teeth a row of four
short stiff bristles, inferior margin with four teeth, middle two small
and subequal ; labium brown, wider than long, tip rebordered ; maxillae
more than twice as long as labium;, sternum triangular, pale, with a
dark margin ; abdomen oval, convex, widest at base, pale, mottled with
brown, on posterior half the brown forms indistinct cross bars, venter
pale, with a large dark spot in middle; legs, 1-2-4-3, anterior pairs
much longer, covered with short hairs, pale, with brown rings most dis-
tinct on ventral side, no ventral femoral spines, spines, I pair, femur,
prolateral, 1, patella, 1 at tip, tibia, dorsal, 2, basal and median, pro-
lateral, 0, retrolateral, 2, near tip, ventral, 0, metatarsus, 0, II pair,
femur, 0, patella, 1 at tip, tibia, dorsal, 2, prolateral, 1, retrolateral, 1,
ventral, 0, metatarsus, 0, III pair, femur, a basal, retrolateral, row of
5 long, colorless bristles; cpigynvm, dark reddish-brown, heavily
chitinized, area wider than long, a wide transverse dark plate, below a
broad median septum, openings probably small depressions below
upper margins of plate, in anterior portion below skin, two oval oblique
spermathecae.

Holotype 9 Haiti; La Visite, 6,000-7,000 feet, 16-23 September
1934, (Darlington).

392 bulletin: museum of comparative zoology

PSEUDOMETA CORNUTA spec. IIOV.

Figures 26, 27, 28

Male. Length, 3.0 mm., ceph. 1.8 mm., abd. 2.0 mm.

Cephalothorax a dull yellow, cephalic portion marked off by heavy
gray lines, higher than thoracic, thoracic depression deep, but of in-
definite shape; eyes cover anterior margin, anterior row recurved,
eyes subequal, a.m.e. separated by little more than a diameter and
from a. I.e. by a little more, posterior row very slightly recurved, lateral
eyes touching, p.m.e. little larger than a.m.e., separated by a diameter
and from p. I.e. by a little more; quadrangle same width in front as be-
hind, higher than wide ; clypeus more than a radius of a.m.e. ; mandibles
vertical, long, basal portion swollen, not smooth, distinct longitudinal
ridge on outer margin, no boss, inner margin on distal half with three
long colorless bristles that interlace with bristles of opposite margin,
fang groove slightly oblique, superior margin with three small teeth,
inferior margin with two teeth; labium wider than long, rebordered;
maxillae more than twice as long as labium, tips widened ; sternum pale,
triangular, as wide as long, carried between IV coxae in a broad lobe ;
abdomen oval, pale yellow with scattered white blotches, very few
colorless hairs and long colorless bristles, venter pale; legs, 1-2-4-3,
little darker than abdomen, with colorless hairs, anterior pairs very
long, spines few and dark, no spines on metatarsi, III and IV femora,
retrolateral basal row of long colorless bristles; palpus as long as
cephalothorax, patella with one long bristle, tibia with an equally long
bristle, base of cymbium divided into a long, straight black process
and a smaller hook directed forward, a thin leaf-like apophysis about
the middle, best seen in a dorsal view; the most conspicuous part of
the palpus is the large smooth black process at the base with a large
posterior part ending in a rather slender tip and a much smaller an-
terior portion, all parts are heavily chitinized; embolus a small black
circle at the tip, coiled with the conductor.

Female. Length, 4.0 mm., ceph. 1.7 mm., abd. 2.5 mm.

Cephalothorax and eyes as in male; mandibles vertical, no long bristles
on inner margins as in male, fang groove slightly oblique, superior
margin with three teeth, inferior with two teeth; mouth parts and
sternum same as in male; abdomen a broad oval, yellowish gray,
covered with pale blotches, venter pale; legs, 1-2-4-3, pale, with faint
narrow rings at tips and middle of tibiae and metatarsi, very few spines
but many fine hairs, spines, I pair, femur, prolateral, 3, patella, 1 at
tip, tibia, dorsal, 1, retrolateral, 2, ventral, 0, metatarsus, 0, II pair,

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 393

femur, 0, patella, 1 at tip, tibia, dorsal, 1, prolateral, 2, ventral, 0,
metatarsus, 0, III pair, femur, a retrolateral row of colorless bristles
posterior pairs with very few spines ; epigynum, large, area wider than
long, a pale recurved tip, twice as long as wide, with very dark areas
each side separated by half a diameter.

Holotype cf Dom. Rep.; Cordillera Central, Loma Viega, south of
Constanza, 6,000 feet, August 1938, (Darlington)

Allotype 9 Dom. Rep.; Cordillera Central, Loma Viega, south of
Constanza, 6,000 feet, August 1938, (Darlington)

Paratypes cf 9 Dom. Rep.; Loma Viega, 6,000 feet, August 1938,
(Darlington)

Paratypes c? 9 Dom. Rep.; Loma Rucilla Mountains, north of
Cordillera Central, 5,000-8,000 feet, June 1938, (Darlington)

Paratypes c? 9 Dom. Rep. ; Cordillera Central, rain forest, north of
Valle Nuevo, 6,000+ feet, August 1938, (Darlington)

The male of Pseudometa cornuta has an enormous hook at the base
of the cymbium and a thin leaf-like process at the middle of the cym-
bium on the dorsal side, this is not found in the Central American
species. The embolus and conductor are very small. The three pairs
of bristles on the inner margin of the mandibles is unusual.

Pseudometa linguiformis (Frang.)

Figure 32

Meta linguiformis Franganillo, 1930, p. 20; 1936, p. 94, fig. 47. " 9 Cuba;

Sierra Maestra"
Pseudometa linguiformis Bryant, 1940, p. 353, fig. 128.

Female. Length, 4.5 mm., ceph. 2.0 mm., abd. 3.0 mm. long, 2.6
mm. wide.

Cephalothora.v pale yellow, cephalic portion high and outlined with
gray, anterior margin two-thirds the greatest width, no thoracic
groove; eyes do not cover anterior margin, anterior row recurved,
a.m.e. separated by fully a diameter, slightly larger than a.l.e., pos-
terior row very slightly recurved, p.m.e. largest of the eight, separated
by slightly less than a diameter and from p. I.e. by a diameter and a
half, lateral eyes contiguous and slightly raised; quadrangle little wider
in front than behind and higher than wide; clypeus less than a diameter
of a.m.e. and retreating; mandibles pale, geniculate, no boss, superior
margin with three teeth, inferior margin with three smaller teeth, inner
margin above fang groove with many dark granules; labium dark

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brown, wider than long, tip rebordered ; maxillae more than twice as
long as labium, tip dark and transverse; sternum dark brown, triangu-
lar, convex, as wide as long, sides emarginate, ending in a truncate lobe
in front of IV coxae; abdomen a broad oval, base high, dull brown,
mottled with paler dots, a pale red median stripe on posterior half,
venter with a mottled rectangle with a distinct dark spot in center,
two diagonal stripes on sides; legs pale, with no dark rings, anterior
pairs very long, spines black, spines, I pair, femur, dorsal, 0, prolateral,
3, retrolateral, 2 small near tip, ventral, 0, two rows of black dots, very
few hairs, patella, dorsal, 1 at tip, tibia, dorsal, 2, basal and median,
prolateral, 3, retrolateral, 3, ventral, 0, few short hairs and a few black
dots, metatarsus, 0, but rows of black hairs, II pair, femur, prolateral,
2, patella, dorsal, 1 at tip, tibia, dorsal, 2, prolateral, 1, retrolateral, 1,
ventral, 0, metatarsus, 0, posterior pairs, spineless except for spine at
tip of patellae and dorsal basal spine on tibiae, III femur, two ventral
rows of long colorless bristles; epigynum area wider than long, the
median scape short, triangular and turned forward, with heavily
chitinized margins each side, spermatheca widely separated anterior
to openings, anterior to median scape are two depressions.

9 Haiti; Kenskoff, 4,500-5,500 feet, 2 September 1934, (Darlington)

Pseudometa obscura spec, now
Figure 30

Female. Length, 4.3 mm., ceph. 2.0 mm., abd. 2.6 mm. long, 2.3
mm. wide.

Cephalothorax dull yellow, cephalic portion high, veined with dark
gray, with the heaviest markings near posterior end, two-thirds as wide
as long, anterior margin more than half the greatest width, no groove
but a depression; eyes do not cover anterior margin, anterior row re-
curved, a.m.e. separated by fully a diameter and from a. I.e. by less,
posterior row very slightly recurved, eyes equidistant, heavily ringed
with black, p.m.e. slightly larger than a.m.e., separated by about a
diameter, lateral eyes touching on a low tubercle, p. I.e. larger than
a. I.e.; quadrangle slightly wider behind than in front and higher than
wide; clypeus about as high as diameter of a.m.e.; mandibles vertical,
no boss, superior margin of groove with three teeth and a parallel row
of three bristles, inferior margin with three smaller teeth; labium pale,
wider than long; maxillae more than twice as long as labium; sternum
pale, longer than wide, (3.0 : 2.5) ; abdomen triangular, nearly as wide
as long, dorsum flattened, dull grayish-brown, with small yellow

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 395

flecks and two widely divergent rows of dark spots on posterior half,
the largest about the middle of the dorsum, venter dull brown ; legs,
1-2-4-3, anterior pairs much longer, same color as cephalothorax, with
faint dark rings at middle and tips of anterior tibiae, and tips of an-
terior metatarsi, very few spines, I pair, femur, prolateral, 3, from a
raised base, patella, 1 small spine at tip, tibia, prolateral, 2 small, basal
and distal, metatarsus, 0, III femur with a retrolateral row of long
colorless bristles, posterior tibiae with dorsal basal spine; epigynum,
chitinized area wider than long, openings large transverse, directly
above fold, anterior half a confused mass of dark oval sacs beneath
the surface.

Holotype 9 Dom. Rep.; Mt. Diago de Ocampo, north range,
3,000-4,000 feet, July 1938, (Darlington)

Subfamily NEPHILINAE

Genus Nephila Leach 1815
Nephila clavipes (Linn.)
Aranea clavipes Linnaeus, 1867, p. 1034, no. 27. " 9 in America"

9s Haiti; Diquini, November 1912, (Mann)
9 Haiti; Grande Cayemitte Island, 1 August 1927, (Eyerdam)
9 Haiti; Cap Haitien, March 1934, (Utawana Exped.)
9 Dom. Rep. ; Constanza, 3,000-4,000 feet, August 1938, (Darl-
ington)
9 Haiti; Port-au-Prince, July 1941, (Audant)

Subfamily TETRAGNATHINAE

According to some recent arachnologists, the old sub-family Tetrag-
nathinae has been split into Tetragnathinae and Glenognathinae based
upon the position of the opening of the posterior spiracle. Each is sup-
posed to have a row of trichobothria on the fourth femur. Other
workers following Menge, (1866), have raised the sub-family to family
rank.

While only five genera of this sub-family have been seen from His-
paniola, it is sufficiently evident that the position of the opening of the
spiracle has little systematic value for the group. Again, the presence
or absence of trichobothria as a character is difficult to use, more par-
ticularly on very small spiders, since trichobothria are often colorless,
and their base is little differentiated from the bases of surrounding
hairs and bristles.

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Simon's division of the sub-family into two groups seems equally
difficult to use. He separates the Tetragndtheae from the Pachygnatheae
by the relative width and length of the labium and straight or inclined
maxillae. With the Hispaniola material, these characters are not con-
stant and can not be used to divide the Tetragnathinae into two groups,
so the five genera are treated here as a unit. The male palpi of all are
similar, with a narrow paracymbium parallel to the cymbium, a large
spherical bulb only partly covered by the cymbium, and embolus and
conductor at tip of the bulb. The mandibles of the male may be diver-
gent, porrect, geniculate or vertical.

Key to Genera

Males

1. Opening of posterior spiracle a transverse slit midway between genital

fold and spinnerets Glenognatha

Opening of posterior spiracle close to spinnerets 2

2. Second femur with a retrolateral specialized spine from a raised base ....

Agriognatha
Second femur with no specialized spine 3

3. Mandibles vertical, only slightly divergent 4

Mandibles divergent or porrect 5

4. Superior margin of fang groove with three or four small teeth, inferior

margin with granules, opening of posterior spiracle not chitinized

Antillognatha

Only small teeth on fang groove, opening of posterior spiracle a transverse

chitinized slit, divided by a median septum Mimognalha

5. A large spur above base of fang, usually many teeth on both margins of

fang groove Tetragnatha

No spur above at base of fang, three sharp teeth on superior margin of
groove, four small teeth on inferior margin Hispanognatha

Females

1. Opening of posterior spiracle midway between genital fold and spinnerets

Glenognatha
Opening of spiracle near spinnerets 2

2. Opening of spiracle divided Mimognalha

Opening of spiracle entire 3

3. Small spiders with an oval abdomen Hispanognatha

Large spiders with a long, cylindrical abdomen 4

4. Fang groove with many graduated teeth on both margins .... Tetragnatha
Only three or four teeth on upper margin of fang groove Agriognatha

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 397

Genus Agriognatha O.P.-Cambridge 1896

The genus Agriognatha was based by O.P.-Cambridge in 1896,
on the species bella from Costa Rica. At the same time, he realized
that a species that he had described previously as Tetragnatha pachy-
gnathoides belonged to the same genus. Later, P.O. P. -Cambridge
in revising the genus, added one more species, Argyroepeira Irpida
O.P.-Cambridge. At this time, it was found that the genus Cyrtognatha
Keyserling, 1881, was the same but that the name was preoccupied by
Faldermann, 1835, in Coleoptera. The type of Cyrtognatha is nigri-
vittata Keys, from Peru and was described from a male. The abdomen
of this species extends beyond the spinnerets and the mandibles are
widely divergent with a large tooth or cusp above at the base of the
fang. Later, Simon described a male and female of this genus from
St. Vincent and Franganillo has described two more species from
Cuba giving little more than the bare generic characters.

No mention is made of the trichobothria on the posterior femora
in any of these species. This is not surprising, as they are colorless
and difficult to see except under high magnification. In A. simoni
Bryant from Cuba and A. rucilla spec, nov., the trichobothria are
short and colorless and form a median ventral row on the third and
fourth femora. In A. argyra spec. nov. and A. espanola spec. nov.
they are in a basal prolateral row on the third and fourth femora and
are much longer in the female than in the male. All three species from
Hispaniola have similar mandibles with the same number of teeth
on the fang groove. In the males, the specific difference is in the size
and the palpi.

In all species, the opening of the posterior spiracle is close to the
spinnerets.

Agriognatha argrya spec, nov
Figure 35

Male. Length, 4.7 mm., ceph. 2.0 mm., abd. 3.1 mm., spread of
mand. 2.2 mm.

Ccphalothorax pale brown, darker about margins and in median
area, low and flat, anterior margin narrowed, groove very near posterior
margin and punctiform, a pair of deep pits midway between lateral
eyes and groove and in line between eyes and groove; eyes cover
anterior margin, in three groups, lateral eyes touching, on a tubercle
higher in front than behind, anterior row strongly recurved, a.m.e.

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separated by less than a diameter, posterior row slightly recurved,
p.m.e. little larger than a.m.e., separated by little more than a radius;
quadrangle higher than wide and slightly wider in front; clypeus
almost wanting below a.m.e.; mandibles vertical, strongly divergent,
so that greatest width is more than length of cephalothorax, brown,
smooth, a large cusp over base of fang, suggestive of Pachygnatha,
fang groove horizontal, superior margin with three teeth, one very
near median margin, inferior margin with four teeth, the two middle
teeth quite close together, fang longer than groove, sinuous, with
tooth about middle, tip abruptly bent; labium brown, wider than long
with tip rebordered; maxillae brown, more than twice as long as
labium, sides parallel, tip abruptly widened; sternum pale brown,
triangular, as wide as long, ending in a slender point between IV
coxae, convex, excavate opposite III and IV coxae; abdomen cylin-
drical, pale, with many silvery spots, a faint gray median stripe with
irregular margins on posterior half, venter pale, opening of spiracle
close to spinnerets; legs, 1-2-4-3, anterior pairs very long, pale, with
tips slightly darker, spines, I pai,r, femur, ventral, row of 3 short
spines near base, prolateral, 6 long black spines, retrolateral, 4 long
spines and an irregular group of short spines that are not the same on
the right and left legs, tibia, with few short spines, metatarsus longer
than tibia, with a row of small spicules on basal two-thirds, II pair,
femur, ventral, 2 short spines near base, retrolateral, an irregular row
of long spines ending with a slender specialized spine near tip, this
spine from a raised base, not as long and heavy as in most species
of the genus, rubs against a groove on the patella, a very small median
spine at apex easily overlooked, metatarsus little longer than tibia,
with smaller spicules than on I pair, III and IV pairs much shorter
than anterior pairs, with scattered spines and a basal row of 4 very
long colorless trichobothria on dorsal prolateral side of femora, longer
on IV femur than on III; palpus pale, about as long as cephalothorax,
trochanter long, femur very long and slender, patella and tibia of equal
length, paracymbium very slender, with tip slightly enlarged, with
no hairs, scarcely reaches middle of bulb, bulb a flattened sphere
with embolus and conductor at top.

Female. Length, 6.5 mm., ceph. 3.0 mm., abd. 4.0 mm.

Cephalothorax same as in male but not as strongly colored, and the
pair of pits not as distinct; eyes same as in male but p.m.e. not quite as
close together; clypeus almost wanting below a.m.e.; mandibles same
color as cephalothorax, vertical, strongly geniculate, no boss, fang
groove oblique, superior margin with three strongly chitinized teeth,

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 399

inferior margin with four teeth, second from base of fang very small,
fang evenly curved; labium, maxillae and sternum same as in male;
abdomen cylindrical, twice as long as wide, widest at posterior end,
pale brown with many silvery spots and two pairs of brown spots above
spinnerets, venter pale brown with scattered silvery spots; legs,
1-2-4-3, anterior pairs very long and the dark bands more distinct
than on posterior pairs, spines scattered, no cusps or spines on anterior
metatarsi, no specialized spine on II femur, trichobothria of prolateral
row on posterior femora longer than in male; epigynum a curved
chitinized fold between book lungs, that shows no structure.

Holotype a71 Dom. Rep.; Cordillera Central, rain forest north of
Valle Nuevo, 6,000+ feet, August 1938, (Darlington)

Allotype 9 Dom. Rep.; Cordillera Central, rain forest north of
Valle Nuevo, 6,000+ feet, August 1938, (Darlington)

Paratype cf Dom. Rep.; Loma Rucilla Mountains, north of Cor-
dillera Central, 5,000-8,000 feet, June 1938 (Darlington)

Paratype d" 9 Dom. Rep. ; foot hills of Cordillera Central, south
of Santiago 2,000-5,000 feet June 1936, (Darlington)

Agriognatha espanola spec. nov.
Figure 34

Male. Length, 3.4 mm., ceph. 1.0 mm., abd. 2.5. mm.

Cephahthorax pale brown, median area darker, anterior margin
much narrowed, cephalic portion slightly higher than thoracic, groove
very near posterior margin and punctiform, a pair of grooves midway
between lateral eyes and thoracic groove ; eyes cover entire width of
head, lateral eyes touching, anterior row recurved, eyes subequal,
a.m.e. separated by more than a diameter and from a. I.e. by more than
two diameters, posterior row straight, p.m.e. slightly larger than
a.m.e. and surrounded by a much wider dark ring; quadrangle higher
than wide and the same width in front as behind ; clypeus equals about
a radius of a.m.e. ; mandibles dark brown, basal portion roughened and
distal half narrowed, strongly divergent, two-thirds as long as cephalo-
thorax, no boss, and a very small' tooth or cusp over base of fang, fang
groove horizontal, poorly defined, superior margin with two widely
separated sharp teeth and a few long bristles, inferior margin with four
widely separated teeth, all smaller than those on opposite margin, fang
longer than groove, sinuous, with tip abruptly bent; labium dark
brown, slightly wider than long, tip strongly rebordered; mJasdllae more

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than twice as long as labium, sides parallel and tips slightly widened ;
sternum light brown, triangular, about as wide as long, with tip pointed
between IV coxae, only slightly excavate between coxae; abdomen
cylindrical, silvery, with three pairs of divergent black spots on pos-
terior third, venter with two parallel silvery stripes and silvery spots,
opening of spiracle directly anterior to spinnerets; legs, 1-2-4-3, an-
terior pairs very long, pale brown with ends of anterior tibiae darker,
spines, all patellae with a long strong spine at tip, I pair, femur, ventral,
4 black basal spines, all longer than the diameter of the joint, a retro-
lateral row of 11 shorter spines from base to tip, tibia with a few lateral
and ventral spines, metatarsus longer than tibia, with a ventral row
of 13-14 short dark cusps or spicules, tarsus about two-thirds length
of metatarsus, II pair, femur with ventral row of 8 short spines on
basal third, retrolateral row of 11 short spines from base to tip, pro-
lateral row of longer spines, a slender retrolateral spine from a raised
base near tip which rubs against a ventral ridge on patella, tibia with
a few lateral and ventral spines, tibia longer than metatarsus, meta-
tarsus with ventral row of cusps smaller than on I pair, posterior pairs
much shorter, no ventral spines on femora, a short prolateral basal
row of long trichobothria on III femur; palpus pale, shorter than
cephalothorax, patella and tibia of equal length, each about diameter
of joint, paracymbium very slender, extending beyond middle of the
bulb and tip expanded, cymbium slender, sides almost parallel, with
tip curved in towards bulb, bulb almost spherical, with conductor and
embolus in an elliptical coil, ending with slender spine-like processes,
the apical longer and more curved than the preapical.

Female. Length, 5.0 mm., ceph. 2.0 mm., abd. 3.0 mm. long, 20.
mm. high.

Cephalothorax very low, not as much narrowed as in male, coloring
same as male, groove in a deep depression very near posterior margin ;
eyes same as in male; mandibles brown, vertical, geniculate, no boss,
fang groove oblique, superior margin with two teeth, inferior margin
with three teeth, fang evenly curved; labium and maxillae same as in
male ; sternum dark brown, convex, triangular, as wide as long; abdomen
pale, oval, with a large dorsal tubercle posterior to middle, a broad
dark median stripe from base to spinnerets, sides dull with small sil-
very spots and dark stripes, venter with a broad median dark stripe
from pedicel to spinnerets, with a pale stripe each side, two silvery
spots each side of spinnerets, opening of posterior spiracle obscure;
epigynum a simple curved fold; legs, 1-2-4-3, brown, darker at tips of
joints, pale ventral spots at tip of anterior femora and patellae quite

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 401

conspicuous, anterior pairs very long, no row of ventral spines on
femora, prolateral row of 3 strong spines near base on first femur, no
cusps on tibia or metatarsus, II and III metatarsi and tarsi with pro-
lateral row of short stiff bristles, much heavier on III pair, III and IV
femora with prolateral row of trichobothria on basal half of joint.
Holotype & Haiti; Kenskoff, 3,000-4 000 feet, 1 May 1935, (Roys)
Allotype 9 Haiti; Kenskoff, 3,000-4,000 feet, 1 May 1935, (Roys)
Paratypes c? 9 Haiti; Kenskoff, 3,000-4,000 feet, 1 May 1935,
(Roys)
Paratype d71 Haiti; Dame-Marie, 1941, (Audant)
The female of Agriognatha cspanola quite unlike others in the genus,
has a large tubercle on the abdomen.

Agriognatha rucilla spec, now
Figure 33

Male. Length, 5.0 mm., ceph. 2.1 mm., abd. 3.5 mm., spread of
mand. 3.5 mm.

Cephalothorax brown, darker about margin and in median area, low
and flat, anterior margin narrowed, thoracic groove very near posterior
margin and punctiform, faint lateral grooves from margin to near
thoracic groove; eyes cover entire margin, in three groups, lateral eyes
touching and subequal, anterior row recurved, a.m.e. separated by a
full diameter, posterior row straight, p.m.e. larger than a.m.e., sepa-
rated by about a radius; quadrangle higher than wide, and wider in
front; clypeiis below a.m.e. not as high as diameter of a.m.e. ; mandibles
pale brown, widely divergent, so that greatest spread is more than
length of cephalothorax, base slightly swollen, a large outward turning
cusp at base of fang, groove horizontal, superior margin with three
sharp teeth, one very near median margin, inferior margin with four
sharp teeth, two middle close together, fang longer than groove,
slender, a large tooth about middle on left fang and a swelling at same
place on right, tip abruptly bent; labium brown, wider than long, tip
rebordered; maxillae more than twice as long as labium, sides parallel,
tip only slightly widened; sternum brown, triangular, slightly longer
than wide, emarginate opposite III and IV coxae ; abdomen cylindrical,
dull gray, with median dark stripe from base to spinnerets, margins
much darker, each side thickly covered with silvery spots, venter
mottled with silvery spots and a pair of vague dark spots about middle,
inconspicuous opening of posterior spiracle; legs, 1-2-4-3, pale with

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vague darker rings on femora and tibiae, anterior pairs longer, spines,
I pair, femur, ventral, a median row of 6 short spines near base,
scattered prolateral, retrolateral and dorsal spines, tibia, scattered
long black spines, metatarsus longer than tibia, with a few ventral
spicules, II pair similar to I pair but fewer median ventral spines on
femur and a retrolateral clasping spine from a raised base near tip,
metatarsus longer than tibia, III metatarsus with prolateral and retro-
lateral rows of bristles, III and IV femora with median ventral row of
very short trichobothria, so small that they are easily overlooked;
palpus pale, about as long as cephalothorax, trochanter about one-
* third as long as femur, tibia little longer than patella, paracymbium
very slender, tip much widened, not reaching middle of bulb, cym-
bium with parallel sides, hairy, bulb, a flattened sphere, with embolus
and conductor at tip.

Female. Length, 5.5 mm., ceph. 2.1 mm., abd. 4.0 mm.

Ccphaloihorax, coloring same as male, anterior margin not as much
narrowed, low and flat, thoracic groove not as near posterior margin
as in male; eyes same as in male; mandibles brown, vertical, geniculate,
no boss, fang groove oblique, superior margin with three sharp teeth,
inferior margin with four large sharp teeth, fang evenly curved ; labium,
maxillae and sicrvum same as in male; abdomen cylindrical, more than
twice as long as broad, base slightly bifid, with a median dark herring-
bone stripe with darker margins, sides with silvery spots, venter with
a broad dark stripe, abruptly widened about middle, with a narrow
stripe of silvery spots each side, openings of spiracle obscure, spinnerets
at end of abdomen; legs, 1-2-4-3, much broken, brown, femora and
tibiae with wide dark rings, spines long and black, no ventral row of
short spines on anterior femora, and trichobothria very inconspicuous
in ventral row on III and IV femora; epigynum, a simple curved fold
showing no structure.

Holotype cf Dom. Rep.; Loma Rucilla Mountains, north of Cordil-
lera Central, 5,000-8,000 feet, June 1958, (Darlington)

Allotype 9 Dom. Rep.; Loma Rucilla Mountains, north of Cordil-
lera Central, 5,000-8,000 feet, June 1938, (Darlington)

Paratypes 2d1 Dom. Rep.; Loma Rucilla Mountains, 5,000-8,000
feet, June 1938, (Darlington)

The two male paratypes are much smaller than the holotype but
the teeth are the same on the fang groove and the tooth is on the fang.

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 403

Genus Antillognatha gen. nov.

Cephalothorax low; eyes, anterior row strongly recurved, posterior
row almost straight, lateral eyes on separate tubercles and separated
by a line, p.m.e. largest of the eight; quadrangle longer than wide;
mandibles vertical, only slightly divergent, upper margin of fang
groove with three or four very small teeth, lower margin with a row
of granules; labium wider than long, sides almost parallel, tip rebor-
dered; maxillae narrow, fully twice as long as labium; abdomen oval,
more than half as wide as long, opening of posterior spiracle very near
spinnerets and not chitinized. Male only known.

Genotype Antillognatha lucida spec. nov.

Antillognatha differs from Hispanognatha by the lateral eyes almost
touching, the small granules on lower margin of the fang groove, and
the much shorter abdomen. It differs from Dyschiriognatha and Glenog-
natha by the granules on the fang groove, and the opening of the
posterior spiracle. It differs from Pachygnatha by the width of the
labium, the very small teeth on the fang groove and the small size.

Antillognatha lucida spec. nov.
Figure 31

Male. Length, 1.9 mm., ceph. 1.0 mm., abd. 1.1 mm. long, 0.6 mm.
wide.

Cephalothorax pale brown, rather low, anterior margin not much
narrowed, thoracic depression quite near posterior margin and trans-
verse; ryes cover anterior margin, anterior row strongly recurved,
a.m.e. separated by a diameter and a little more from a. I.e., posterior
row same length as anterior, almost straight, eyes equidistant, lateral
eyes on separate tubercles that touch at base, p.m.e. separated by more
than a diameter, largest of the eight; quadrangle longer than wide, and
wider behind than in front; elypeus little higher than diameter of
a.m.e.; mandibles vertical, only slightly divergent, distal half granular,
with a small hook above base of fang, fang groove oblique, superior
margin with three or four minute teeth and a row of colorless bristles,
inferior margin with a row of minute granules, fang long, slender and
sinuous; labium dark brown, wider than long, tip rebordered and about
as wide as base; maxillae fully twice as long as labium, narrow, tips
not widened, slightly inclined; sternum dark brown, triangular, as wide
as long, convex, carried between II and III coxae, ending in a broad

404 bulletin: museum of comparative zoology

lobe between IV coxae, so that coxae are separated by almost a di-
ameter, posterior coxae globular; abdomen oval, two-thirds as wide as
long, shining, no hairs, paler brown than cephalothorax, with a pair
of widely separated lines of silvery spots from base to spinnerets,
venter brown, genital fold only slightly posterior to openings of book
lungs, opening of posterior spiracle directly anterior to spinnerets and
not chitinized; legs, 1-2-4-3, paler than cephalothorax, no spines and
very few hairs, I pair very long, tarsus short, IV femur with a dorsal
row of trichobothria ; palpus not as long as cephalothorax, bulb very
large, both patella and tibia as long as diameter of joint, paracymbium
very slender, not reaching tip of bulb, tip widened and ending in a
truncate lobe, bulb a much flattened sphere, cymbium long and
slender, extending beyond the bulb, conductor and embolus form a
narrow loop at top of bulb with long straight free ends which rest
against the tip of cymbium.

Antillognatha Incida is smaller than Mimognatha fdxi (McCook)
found in North and Central America and some of the islands of the
Caribbean. The opening of the posterior spiracle is entire, not chiti-
nized and is very near the spinnerets, the labium has almost parallel
sides and the embolus and conductor form a much larger loop at the
tip of the bulb.

Genus Glenognatha Simon 1887

The genus Glenognatha was based by Simon on a small spider from
Arizona and named for Mr. J. H. Emerton. Unfortunately, this species
is not in the museum collection. The genus differs from Paehygnatha
by the anterior legs that are much longer than the posterior, (in
Paehygnatha there is little difference in the length of legs), the longer
labium, and the position of the posterior spiracle which is midway
between the genital fold and the spinnerets and opens by a transverse
slit with heavily chitinized margins.

The small spider figured by McCook, (1893), as Theridion foxi was
recognized by Banks in 1929 as his Mysmena bulbifera that had been
placed in the genus Glenognatha by Barrows in 1924, who found the
spider in its web. Banks, (1929), erected the genus Mimognatha for
it, separating it from Glenognatha by the small mandibles, that are not
divergent, and with only small teeth on the fang groove. The opening
of the spiracle also is a chitinized transverse slit but it is divided by a
median septum and is very near the spinnerets. The third femur has
two parallel ventral rows of trichobothria.

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 405

It is not impossible that the species from St. Vincent described by
Simon as Dyschiriognatha montana is a Glenog?iatha, as the male is
described as with very long anterior legs and the genus Dyschiriognatha
is based on a species from Sumatra in which the legs vary little in
length.

Glenognatha mira spec. nov.

Female. Length, 3.0 mm., ceph. 1.2 mm., abd. 1.7 mm.

Cephalothorax dark brown, cephalic portion elevated, thoracic de-
pression one-third from posterior margin, a transverse pit; eyes cover
anterior margin, anterior row strongly recurved, a.m.e. largest of the
eight, separated by fully a diameter and from a. I.e. by a little less,
posterior row straight, lateral eyes touching on a common tubercle,
p.m.e. little smaller than a.m.e., separated by little more than a
diameter and from p. I.e. by more than a diameter more; quadrangle
wider in front, and as high as wide ; clypcus higher than quadrangle, con-
cave, with a distinct groove midway between a.m.e. and margin; man-
dibles brown, vertical, slightly roughened, upper margin of groove with
three equidistant teeth, lower margin with four small teeth; labium
dark brown, wider than long, and wider at base than at tip; maxillae
twice as long as labium, slightly inclined, sides parallel; sternum dark
brown, triangular, longer than Avide, convex, carried between II and III
coxae, emarginate opposite III and IV coxae, ending in a broad lobe
between IV coxae, posterior coxae globular ; abdomen globular, dorsum
infuscate, a few silvery dots, sides silvery, venter convex, with a broad
dark stripe from pedicel to spinnerets and a pair of widely separated
silvery spots about middle, opening of posterior spiracle a strongly
chitinized slit midway between epigynum and spinnerets; legs 1-2-4-3,
anterior pairs very long, pale brown, spineless, but rows of hairs, I tibia
longer than I metatarsus, III femur, two parallel ventral rows of
trichobothria from base which disappear before reaching tip;
epigynum a procurved line ending in silvery spots, midway between
pedicel and spinnerets.

Holotype 9 Haiti ; Miragoane, 2 November 1934, (Darlington)

Genus Hispanognatha gen. nov.

Cephalothorax oval, low; both rows of eyes recurved, lateral eyes on
separate tubercles ; quadrangle narrowed in front and as high as wide
behind; mandibles divergent, two-thirds as long as cephalothorax,

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upper margin of fang groove with three sharp separated teeth, lower
margin with four smaller teeth ; labium wider than long, sides parallel
with tip strongly rebordered; maxillae more than twice as long as
labium, narrow, slightly inclined; abdomen twice as long as wide,
opening of posterior spiracle obscure; legs spineless, anterior pairs
long, a short dorsal basal row of long trichobothria on all femora;
palpus very similar to Pachygnatha. Male only known.

Genotype, Hispanognafha guttata spec. nov.

The genus Hispanognatha differs from Dysehiriagnatha and Glenog-
natha by the opening of the posterior spiracle near the spinnerets, by
the elongate abdomen, and the labium as wide at tip as at the base.
It differs from Pachygnatha by the divergent mandibles, the labium,
the long maxillae, and the elongate abdomen.

HlSPANOGNATHA GUTTATA spec. nOV.

Figure 36

Male. Length, 3.5 mm., ceph. 1.3 mm., abd. 2.4 mm. long, 1.1 mm.
wide.

Cephalothorax dull brown, with darker stripes converging from below
lateral eyes to thoracic groove, a dark marginal line, low and flat,
anterior margin only slightly narrowed, thoracic depression quite near
the posterior margin; eyes cover entire anterior margin, anterior row
recurved, a.m.e. separated by fully a diameter and from a.l.e. by more
than two diameters, a.l.e. slightly smaller than a.m.e., posterior row
slightly longer than anterior, and not as much recurved, eyes equi-
distant, p.m.e. largest of the eight, separated by more than a diameter
and from p. I.e. by fully a diameter and a half, lateral eyes on separate
tubercles that are connected at base, separated by more than a di-
ameter, a.l.e. smaller than p. I.e.; quadrangle narrower in front and
higher than wide behind; clypeus below a.m.e. higher than diameter of
a.m.e. ; mandibles two-thirds as long as cephalothorax, slightly diver-
gent, no boss, with a strong curved tooth that projects forward near
exterior margin about one-third above base of fang, several long bristles
on distal third, fang groove oblique, superior margin with three long
slender teeth, inferior margin with four much smaller teeth, fang as
long as groove, curved, with distinct swelling near basal third; labium
dark brown, wider than long, sides parallel, tip rebordered; maxillae
nearly three times as long as labium, slightly inclined, narrow; sternum
four-fifths as wide as long, triangular, convex, carried between coxae

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 407

and continued in a broad lobe between IV coxae; abdomen light brown,
covered with pale silvery spots, two irregular converging dark lines
from before the middle to spinnerets, about twice as long as wide,
widest at basal third, venter with a wide dark stripe from pedicel to
spinnerets, with silvery spots each side, opening of posterior spiracle
indistinct; legs, 1-2-4-3, anterior pairs very long, pale brown with
femora darker, no spines or cusps but rows of very fine hairs, a few
long trichobothria in a dorsal basal row on all femora; palpus longer
than cephalothorax, femur long and slender, tibia slightly longer than
patella, paracymbium very narrow, sides almost parallel, tip bent,
bulb a flattened sphere with embolus and conductor at tip in a small
circle, free ends of both in a simple curve beyond bulb.

Holotype cf Dom. Rep.; Cordillera Central, Valle Nuevo, south
west of Constanza, 7,000 feet, August 1938, (Darlington)

Genus Tetragnatha Latreille 1804

Tetragnatha elongata Walckenaer

Tetragnatha elongata Walckenaer, 1805, p. 69. "Bosc manuscrit sur les
araignees de la Caroline, pi. 5, fig. 5."

9 Dom. Rep.; Puerto Plata, July 1941, (Hurst)

Tetragnatha festina spec, now
Figures 38, 39, 40, 41

Male. Length, 7.4 mm., ceph. 2.7 mm., abd. 5.0 mm., mand. 2.6
mm.

Cephalothorax pale brown, no markings; eyes about cover anterior
margin, both rows recurved, space between lateral eyes less than be-
tween median, lateral eyes on separate tubercles; mandibles porrect,
almost as long as cephalothorax, dorsal spur pointed with a large
ventral tooth, the first tooth at base of fang not large and directed
forward, second tooth larger than first and directed upward, followed
by two teeth, then a space and eight graduated teeth, the last only a
granule, only six are seen from dorsal side, inferior margin with a
shorter row of eleven graduated teeth, fang with a small ventral tooth
near middle and thickened near base; labium longer than wide, tip re-
bordered; sternum, triangular, carried between I and II and II and III

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coxae; abdomen cylindrical, nearly four times as long as wide, spin-
nerets at tip, pale brown with paler flecks, no distinct markings; legs,
1-2-4-3, first pair very long, few spines, a few tricobothria at base of
IV femur; palpus as long as cephalothorax, tibia one and a half times
as long as patella, paracymbium with the usual chitinized lobe about
middle, with tip bifid.

Female. Length, 11.5 mm., ceph. 2.6 mm., abd. 9.0 mm., mand.
2.1 mm.

Cephalothorax and eyes as in male; mandibles porrect, but not as
long as in male, thicker at base and more divergent, no dorsal spur, two
cone-shaped teeth on superior margin near base of fang, followed by
a space and then eight graduated teeth as in male, inferior margin
with eleven graduated teeth from fang to base, fang long and sinuous
but with no ventral tooth as in male, a thickened area near base is
produced in a dorsal tooth; labium and sternum as in male, more
trichobothria on IV femur than in male; epigynum a small curved slit
posterior to openings of the book lungs.

Holotype cf Dom. Rep.; foot hills of Cordillera Central, south of
Santiago, 1,000-3,000 feet, June 1938, (Darlington)

Allotype 9 Dom. Rep.; foot hills of Cordillera Central, south of
Santiago, 1,000-3,000 feet, June 1938, (Darlington)

Paratype c? Haiti; Diquini, November 1912, (Mann)

Tetragnatha festina is separated from other species of the genus
found in Hispaniola, in the male, by the second tooth from the base
of the fang that is directed upward and outward, from the plane of
the mandible, and in the female, by the long sinuous fang with a dis-
tinct tooth near the base.

Tetragnatha haitiensis spec. nov.
Figure 37

Female. Length without mand., 8.0 mm., ceph. 2.6 mm., abd. 5.6
mm., mand. 2.4 mm.

Cephalothorax brown, with darker brown shading, quite flat, thoracic
depression very near posterior margin; eyes, anterior row recurved,
a.m.e. separated by about a diameter and slightly larger than p.m.e.,
posterior row very slightly recurved, p.m.e. separated by more than
a diameter, less space between lateral eyes than between median eyes;
quadrangle longer than wide, and slightly narrower in front; mandibles
slender, divergent, superior margin, no dorsal spur, one sharp tooth

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 409

at base of fang, followed by a space, then a small tooth directed for-
ward, followed by six equally spaced, graduated teeth, inferior margin
with a very long tooth or lobe at base of fang, directed forward, as
seen from dorsal side, a short heavily chitinized tooth at inner base
of the fang, followed by a space, then two small subequal teeth and a
row of 10-12 graduated teeth; fang not as long as groove but extending
the length of the rows of teeth, sinuous, with a ventral tooth about
middle; sternum longer than wide, and carried between I and II coxae
and between II and III coxae; abdomen, base not bifid, pale brown,
with a pair of wavy darker stripes from base to spinnerets; legs,
1-2-4-3, brown, I pair very long, spines irregular, III and IV femora
with a prolateral row of short trichobothria best seen from ventral side.
Holotype 9 Haiti; Ennery, 7 September 1934, (Darlington)
Paratype 9 Haiti; Grand Riviere, January 1913, (Mann)
Paratype 9 Haiti; Kenskoff, 4,300 feet, 3 May 1935, (Roys)
Tetragnatha haitiensis belongs near T. antiUana Simon. The latter
species has evidently been misidentified by Seely, (1929, p. 105) and
F.O.P.-Cambridge, in the Biol. Centr. Amer., 1903, 2: 433, pi. 41,
figs. 5, 6, the specimens that they describe and figure do not have
the two subequal but strongly divergent teeth near the base of the
fang on the dorsal side of the male mandible and the single isolated
tooth on the lower margin of the female mandible that Simon de-
scribes. However, the female described by Seely and Cambridge, has
the long tooth or lobe that is directed forward at the base of the fang
on the lower side.

Tetragnatha haitiensis known only from the female, differs from
T. antiUana Simon, by the smaller size, the long sinuous fang with a
tooth on the ventral side, the two rather widely separated teeth,
instead of one, and the more numerous teeth on the lower margin of
the fang groove. Both have the long lobe or tooth at the base of the
fang on the ventral side that projects forward.

Tetragnatha orizaba (Banks)

Eugnatha orizaba Banks, 1898, p. 248, pi. 15, fig. 16. "one pair from Mt.
Orizaba"

2 cf Dom. Rep.; Puerto Plata, July 1941, (Hurst)
2 cf 1 9 Dom. Rep. ; San Jose de las Matos, 1,500+ feet, June 1938,
(Darlington)

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Tetragnatha pallescens F.O.P.-Cambridge

Tetragnaiha pallida Banks, 1892, p. 51, pi. 5, figs. 88, 88a. "2 c? Upper Cayuga

Lake Basin" nee pallida O. P. -Cambridge, 1889.
Tetragnatha pallescens F.O.P.-Cambridge, 1905, p. 436

9 Haiti; Port-au-Prince, 1-5 September 1934, (Darlington)
cf 9 Haiti; Trou Caiman, 4 September 1934, (Bates)

Tetragnatha tenuissima O.P. -Cambridge

Tetragnatha tenuissima O.P.-Cambridge, 1889, p. 9, pi. 1, figs. 1, 2. " d1 9
Guatemala, upper road to Chichochee near Coban, Tomahu; Costa Rica;
Panama."

9 Haiti; hills near Port-au-Prince, 2,000 feet, 2 October 1934,

(Darlington)
9 Dom. Rep.; Puerto Plata, April 1941, (Hurst)

Subfamily THERIDIOSOMATINAE
Key to Genera

1 . Abdomen much wider than long Allotot.ua

Abdomen globose * 2

2. A.M.E. largest of the eight, sternum triangular Wendilgarda

P.M.E. largest of the eight, sternum rectangular Ogulnius

Genus Allototua gen. nov.

Cephalothorax as wide as long, slightly convex, no thoracic groove,
anterior margin much narrowed; eyes, anterior row recurved, a.m.e.
diurnal, separated by a diameter and a little nearer a. I.e., posterior
row straight, eyes equidistant, p.m.e. largest of the eight, lateral eyes
on a common tubercle and almost touching; quadrangle wider than
long, as wide behind as in front; clypeus as high as quadrangle; labium
fused to sternum, wider than long; maxillae very broad, inclined over
labium; sternum wider than long, convex, posterior margin little nar-
rower than anterior; abdomen wider than entire length of spider,
widest posterior to middle, does not extend over cephalothorax; legs
rather short, no spines, with a distinct bristle at tip of patella and a
median dorsal bristle on posterior tibia; epigynum small. Male not
known .

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 411

Genotype Allototua guttata spec, no v.

Allototua differs from Theridiosoma by the very wide sternum, equi-
distant eyes of the posterior row and the bristles at the tip of the
patellae; it differs from Totua Keyserling 1891, by the shorter legs,
fewer bristles on tibiae and the eyes of the posterior row; from Ogulnius
O.P.-Cambridge, 1882, by the very wide abdomen that does not ex-
tend over the cephalothorax, the higher clypeus, and the eyes of the
anterior row.

Ogulnius Cambridge, 1882, is based on a very small spider, (1/24
inch = 0.5 mm.), with a globular abdomen that extends far over the
cephalothorax. Keyserling, Theridiidae, 2: 249, has placed in the
genus two more species, both known only from females from Southern
Brazil. Both are 2.0 mm. long. One is described with a strong bristle
at the tip of the patellae and a dorsal basal bristle on tibiae. The
genus Totua was described by Keyserling in the Brasilianische Spin-
nen, 1891, (p. 216) but was omitted by Simon in the Histoire Naturelle
des Araignees. Totua is also from southern Brazil. The quadrangle of
eyes is narrower in front, clypeus lower than quadrangle, abdomen
oval, and not extending over cephalothorax, legs long, with a bristle
at tip of patellae and two bristles on tibiae.

Allototua guttata spec, no v.
Figures 42, 45

Female. Length, 2.0 mm., ceph. 0.7 mm., abd. 1.6 mm. long 2.0
mm. wide.

Cephalothorax pale yellow with a distinct marginal black stripe and
three pairs of small black spots inside margin, no thoracic groove,
moderately convex, a pair of very long bristles directed forward on
thoracic slope, about as wide as long, anterior margin very narrow;
eyes cover anterior margin, anterior row recurved, a.m.e. diurnal,
strongly convex, separated by a diameter and from a. I.e. by a little
less, posterior row almost straight, eyes equidistant, p.m.e. largest of
the eight, rather flat, separated by a little less than a diameter, lateral
eyes almost touching, on a common tubercle; quadrangle as wide be-
hind as in front and wider than high; clypeus as high as quadrangle;
mandibles pale and weak, vertical, fang groove horizontal; labium pale,
fused to sternum, wider than long; maxillae pale, twice as long as
labium, broad, inclined over labium, so that tips almost meet; stertium
pale, with many gray dots irregularly placed, strongly convex, wider

412 bulletin: museum of comparative zoology

than long, with a few hairs, posterior margin fully two-thirds the an-
terior, so that IV coxae are separated by more than a diameter;
abdomen as wide as entire length of spider, not extending over cephalo-
thorax, widest posterior to middle, basal half very high, pale, with
irregular dark blotches and smaller cream-colored spots, distal third
slopes rapidly, with two parallel transverse rows of dark spots, followed
by three longitudinal pairs of dark dots, venter pale, two distinct
pairs of dark spots just above spinnerets; legs, 1-2-4-3, rather short,
pale, with paired lateral dark spots on femora, patellae and tibiae,
many long colorless hairs, and a long bristle at tip of patellae, a dorsal
basal bristle on III and IV tibiae; epigynum rather small, a pale oval
lobe, with heavily chitinized margins.

Holotype 9 Dom. Rep. ; Loma Viega, Cordillera Central, south of
Constanza, 6,000 feet, August 1938, (Darlington)

Genus Ogulnius O.P.-Cambridge 1882

Ogulnius fulvus spec. nov.
Figures 46, 47

Female. Length, 1.4 mm.

Cephalothorax pale yellow, about as wide as long, cephalic portion
very high, with a few long bristles posterior to eyes on margins,
three pairs of dark elongated radial striae, abdomen extends so far
over cephalothorax obscuring the thoracic groove; eyes raised on a
turret, anterior row recurved, a.m.e. diurnal, largest of the eight, sepa-
rated by less than a line, a. I.e. smaller, and separated from a.m.e. by
almost a diameter of a.m.e., posterior row slightly procurved, lateral
eyes touching, p.m.e. separated by more than a diameter; quadrangle
narrower in front and higher than wide; labium wider than long,
not fused to sternum, tip narrowed; maxillae almost twice as long as
labium, inclined, very broad; sternum as wide as long, convex, not
carried between coxae, posterior margin two-thirds as wide as anterior,
with a circular pit each side of labium; abdomen globose, higher than
long, covering the thoracic portion of the cephalothorax, to the ocular
tubercle, pale brown with a few paler blotches, a pair of dark spots
directly posterior to anterior muscle spots, smooth, with very few
hairs, anterior muscle spots close together, posterior muscle spots
widely separated, venter pale, colulus present; legs 4-1-2-3, pale
yellow, quite short and stout, a long bristle at tip of patellae and
middle of tibiae, I tibia enlarged and little longer than patella; epigy-

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 413

num very near to pedicle, a broad curved chitinized slit, showing no
structure.

Holotype 9 Dom. Rep.; foot hills of Cordillera Central, south of
Santiago, 2,000-5,000 feet, June 1938, (Darlington)

Paratype 9 Dom. Rep.; San Jose de las Matas, 1,500 feet, June
(1938, (Darlington)

The genus Ogulnius was based by Cambridge in 1882, on a species
from the Amazon known only from a female. Later, Simon has added
to the genus, one species from Ceylon and one from St. Vincent.
Cambridge has several figures which show that fulvus agrees with
the genotype in the three pairs of diagonal dark marks on the margin
of the cephalothorax and a larger pair directly posterior to the p.m.e.
but the a.m.e. are much longer than the figure of the genotype. The
male is not known.

Genus Wendilgarda Keyserling 1886
Wendilgarda theridionina Simon

Vendilgarda theridionina Simon, 1895, 1, p. 919, fig. 986. 'V 9 Venezuela;
San Esteban"

9 Haiti; Grande Anse, (Uhler)
<? 9 Dom. Rep.; Puerto Plata, 30 August 1938, (Darlington)
9 Dom. Rep. ; Cordillera Central, Constanza, 3,000-4,000 feet,
August 1938, (Darlington)
This species was first described from Venezuela and has since been
found widely distributed among the islands of the Caribbean. Petrun-
kevitch has a full description of both sexes in his Spiders of Porto Rico,
1930, p. 297. With strong direct light, small circular pits are seen on
the sternum, each side of the labium. These are larger and more dis-
tinct in the female than in the male.

Subfamily GASTERACANTHINAE

Key to Species

1 . Head elevated, abdomen extending over thorax, wider than long 2

Head not elevated, abdomen not extending over thorax, longer than

wide 3

2. Abdomen with 2 pairs of spines Gasteracantha tetracantha

Abdomen with 3 pairs of spines Gasteracantha cancriformis

3. Abdomen with 2 pairs of spines, posterior pair very long, sharp and

divergent Micrathena armata

Abdomen with 3 pairs of spines, all relatively short, 2 posterior pairs,
swollen, with sharp tips, subparallel Micrathena similis sp. n.

414 bulletin: museum of comparative zoology

Genus Gasteracantha Sundevall 1833
Gasteracantha cancriformis (Linn.)

Aranea cancriformis Linnaeus, 1758, p. 624. " 9 Jamaica"
Gasteracantha hexacantha Banks, 1903, p. 341.
Gasteracantha sexserrata Banks, 1903, p. 341.

A common spider found in all parts of Haiti and the Dominican
Republic.

Gasteracantha tetracantha (Linn.)

Aranea tetracantha Linnaeus, 1767, p. 1037. " 9 St. Thomas"
Gasteracantha hilaris Banks, 1903, p. 341.

3 9 Haiti; Port-au-Prince, (Crew), Banks Coll.

Genus Micrathena Sundevall 1833

MlCRATHENA MILITARIS (Fabr.)

Aranea militaris Fabricius, 1775, p. 434, [ 9 1 "in America"

Aranea armata Olivier, 1791, 4, p. 202. "no locality"

Plectana armata Walckenaer, 1841, 2, p. 179, pi. 22, fig. 1. "9 Nouveau-

Monde — Archipel occid. — Saint Dominque."
nee Micrathena sexspinosa Reimoser, 1919, p. 126.

This species must be very common on the island from the number
found by collectors. Walckenaer figures a female under the name
Plectana armata. The- anterior pair of spines are about one-third from
the base and are sharp, black, vertical, close together and so little
divergent that they are almost parallel. The posterior spines are much
longer and are strongly divergent In a few specimens there is a third
pair of spines on the lateral margins midway between the anterior and
posterior pairs. These are short and little more than cusps in
most specimens.

Reimoser in his revision of the genus, places Plectana armata Walck-
enaer as a synonym of Micrathena sexspinosa Halm. This is an error
as in the latter species, the posterior spines are thick, not sharply
pointed, only a little divergent, and the first and second pairs of spines
are about subequal and the middle pair of spines is not on the margin.
The figure of the epigynum also shows that the middle section is broad.

Micrathena militaris is probably confined to the islands of the
Caribbean. Petrunkevitch reports it from the Virgin Islands and
Puerto Rico and it is not uncommon in Cuba. Specimens are in the
museum collection from at least a dozen localities in Hispaniola.

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 415

MlCRATHENA SIMILIS spec. nOV.

Figures 8, 43, 44

Female. Length, 5.0 mm., ceph. 2.0 mm., abd. 4.0 mm. long, 4.0
mm. wide at posterior margin.

Cephalothora.r pale brown, cephalic portion fairly high, thoracic
groove punctiform, each side on thoracic portion, midway between
groove and margin, three small depressions; eyes in three groups,
lateral eyes subequal, touching and on extreme margin, quadrangle of
median eyes higher than wide, slightly narrower in front, p.m.e. a little
larger than a.m.e., and separated by less than a diameter, a.m.e. sepa-
arated by a scant diameter; clypens almost wanting below a.m.e. and
below lateral eyes equal to more than a diameter of a.m.e.; mandibles
brown, vertical, no boss, three teeth on inferior margin of fang groove;
libium wider than long, tip rounded; maxillae twice as long as labium,
sides parallel; sternum dark brown, iridescent, strongly convex, pen-
tagonal, with sides almost parallel, as wide as long with a distinct
swelling opposite the first three pairs of coxae; abdomen as long as
wide, much narrowed at base, with three pairs of spines, basal pair
small, with blunt tips, extending far over cephalothorax, other two
pairs of spines at posterior end, one above the other, distal pair slightly
larger, rather short, with sharp points, sequilla best seen in figure, the
two pairs of muscle spots form a quadrangle longer than wide, height
ot abdomen above spinnerets, twTo-thirds length of abdomen, sides
creased, dorsum pale w7ith sides and distal area black, venter black;
legs, spineless and rather short ; epigynum, area wider than long, a small
median finger protrudes from plane of abdomen, with tip colorless and
much narrowed, each side above fold are circular openings.

Holotype 9 Dom. Rep. ; Puerto Plata, 30 August 1938, (Darlington)

Paratype 9 Dom. Rep. ; Mt. Diego de Ocampo, north range, 3,000-
4,000 feet, July 1938, (Darlington)

Micrathena similis is closely related to M. eubana (Banks). Both
have three pairs of spines in similar positions, but in eubana the two
posterior pairs of spines are not parallel, the spines are much sharper
and the abdomen usually has a large dark median spot which is lacking
in similis. The epigynums are quite unlike. Both species have a punc-
tiform thoracic groove with the three pairs of depressions each side on
the thoracic portion and a pentagonal sternum as wide as long.

416 bulletin: museum of comparative zoology

BIBLIOGRAPHY

Banks, Nathan

1892. The Spider Fauna of the Upper Cayuga Lake Basin. Proc. Acad.

Nat. Sci. Phila., pp. 11-81, pis. 1-5.
1898. Arachnida from Baja California and other Parts of Mexico.

Proc. Cal. Acad. Sci., 3rd. ser., 1, pp. 205-296, pis. 13-17.
1903. A List of Arachnida from Hayti, with descriptions of New Species.

Proc. Acad. Nat. Sci. Phila., 65, pp. 340-345, pi. 15.

BryanTj Elizabeth B.

1940. Cuban Spiders in the Museum of Comparative Zoology. Bull.
Mus. Comp. Zool., 86, pp. 249-532, pi. 1-22.

Cambridge, F. O. P.-

1897-1905. Arachnida; Araneidae and Opiliones. 2, pp. XII +610, 54
pis. Biologia Centrali-Americana.

Cambridge, O.P.-

1882. On new Genera and Species of Araneidae. Proc. Zool. Soc. London,

pp. 423-442, pis. 29-31.
1889-1902. Arachnida; Araneidea, 1, pp. XV 317, 39 pis. Biologia

Centrali-Americana.

Dahl, Friedr.

1914. Die Gasteracanthen des Berliner Zoologische Museums und deren
geographische Verbreitung. Mitt. Zoolog. Mus. Berlin, 7, pp.
pp. 235-501, figs.

Fabricttjs, J. C.

1775. Systema Entomologiae, etc. Aranea, pp. 431-439.

Forskal, P.

1775. Descript. Anim.

Franganillo, Pelegrin Balboa, S. J.

1930. Mas Aracnidos Nuevos de la Isla de Cuba. Mem. Inst. Nat. de
Investigue. Cientif. Habana, 1, pp. 47-99, 21 figs. (A reprint or
reissue by the author is separately paged, beginning with page 1.)

Hentz, N. M.

1847. Descriptions and Figures of the Araneides of the United States.

Journ. Boston Soc. Nat. Hist., 5, pp. 444-478, pis. 23-24, and

30-31.
1850. (continuation), 6, pp. 18-35, pis. 3-4.
1875. The Spiders of the United States. Occ. Papers Boston Soc. Nat.

Hist., 2, pp. XIII + 171, pis. 1-21.

BRYANT: THE ARGIOPIDAE OF HISPANIOLA 417

Keyserling, Graf Eugen von

1864. Beschreibungen neuer und wenig bekannter Arten aus der Familie
Orbitelae, etc. Sitz-Bericht Isis, Dresden, (Jhg. 1863), pp 63-98
119-154, pis. 1-7.

1865. Beitrage zur Kenntnis der Orbitelae. Verh. z.-b. Ges. Wien, 15,
pp. 799-856, pis. 18-21.

1884. Neue Spinnen aus Amerikas. Ibid., 34, pp. 489-534, pi. 13.
1884-1886. Die Spinnen Amerikas. vol. 2, Theridiidae, in 2 parts, pp.

1-222, pis. 1-10, and pp. 1-295, pis. 11-21.
1891. Ibid., Brasilianische Spinnen. vol. 4, pp. 1-278, pis. 1-10.

Koch, Ludwig

1875. AegyptischeundAbyssin.,Arachniden, ges. vonJickeli. (Nurnberg)

Linnaeus, Carolus

1758. Systema Naturae, etc. Editio X, reformata.
1766-68. Systema Naturae, etc. XII, reformata.

Marx, George

1889. Catalogue of the described Araneae of temperate North America.
Proc. U. S. Nat. Mus., 12, pp. 497-594.

McCook, Henry C.

1887. Note on Cyrtophora bifurca n.sp. and her Cocoons. Proc. Acad.

Nat. Sci. Phila., p. 342.
1889-1893. American Spiders and their Spinning Work. In 3 volumes
published by the author, Philadelphia, vol. 1, 1889, vol. 2, 1860,
vol. 3, 1893.

t
Olivier, Guill. Ant.

1791. Araignee. Encyclopedie Mdthodique. pp. 173-240, pis. 256-261

Petrunkevitch, Alexander

1911. A Synonymic Index-Catalogue of Spiders of North, Central, and

South America, etc. Bull. Amer. Mus. Nat. Hist., New York, 29,

pp. 1-791.
1926. Spiders from the Virgin Islands. Trans. Conn. Acad. Arts Sci.,

28, pp. 21-78, 28 figs."
1930. The Spiders from Porto Rico. Ibid., 30, pp. 159-355, figs.

Reimoser, Eduard.

1917. Die Spinnengattung Micrathcna Sundevall. Verh. z.-b. Ges.
Wien, 67, pp. 73-160, pis. 1-9.

Seeley, Ralph M.

1938. Revision of the Spider Genus Tetragnatha. Bull. N. Y. State
Mus., Albany, no. 298, pp. 99-150, pis. 1-4.

418 bulletin: museum of comparative zoology

Simon, Eugene

1892-1903. Histoire Naturelle des Araignees. 2nd. edition, Paris, 2 vols.,

Vol.1, 1892, pp. I-VII, 1-1084, 1098, figs.; vol. 2, fasc. 1, 1897;

pp. 1-192, figs. 1-200, fasc. 2, 1898, pp. 193-380, figs. 201-384,

fasc. 3, 1901, pp. 381-668, figs. 385-792, fasc. 4, 1903, pp. 669-

1080, figs. 793-1122.
1897. On the Spiders of St. Vincent. Part III. Proc. Zool. Soc. London,

pp. 860-890.

Walckenaer, Baron deC. A.

1802. Faune Parisienne, 2 volumes, Paris. Aranea in vol. 2, pp. 187-250.
1805. Tableau des Araneides. Paris, XII + 88 pages, 9 pis.
1837-1847. Histoire Naturelle des Insects Apteres. 4 volumes, Paris.
Atlas with 52 plates.

PLATES

PLATE 1

Bryant — Argiopidae of Hispaniola

PLATE 1

Fig. 1. Aranea(?) crewi (Banks); right palpus, retrolateral view.

Fig. 2. Aranea{7) hispaniola Bryant; epigynum.

Fig. 3. AraneaC?) crewi (Banks); right palpus, prolateral view.

Fig. 4. Eustala perdita Bryant; epigynum.

Fig. 5. Eustala bisetosa Bryant; left palpus, prolateral view.

Fig. 6. Eustala bisetosa Bryant; epigynum.

Fig. 7. Eustala vegeta (Keys.); left palpus, prolateral view.

Fig. 8. Micrathena similis Bryant; epigynum, lateral view.

Fig. 9. Eustala perdita Bryant; left palpus, prolateral view.

Fig. 10. Drexelia minor Bryant; epigynum.

Fig. 11. Eustala delasmata Bryant; left palpus, prolateral view.

cl. = clavis; emb. = embolus; para. =paracymbium; unc. =unca.

BULL. MUS. COMP. ZOOL.

Bryant. Argiopidae of Hispaniola. Plate 1

PLATE 2

Bryant — Argiopidae of Hispaniola

PLATE 2

Fig. 12. Metazygia manni Bryant; left palpus, prolateral view.

Fig. 13. Metazygia manni Bryant; left mandible, front view.

Fig. 14. Parawixia darlingtoni Bryant; left palpus, prolateral view.

Fig. 15. Metargyra maculata Bryant; epigynum.

Fig. 16. Parawixia darlingtoni Bryant; paracymbium.

Fig. 17. M angora striatipes Bryant; left palpus, prolateral view.

Fig. 18. M angora striatipes Bryant; epigynum.

Fig. 19. Neosconella parva Bryant; epigynum.

Fig. 20. Metepeira inerma Bryant; epigynum.

Fig. 21. Parawixia darlingtoni Bryant epigynum.

Fig. 22. Meta (?) blanda Bryant; epigynum.

Fig. 23. Metazygia manni Bryant; epigynum.

Fig. 24 Neosconella parva Bryant; epigynum, lateral view.

Fig. 25. MetaC?) hotteiensis Bryant; epigynum.

BULL. MUS. COMP. ZOOL.

Bryant. Argiopidae of Hispaniola. Plate 2

PLATE 3

Bryant — Argiopidae of Hispaniola

PLATE 3

Fig. 26. Pseudometa comuta Bryant; left palpus, ventral view.

Fig. 27. Pseudometa comuta Bryant; epigynum.

Fig. 28. Pseudometa comuta Bryant; left palpus, dorsal view.

Fig. 29. Pseudometa conspersa Bryant; epigynum.

Fig. 30. Pseudometa obscura Bryant; epigynum.

Fig. 31. Antillognatha lucida Bryant; left palpus, lateral view.

Fig. 32. Pseudometa linguiformis (Frang.); epigynum.

Fig. 33. Agriognatha rucilla Bryant; left palpus, ventral view.

Fig. 34. Agriognatha espanola Bryant; left palpus, lateral view.

Fig. 35. Agriognatha argyra Bryant; left palpus, lateral view.

BULL. MUS. COMP. ZOOL.

Bryant. Argiopidae of Hispaniola. Plate 3

PLATE 4

Bryant — Argiopidae of Hispaniola

PLATE 4

Fig. 36. Hispanognatha guttata Bryant; left palpus, lateral view.

Fig. 37. Tetragnatha haitiensis Bryant; female, left mandible, ventral view.

Fig. 38. Tetragnatha festina Bryant; left palpus, embolus.

Fig. 39. Tetragnatha festina Bryant; left palpus, lateral view.

Fig. 40. Tetragnatha festina Bryant; female, right mandible, dorsal view.

Fig. 41. Tetragnatha festina Bryant; male, right mandible, dorsal view.

Fig. 42. Allototua guttata Bryant; epigynum.

Fig. 43. Micrathena similis Bryant; epigynum.

Fig. 44. Micrathena similis Bryant; female, dorsum.

Fig. 45. Allototua guttata Bryant; female, cephalothorax.

Fig. 46. Ogulnius fulvus Bryant; female, face.

Fig. 47. Ogulnius fulvus Bryant; female, lateral view.

BULL. MUS. COMP. ZOOL

Bryant. Argiopidae of Hispaniola. Plate 4

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCV, No. 5

A REVISION OF THE GENUS EPICAUTA

IN AMERICA NORTH OF MEXICO

(COLEOPTERA, MELOIDAE)

By Floyd G. Werner

Biological Laboratories
Harvard University

With Seven Plates

CAMBRIDGE, MASS. U. S. A.
PRINTED FOR THE MUSEUM

May, 1945

PUBLICATIONS
OF THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

The Bulletin and Memoirs are devoted to the publication of
investigations by the Staff of the Museum or of reports by spec-
ialists upon the Museum collections or explorations.

Of the Bulletin, Vols. I to XCIV, and Vol. XCV, No. 1, 2, 3, 4
and 5 have appeared and of the Memoirs, Vol. I to LV.

Publication of Memoirs ceased with Vol. LV.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. XCV, No. 5

A REVISION OF THE GENUS EPICAUTA

IN AMERICA NORTH OF MEXICO

(COLEOPTERA, MELOIDAE)

By Floyd G. Werner

Biological Laboratories
Harvard University

With Seven Plates

CAMBRIDGE, MASS. U. S. A.
PRINTED FOR THE MUSEUM

May, 1945

No. 5. — A Revision of the Genus Epicauta in America North of Mexico1

By Floyd G. Werner

INTRODUCTION

Since the time of C. V. Riley, every textbook of entomology has
devoted space to one or more species of Epicauta, as an example of
hypermetamorphosis, and as a garden pest. Every beginner knows
Epicauta marginata, the "margined blister-beetle" and its grey form
cinerea. It seems amazing that nobody has ever pointed out that the
margined blister-beetle of potatoes does not have a grey form over
most of its range. True, in most localities one may collect margined
beetles with a few grey individuals among them but they occur on
Clematis, not potatoes, and are actually of a species distinct from that
found on potatoes. Over the western part of the United States and in
Mexico, five species have been known as cinerea. Yet all are perfectly
distinct and one is only remotely related to it. Coming back to mar-
ginata, we find that Fabricius used the name for an insect from the
Cape of Good Hope, later for a North American species.

Oversights like these do not speak well for the work of field men and
collectors alike: Perfectly distinct species are confused in the field and
in the collections because superficially they resemble each other.
Everyone seems to assume that since the insect is large and common
it is well known.

This revision is offered to the field entomologist as a preliminary
work. I have attempted to straighten out the nomenclatorial problems
and to define the known species as well as I could from museum ma-
terial. It is my hope that it will form a basis for a more exact knowl-
edge of this economically important group.

ACKNOWLEDGEMENTS

Originally this revision was presented as a thesis for honors at grad-
uation at Harvard College, the work being done under the supervision
of Dr. Frank M. Carpenter. Much of the work then and later has been
done in the M. C. Z. and with the Museum collection. Mr. Banks and
Dr. Darlington have been extremely helpful in extending favors and
giving advice. During the progress of the work, it was discovered that
Dr. William H. Anderson of the National Museum had been working
up a revision also. Dr. Muesebeck and he decided to let me finish the
job. Dr. Anderson and Mr. H. S. Barber of the National Museum

i Published with the aid of a special gift from Mr. George R. Agassiz.

422 bulletin: museum of comparative zoology

generously contributed their notes and observations and gave me free
access to the collections. During the course of the work, the following
museums have been visited, the collections and types examined : U. S.
National Museum, Academy of Natural Sciences at Philadelphia,
American Museum of Natural History, Field Museum of Natural
History, Cornell University collection, Carnegie Museum, Purdue
University and the collection of W. S. Blatchley, and Ohio State
College. The curators have everywhere been extremely free with
favors. In addition, the collections of Colorado State College and of
the Iowa Insect Survey were sent for examination.

DISCUSSION OF CHARACTERS USED

The tribe Epicautini includes all of the Meloidae with a patch of
cinerous pubescence on the inside of the anterior femora. All of our
species have this patch in a shallow excavation. In a few it extends
the length of the femur.

The relationship of the species and delimitation of genera within the
tribe is somewhat complicated. On the basis o? one character, the
groups form one way; on another, they form another way. The only
group the author has been able to delimit satisfactorily is the genus
Pleuropompha, with two North American species. It has strongly
raised costae on the elytra, scale-like pubescence in part and has
several of the intermediate segments of the antennae, not the basic,
elongated and modified in the male.

Henous and Nomaspis were originally described as being related to
Meloe because they are wingless and the elytra are shortened. They
have a cinereous patch on the anterior femora and fit into Epicauta
very well, in the group with slender, tapered antennae. Winglessness
is not a sufficient character for separating them in my estimation.

Causima includes a single South American species, vidua, which
differs from Epicauta only in being heavier and having the elytra
slightly shortened. If the genus were to be maintained, a whole series
of intermediate species would have to be taken into account. At best
it would be a separation on a very trivial character.

Isopentra was described as an Old World genus and would include
the species with filiform to ensiform antennae and with two spurs on
the anterior tibiae of the male. It is hard to maintain because one
must arbitrarily limit it. Many species have antennae on the border-
line between slightly broadened and ensiform. Some of these have
ensiform antennae in the male and not in the female. The Old World

WERNER: NORTH AMERICAN EPICAUTA 423

group is uniform but no good characters appear to separate it from
such species as oregona and ensifarmis in our fauna.

Gnathospasta was described to include one species, mimetica, with
deeply incised labrum, elongated mandibles and modified maxillae.
One other Mexican species was added to it. In this paper several
species are described, which make it difficult to maintain the genus.
Calcarata has the mouth characters but falls into a very different group
on the basis of the antennae. Mimetica has the antennae slender and
tapered, with the first segment somewhat elongated and excavated
externally at the tip. Uniforma and alpina have antennae approaching
mimetica and also mouthparts, especially alpina. Calcarata has heavy,
uniform antennae and would fall into Group C of Horn's tables. So
Gnathospasta can be sunk under Epicauta on the basis of both connect-
ing species and duplications of diagnostic characters in an unrelated
species.

The genus Macrobasis, defined as including all species with the second
segment of the antennae not shortened, is a very heterogeneous group.
It includes almost all of the species with the first segment of the
antennae elongated in the male. But not all. Longicollis, linearis and
maculifera have the first segment elongated in the male but have a
short second segment. An arbitrary division is made between tenella
in Macrobasis and merkeliana in Epicauta, two species otherwise
closely related. Segmenta, immaculata and some others do not have the
first segment elongated in the male.

Apterospasta was created by Le Conte to include the form of seg-
vienta with short wings and somewhat bulbous elytra, which he called
valida. It seems to be just a form of a species that would be placed in
Macrobasis and the genus has long been sunk under it.

It is inconvenient to have a genus very large and not subdivided,
especially when it is World-wide in distribution as is Epicauta. Horn
divided the North American species of Epicauta proper into three
groups, "A" which included the species with fairly short uniform
antennae, "B" which contained species with somewhat tapered
antennae and modified heads, "C" which contained the species with
filiform to ensiform antennae. Group "C" would contain all of the
other genera here sunk under Epicauta. It also contains all of the
species of Epicauta as now constituted outside of North America.

Groups "A" and "B" stand as recognizable units. Group "C" is
intimately associated with Macrobasis and the rest of the old genera
which fall into the Epicautini. Division on the basis of antennal char-
acters is not natural, as pointed out under the discussion of Macrobasis.

424 bulletin: museum of comparative zoology

The presence of one spur or two spurs on the anterior tibiae of the
male is one possibility for division. Some species of the old group "C",
some of Macrobasis with straight first antennal segment in the male
and some with curved first antennal segment fall into each group.
There is no appearance of homogeneity in either.

Late in my work on the group I found a character which seems to
follow a more natural grouping of species than has as yet emerged. On
the inner side of the apex of the posterior tibiae of the male, there is
a short row of stout teeth.

They occur in all of the Macrobasis group with curved first antennal
segment in the male except longicollis, albida and texana. In addition
they occur in the species of Epicauta with a suggestion of the curved
first segment: uni forma, alpina and mimetica. Beside that they occur
in alastor and ingrata in Epicauta and in lauta and tenella among the
species formerly included as borderline species in Macrobasis. They
occur in polingi, liebecki and arizonica, which have a straight first
segment in the male. They do not occur in the rest of the species of
group "C", in immaculata and segmenta of the borderline species of
Macrobasis, in the species of Macrobasis with straight first antennal
segment in the male, except for the polingi group, nor in longicollis,
albida and texana with curved first antennal segment.

Following is an attempt to group the species on the basis of the
characters mentioned. These groups seem to represent natural units.

A. Posterior tibiae of male without apical teeth.
B. Antennae tapering toward apex.
C. Head or legs not modified.

D. Anterior tibiae of male with two spurs.

E. Second segment of antennae not more than half as long
as third (corvina, funebris, pensylvanica, conferta, parva,
fallax, albolineata, brunnea, cinerea, fissilabris, solani,
floridensis, ruidosana, punctipennis, balli, oregona, ensi-
formis, stuarti, lemniscata, vitlata, occidenialis.)
EE. Second segment of antennae more than half as long as
third.

F. First segment of male antennae straight, {immaculata,
segmenta, sublineata, flavocinerea, fabricii, murina,
subglabta, tenuis, excors, tenuilineata, (languida?) .)
FF. First segment of male antennae curved (albida, tex-
ana, longicollis).
DD. Anterior tibiae of male with one spur, (abadona, insignis,
nigritarsis, crassitarsis, pedalis, maculata, normalis, nogales,
phoenix, pardalis, andersoni, magnomaculata, ventralis).

WERNER: NORTH AMERICAN EPICAUTA 425

CC. Pronotum with a single median tubercle (excavatifrons).

CCC. Head usually modified. Pubescence usually short and dense.

Some species have anterior lateral black spots on the abdominal

sternites. (caviceps, impressifrons, rileyi, straba, rehni, wheeleri,

diversipubescens, aspera).

CCCC. Middle and posterior legs of male flat behind and curved, with

fringes of long pubescence, (calif or nica, alphonsii).
BB. Antennae uniform or almost uniform in thickness, rather short (fer-
ruginea, fortis, strigosa, baksii, callosa, sericans, atrata, pruinosa,
immerita, piceiventris, sanguinicollis, puncticollis, oblita, barberi,
kansana, calcarata, heterodera) .
AA. Posterior tibiae of male with a row of stout apical teeth. Fig. 8.
G. Anterior tibiae of male with two spurs.

H. First segment of antennae straight (ingrata, tenella, merkeliana) .
HH. First segment of antennae with an external apical excavation,
especially in the male, (uniforma, alpina, mimetica).
GG. Anterior tibiae of male with one spur (none in alastor).

I. First segment of anterior tarsi of male at least as long as second,
straight. First segment of male antennae large, curved, {pur-
purea, torsa, atrivittata) .
II. First segment of anterior tarsi of male short, contorted.
J. First segment of antennae not modified in male (alastor.)
JJ. First segment of antennae elongated and modified in male.
K. First segment of male antennae straight, not excavated
(lauta, polingi, arizonica, liebecki).
KK. First segment of male antennae curved and excavated
externally near the tip. (ochrea, gissleri, parkeri, virgulata,
linearis, maculifera.)

Tribe Epicautini Denier, 1935, Rev. de la

Soc. Ent. Arg. 7:151

Epicauta Dejean

Epicauta Dejean, 1833, Catalogue des Coleopteres:224. Genotype: Lytta
erythrocephala Pallas, 1776, Reise russ. Reich 1 App.:14 (new designation
to concur with designation as genotype of Epicauta Redtenbacher, 1845,
Gatt. deutsch. Kafer-Fauna:133 by Wellman, 1910, Canadian Entomolo-
gist 42:396.)

Causima Dejean, 1837, Catalogue des Coleopteres, 3rd Ed. :248. Genotype:
Cantharis vidua Klug, 1825. (new designation to concur with designation
as genotype of Causima Lacordaire, 1859 by Wellman, 1910, Canadian
Entomologist 42:394.)

426 bulletin: museum of comparative zoology

Henous Haldeman, 1852, Stansbury's Expl. and Surv. Great Salt L. Valley:
377. Genotype : Henous techanus Haldeman, loc. cit. (monobasic) = Meloe
confertus Say, 1824, Journ. Acad. Nat. Sci. Phila. 3:281.

Isopentra Mulsant, 1858, Opusc. Ent. 8:106. Genotype: Lytta megalocephala
Gebler, 1817, designated by Wellman, 1910, Canadian Entomologist
42:396.

Macrobasis Le Conte, 1858, Journ. Acad. Nat. Sci. Phila. (2) 4:39. Genotype:
Lytta albida Say, 1823, Journ. Acad. Nat. Sci. Phila. 3:305, designated by
Wellman, 1910, Canadian Entomologist 42:396.

Apterospasta Le Conte, 1858, Journ. Acad. Nat. Sci. Phila. (2) 4:39. Geno-
type: Apterospasta valida Le Conte, loc. cit. (monobasic).

Nomaspis Le Conte, 1866, Smiths. Misc. Coll. 6 no. 167, 2nd Ed.:156. Geno-
type: Meloe parvus Haldeman, 1852, Stansbury's Expl. and Surv. Great
Salt L. Valley: 377, (monobasic.)

Gnathospasta Horn, 1875, Trans. Am. Ent. Soc. 5:154. Genotype: Gnatho-
spasta mimetica Horn, loc. cit. (monobasic).

Key to the Epicautini of North America, North of Mexico

Elytra with several strongly raised costae. Third to fifth antennal segments
somewhat elongated. Body with some scale-like hairs.

Genus Pleuropompha Le Conte

Elytra without raised costae. Third to fifth antennal segments not elon-
gated. Pubescence not scale-like. Genus Epicauta Dejean

Genus Pleuropompha Le Conte

With four raised costae on each elytron, costata (Le Conte)
With three raised costae on each elytron, tricostata Werner

Genus Epicauta Dejean

1. Second segment of antennae half as long as third or shorter 2

Second segment of antennae two-thirds as long as third or longer . 86

2. (1) Pubescence cinereous, denuded in a number of spots, at least on
the elytra 3

Pubescence not denuded in small spots 19

3. (2) Anterior tibiae with a single strong, incurved spur ( d" c?) 4

Anterior tibiae with two slender, spiniform spurs 12

4. (3) With a broad black mark apically on the abdominal sternites 5

Without a median mark on the abdominal sternites 6

WERNER: NORTH AMERICAN EPICAUTA 427

5. (4) 18 mm. or longer. Spots on elytra very irregular in size, some large
but not confluent. A pair of small denuded callosities on the pronotum.
California magnomaculata Martin cf

15 mm. or less. Spots on elytra small, quite regular in size. Pro-
notum without callosities ventralis sp. n. c?

6. (4) One or both pairs of palpi expanded 7

Palpi not expanded 9

7. (6) Both pairs of palpi very broad and flat. Spots on body large and
usually partly confluent andersoni Werner d1

Only maxillary palpi expanded 8

8. (7) Maxillary palpi with last segment almost orbicular. Elytra not
truncate maculata (Say) cf

Maxillary palpi broad but last segment triangular. Elytra truncate.
Nogales and Tucson, Arizona nogales Werner c?

9. (6) First segment of anterior tarsi narrow at base, without pad except
on apical fourth normalis Werner c?

First segment of anterior tarsi with pad extending almost its full
length 10

10. (9) Spots very large, confluent, shiny. Head with a number of deep
punctures pardalis Le Conte <?

Spots smaller. Head without deep punctures 11

11. (10) Spots on elytra fairly large, on rest of body small. Elytra swollen
toward the apex, complete. Phoenix, Arizona phoenix Werner c?

Spots on elytra small. Elytra truncate, exposing two or three
abdominal tergites. Maxillary palpi expanded, but not conspicuously.
Nogales and Tucson, Arizona nogales Werner d*

12. (3) Elytra brown, with small denuded spots. Rest of body black,
without spots punctipennis Werner

Elytra of same color as rest of body 13

13. (12) Pubescence short and fairly sparse, exposing surface. Denuded
spots small. Antennae ensiform, especially in the male, very slender at
apex. Fig. 16 oregona Horn

Pubescence dense, obscuring surface. Antennae not very slender at
apex 14

14. (13) Abdominal sternites with anterior lateral black spot 15

Abdominal sternites without anterior lateral black spot 16

15. (14) 18 mm. or longer. Spots on elytra very irregular in size, some large
but not confluent. A pair of denuded callosities on pronotum. Cali-
fornia magnomaculata Martin 9

15 mm. or less. Spots on elytra smaller, quite regular in size. No
callosities on pronotum. Nebraska ventralis sp. n. 9

16. (14) Spots small to medium, not confluent 17

Spots large, in part confluent. Females of andersoni Werner and
pardalis Le Conte. The former has more slender antennae. Compare
with males for safe determination.

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17. (16) Elytra not swollen or truncate. Spots more or less uniform over
body and elytra. Females of maculata (Say) and normalis Werner. Asso-
ciate with males from same locality.

Elytra swollen or truncate. Arizona 18

18. (17) Elytra swollen behind. Spots on elytra large, on body small.
Phoenix, Arizona -phoenix Werner 9

Elytra truncate, exposing two or three abdominal tergites. Spots
small. Nogales and Tucson, Arizona nogales Werner 9

19. (2) Elytra with one or more vittae, either structural or formed by
denser pubescence, or by differently colored pubescence 20

Elytra without vittae, even indistinct 31

20. (19) Vittae marked on elytra as well as in pubescence 21

Vittae marked in pubescence only 24

21. (20) Outer vitta very narrow, not over half as broad as the one next
to it abadona Skinner

Outer vitta at least as broad as one next to it 22

22. (21) Outer edge of anterior tibiae and tarsi smooth, denuded, shiny.
Antennae flattened and broadened toward the middle. Fig. 10.

occidenlalis Werner

Anterior tibiae and tarsi not shiny, moderately densely pubescent.

Antennae not broadened. Fig. 9 23

23. (22) Outer two vittae of elytra fused or partly fused, at least in some
specimens of a series. Northern and central states vittata (Fab.)

Outer two vittae never fused or partly fused. Southern states,
extending north to Nebraska west of the Mississippi. . .lemniscata (Fab.)

24. (20) Head and pronotum rufous. Elytra black. Vitta formed by denser
cinereous pubescence sanguinicollis Horn

Head and pronotum not both rufous 25

25. (24) Head rufous. Pronotum black or dark brown. Vittae formed by
darker pubescence. Fig 18 atrata (Fab.) in part

Head and pronotum of same color 26

26. (25) Ground color black 27

Ground color mainly tan. Pubescence denser on a single line on
each elytron, producing an indistinct vitta 28

27. (25) Three white lines on each elytron. Southwestern states.

albolineata (Duges)

Pubescence cinereous to tan with two dark brown or black vittae.

Gulf and, rarely, Atlantic states strigosa (Gyll.)

28. (26) With scutellar and humeral black or dark brown spot on elytra.
First segment of male antennae elongated, curved, excavated externally
at the apex. Fig. 53 virgidata (Le Conte)

No scutellar or humeral spot 29

29. (28) 14 mm. or longer. Narrower than the following two species. Ante-
tior tibiae of male without trace of spur. Fig. 12, 13 alastor Skinner

12 mm. or less. Moderately stout species 30

WERNER: NORTH AMERICAN EPICAUTA 429

30. (29) Ground color of head and pronotum black, of elytra tan. Elytra
blackish across base, where normally covered by the base of the pro-
notum. Fig. 15 insignis Horn

Ground color uniformly tan to brown, with pronotum slightly
clouded. Elytra not darker across base. Mesotarsi of male much swollen,
denuded crassitarsis Maydell

31. (19) Pronotum glabrous. Rest of body densely clothed with cinereous to
ferrugineous pubescence heterodera Horn 9

Pronotum not glabrous 32

32. (31) With a posterior median black spot on at least the third to fifth
abdominal sternites 33

No spot on midline of abdominal sternites 38

33. (32) Pubescence on pronotum rufous, on rest of body cinereous.

wheeleri Horn
Pubescence cinereous to luteous over all of body 34

34. (33) An impression on the inner margin of the eye 35

No impressions on head near the eyes 36

35. (34) With a rounded pit on the inner margin of the eye. Pubescence
luteous rileyi Horn

With a shallow impression on inner margin of the eye and with a
pair of prominent occipital callosities. Pubescence cinereous.

caviceps Horn

36. (34) Pronotum with a black spot at the anterior angles, often one at the
posterior angles. Pubescence luteous diversipubescens Maydell

No spots at angles of pronotum 37

37. (36) Brown, with cinereous pubescence. Abdominal sternites broadly
black-pubescent apically. Scutellar and humeral spots of elytra con-
nected by a dark area across the base nigritarsis (Le Conte)

Black, with dense cinereous pubescence. Spots on abdominal ster-
nites rounded, small. Scutellar and humeral spots not connected.

aspera Werner

38. (32) Pubescence orange to rufous. Each elytron with two large black
spots stuarti Le Conte

Not so marked 39

39. (38) Black. Pubescence black over body, white to cinereous on the
suture and margins of the elytra 40

Elytra not black with pale pubescence on the suture and margins .44

40. (39) Eyes strongly slanted and pointed inwardly straba Horn var.

foxi Van Dyke
Eyes rounded inwardly 41

41. (40) Less than 10 mm. Head and pronotum shiny. Pale pubescence long
and moderately sparse. Some long pale pubescence on the anterior legs.
Male with middle and posterior femora curved and flattened behind,
margined with long pale pubescence alphonsii Horn

Over 10 mm. Head and pronotum not shiny. Legs not as in
alphonsii 42

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42. (41) Elytra narrowly and sharply margined with white pubescence.
Pale pubescence below long and silky. Antennae slender. Southwestern
mountains raidosana Fall

Elytra more broadly and less sharply margined. Pale pubescence
below short 43

43. (42) First segment of antennae swollen, especially in the male. Inter-
mediate segments thickened, especially in male. Posterior tibial spurs
slender, spiniform. Fig. 3, 4 cinerea (Forst.)

First segment of antennae not swollen. Intermediate segments,
longer, only slightly thickened. Posterior tibial spurs broadened. Fig. 5
solani sp. n.

44. (39) With uniform black pubescence, at least above 45

Pubescence mainly cinereous to luteous 60

45. (44) Outer spur of posterior tibiae somewhat broadened, never spiniform
or sticklike 46

Outer spur of posterior tibiae spiniform or sticklike 49

46. (45) Elytra somewhat bulbous. Pronotum with a V-shaped notch in the
middle of the base. Scutellum very narrow. Wings absent, .conferta (Say)

Not so 47

47. (46) Inner spur of anterior tibiae noticeably longer and stouter than the
outer. Antennae slender funebris Horn

Inner spur of anterior tibiae not stouter than the outer 48

48. (47) Visible portion of scutellum very small. Second segment of antennae
moderately slender. Antennae tapering toward apex. Fig 14.

pensylvanica (DeG.)

Visible portion of scutellum normal in size. Second segment of

antennae fairly stout, with longer pubescence. Antennae not tapering

noticeably toward apex. Head black or red. Fig. 18 atrata (Fab.)

49. (45) Eyes produced obliquely inward in an acute angle straba Horn

Eyes rounded inwardly 50

50. (49) Antennae flattened, strongly ensiform. Entirely black.

ensiformis Werner
Antennae not strongly ensiform 51

51. (50) Pubescence, at least in part, erect, long and silky 52

Pubescence short, recumbent, sometimes very sparse 57

52. (51) Some of pubescence on femora white californica Werner

All of pubescence on legs, except for the characteristic cinereous
patch on the anterior femora, black 53

53. (52) Antennae very slender apically fallax Horn

Antennae almost uniform in thickness or thickened apically, or
slightly more slender apically 54

54. (53) Tarsal claws curved from base. Head and pronotum shiny, with
scattered deep punctures barberi Werner

Tarsal claws straight for the basal three-fourths 55

WERNER: NORTH AMERICAN EPICAUTA 431

55. (54) Antennae more slender toward apex than at the middle. Fig. 23.

oblita (Le Conte)

Antennae slightly thickened apically, with the apical segments

distinctly longer than broad 56

56. (55) Tarsi of normal thickness. Long pubescence sparse to very dense.
California to North Dakota. Fig. 22 puncticollis (Mann.)

Tarsi very slender. With only a few scattered long hairs. Kansas.

kansana Werner

57. (51) Pubescence almost absent above, invisible macroscopically 58

Pubescence visible above, quite dense 59

58. (57) Elytra short, truncate, leaving several abdominal tergites exposed.
Shiny, with scattered large punctures parva (Hald.)

Elytra entire. Punctures small to medium .... piceiventris Maydell

59. (57) Small to medium in size. First segment of antennae stout. Antennae
thickened toward middle. Lake Superior to Hudson's Bay. Fig. 4, 5.

fissilabris (Le Conte)

Large. First segment of antennae not stout. Antennae not thickened

toward middle. Texas and Great Plains corvina (Le Conte)

60. (44) With a tubercle on middle of pronotum excavatifrons Maydell

No tubercle on middle of pronotum 61

61. (60) Tibial spurs all heavy, curved, blunt calcarata Werner

Tibial spurs of at least anterior and middle legs slender, spiniform.62

62. (61) Body color black, legs luteous. Pubescence sparse 63

Not so 64

63. (62) Labrum not deeply excised. Mandibles normal. Antennae ensiform
in male pedalis Le Conte

Labrum excised. Mandibles long and slender balli sp. n.

64. (62) With black markings at base of elytra . .• 65

No black markings at base of elytra 72

65. (64) With dark humeral and scutellar spot 66

No scutellar and humeral spot. Black across base 68

66. (65) Spots connected across base of elytra. Dark. Abdominal sternites
with black pubescence across apices nigritarsis Le Conte

Spots not connected across base of elytra. No dark markings below.

67. (66) Under 12 mm. Antennae very slender. Margins of elytra usually
with denser pubescence. Fig. 17 ingrata Fall

Over 15 mm. Antennae moderately stout, with first segment
elongate, cylindrical and curved in male. Fig. 28, 29.

longicollis (Le Conte)

68. (65) Head and pronotum black. Elytra tan. Fig. 15 insignis Horn

Concolorous 69

69. (68) Pubescence brownish. Pad of first segment of male anterior tarsi
broad. Fig. 6, 7 '. brunnea Werner

Pubescence cinereous 70

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70. (69) Outer spur of posterior tibiae broadened. Antennae slender, the
first segment not enlarged. Fig. 5 solani sp. n.

Outer spur of posterior tibiae slender 71

71. (70) First segment of antennae thickened. Intermediate segments
heavy. See Fig. 3, 4 cinerea (Forst.)

First segment of antennae not greatly thickened. Intermediate
segments only moderately thickened. Fig. 1,2 floridensis Werner

72. (64) Labrum bifid. Mandibles long and slender miwfh'm(Hom)

Labrum not deeply excised. Mandibles barely exceeding labrum .73

73. (72) Antennae tapered apically, becoming slender 74

Antennae not tapered apically 77

74. (73) Pubescence cinereous, sparse. Head notched on the occiput.

impressifrons Van Dyke
Pubescence luteous to tannish-cinereous. Head not deeply notched.

75. (74) Moderately stout species. Pronotum quadrate. Pubescence dense,
luteous. A weak ridge from inner margin of the eyes obliquely backward
to near the occiput rehni Maydell

Fairly slender to slender. No ridges on head 76

76. (75) Head subquadrate uniforma Werner

Head narrowly triangular alpina Werner

77. (73) A pair of denuded callosities on pronotum callosa Le Conte

No denuded callosities on pronotum 78

78. (77) Body brown. Legs paler ferrugineous batesii Horn

Body and legs of same color 79

79. (78) Apical antennal segments at least as broad as long. Fig. 20;

fortis Werner
Apical antennalsegments longer than broad 80

80. (79) Pronotum flat, with very short but dense pubescence.

heterodera Horn c?
Pronotum at least partly bulged 81

81. (80) Outer posterior tibial spur broad 82

Outer posterior tibial spur slender, spiniform or sticklike 83

82. (81) Third segment of antennae as long as first. Fig. 19.

ferruginea (Say)
Third segment of antennae almost as long as first two. Fig. 18.

a form of atrata (Fab.)

83. (81) Median suture of pronotum deep, shiny 84

Median suture of pronotum shallow and indistinct 85

84. (83) Pubescence only partly concealing surface pruinosa Le Conte

Pubescence almost completely concealing surface. Ammerita Walker

85. (83) Pronotum large and bulged. Pubescence cinereous. Fig. 21.

sericans Le Conte
Pronotum moderate in size. Pubescence ferrugineous . strigosa (Gyll.)

87.

(86)

88.

(87)

89.

(88)

90.

(89)

WERNER: NORTH AMERICAN EPICAUTA 433

86. (1) Second segment of antennae distinctly longer than third 87

Second segment of antennae equal to third or slightly shorter . . 108
Basal two segments of antennae not markedly paler than rest . . 88
Basal two segments of antennae luteous, rest brown or black. .107

Uniformly colored above 89

With stripes or numerous denuded spots 105

Body color black or very dark brown. Elytra not paler 90

Body color luteous to brown 102

First segment of antennae attaining occiput ( d1 <?) 91

First segment of antennae reaching beyond eye in some but not
coming near the occiput 96

91 . (90) One spur on anterior tibiae 92

Two spurs on anterior tibiae 94

92. (91) First segment of antennae much curved, S-shaped. Fig. 45.

torsa (Le Conte) d"
First segment of antennae straight 93

93. (92) Second segment of antennae as long as first. Palpi dark

excors (Fall) c?

Second segment three-fourths as long as first. Palpi luteous. Fig.

34 tenuis (Le Conte) d"

94. (91) Second segment distinctly shorter than the following three. Fig.
47 fabricii (Le Conte) d1

Second segment as long as next three or just perceptibly shorter.
Fig. 49 95

95. (94) Pubescence pale, giving color grayish tinge . .murina (Le Conte) d1

Pubescence dark, very sparse, not affecting color . . subglabra (Fall) d1

96. (90) Second segment of antennae equal to first or slightly longer 97

Second segment of antennae distinctly shorter than first 98

97. (96) Palpi luteous tenuis (Le Conte) 9

Palpi dark excors (Fall) 9

98. (96) With a fringe of long hairs on the middle and posterior femora.
First segment of antennae reaching hind margin of eye. Second half as
long as first. Fig. 51 flavocinerea (Blatchley) d"

Pubescence on femora short. First segment of antennae distinctly
short of hind margin of eye. Second distinctly more than half as long
as first 99

99. (98) Pubescence short, dark, sparse, not affecting color. Fig. 50.

subglabra (Fall) 9

Pubescence cinereous, visible macroscopically 100

100. (99) Pubescence quite dense, concealing most of surface. Second seg-
ment of antennae about equal to third. Fig. 48 .fabricii (Le Conte) 9 -
Pubescence sparse, causing a gray color. Second segment of antennae
longer than third 101

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101. (100) Second segment of antennae four-fifths as long as first. Third and
following two-thirds as long as second. Fig. 50. . . .murina (Le Conte) 9

Second segment of antennae three-fifths as long as first. Third and
following three-fourths as long as second. Fig. 46. torsa (Le Conte) 9

102. (89) Anterior tibiae with two spurs. First segment of anterior tarsi
longer than second languida (Horn) d"

Anterior tibiae with one spur. First segment of anterior tarsi
shorter than second, contorted 103

103. (102) Elytra distinctly paler than head and pronotum. First two seg-
ments of antennae equal to rest. Fig. 37 polingi Werner d1

Elytra not paler than rest of body 104

104. (103) Entirely dusky brown. Pubescence denser on margins of elytra,
forming an indistinct pale border. First two segments of antennae
longer than rest. Fig. 38 licbecki Werner cf

Entirely pale tan. First two segments of antennae shorter than
rest. Fg. 39 arizonica Werner d"

105. (88) With denuded spots maculifera (May dell) d"

With vittae on elytra 106

106. (105) Four fine dark lines on each elytron. Fig. M .tenuilineata (Horn) d"

Distinct pale line on each elytron. Fig. 44 . . . .-purpurea (Horn) d1
Two wide black stripes on each elytron. Fig. 24.

atrivittata (Le Conte) d1
An indistinct pale line on each elytron parkeri Werner d"

107. (87) Abdominal sternites with black marks. Fig. 25. . albida (Say) d1

Abdominal sternites without black marks. Fig.J27.

texana Werner a*

108. (86) With numerous denuded spots maculifera (Maydell) 9

Without denuded spots 109

109. (108) With vittae on the elytra 110

Elytra without vittae 112

110. (109) Fine black lines on pale background Ill

Light line on black elytra purpurea (Horn) 9

Two heavy black vittae atrivittata (Le Conte) 9

An indistinct pale line on each elytron parkeri Werner 9

111. (110) Four lines on each elytron, sometimes with suffusions from them.

Base of antennae not paler sublineata (Le Conte)

Three lines on each elytron. Basal two segments of antennae
luteous. Fig. 27 ... oUnda (Say) 9

112. (109) Ground color black U3

Ground color at least in part brown to luteous 121

113. (112) Elytra entirely black-pubescent. Hind margin of pronotum with
white pubescence. Fig. 30, 31 segmenta (Say)

Elytra not entirely black-pubescent 114

114. (113) Black or dark gray, with paler margins on the elytra 115

Margins of elytra not paler 117

WERNER: NORTH AMERICAN EPICAUTA 435

115. (114) Apices of abdominal sternites white sublineata (Le Conte)

Abdominal sternites uniform in color 116

116. (115) Head narrowly triangular. An indistinct paler line down middle
of each elytron, usually parkeri Werner

Head subquadrate. A stouter species. No pale line on elytra. Fig.
43 gissleri{}iom)

117. (114) Antennae with first two segments luteous 118

Antennae uniform in color 119

118. (117) Abdominal sternites with black marks albida (Say) 9

Abdominal sternites without black marks texana Werner 9

119. (117) Second segment of antennae as long as third. Fig. 32, 33.

immaculate/, (Say)
Second segment of antennae distinctly shorter than third 120

120. (119) Pubescence very dense. Most specimens over 15 mm. Fig. 29.

longicollis (Le Conte ) 9

Pubescence moderately dense. Surface visible. Not over 13 mm.

Fig. 52 flavocinerea (Blatchley) 9

121. (112) Head and pronotum pale. Elytra black, with sparse cinereous
pubescence 122

Elytra brown or ferrugineous 123

122. (121) Second segment of antennae slightly shorter than third. Fig. 35.

tenella, (Le Conte)
Second segment of antennae as long as third . . merkeliana (Horn)

123. (121) Elytra paler than head and pronotum, which are black or very
dark brown. Fig. 40 polingi Werner 9

Elytra not noticeably paler than rest of body 124

124. (123) Brown. Margins of elytra paler, due to denser pubescence.

liebecki Werner 9
Entirely luteous to ferrugineous 125

125. (124) Antennae noticeably moniliform, ferrugineous. Fig. 42.

ochrea (Le Conte)
Antennae not at all moniliform, black 126

126. (125) Some cinerous pubescence on first two antennal segments.

arizonica Werner 9

Basal antennal segments shiny, with no cinereous pubescence.

Fig. 36 lauta (Horn)

Epicauta abadona Skinner

Epicauta abadona Skinner, 1904, Entomological News 15:217.
Epicauta mutchleri Maydell, 1934, Trans. Am. Ent. Soc. 60:331.
Epicauta abandona Maydell, 1934, loc. cit (lapsus calami).

Length : 9 to 16 mm. Elongate, slender. Tan, rather densely clothed
with pale tan pubescence, except for a pair of narrow longitudinal

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brown or black lines on the pronotum and two incomplete and a third
much less distinct black vittae on the elytra. Posterior tibiae of the
male with very broad spoon-shaped spurs. Maydell's mutchleri is
based on this sexual difference.

Head subquadrate, partly denuded and shiny in the male. Median
suture only feebly impressed. Eyes prominent, moderately broad.
Antennae fairly long, three times as long as an anterior tibia, slender,
somewhat flattened. First segment reaching to the middle of the eye,
fairly stout ; second two-thirds as long as the first, narrow ; third about
two and one-third times as long as the second; fourth and following as
long as the first and gradually decreasing in thickness. Pronotum one-
third longer than broad, campanuliform, with a fairly broad but
shallow median longitudinal impression. Suture very faintly impressed,
inconspicuous. Elytra with a narrow stripe arising near the scutellum
and extending to near the apex, another of equal width from the
humerus almost to the apex, and a third, much narrower, just outside
the latter. Uniformly tannish below. Anterior tibiae of the male
with a single heavy, incurved spur. Hind tibial spurs of the male
broad, flat, spoon-shaped. Hind spurs of the female also flattened
but not conspicuously broadened. First segment of the anterior tarsi
of the male slightly elongated but not otherwise modified.
Type: Holotype c? A.N.S.P. No. 8068, examined.
Type locality: Phoenix, Arizona.
of mutchleri Maydell

Type: Holotype d\ A.M.N.H., examined.
Type locality: Phoenix, Arizona.
Additional localities: Arizona: Tucson, Avondale Ranch — Agua

Fria R. Mexico: Mazatlan — Sinaloa. Occurs in August.

Epicauta alastor Skinner
Fig. 12, 13

Epicauta alastor Skinner, 1904, Entomological News 15:217.

Length: 8 to 14 mm. Elongate, slender. Light to dark brown,
clothed with tannish-cinereous pubescence, which is sparse but for
a. narrow stripe on the elytra from humerus to apex, and the mar-
gins, which are more densely pubescent. The form of pubescence
of insignis Horn is very similar but that species is shorter and
stouter, has the head and pronotum black or dark brown and has

WERNER: NORTH AMERICAN EPICAUTA

437

black pubescence at the base of the elytra. Crassitarsis Maydell
also has similar pubescence but it also is much shorter and stouter.
Head suboval. Suture fine. Eyes very large, conspicuous, quite
broad. Antennae, especially in the male, subensiform; three and
one-half times as long as an anterior tibia. First segment moder-
ately stout, reaching to the middle of the eye; second half as long
as the first, more slender; third and following flattened, the third
two and one-half times as long as the second; fourth and following
equal in length, four-fifths as long as the third. The fourth seg-
ment is the broadest, the rest gradually decreasing in breadth
toward the apex. Pronotum one-third longer than broad, campan-
uliform, with a distinct suture and basal impressed line. Below,
uniformly colored as above. Anterior tibiae of the male without
any trace of a spur. Outer posterior tibial spur flattened, stick-
like; the inner slender, spiniform. First segment of the anterior
tarsi of the male short, contorted, but not denuded.
Type: Holotype <? , A.N.S.P. No. 8084, examined.
Type locality: Florence, Arizona.

Additional localities: Colorado: Ft. Collins (one specimen in
Colo. State College). Texas: Presidio. Arizona: Yuma,
Florence Jet., Avondale Ranch — Agua Fria R., Aug. 7, 17,
Texas Pass — Dragoon Mts., July 21, 17, Phoenix. California:
Imperial Co., El Centro, Yuma.

At first sight, there seems to be a difference between the Arizona
and California specimens in that the latter are darker. But Ari-
zona specimens are sometimes dark and California specimens light.

Epicauta insignis Horn
Fig. 15
Epicauta insignis Horn, 1885, Trans. Am. Ent. Soc. 12:110.

Length: 7 to 10 mm. Rather stout, black, elytra and legs ferru-
gineous. Entirely clothed with rather sparse tannish pubescence,
which is a little denser on an indistinct median line and the mar-
gins of the elytra. Crassitarsis Maydell has a ferrugineous pro-
notum and a very distinct line of denser pubescence on the elytra.

Head suboval, sparsely and finely punctured, shiny. Suture dis-
tinctly impressed. Antennal calluses denuded, smooth, shiny.
Eyes prominent, transverse, moderately narrow. Antennae black,
two and three-fourths times as long as an anterior tibia in the male,

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two and one-fourth in the female, ensiform in the male, subensiform
in the female. First segment reaching two-thirds across the eye;
second short, small, one-third as long as the first; third as long as
the first, flattened, the broadest segment; rest a little shorter than
the third, gradually narrower toward the tip, flattened. Last seg-
ment of the maxillary palpi broader in the male than in the female.
Pronotum subquadrate, one-fourth longer than broad. Suture
distinct but not deeply impressed. Surface as on the head. Black
below; legs ferrugineous. The tips of the femora are a dark brown.
Anterior tibiae of the male with a single short, incurved spur.
Posterior tibial spurs flattened but not very broad, spiniform.
Type: Lectotype d\ new designation, A.N.S.P. No. 8097, exam-
ined.
Type locality: Arizona.

Additional localities: Texas: "Tex." - - Eddy Coll. New Mexico:
Socorro Co. Arizona: Baboquivari Mts., Nogales, Palmerlee,
Huachuca Mts.

Epicauta nigritarsis (Le Conte)

Lytta nigritarsis Le Conte, 1853, Proc. Ac. Nat. Sci. Phila. 6:340.
Epicauta nigritarsis, Horn, 1873, Proc. Am. Phil. Soc. 13:100.

Length: 7 to 13 mm. Tan above, with rather dense cinereous
pubescence. Brown humeral and scutellar spots on the elytra,
narrowly connected across the base where not normally visible.
Below, mainly black, with cinereous pubescence. Legs tan, with
apices of the segments brown. Easily distinguished from the other
tan Epicautae because the head and pronotum are of the same
color as the elytra.

Head subquadrate. Suture distinctly impressed, supplemented
by a narrow denuded line. Antennal calluses not raised, small,
denuded, shiny. Eyes rather small, narrow, transverse. The
antennae are quite slender, reaching the basal third of the elytra,
two and one-fourth times as long as an anterior tibia. First seg-
ment somewhat thickened, reaching two-thirds across the eye;
second slender, half as long as the first; third slightly longer than
the first two, slender; fourth and following two-thirds as long as
the third, decreasing slightly in thickness and just perceptibly in
length. Pronotum quadrate, slightly broader than long. Median
suture distinct on the middle of the disc, less distinct toward the
base and apex. Basal impressed line distinct but covered with

WERNER: NORTH AMERICAN EPICAUTA 439

pubescence. Elytra a little broader behind than at the humerus.
Margins sometimes slightly paler from the denser pubescence.
Posterior border of the third to fifth abdominal sternites with
black pubescence; sixth mostly black; anterior lateral margin of
the fourth and fifth with a black spot. These black markings have
heretofore passed unnoticed because of the dark color of the under-
side. Anterior tibiae of the male with a single short, usually in-
wardly directed spur (not two as stated by Horn, 1873). Posterior
tibiae with two slender, sticklike spurs, the inner long, the outer
shorter.

Type : Lectotype 9 , new designation, M. C. Z. No. 4998, examined.
Type locality: Mexican Boundary, Texas.

Additional localities: Texas: New Braunfels, Brownwood,
Brownsville, Big Springs, Knipps, Del Rio, Harris Co., Dallas,
Uvalde. New Mexico: Tucumcari. Occurs in June and July.
Apparently abundant when it occurs.

Epicauta crassitarsis Maydell
Epicauta crassitarsis Maydell, 1935, Journ. Wash. Acad. Sci. 25:72.

Length: 7 to 9 mm. Light to dark ferrugineous, clothed with
tannish-cinereous pubescence, moderately densely but for the
more densely clothed margins of the elytra. Short and rather
stout. The pronotum and the elytra are of the same color.

Head suboval, black on the front and vertex, ferrugineous be-
hind, rather coarsely but shallowly punctured. Suture distinctly
impressed, complete. Antennal calluses not denuded except for a
very small area at the base of the antennae. Eyes prominent,
transverse, moderately broad. Antennae moderately long, slightly
flattened, tapering, reaching to the basal eighth of the elytra, two
and one-half times as long as an anterior tibia, black but for the
first and second segments. Middle thicker in the male than in the
female. First segment short, rather stout, reaching to the middle
of the eye; second small, two-fifths as long as the first; third one-
fourth longer than the first; fourth the thickest segment, the fourth
and following approximately as long as the first; the segments
beyond the fourth decrease noticeably in thickness in the male,
less so in the female. Pronotum subquadrate, one-fourth longer
than broad; sides parallel for the basal four-fifths, then converging
abruptly. Ferrugineous but for an obscure interrupted darker band

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near the apex. Median suture and basal impressed line distinct,
with a shallow median basal impression. Scutellum rather small.
Elytra denuded across the base, where covered by the pronotum.
Abdominal sternites darker toward the apex; or mainly darker
below. Anterior tibiae of the male with a single, incurved, spini-
form spur. Middle tarsi of the male strangely modified. Glabrous
but for the pads. First segment stout, inflated, thicker by one-
fourth than the apex of the tibia. Rest much smaller than the first,
decreasing in thickness, the last not at all inflated. Posterior tibial
spurs broad, flattened, slightly pointed.
Type: Holotype c? , U.S.N.M., examined.

Type locality: Tempe, Arizona, Sept. 7, 1933, K. B. McKinney.
Additional records: Tempe, Arizona, Sept. 20, 1934, on alfalfa
K. B. McKinney (U.S.N.M.).

Epicauta pedalis Le Conte

Epicauta pedalis Le Conte, 1866, Smiths. Misc. Coll. 6 no. 167, 2nd ed.: 157.
Horn, 1873, Proc. Am. Phil. Soc. 13:99.

Length: 6 to 9 mm. Black, rather sparsely clothed with cinere-
ous pubescence. Legs bright ferrugineous. More slender than the
others in this group.

Head triangular, with scattered shallow punctures. Suture finely
but distinctly impressed. Antenna! calluses small, denuded, shiny.
Eyes moderately prominent, transverse, fairly narrow. Antennae
of male ensiform, of female quite slender, reaching two-thirds the
length of the elytra in the male, to the basal third in the female.
First segment rather stout, reaching three-fourths across the eye;
second short, half as long as the first; third almost as long as the
first two together; rest subequal, two-fifths as long as the third,
decreasing in thickness from the fourth. Pronotum one-fourth
longer than broad, subcampanuliform. Median suture distinctly,
deeply impressed. Basal impressed line also distinct but not as
deep. Elytra without markings. Below, of the same color as
above. Anterior tibiae of the male with a single short, incurved,
spiniform spur. Anterior tarsi of male not modified. Posterior
tibial spurs slender, sticklike.

Type: Lectotype 9 , new designation, M. C. Z. No. 4997, examined.
Type locality: Lower California.
Additional localities: Arizona: Tucson.

WERNER: NORTH AMERICAN EPICAUTA 441

Epicauta maculata Say

Lytta maculata Say, 1823, Journ. Ac. Nat. Sci. Phila. 3:298. Le Conte, 1853,

Proc. Ac. Nat. Sci. Phila. 6:340.
Lytta conspersa Le Conte, 1853, loc. cit. , nee Curtis, 1845.
Epicauta maculata, Horn, 1873, Proc. Am. Phil. Soc. 13:100.

Length: 6 to 12 mm. Black, quite densely clothed with olive-
cinereous to cinereous pubescence. With denuded spots of varying
extent, which do not usually run together. They are generally dis-
tinct on the elytra but may be small on the head and pronotum.
Male with the maxillary palpi large, flattened, the labial palpi
small.

Head subquadrate. Suture distinctly impressed. Antennal cal-
luses small, denuded, shiny. Eyes moderately prominent, quite
small, transverse, fairly narrow. Maxillary palpi broad and flat
in the male, the last segment being almost orbicular in outline,
shiny, with very short scattered brown pubescence below. Labial
palpi small. Antennae about twice as long as an anterior tibia,
quite slender, tapering slightly toward the tip. First segment
moderately stout, reaching half-way across the eye; second narrow,
two-thirds as long as the first; third as long as the first two together;
fourth and following four-fifths as long as the third, decreasing
gradually in thickness. Pronotum quadrate, rather rotund.
Median suture finely impressed, supplemented on the middle of the
disc by a narrow denuded line. Basal impressed line distinct.
Anterior tibiae of the male with a single short, stout, incurved
spur. First segment of the anterior tarsi not modified in the male,
with the pad extending almost its full length. Posterior tibial
spurs slender, spiniform, the inner longer.

One male from Oklahoma City, Okla., in the U.S.N.M. has two
spurs on the anterior tibjae, as in the female. Several cases of this
sort have occurred in other species also, but they are of such rare
occurrence that listing them in the key to species would only make
the keys longer, adding little to their usefulness.
Type: Say's type lost. Neotype cT, and neoparatypes in the

M. C. Z.
Type locality: "Inhabits Missouri and Arkansas . . . not uncom-
mon near the Rocky Mountains." Neotype locality: Indianola,
Nebraska — A. P. Morse,
of conspersa. Le Conte

Type: Lectotype d\ new designation, M. C. Z. No. 5000, examined.
Type locality: Missouri Territory.

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Additional localities: Common from the Rockies to western Iowa
and Missouri, and from North Dakota to Wickenburg and
Palmerlee, Arizona and Texas. Probably extends some distance
into Mexico. Occurs in June and July.

Epicauta nogales Werner
Epicauta nogales Werner, 1944, Psyche 50:65.

Length: 7 to 9 mm. Close to maculata (Say). The male has the
labial palpi flattened, broad but the last segment is not as broad as
in the male of maculata. The elytra are short and broad, leaving
the tips of the abdomen broadly exposed. The elytra are more
separately rounded than in any of the other species in the group.
The spots on the body and elytra are small. Anterior tarsi of the
male not modified, the pad of the first segment extending almost
its full length.

Type: Holotype d\ M. C. Z. No. 26085, examined.
Type locality: Nogales, Arizona.
Additional localities: Arizona: Tucson. Occurs from July to

September.

Epicauta normalis Werner

Epicauta normalis Werner, 1944, Psyche 50:65.

Length: 6 to 12 mm. This species replaces maculata in the Rocky
Mountains and westward to the Pacific, except in Arizona, where
nogales and phoenix are found. The species is as variable as
maculata as to size of spots but differs from that species in having
the maxillary palpi small in the male, not shiny and by having the
first segment of the anterior tarsi of the male narrow, slightly
outcurved, with the pad absent from at least the basal three-
fourths. More of the specimens have cinereous pubescence than in
maculata but a few have olive-cinereous.
Type: Holotype d\ M. C. Z. No. 26080, examined.
Type locality: Bridgeport, California.

Additional localities: California: Shasta Valley, Eagle ville.
Oregon: Echo, Burns. Washington: Yakima Valley. Nevada:
Paymaster Canon - - Lone Mt. - - Esmeralda Co., Reno. Idaho :
Ruhl. Wyoming: Green River City, Laramie. Montana:
Helena. Utah: Fillmore, American Fork, Duchesne Co., Jor-
dan R., Provo, Indianola, Ogden, Colorado: Manitou, La
Veta, Colorado Springs, Windsor, Ft. Collins. Arizona:
Chiricahua Mts., Huachuca Mts.

WERNER: NORTH AMERICAN EPICAUTA 443

Epicauta phoenix Werner
Epicauta phoenix Werner, 1944, Psyche 50:66.

Length: 7 to 12 mm. Close to metadata (Say). Easily distin-
guished from the others of the group by the more convex form of
the elytra, especially toward the apex, the slender antennae and
by the small palpi of the male. The antennae tend to be shorter
than in the others. The spots on the elytra are moderately large,
but distinct from each other. The spots on the head and pronotum
are small and indistinct. Below, they are small, but larger than on
the head and pronotum.

Type: Holotype d\ M. C. Z. No. 26070, examined.
Type locality: Phoenix, Arizona. Apparently fairly abundant

at the type locality.

Epicauta pardalis Le Conte

Epicauta pardalis Le Conte, 1866, Smith. Misc. Coll. 6 no. 167,2nd ed.: 157;
Horn, 1873, Proc. Am. Phil. Soc. 13:99. Champion, 1892, Biol. Cent. -Am.
Coleop. 4 pt 2:414.

Length: 7 to 12 mm. Black, shiny, quite densely clothed with
cinereous pubescence except for fairly large round denuded spots,
which are in part confluent. These spots are very smooth and shiny
with only a few very small punctures. Differs from andersoni
Werner by its thicker antennae and by the small palpi of the male.

Head triangular. Suture distinctly impressed. Antennal cal-
luses small, denuded, shiny. Eyes rather small, transverse, quite
narrow. Neither pair of palpi enlarged in the male. Antennae
quite slender but not as slender as in andersoni, reaching to the
basal eighth of the elytra, twice as long as an anterior tibia. First
segment rather stout, reaching to the middle of the eye ; second half
as long as the first, small; third just shorter than the first; first
three segments with some cinereous pubescence behind; fourth to
last three-fifths as long as third, gradually tapering from the fourth.
Pronotum as broad as long. Sides parallel for the basal three-
fourths, then converging abruptly. Anterior tibiae of male with a
single short, incurved spur. Male anterior tarsi with the first seg-
ment a little broader than in the female but not otherwise modified.
Inner spur of posterior tibiae quite long, spiniform; outer shorter,
flattened, sticklike.

Type: Lectotype 9 , new designation, M. C. Z. No. 4999, examined.
Type locality: Valley of the Gila.

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Additional localities: Arizona: Ft. Grant, Tucson. New Mexico:
Silver City, Faywood. Texas: Marfa. Coahuila: Saltillo.
Durango: Descubridora.

Epicauta andersoni Werner
Epicauta andersoni Werner, 1944, Psyche 50:66.

Length: 7 to 11 mm. Practically identical with pardalis Lee, but
the differences are constant. The antennae are more slender than
in that species, but both must be seen at the same time when this
character is used. The male palpi provide the main difference.
Both pairs are enlarged, flattened, shiny below, with scattered
punctures. The last segment of the labial palpi is almost circular
in outline.

Type: Holotype'cf, U.S.N.M., examined.
Type locality: Gallo Springs, New Mexico.
Additional localities: Arizona: Flagstaff. Texas: Childress.

Kansas: "Ks" Dietz Coll. M. C. Z. Taken in June and August.

Not common.

Epicauta ventralis sp. n.

Length: 9 to 11 mm. Another of the maculata species. Similar
to magnomaculata Martin in that the male has a brown mark
across the abdominal sternites and a small anterior spot on the
edge of the sternites. Differs from that species by its smaller size,
even punctures and brown appendages. Black, elytra dark brown
or black.

Head subquadrate, densely punctured, with a few small denuded
spots. Median suture distinct. Antennal calluses small, denuded.
Eyes fairly small, narrow, transverse. Lower part of front, clypeus
and labrum dark brown. Palpi light brown, not enlarged in the
male. Antennae as in maculata but with the first segment stouter
and the pubescence on the first two segments more erect. Pubes-
cence on first segment cinereous behind. Antennae reaching to
basal third of elytra, two and one-eighth times as long as an
anterior tibia. First segment moderately stout, reaching three-
fourths across the eye; second slender, half as long as first; third
as long as first but slender; fourth and following two-thirds as long
as third, decreasing just perceptibly in thickness. Pronotum
quadrate, with a well-marked median suture. Basal impressed
line obscured by pubescence. Spots small. Elytra with spots

WERNER: NORTH AMERICAN EPICAUTA 445

moderately large, uniform. Second to fifth abdominal sternites
with a transverse brown mark. Apex of sixth brown. Legs longer
than in the other maculate species, except for magnomaculata.
Anterior tibiae of male with a short incurved spiniform spur. Male
anterior tarsi as in normalis. Posterior tibial spurs longer and
more slender than in the other maculate species.
Type: Holotype d\ Walsenburg, Colorado Aug. 4, 1934, J. F.

Schuett (Field Mus.). Allotype 9, eutopotypical (F.M.N.H.).

Paratypes: 4 eutopotypical (F.M.N.H.), 2 eutopotypical

(M. C. Z. No. 26633).

Epicauta magnomaculata Martin

Epicauta magnomaculata Martin, 1932, Pan Pacific Ent. 8:169.

Length: 14 to 16 mm. Black, densely clothed with appressed
pale cinereous pubescence except for a number of denuded spots,
which are large on the elytra, small and scattered on the body.
Easily distinguished from the other maculate species by the
elongate, more slender form and by the possession of a pair of
small denuded callosities on the pronotum. The male has large
black median abdominal spots.

Head subtriangular. Suture distinct, augmented by a narrow
smooth denuded area. Antennal calluses small, denuded, shiny.
Surface of head densely punctured, with the intervals smooth and
shiny. Eyes fairly small, transverse, excavated, unusually squarely
cut on the internal margin. Antennae of quite a distinctive form.
They are short, rather stout at base, and tapering to the tip; twice
as long as an anterior tibia and reaching just beyond the base of
the elytra. First segment moderately stout but not thickened
greatly at the tip, reaching three-fourths across the eye; second
two-fifths as long, quite stout; third twice as long as second; fourth
three-fifths as long as the third; fifth to last slightly longer than
fourth, decreasing gradually in thickness. The first four segments
have some cinereous pubescence behind. Pronotum subcampanuli-
form, one-fourth longer than broad. Sides almost parallel for the
basal four-fifths. Median suture distinct from the base three-
fourths to the apex, supplemented in the middle of the disc by a
narrow denuded area. Basal impressed line distinct. Scutellum
cinereous-pubescent. Base of elytra black. Spots on the elytra
varying in size but in all specimens examined larger and more
irregular in size than in the other immaculate species. Abdominal

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sternites with an anterior lateral black mark. Male with posterior
median black marks. Anterior tibiae of male with a single short,
incurved spur. Posterior tibial spurs flattened, sticklike, almost
equal. Tips of femora, tips and inner side of tibiae, tarsi black.
Type: in Martin Coll.

Type locality: Ballart, Inyo Co., California. Panamint Valley.
Additional localities: Stove Pipe Well, Death Valley, California.

Epicauta corvina (Le Conte)

Lytta corvina Le Conte, 1858, Journ. Ac. Nat. Sci. Phila. (2) 4:21.
Cantharis nigerrima Duges, 1869, La Naturaleza 1:162; 1879, op. cit. 4:66.
Epicauta corvina, Horn, 1873, Proc. Am. Phil. Soc. 13:96, 102.

Length: 13 to 24 mm. A large stout black species, with rather
dense but short and closely appressed pubescence. Larger than
the other black species of our fauna and distinguished from them
by its slender posterior tibial spurs. Entire surface densely but
finely punctured.

Head subtriangular. Suture distinct, but not conspicuous. Eyes
not prominent, moderately broad, oblique. Antennae rather
slender, tapering, reaching the basal fifth of the elytra and twice
as long as an anterior tibia. First segment stout toward the apex,
reaching two-thirds across the eye; second small, half as long;
third slightly longer than first, slender; fourth and following two-
thirds as long as the third, gradually decreasing in thickness. The
third segment and beyond are very slightly flattened. Last segment
a little over two-thirds as thick as the fourth. Pronotum sub-
quadrate, stout, almost as broad as long. Median suture absent or
extremely indistinct. Basal impressed line distinct. A midbasal
impression present, distinct. There is also a pair of very indistinct
longitudinal impressions on the disc, before the middle. Exposed
portion of the scutellum smaller than normal. Elytra with four
indistinctly raised lines. Tibial spurs all spiniform, often broken
off. Male anterior tibiae not modified.

Type: Lectotype 9, new designation, M. C. Z. No. 5097, exam-
ined.
Type locality: Valley of the Gila.

Additional localities: Kansas: Ellis, Grove Co., Ft. Hayes.
Colorado: Ft. Collins. Texas: Alpine. New Mexico: Koehler.
Arizona: Flagstaff. Occurs from July to September.

WERNER : NORTH AMERICAN EPICAUTA 447

Epicauta funebris Horn
Epicauta funebris Horn, 1873, Proc. Am. Phil. Soc. 13:102.

Length: 11 to 18 mm. Large, black, sparsely clothed with short
black pubescence. Easily distinguished from corvina (Lee.) by
its smaller size, narrower form and broad posterior tibial spurs.
It could be confused with large specimens of pcnsylvanica (De. G.)
but is more slender and has the inner spur of the anterior tibiae
noticeably longer and stouter than the outer.

Head subquadrate. Surface rather densely and deeply punc-
tured, with the intervals finely granulose so that the surface
appears dull. Median suture distinct, continued deeper below the
level of the eyes. Antennal calluses only slightly elevated, denuded,
smooth. Antennae almost uniform in thickness, two and three-
fourths times as long as an anterior tibia and reaching the basal
fourth of the elytra. They are slightly stouter in the male than
in the female. First segment moderately slender, reaching three-
fourths across the eye; second slender, half as long; third almost
as long as the first two; fourth three-fourths as long as the third;
rest decreasing slightly in length and thickness. Pronotum sub-
quadrate, one-fifth longer than broad. Sides parallel for the
basal four-fifths, then converging abruptly. Median suture dis-
tinct but not deep. Basal impressed line distinct. The pronotum
is noticeably narrower than the head or elytra. Scutellum small,
as in pcnsylvanica. Anterior tibiae with two spiniform spurs in
both sexes, the inner being somewhat longer and stouter than the
outer, especially in the male. Anterior tarsi of male not modified.
Posterior tibial spurs flattened, the outer more blunt than the
inner, both moderately broad.
Type: Lectotype c? , new designation, Horn Coll. A.N.S.P. No.

8027, examined.
Type locality : Texas.
Additional localities : Kansas : Garden City, Wichita. Oklahoma :

Webster. Texas: Columbus, Victoria, Dallas. New Mexico:

Albuquerque.

Epicauta pensylvanica (De Geer)
Fig. 14
Cantharis pensylvanica De Geer, 1775, Mem. pour servir a l'hist. des Ins. 5:15.
Meloe nigra Woodhouse, 1800, Med. Repos. 3:213.
Lytta pennsylvanica, Le Conte, 1853, Proc. Ac. Nat. Sci. Phila. 6:329.
Lytta morio Le Conte, 1853, op. cit.: 447.
Epicauta pennsylvanica, Horn, 1873, Proc. Am. Phil. Soc. 13:102.

448 bulletin: museum of comparative zoology

Length: 6 to 12 mm. Dull black, sparsely clothed with very
short black pubescence, which is not ordinarily visible macroscopi-
cally. Outer posterior tibial spur broad. Wings fully developed.

Head broadly triangular. Surface rather densely and deeply
punctured, the intervals densely punctulate. Median suture fine
but distinct. Antennal calluses small, smooth, shiny. Eyes
moderately prominent, transverse, excavated. Antennae a little
more than twice as long as an anterior tibia, reaching to the basal
fifth of the elytra. First segment stout, especially in the male,
reaching half way across the eye; second small, half as long; third
as long as the first but more slender; fourth two-thirds as long as
the third; fifth and following slightly shorter than the fourth.
Segments 3-11 gradually decrease in thickness, but only slightly.
The intermediate segments are thicker in the male than in the
female. The last segment is half as thick as the third in the male,
two-thirds in the female. Pronotum quadrate, slightly broader
at the front angles. Median suture distinct, augmented by a
shallow median impression. Basal impressed line distinct, deep.
Visible portion of scutellum very small. The small scutellum serves
to distinguish pensylvanica from pure black atrata specimens
at a glance. Male anterior legs not modified. Posterior tibial
spurs flattened, pointed, the outer broader. Neither is narrow
enough to be called "sticklike."
Type : Present location unknown to author.
Type locality : Pennsylvania.
Lytta morio Le Conte

Type: Lectotype 9 , new designation, M. C. Z. No. 5098, examined.
Type locality: Texas.

Additional localities: From Maine to Florida, west to Montana
and Texas, apparently more common in the northern portion
of its range. Occurs abundantly in the fall on flowers of golden-
rod (Solidago) and other bright Compositae. Has been recorded
as damaging potatoes and beets.

Epicauta conferta (Say)

Meloe confertus Say, 1824, Journ. Acad. Nat. Sci. Phila. 3:281.

Henous techanus Haldeman, 1852, Stans. Exp. to Gr. Salt L. : 377, pi. 9 fig.

12-14.
Henous confertus, Le Conte, 1853, Proc. Ac. Nat. Sci. Phila. 6:330. Champion,

1892, Biol. Cent.-Am., Coleop. 4 pt. 2:368. Van Dyke, 1928, U. Calif.

Publ. Ent. 4:411, pi. 15, fig. 1.

WERNER: NORTH AMERICAN EPICAUTA 449

Length: 8 to 15 mm. Black, sparsely clothed with short black
pubescence, which is invisible to the unaided eye. Upper surface
densely punctured. Elytra short, leaving the tip of the abdomen
exposed, separately rounded. Wingless.

Head subquadrate. Punctures sparser below the level of the
eyes. Median suture distinct. Antennal calluses large, denuded,
smooth. Eyes moderately narrow, transverse. Antennae slightly
more than twice as long as an anterior tibia, reaching to the basal
sixth of the elytra. First segment rather stout, reaching half way
across the eye; second half as long; third as long as first two;
fourth equal to third; fifth and sixth slightly longer than fourth;
remaining slightly longer than sixth. The antennae taper grad-
ually from the third segment. Pronotum quadrate, as broad as
long. Sides converging at the apical fifth. Base somewhat exca-
vated before scutellum. Median suture distinct for a short distance
on the middle of the disc. Basal impressed line distinct. Scutellum
slender, lance-shaped. Elytra appearing inflated. They are not
connate. Poorly mounted specimens may have them overlapping
but fresh ones never do. Male anterior tibiae not modified. Inner
spur of posterior tibiae spiniform, outer flattened but not very
broad.

Type: Say's type lost. Neotype 9 and neoparatypes in M. C. Z.
Type locality: "Council Bluffs, Iowa and near the Rocky Moun-
tains." Neotype locality: Dallas, Texas.
Type of techanus Haldeman: Present location unknown. May

be lost.
Type locality: Texas, probably Fort Gates.

Additional localities: From Iowa to Arkansas, west to the Rockies.
Common.

Epicauta parva (Haldeman)

Meloe parvus Haldeman, 1852, Stans. Exp. to Gr. Salt L. : 377.

Meloe parvulus Haldeman, 1853, Proc. Ac. Nat. Sci. Phila. 6:40.

Nomaspis parvulus, Le Conte, 1866, Smiths. Misc. Coll. 6 no. 167, 2nd ed.

:156.
Henous parvulus, Van Dyke, 1928, U. Calif. Publ. Ent. 4:411, pi. 15, fig. 2.

Length: 8 to 12 mm. Black, almost glabrous above and very
sparsely pubescent below. Elytra short, not meeting on the
midline, leaving almost half of the abdomen exposed. Elytra
rugose and shiny; head and pronotum with deep punctures.

Head subtriangular, rather large, with numerous deep uneven

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punctures. Median suture fine, sometimes obliterated. Antennal
calluses moderate in size, somewhat elevated, smooth. Eyes not
prominent, fairly narrow, excavated. Antennae reaching to the
basal fourth of the elytra, one and one-half times as long as an
anterior tibia. First segment stout, reaching almost to the hind
margin of the eye in the male, two-thirds across the eye in the
female; second small, two-fifths as long as the first; third as broad
and almost as long as the first; fourth and following as short as
the second, decreasing gradually in thickness; last segment twice
as long as next to last. Segments three to five have their tips
obliquely truncated. Intermediate segments broader in the male
than in the female. Pronotum quadrate, one-fourth broader than
long. Median suture fine, indistinct. Basal impressed line distinct.
There is a broad impression on the middle of the disc. Surface
as on the head but not so deeply punctured. Scutellum small, of
normal shape. Elytra somewhat broadened behind, never meeting
on midline. First segment of anterior tarsi of male excavated
ventrally at the base, broad at the apical half. Posterior tibial
spurs moderately slender, the inner spiniform, the outer sticklike.
Type: Present location unknown.
Type locality: "Route to Santa Fe — Kearns"
Additional localities: Colorado: Flagler. Nebraska: McCook.
Kansas: Garden City. Wyoming: Cheyenne.

Epicauta fallax Horn
Epicaiita fallax Horn, 1885, Trans. Am. Ent. Soc. 12:111.

Length: 8 to 13 mm. Black. More slender than the other
black species. Head and pronotum shiny, with scattered shallow
but distinct punctures. Elytra densely punctured.

Head subquadrate. Median suture very fine. Antennal calluses
slightly raised, smooth. Clypeus and labrum more densely punc-
tured than the rest of the head, the clypeus densely punctulate
besides. Eyes prominent, quite broad, transverse, excavated.
Antennae reaching to just beyond the base of the elytra, one and
seven-eighths as long as an anterior tibia, filiform, slightly thick-
ened at the third segment. First segment stout, reaching to the
middle of the eye; second small, a little more than half as long as
the first; third slightly longer than the first and almost as thick;
fourth and following five-sevenths as long as the third, decreasing
gradually in thickness. Last segment three-fourths as thick as the

WERNER: NORTH AMERICAN EPICAUTA 451

third. Pronotum half again as long as wide, narrowly campanuli-
form. Median suture fine, indistinct. Basal impressed line distinct.
Elytra, as well as the underside and lower side of head, loosely
clothed with black pubescence. Each elytron with four slightly
raised lines, two from the base and two from the humerus. Male
anterior legs not modified. Posterior tibial spurs slender, stick-like,
quite long.
Type: Lectotype cf, new designation, Horn Coll., A.N.S.P. No.

8099, examined.
Type locality: Owen's Valley, California.

Additional localities: California: Independence, July 17 (Wick-
ham). Nevada: "Nev." Eddy Coll.

Epicauta albolineata (Duges)

Cantharis albolineata Duges, 1877, La Naturaleza, 4:64.

Epicauta albolineata, Duges, 1889, Anales del Museo Michoacana :84.

Epicauta duplicata Casey, 1891, Ann. N. Y. Ac. Sci. 6:172.

Epicauta albolineata, Champion, 1892, Biol. Cent.-Am., Col. 4 pt. 2:416.

Length: 10 to 12 mm. Stout; black, with pubescence cinereous
except for the black occiput divided by a median cinereous line, a
pair of elongated black spots on the pronotum. Elytra black;
margins cinereous, as well as a strong line from humerus to near
the apex, another inside it, arising from it near the base and united
with it at the apex. There is often an indistinct cinereous line just
outside the humeral line.

Head broadly triangular. Median suture distinct but partly
covered by pubescence in most specimens. Eyes small, narrow,
oblique, excavated. Antennae stout, rather short, of almost uni-
form thickness, about two and one-third times as long as an anterior
tibia. First segment stout, reaching to the middle of the eye;
second half as long; third as long as first but more slender; fourth
to tenth a little more than half as long as third ; last as long as third.
In the male, segments three to six slightly swollen, shiny. Pronotum
as broad as long, heavy, with the sides parallel for the basal four-
fifths, then converging abruptly. Median suture and basal im-
pressed line distinct. In addition, there is a narrow median
impression. Visible portion of scutellum very small, covered with
cinereous pubescence. Pubescence cinereous below. Pubescence
of femora and tibiae also cinereous in part. Anterior femora of
male denuded, especially below; anterior tibiae denuded but for

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a tuft of cinereous pubescence externally at the base. Anterior
tibial spurs of male shorter and stouter than in the female. Inner
spur of posterior tibiae slender, spiniform, outer broad, flattened,
obtuse.

Type: probably in Mexico City — Duges collection, not examined.
Type locality : Guanajuato.
of duplicata Casey.

Type: Casey Coll., U.S.N.M., examined.
Type locality: Arizona.

Additional localities: Arizona: Sta. Rita Mts. 5-8000 ft. Snow-
July (Fall Coll.).

Epicauta uniforma Werner
Epicauta uniforma Werner, 1944, Psyche 50:67.

Length: 8 to 12 mm. Black, uniformly and quite densely
clothed with dull ferrugineous to golden-ferrugineous pubescence.
Close to alpina Werner but the head is not narrowly triangular;
the first antennal segment of the male reaches only to the hind
margin of the eye and the first segment is not covered entirely with
golden-ferrugineous pubescence. Pubescence a little coarser and
sparser.

Head subquadrate, deeply and quite densely punctured, with the
intervals finely and densely punctulate, appearing dull. Median
suture distinct. Antennal calluses small, not denuded. Eyes
moderately prominent, moderately narrow, excavated. Antennae
reaching to the middle of the elytra, two and one-half times as
long as an anterior tibia. First segment of male antennae moder-
ately slender with a just perceptible excavation externally near the
tip, reaching the hind margin of the eye; of the female reaching
half-way across the eye and not excavated; second half as long as
the first; third one-third longer than second; fourth and following
four-fifths as long as third, decreasing gradually in thickness.
Pronotum one-third longer than broad, campanuliform. Median
suture distinct to one-fourth from apex. Basal impressed line
distinct. Male anterior legs not modified. Inner posterior tibial
spur flattened, spiniform, outer flattened, slightly broader apically
but pointed.
Type: Holotype cf, M. C. Z. No. 26,581, examined. (Erroneously

listed as 9 in original paper.)
Type locality: Ramah, Arizona.

WERNER: NORTH AMERICAN EPICAUTA 453

Additional localities: Arizona: Baboquivari Mts., Patagonia
Mts., Tombstone, Douglass, Tucson, Nogales, Picacho. Texas:
Davis Mts., Odell, Chisos Mts. Colorado: La Junta, Hoehne.

Epicauta alpina Werner
Epicauta alpina Werner, 1944, Psyche 50:67.

Length: 10 to 12 mm. Black, elytra tan to brown, the whole
densely clothed with appressed golden-ferrugineous pubescence.
Quite narrow. Head narrowly triangular, longer than normal.

Head rather narrowly triangular, especially in the male. Median
suture generally not visible but may be indicated on the occiput.
Antennal calluses not denuded. Eyes not very prominent, oblique,
narrow, excavated. Antennae reaching to the basal third of the
elytra, slender. First segment of male reaching one-fourth its
length beyond the eye, slender, slightly excavated externally near
the tip, clothed mainly with golden pubescence; of female, reaching
to hind margin of eye and not excavated; second segment of male
half as long as first and with a little golden pubescence behind; of
female, two-fifths as long as first, with or without golden pubescence;
third and following almost equal in thickness, tapering only slightly.
Third one and one-third times as long as second; rest slightly shorter.
Pronotum two-fifths longer than broad, campanuliform. Median
suture and basal impressed line distinct. There is a small midbasal
impression. Elytra without markings of any kind, even at base. Male
anterior legs not modified. Inner posterior tibial spur flattened-spini-
form, outer flattened, a little broader toward apex. Legs without dark
markings.

Type: Holotype cf, M. C. Z. No. 26061, examined.
Type locality: Alpine, Texas, 4400-6000 ft.
Additional localities: New Mexico: Deming.

Epicauta mimetica (Horn)
Gnathospasta mimetica Horn, 1875, Tr. Am. Ent. Soc. 5:154.

Length: 8 to 13 mm. Black or dark brown, rather densely and
evenly clothed with brownish-cinereous pubescence, which is easily
rubbed off. Labrum deeply incised. Mandibles long, meeting at tip.

Head subtriangular. Pubescence sparse. Median suture dis-
tinct. Antennal calluses quite long and narrow, extending beyond

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the inner margin of the eye. Eyes moderately prominent, oblique,
rather narrow, excavated externally near the tip. Antennae
slender, reaching to the basal third of the elytra. First segment of
male antennae rather slender, reaching one-fifth its length beyond
the eye, with a shallow excavation externally near the tip, of female
reaching four-fifths across the eye and without excavation ; second
half as long as third, which is as long as the first in the female, two-
thirds as long in the male; fourth and following four-fifths as long
as third, decreasing very gradually in thickness. Pronotum rather
small, as broad as long. Sides parallel for basal three-fourths, then
converging rather abruptly. Median suture lacking. Basal
impressed line distinct. Elytra without markings. Inner spur of
anterior tibiae of male long and stout, outer normal. Both spurs
normal in female. Posterior tibial spurs flattened, inner spiniform,
outer broader but still spiniform.
Type: Holotype d", A.N.S.P., examined.
Type locality: Arizona.

Additional localities: Arizona: Riley Coll. (U.S.N.M.). Texas:
Davis Mts. 1X-20-38 (Ohio State).

Epicauta brunnea Werner
Fig. 6, 7
Epicauta brunnea Werner, 1944, Psyche 50:67.

Length: 9 to 16 mm. Black, the elytra brown, the whole quite
densely clothed with appressed ferrugineous pubescence. Elytra
darker across the base. First segment of anterior tarsi of male
broad across the pad.

Head subquadrate. Median suture distinct. Antennal calluses
very small, denuded. Eyes prominent, transverse, quite broad.
Antennae reaching the basal third of the elytra, two and one-fourth
times as long as an anterior tibia. First segment in male fairly
stout toward tip, reaching half-way across the eye; in the female
more slender and reaching two-fifths across the eye; in both sexes
there are a few ferrugineous hairs toward the tip. Second segment
small, two-fifths as long as the first in the male, slightly more than
two-fifths in the female; third about as long as the first two; fourth
and following subequal, four-fifths as long as third. The fifth seg-
ment is the broadest. The intermediate segments of the antennae
are broader in the male than in the female. Pronotum subquadrate,

WERNER: NORTH AMERICAN EPICAUTA 455

one-sixth longer than broad. It is broadest four-fifths from the
base, the sides tapering gradually to the base and abruptly to the
apex. Median suture and basal impressed line distinct, not denuded.
Scutellum with ferrugineous pubescence. Male anterior tibiae not
modified. First segment of male anterior tarsi broadened, so that
the "sole" is about twice as broad as that of the second segment.
In the female, it is just slightly broader than that of the second.
Inner posterior tibial spur long, sticklike, outer sticklike but
shorter.

Type: Holotype d\ M. C. Z. No. 26604, examined.
Type locality: Alpine, Texas, 6000 ft.

Additional localities: Texas: Brewster Co., Fort Davis, Sabinal.
Arizona: Sta. Cruz Village-Cobabi Mts., Superior, Douglass,
Capitan Mt., Globe, San Carlos, Canon Lake, Gila Valley,
Whiteriver. Colorado: Denver. Occurs from mid-July to mid-
September.

Epicauta ingrata Fall
' _ Fig. 17-

Epicauta ingrata Fall, 1907, Trans. Am. Ent. Soc. 33:258.

Length: 6 to 9 mm. Black; densely and evenly clothed with
appressed cinereous pubescence. Antennae very long and slender,
reaching beyond the middle of the elytra.

Head subtriangular. Median suture fine, supplemented by a
narrow denuded line. Antennal calluses large. Eyes moderately
prominent, rather narrow. Antennae very long and slender.
First segment just surpassing the eye; second half as long as first;
third almost as long as first two; rest three-fifths as long as third,
decreasing in thickness from the fourth. Pronotum small, quad-
rate, a little longer than broad. Median suture distinct. Elytra
with black or dark brown scutellar and humeral spot. Pubescence
often denser on the margins of the elytra. Male anterior legs not
modified. Posterior tibial spurs both slender, spiniform.
Type: Holotype d\ M. C. Z. No. 24294, examined.
Type locality: Cloudcroft, New Mexico., 9000 ft.
Additional localities: New Mexico: Sante Fe Co., Sandia Mts.,
'Coolidge. Arizona: Flagstaff, Mormon Lake, Globe, White

Mts., Sierra Ancha Mts. Colorado: Colorado Springs. Occurs

in June and July.

456 bulletin: museum of comparative zoology

Epicauta cinerea (Forster)
Fig. 3, 4

Meloe cinereus Forster, 1771, Cat. Animals of N. Am. :62.
Meloe clematides Woodhouse, 1800, Med. Repository 3:213.
Epicauta marginata auct., nee Fabricius, (in part).

Length: 9 to 13 mm. Black, quite densely clothed with ap-
pressed pubescence. Two forms occur. One is entirely cinereous
except for the base of the elytra. The other has the elytra black
except on the margins and a pair of black marks on head and
pronotum. Mr. H. S. Barber has found the cinereous form com-
posing about 1% of the population near Washington. It is dis-
tinguished from the common potato species, solani, by its thick an-
tennae, short, stout first antenna! segment and by the rather slender,
spiniform posterior tibial spurs. This species appears to be confined to
species of Clematis in its food habits.

Head subquadrate, minutely granulose, quite densely but not
deeply punctured. Median suture distinct. Eyes small, trans-
verse, rather narrow. Antennae reaching to the basal fourth of
the elytra, two and one-third times as long as an anterior tibia.
First segment stout, reaching one-third across the eye; second
three-fourths as long, narrow; third just shorter than first two;
fourth as long as first, the stoutest of the median segments; rest
gradually decreasing in length and thickness. Pronotum subquad-
rate, slightly broader apically. Median suture and basal impressed
line distinct, the basal the deeper. Tips of femora, tibiae and most
of tarsi black. Male anterior legs not modified. Hind tibiae with
spurs spiniform, quite slender but outer broader.
Type: Present location unknown to author.
Type locality: North America.
Additional locality: Common in the coastal states from New

Hampshire to South Carolina and more less sparingly west to

Iowa. Occurs from June to September.

Epicauta fissilabris (Le Conte)

Lytta fissilabris Le Conte, 1850, Agassiz L. Sup. 4:232; 1853, Proc. Acad. Nat.

Sci. Phila. 6:339.
Epicauta fissilabris, Horn, 1873, Proc. Am. Phil. Soc. 13:102.

Length: 9 to 13 mm. Black, with short black pubescence, as in
corvina. First segment of antennae much stouter and pronotum
less bulged than in that species. Structurally resembles cinerea.

WERNER: NORTH AMERICAN EPICAUTA 457

Head subtriangular. Median suture fine. Antennal calluses
small, denuded, smooth. Eyes moderately prominent, fairly nar-
row. Antennae reaching to basal third of elytra, one and one-half
times as long as an anterior tibia. First segment stout, short,
reaching one-third across the eye; second slender, three-fourths as
long as first; third twice as long as second; fourth and following
two-thirds as long as third, decreasing gradually in thickness.
Pronotum one-sixth broader than long. Sides straight, widest four-
fifths from base, then converging abruptly. Median suture dis-
tinct except on basal fifth. Basal impressed line distinct. Midbasal
impression not strong, pronotum not bulged as they are in corvina.
Male anterior legs not modified. Outer spur of posterior tibiae
sticklike, the inner spiniform. The labrum is incised, but no more
than in cinerea and only slightly more than in corvina.
Type: Lectotype 9, new designation, M. C. Z. No. 5096, exam-
ined.
Type locality: Lake Superior.

Additional localities: Hudson's Bay Terr. (Le Conte Coll.).
Manitoba: Aweme (Criddle VII-07).

Epicauta solani sp. n.

Fig. 5

Epicauta marginata auct., nee Fabricius (in part).
Epicauta cinerea auct., nee Forster (in part).

Length: 6 to 16 mm. Markings, form, color same as in cinerea
but with differences in the antennae and tibial spurs. An entirely
cinereous form exists but is rare. Only a very few have been seen
by the author. They occur in the western part of its range.

Antennae much more slender than in cinerea. First segment
rather slender, reaching half-way across the eye. Cinerea has the
first segment very stout by comparison. Second half as long as
first, slender; third a little longer than first, slender; fourth to last
two-thirds as long as third. The sixth segment is the thickest but
the intermediate segments are not conspicuously thickened as
they are in cinerea. Inner posterior tibial spur flattened, pointed,
outer broader end with a blunt tip. They are decidedly more
slender in cinerea.

This is the common margined blister beetle of potatoes. The
name marginata cannot be used for it because Fabricius used the

458 bulletin: museum of comparative zoology

name first for an insect from the Cape of Good Hope and later
applied it to one of our North American margined species.

Occurs abundantly from the Atlantic coast to the Rocky Mts.,
from July to September, often doing considerable damage to
potatoes and occasionally to other soft garden plants.

Large numbers of this species have been examined. The author ex-
pected during the course of the work that Mr. H. S. Barber, who first
noticed the differences between cinerea and solani, would describe this
species. As a result only one short set is designated as type series.

Type: Holotype & (M. C. Z.).

Type locality: Norwood, Pennsylvania VII-31 Liebeck Coll.

Allotype 9, eutopotypical (M. C. Z.), Paratypes: lcf , 1 9 topo-
typical VII-24 Liebeck Coll. (M. C. Z.).

Epicauta floridensis Werner
Fig. 1,2 .
Epicauta floridensis Werner, 1944, Psyche 60:68.

Length: 6 to 11 mm. Resembles the cinereous form of cinerea
but is more slender and has the pubescence sparser. The antennae
are intermediate in form between cinerea and solani. The posterior
tibial spurs are slender, as in solani.

Head subquadrate, moderately densely punctured and punctu-
late, except along the median suture. Median suture distinct.
Antennal calluses small, slightly elevated, denuded, smooth. Eyes
moderately prominent, fairly narrow. Antennae reaching to the
basal fourth of the elytra, two and one-third times as long as an
anterior tibia. First segment fairly stout, reaching almost to hind
margin of eye in the male, halfway across in female; second half as
long as first ; third as long as first, narrow at base and fairly broad
at apex. The first three segments have some cinereous pubescence
behind. The fourth is the broadest segment, and is three-fourths
as long as third. Rest decreasing gradually in breadth and just
perceptibly in length. The intermediate segments of the female
antennae are not as thick as in the male. Pronotum subquadrate,
one-fourth longer than broad. Median suture and basal impressed
line distinct. Elytra uniformly clothed. Anterior legs of male not
modified.

Type: Holotype d\ M. C. Z. No. 26067, examined.
Type locality : Sebring, Florida.

WERNER: NORTH AMERICAN EPICAUTA 459

Additional localities: Florida: Croom, Cleveland, Capron, Bis-
cayne, Tampa, Cutler, Jacksonville, St. Nicholas, Lake Istopoga.
Mississippi: Lucedale. New Jersey: Da Costa.

Epicauta ruidosana Fall
Epicauta ruidosana Fall, 1907, Trans. Am. Ent. Soc. 33:258.

Length: 9 to 13 mm. This species has been confused with
"marginata" in collections but it is very distinct. The cinereous
margins of the elytra are narrow and very sharply defined. The
cinereous pubescence below is sparse and long.

Head subtriangular. Surface quite densely and deeply punc-
tured, sparsely punctulate. Median suture strong. Antennal
calluses small, not distinctly elevated. Eyes moderately prominent,
fairly narrow. The surface of the body is more densely but shal-
lowly punctured, much more punctulate than the head. Pubes-
cence of head and thorax pale, long, sparse, densest on under side
and along midline. Antennae without pale pubescence. They are
rather loosely jointed and reach to the basal third of the elytra.
First segment reaching to middle of eye, quite stout; second half
as long as first, small; third slender, one-fourth longer than the
first two; fourth and following two-thirds as long as third, slender,
especially toward the tip. Pronotum quadrate, slightly broader
than long, appearing flattened on the disc. Median suture and
basal impressed line distinct. A narrow median impression supple-
ments the suture. Pubescence concentrated in this groove and on
the posterior margin. Elytra with short black pubescence, mar-
gined with short cinereous to tannish pubescence. Scutellum also
cinereous pubescent. First segment of anterior tarsi of male slightly
longer than in the female. Posterior tibial spurs slender, spiniform,
the outer longer.

Type: Holotype 9 , M. C. Z. No. 24295, examined.
Type locality: N. Fork of Ruidoso R., White Mts., Arizona.
Additional localities: Arizona: Ida's Peak — Chiricahua Mts.,

7-8000 ft. Texas : Casa Grande— Chisos Mts., 7000 ft. Occurs in

June and July.

Epicauta punctipennis Werner

Epicauta punctipennis Werner, 1944, Psyche 50:68.

Length: 10 to 12 mm. Black or dark brown, elytra ferrugi-
neous. Rather densely clothed with tannish-cinereous pubescence.

460 bulletin: museum of comparative zoology

Elytra with small, sparsely scattered denuded spots. There are
no denuded spots on the rest of the body.

Head subquadrate, densely but quite shallowly punctured,
densely punctulate. Antennal calluses moderate in size, denuded,
shiny. Eyes small, narrow, transverse. Antennae twice as long as
an anterior tibia. First segment moderately stout, reaching half-
way across the eye, with cinereous pubescence behind ; second two-
thirds as long as first, quite slender, also with some cinereous
pubescence behind; third a little longer than first, the thickest
segment; fourth and following three-fifths as long as third, decreas-
ing gradually in thickness. The apical segments are quite slender.
Palpi of male smooth beneath, with the apical segments slightly
expanded. Sculpture of pronotum similar to that of head. Median
suture fine, indistinct near base and apex. Basal impressed line
fine. Elytra with brown pubescence at very base. Scutellum with
cinereous pubescence. Elytra with sixty to seventy small denuded
spots, evenly scattered. Underside dark. All but tips of femora,
tibiae and first tarsal segment ferrugineous. Male anterior legs not
modified. Posterior tibial spurs slender, the inner spiniform, the
outer sticklike.

Type: Holotype 9 , M. C. Z. No. 26159, examined.
Type locality: Columbus, Texas, May 24, 1879, Le Conte Coll.

Allotype' <?, from "Tex." Liebeck Coll.

Epicauta balli sp. n.

Length: 6 to 8 mm. Black, sparsely clothed with moderately
long cinereous pubescence. First segment of antennae, mouth
parts and legs pale ferrugineous.

Head rather elongate in aspect, densely and deeply punctured.
Median suture distinct. Antennal calluses smooth. Eyes moder-
ately broad, fairly coarsely granulated.. Labrum narrow, deeply
and broadly excavated. Mandibles slender. Maxillae similar to
those of mimetica, the palpi long and slender. First segment of
antennae of male stout, cylindrical, reaching two-thirds across the
eye, of female reaching half-way across; second about two-fifths as
long as first; third twice as long as second in male, not quite twice
in female; rest slightly shorter than third, narrowing gradually.
Pronotum a little longer than broad, with the sides parallel for the
basal three-fourths, then converging abruptly. Median suture
distinct. Pronotum and elytra coarsely and deeply punctured.

WERNER: NORTH AMERICAN EPICAUTA 461

Spurs of anterior tibiae of male stouter than in the female. Pos-
terior tibial spurs slender, spiniform.

Mr. Frank H. Parker intended to name this species in honor of
the collector but has allowed me to describe it here.
Type: Holotype d71, Calif. Acad. Sci.

Type locality: Patagonia, Arizona, Jul. 28, 33. E. D. Ball.
Paratypes: Arizona: Nogales, 1 Aug. 24, 39, R. H. Crandall

(Crandall Coll.), 2 Sept. 22, 06 (USNM).

Epicauta oregona Horn

Fig. 16

Epicauta oregona Horn, 1875, Trans. Am. Ent. Soc. 5:153.

Length: 7 to 11 mm. Black, sparsely clothed with rather long
cinereous pubescence and with numerous small denuded spots.
Antennae subensiform, especially in the male. Some of the seg-
ments are flat even in the female. This character easily dis-
tinguishes the species from the other maculate species.

Head subquadrate, densely and rather deeply punctured.
Median suture distinct. Antennal calluses denuded, shiny, with a
strong impression near the inner edge. Eyes fairly prominent,
quite narrow. Antennae a little more than two and one-half times
as long as an anterior tibia. First segment stout, reaching to the
middle of the eye; second small, half as long as first; third almost
as long as first, flat, quite broad in male; fourth to tenth three-
fourths as long as the third, the fourth very broad in the male.
The segments near the apex are very narrow. Last segment one-
third longer than tenth. Pronotum subquadrate, one-fourth longer
than broad. Sides parallel for basal four-fifths, then converging
abruptly. Median suture distinct, supplemented by a narrow
median impression. Basal impressed line distinct. Scutellum quite
small. Elytra denuded at very base. Below with denuded spots
sparser than above. Male with long pubescence on base of the
femora but with the legs not otherwise modified. Posterior tibial
spurs slender and spiniform.

Type: Lectotype d\ new designation, A.N.S.P. No. 809S, exam-
ined.
Type locality: Oregon.

Additional localities: British Columbia: Martin. Washing-
ton: Yakima City, Ellenburg. Idaho: Winchester. Utah:

462 bulletin: museum of comparative zoology

Trenton. Montana: Perma, Hot -Springs. Colorado: Monte-
zuma. Arizona: Flagstaff. Occurs in June and July. Not
common.

Epicauta ensiformis Werner
Epicauta ensiformis Werner, 1944, Psyche 60:68.

Length: 9 to 10 mm. Black, sparsely clothed with erect black
pubescence. Resembles oblita in general form but is easily dis-
tinguished by its strongly ensiform antennae.

Head subtriangular, shiny, sparsely but deeply punctured. Me-
dian suture very fine. Antennal calluses smooth, fairly large. Eyes
moderately prominent, fairly narrow. Male antennae reaching to
basal fourth of elytra, twice as long as an anterior tibia. First
segment stout, hairy, reaching one-third across the eye; second
very small, one-third as long as first, also hairy; third large, as
long as the first two, broad, somewhat flattened. It is the broadest
segment. Fourth and following two-thirds as long as third, slightly
flattened, tapering gradually to the apical segment, which is very
slender. Segments strongly separated, with the expansion on the
posterior side. Pronotum as broad as long, broadest at base,
tapering gradually halfway to apex, then converging more abruptly.
Median suture fine but distinct. Basal impressed line distinct.
Surface similar to that of head. Male anterior legs apparently not
modified. Posterior tibial spurs long, the inner slightly longer,
slender, spiniform.
Type: Holotype cf , "Cal" with an ink spot on lower portion of

the "a", which may indicate southern California. Liebeck Coll.,

M. C. Z. No. 26079. Paratype: tf, "Cal," Fall Coll. M. C. Z.

Epicauta stuarti Le Conte

Epicauta stuarti Le Conte, 1868, Trans. Am. Ent. Soc. 2:54. Horn, 1873,
Proc. Am. Phil. Soc. 13:101.

Length: 6 to 11 mm. A striking species, looking more like a
Tetraonyx than an Epicauta. Black, densely clothed with long
pubescence. Head black down to level of the eyes, orange beyond.
Pronotum orange with a pair of rounded black spots. Elytra
narrowly orange along suture, black across the base from the
humerus obliquely back to near the middle and with a pair of large
black fasciae behind. Abdomen with middle of sternites denuded,

WERNER: NORTH AMERICAN EPICAUTA 463

smooth. Sides of sternites broadly orange behind. Fifth sternite
with apex entirely orange. Pubescence on anterior legs cinereous,
sparse.

Head broadly triangular. Median suture distinct. Antennal cal-
luses not denuded. Eyes not prominent, moderately narrow,
transverse. Clypeus with pubescence orange, labrum black.
Antennae a little more than twice as long as an anterior tibia,
tapering slightly. First segment reaching to the middle of the eye;
second two-fifths as long as first; third as long as first two; follow-
ing equal, a little shorter than first. Pronotum almost twice as
broad as long, flat, gradually narrowed forward. Midline im-
pressed. A pair of impressions on middle of disc. Scutellum small,
with orange pubescence. Male anterior legs not modified. Poste-
rior tibial spurs flattened, sticklike, the outer slightly broader than
the inner.

Type: Lectotype 9, new designation, M. C. Z. No. 5095, exam-
ined.
Type locality: New Mexico.

Additional localities: Colorado: Crowley, Holly, Berkeley, Mont-
clair. Kansas: taken by Williston.

Epicauta lemniscata (Fabricius)

Fig. 9

Lytta lemniscata Fabricius, 1801, Syst. Eleutheratorum : 279. Le Conte, 1853,

Proc. Acad. Nat. Sci. Phila. 6:341.
Epicauta lemniscata, Horn, 1873, Proc. Am. Phil. Soc. 13:100. ?Champion,
1892, Biol. Cent.-Am., Coleop. 4 pt. 2:415.

Length: 8 to 16 mm. Pale luteous, with subsutural, discal and
submarginal black stripes on the elytra. Pronotum with a pair of
longitudinal black stripes, which are sometimes abbreviated or
broken, or expanded outward. Head with a pair of elongate curved
black marks on the occiput. Below black or partly luteous.

I find it impossible to separate this species satisfactorily from
vittata unless a series from one locality is at hand. No tendency is
shown in the South and West toward a narrowing of the interval
between the two outer black stripes on the elytra. These I have
called lemniscata. In the North this interval is narrowed or obliter-
ated and I call the series vittata. Both of these names are so well
established that they should not be changed until there is absolute

464 bulletin: museum of comparative zoology

proof that there is only one species involved. But they are
extremely close if not identical.

Head subquadrate, minutely rugose, appearing dull. Rest of
body similarly sculptured. Median suture distinct. Antennal
calluses broad, moderate in size, touching the eye. Eyes only
moderately prominent, transverse. Antennae slender, filiform,
slightly flattened but not broadened, reaching to the basal third
of the elytra, two and one-half times as long as an anterior tibia.
First segment stout, reaching two-thirds across the eye; second
slender, three-fifths as long as first; third slightly longer than first
two, slightly flattened; fourth three-fourths as long as third; rest
decreasing slightly in length. Fifth and sixth as thick as fourth,
rest decreasing to a quite slender apex. Pronotum one-third longer
than broad. Sides parallel for the basal three-fourths, then con-
verging gradually. A shallow impression crosses the disc at the
obtuse angle thus formed. Median suture and basal impressed line
distinct. A shallow midbasal impression present. Luteous pubes-
cence forms a median stripe and a pair of lateral stripes outside of
black stripes, independent of color of cuticle. Bases of femora,
tibiae and part of thorax and abdomen usually at least partly
luteous. Male anterior legs not modified. Inner posterior tibial
spur flattened, sticklike, outer flattened and somewhat broader.
Type : present location unknown to author.
Type locality: North America.
Additional localities: Occurs from New Jersey and Florida to

Nebraska and Texas, probably extending far into Mexico.

Abundant in early summer, eating cotton, potatoes and other

soft plants.

Epicauta vittata (Fabricius)

Lytta vittata Fabricius, 1775, Syst. Ent. :260; 1781, Syst. Ins. :328; 1787.
Mant. Ins. :215 (Litta). Le Conte, 1853, Proc. Ac. Nat. Sci. Phila. 6:340,
Staig, 1940, The Fab. Types in Hunterian Coll., Coleop. 2:139, PL 56.

Melos Chapmani Woodhouse, 1800, Medical Repository 3:214.

Epicauta vittata, Horn, 1873, Proc. Am. Phil. Soc. 13:100. Staig, 1940, The
Fab. Types in Hunterian Coll. :139.

Length: 9 to 14 mm. Structurally identical with lemniscata. If
the interval between the outer black stripes on the elytra tends to
be reduced or obliterated in a series, I have referred the whole
series to this species. Occurs from Maine to New Jersey west to

WERNER: NORTH AMERICAN EPICAUTA 465

Iowa. Often a serious pest of potatoes and also feeds occasionally

on many other soft garden plants.

Type: Lectotype, new designation, Hunter Coll., the specimen

described by Staig.
Type locality: "America."

Epicauta occidentalis Werner
Fig. 10
Epicauta occidentalis Werner, 1944, Psyche 50:69.

Length: 10 to 13 mm. Identical with lemniscata except for the
broadened antennae and smooth, shiny outer edge of the anterior
tibiae and tarsi. Lengths of antennal segments same as in lemnis-
cata. First three segments and the last three the same as in that
species. Fourth to eighth much flattened, the sixth the broadest.
Outer half of anterior tibiae and first two tarsal segments denuded,
with only a few scattered punctures, bearing macroscopically
invisible hairs.

No intergrades occur between this species and lemniscata in my
series, although some are from the same locality. Both sexes are
represented. Just what the status of this species is will have to be
determined by field investigation.
Type: Holotype <?, M. C. Z. No. 26069, examined.
Type locality: Cambridge, Nebraska.
Additional localities: Kansas: Abilene. Colorado: Rocky Ford,

Joes, Fort Collins. Texas: Dallas. Louisiana: Baton Rouge,

New Orleans, Harahan, Belt City.

Epicauta ferruginea (Say)
Fig. 19

Lytta ferruginea Say, 1823, Journ. Acad. Nat. Sci. Phila. 3:298. Le Conte

1853, Proc. Acad. Nat. Sci. Phila. 6:341.
Epicauta ferruginea, Horn, 1873, Proc. Am. Phil. Soc. 13:98.

Length: 6 to 9 mm. Black, densely clothed with rather long
ferrugineous pubescence. Outer spur of posterior tibiae broad,
inner slender. Third segment of antennae just surpassing the eye.

Head subtriangular, moderately densely punctured, smooth and
shiny between the punctures. Median suture visible only on the
occiput and there obscured by the pubescence. Antennal calluses

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small, denuded, shiny. Eyes prominent, broad, excavated. An-
tennae rather stout, uniform in thickness, just reaching the base
of the elytra, twice as long as an anterior tibia. First segment
moderately stout, reaching one-fourth across the eye; second small,
half as long as first; third as long as first but more slender; fourth
slightly longer than second; fifth to tenth slightly longer than
fourth; last as long as first. All segments longer than broad.
Pronotum subquadrate, as broad as long, bulging somewhat. The
sides converge three-fourths from the base. Median suture absent
but a narrow median impression is present. Basal impressed line
distinct. Elytra without markings. Some ferrugineous specimens
have the marginal pubescence slightly paler. Pubescence below
sparser than above. Anterior legs of male not modified.
Type: Say's type lost. Neotype 9 and 6 neoparatypes in M. C. Z.
Type locality: "Found on the Missouri by Mr. Nuttall." Neotype

locality: Cambridge, Nebraska.
Additional localities: From Iowa to Idaho, south to Kansas,

Texas, and Arizona. Fairly common. Occurs from July to

September.

Epicauta fortis (Werner)
Fig. 20
Epicauta fortis Werner, .1944, Psyche 50:69.

Length: 6 to 9 mm. This species resembles sericans greatly in
general shape and habitus but is consistently smaller than that
species and has the outer spur of the hind tibiae broad and flat-
tened, and the tarsal claws curved from just beyond the middle.

Black, densely clothed with cinereous pubescence. Surface
thickly punctured but it is usually not visible under the pubescence.
Head subquadrate. Median suture very indistinct. There is a
narrow median smooth area on the front. Eyes only moderately
prominent, broadly oval, excavated slightly. Antennae black, very
short, only one and one-half times as long as an anterior tibia,
thick, thicker toward the apex. First segment stout, reaching one-
fifth across the eye; second half as long as first; third as long as
first. Fourth to tenth equal, two-thirds of the third, increasing
slightly in thickness. Last as long as first. The fourth segment
just reaches the hind margin of the eye. The first three segments
have a few cinereous hairs. Pronotum broad, at broadest part
almost as wide as head, as broad as long, broadly campanuliform.

WERNER: NORTH AMERICAN EPICAUTA 467

Median suture indistinct but there is a narrow median impression,
which is broader and deeper posteriorly. Basal impressed line
distinct. A pair of shallow impressions on the disc just behind the
middle. Male anterior legs not modified.

The characteristics of this species are very similar to those of
ferrugined but it is more often cinereous. The general form is
broader. The antennae are shorter, the eyes broader. The anten-
nae are stouter, more closely articulated, and with some segments
broader than long.

Type: Holotype d\ M. C. Z. No. 26066, examined.
Type locality: Phoenix, Arizona.
Additional localities: Arizona: Tucson, Picacho, Florence. New

Mexico: Las Cruces.

Epicauta strigosa (Gyllenhal)

Lytta strigosa Gyllenhal, 1817, App. to Schoenherr's Syn. Ins. 3:18. Le Conte,

1853, Proc. Acad. Nat. Sci. Phila. 6:341.
Cantharis nigricornis Melsheimer, 1847, Proc. Acad. Nat. Sci. Phila. 3:53.
Epicauta strigosa, Horn, 1873, Proc. Am. Phil. Soc. 13:97.

Length: 6 to 11 mm. Black, rather densely pubescent. But the
black background shows through and affects the general color.
Pubescence ferrugineous to cinereous, usually with black or dark
brown humeral and scutellar stripes, a pair of narrow lines on the
pronotum. Legs black. The only species with which this species
can be confused, in its range, is batesii. Unicolorous specimens
resemble that species but have black legs instead of ferrugineous.

Head subquadrate, rather sparsely and shallowly punctured.
Median suture distinct. Eyes large, broadly oval but not very
prominent, excavated. Antennae black, twice as long as an
anterior tibia, thick, almost uniform in thickness. First segment
reaching one-third across the eye; second small, half as long as
first; third almost as long as first; rest two-thirds as long as third,
rather closely articulated. Pronotum.one-fourth longer than broad.
Basal four-fifths with sides parallel, then converging rather
abuptly. Median suture obscure. A narrow median impression
present. On the sides of the disc are a pair of shallow longitudinal
impressions. Elytra as described above. Suture always pale. Male
anterior legs not modified. Posterior tibial spurs slender, sticklike,
rather long.

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Type: Presumably in Gyllenhal's collection, Zool. Univ. Mus.,
Upsala.

Type locality: Presumed to be North America.

of nigricornis (Melsh.)

Type: specimen in Melsheimer Coll. is hereby designated as lecto-
type (M. C. Z. No. 26675). It is a specimen without stripes.

Type locality: Alabama.

Additional localities: Coastal states from North Carolina to
Mississippi. Particularly abundant in Florida. A few specimens
from Wellesley, Framingham, and Dover, Massachusetts. One
of these has the normal stripes. All the northern specimens are
a little smaller and paler than normal.

Epicauta batesii Horn
Epicauta Batesii Horn, 1875, Trans. Am. Ent. Soc. 5:153.

Length: 6 to 10 mm. Rufo-ferrugineous. Head and underside,
except for legs, darker. Sparsely clothed with tannish pubescence.

Head subquadrate, shiny, deeply and densely punctured.
Median suture moderately deep. Eyes very prominent, globular,
almost round but slightly excavated near the antennae. Antennae
two and one-third times as long as an anterior tibia. First segment
stout, reaching one-fourth across the eye; second half as long as
first; third almost as long as first two, rather slender; rest a little
shorter than first, stout, closely articulated. Pronotum one-third
longer than wide, broadly campanulate, deeply and densely punc-
tured. Median suture feeble; a median impression present basally.
Basal impressed line distinct. A faint margin may be formed on
the elytra by slightly denser pubescence. Male anterior legs not
modified. The cinereous patch on the anterior femora extends
almost the full length. Posterior tibial spurs slender, spiniform.
Type : Lectotype 9 , new designation, A.N.S.P. No. 8092, examined.

From the Dejean Coll.
Type locality: Savannah, Georgia.
Additional localities: Quite common in Florida and extending as

far north as New Jersey. Occurs in fall and winter.

Epicauta callosa Le Conte

Epicauta callosa Le Conte, 1866, Smiths. Misc. Coll. 6 no. 167, 2nd ed: 158.
Horn, 1873, Proc. Am. Phil. Soc. 13:99.

WERNER: NORTH AMERICAN EPICAUTA 469

Length: 7 to 10 mm. Black, densely clothed with tannish to
rufo-ferrugineous pubescence. Rather slender. Pronotum always
with a pair of small, slightly raised, denuded areas just before the
middle.

Head subquadrate, shiny, densely and finely punctured. An-
tennal calluses not denuded. Eyes broad, excavated. Antennae
fairly slender, two and one-half times as long as an anterior tibia.
First segment rather stout, reaching one-third across the eye;
second half as long as first, almost as broad as long; third a little
longer than first, quite slender; rest two-thirds as long as third,
loosely articulated, equal in thickness. Pronotum one-fourth
longer than broad, campanuliform. Surface similar to head.
Median suture distinct and with a basal median impression. Basal
impressed line distinct. Elytra often with the pubescence slightly
paler on the margins. Male anterior legs not modified. Posterior
tibial spurs slender, sticklike, the outer sometimes a little broader.
Type: Lectotype 9, new designation, M. C. Z. No. 4996, exam-
ined.
Type locality: Nebraska.

Additional localities: Nebraska: Edgar 7-28. Kansas: Ellsworth
8-4, Fort Hayes. Oklahoma: Guy man. Texas: Denton Co.
5-27, Flatonia, El Paso, Alpine, Dallas.

Epicauta sericans Le Conte
Fig. 21

Epicauta sericans Le Conte, 1866, Smiths. Misc. Coll. 6 no. 167, 2nd ed :158.
Horn, 1873, Proc. Am. Phil. Soc. 13:98.

Length: 7 to 11 mm. Stout, black, densely clothed with ciner-
eous pubescence. Posterior tibial spurs slender, sticklike. Tarsal
claws curved from the base.

Head broadly triangular, finely and densely punctured. Median
suture not discernible. Eyes moderately prominent, broadly oval,
slightly excavated. Antennae black, uniform in thickness, a little
more than one and one-half times as long as an anterior tibia.
First segment reaching one-fourth across the eye, with cinereous
pubescence; second half as long as first; third a little longer than
first and more slender; fourth to tenth two-thirds of third, closely
articulated, and just perceptibly thicker apically. Last segment
as long as third. Pronotum stout, almost as wide as the head, as

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broad as long, subquadrate. Sides parallel for basal three-fourths,
then converging abruptly. Median suture not visible, replaced by
a narrow median impression. Basal impressed line distinct. Below
uniform except for the black tarsi. Male anterior legs not modified.
Posterior tibial spurs slender, sticklike.

Type: Lectotype 9 , new designation, M. C. Z. No. 4994, exam-
ined.
Type locality: Kansas.

Additional localities: Common from Kansas to Texas but has a
much wider range. I have seen it from Indiana, Alabama,
Mississippi, Iowa, Alberta, Saskatchewan, Utah, Colorado,
Arizona, New Mexico, Texas, Oklahoma, Kansas, and Nebraska.
Occurs mostly in June and July.

Epicauta atrata (Fabricius)
Fig. 18

Lytta atrata Fabricius, 1775, Syst. Ent. :260; 1781, Spec. Ins. 1:329. nee 1787,

Mant. Ins. 1:215, where listed as from Barbary. Staig, 1940, the Fab.

Types in Hunterian Coll. Coleop. 2:135, PI. 55.
Meloe trichrus Pallas, 1789 (See Sherborn-Ann & Mag. Nat. Hist., Feb. 1891),

Icones Ins. (1781) :100, Tab. E, fig. 32 (1782).
Lytta coracina Illiger, 1804, Mag. f. Inskunde 3:171 (given as nom. nov. for

American atrata but it was described first (1775), the Barbary species in

1787.
Lytta convolvuli Melsheimer, 1848, Proc. Acad. Nat. Sci. Phila. 3 (1847) :53.

(as a variety of atrata Fab.). Le Conte, 1853, Proc. Acad. Nat. Sci.

Phila, 6:339.
Epicauta convolvuli, Le Conte, 1866, Smiths. Misc. Coll. 6 no. 167, 2nd ed. :157.

Horn, 1873, Proc. Am. Phil. Soc. 13:97.
Epicauta trichura Wellman, 1910, D. E. Z. 6:24 (This change of names was

unwarranted. Trichrus, as used by Pliny, was feminine, for a three-
colored gem.)

Length: 6 to 12 mm. Black or dark brown, the head sometimes
bright rufous. Pubescence entirely black, black with margins of
elytra cinereous, cinereous with obscure dark discal and humeral
stripes on the elytra, or entirely cinereous. Can be distinguished
in the black form from pensylvanica by its larger scutellum, stouter
second antennal segment and more erect pubescence on the head
and antennae.

Head subquadrate, moderately densely, deeply punctured, the

WERNER: NORTH AMERICAN EPICAUTA 471

intervals finely and densely punctured in the male, smooth in the
female. In a few specimens from Indiana, the head is much ex-
panded behind the eyes. Median suture fine, visible only on the
occiput. Eyes prominent, broad, only slightly excavated. An-
tennae a little over twice as long as an anterior tibia. First segment
stout, reaching to just behind the anterior margin of the eye;
second also stout, two-fifths as long as first; third more slender,
almost as long as first two; rest a little longer than the second,
equal in length and thickness. Pronotum slightly broader than
long. Sides parallel for the basal four-fifths, then converging
abruptly. Surface densely and rather deeply punctured. Median
suture indistinct. An indistinct basal median impression present,
one before the middle and a longitudinal pair on the sides. Scutel-
lum normal in size. Male anterior legs not modified. Inner spur of
hind tibia slender, spiniform, outer somewhat broad, flattened.
Type: The specimen figured by Staig hereby designated lectotype.

Hunterian Coll., Glasgow University.
Type locality: North America.
Additional localities: From Maryland and North Carolina to

Nebraska and Texas. Occurs in early summeronConvolvulaceae.

Epicauta pruinosa LeConte

Epicauta pruinosa Le Conte, 1866, Smiths. Misc. Coll. 6 no. 167, 2nd ed.: 158.
Horn, 1873, Proc. Am. Phil. Soc. 13:98.

Length: 6 to 11 mm. Black, very sparsely clothed with fine
cinereous pubescence so that the surface appears greyish black.
Median suture of head and pronotum conspicuous. Surface
moderately densely punctured, finely punctulate so that it appears
dull.

Head subtriangular. Median suture very distinct, supplemented
by smooth, shiny margins. Antennal calluses small, denuded,
shiny. Eyes fairly prominent, moderately narrow. Pubescence on
under side of the head erect, longer than on the rest of the body.
Antennae almost uniform in thickness, reaching to just beyond
the base of the elytra, about one and two-thirds times as long as
an anterior tibia. First segment stout, reaching almost to the
middle of the eye in the male, one-third across in the female; second
small, half as long as first; third almost as long as first two; fourth
and following three-fourths as long as third. Pronotum broadly

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campanulate, as broad as long. Median suture very distinct and
free of pubescence. Basal impressed line distinct, pubescent. Base
of pronotum notched at middle. Hind margin with denser pubes-
cence. Anterior legs of male not modified. Posterior tibial spurs
both slender, spiniform.

Type: Lectotype 9, new designation, M. C. Z. No. 4995, exam-
ined.
Type locality: Colorado.

Additional localities: Manitoba: Olds. Alberta: Winnipeg.
Washington: Blewett, Signal Peak. Idaho: Lava Hot Springs.
Montana : Maiden City. Wyoming: Bridger Basin. Colorado:
Mt. Manitou, Marshall Pass, Skyway 10,000 ft., Pikes Peak
10,000 ft. Halfway House, Mt. Lincoln, Alma, Little Willow Ck.
- Gun Co.

Epicauta immerita Walker

Epicauta immerita Walker, 1866, Lord's Naturalist Vancouver Is., etc.

Length : 6 to 9 mm. Closely related to pruinosa. The pubescence
is twice as dense and longer so that the color is most cinereous, with
very little black background showing through. The margins of the
elytra are a little more densely pubescent. I can find no other dif-
ferences in the series at hand. In this species, the pubescence is
easily rubbed off so that large black areas may be exposed.
Type: presumably in the British Museum.
Type locality: Vancouver Island.

Additional localities: British Columbia: Cowston. Washing-
ton: Yakima Valley. Oregon: The Dalles, Umatella. Occurs
in June.

Epicauta piceiventris Maydell

Epicauta piceiventris Maydell, 1934, Trans. Am. Ent. Soc. 60:327.
Epicauta elongatocalcarata Maydell, 1934, op. cit. :328.

Length: 8 to 12 mm. Black, with no visible pubescence above.
There is fine, short pubescence in the punctures, visible under some
magnification. Below, the pubescence is fairly short, more dense,
dark brown. Resembles a denuded pruinosa but is undoubtedly
distinct. No signs of abrasions were evident in the series examined
and all of the pruinosa specimens had cinereous, not brown,
pubescence.

Head triangular, densely and deeply punctured, the intervals

WERNER: NORTH AMERICAN EPICAUTA 473

finely punctulate, therefore dull. Median suture distinct. Antennal

calluses small, denuded, smooth. Antennae reaching to the basal

fifth of the elytra, twice as long as an anterior tibia. First segment

moderately stout, reaching two-fifths across the eye; second stout,

half as long as first; third a little longer than first, more slender;

fourth to tenth three-fifths as long as third; last as long as third.

Third to last segments almost equal in thickness. Pronotum

quadrate, slightly broader than long. Sides diverging gradually to

four-fifths from base, then converging abruptly. Disc flattened.

Median suture and basal impressed line distinct. Surface as on

head but more shiny posteriorly. Elytra more shallowly punctured,

more sparsely punctulate, therefore more shiny. Anterior legs of

male not modified. Posterior tibial spurs quite long, slender,

spiniform.

Type: Holotype & , M. C. Z. No. 23499, examined.

Type locality: Utah.

of elongatocalcarata

Type: Holotype 9 , M. C. Z. No. 23498, examined.

Type locality: Atlanta, Idaho.

Additional localities: Montana: Darby 7000 ft., Gallatin Co.
5000 ft., Gallatin Nat. For. Wyoming: National Park, Bridger
Basin. Idaho: Bingham Co. Utah: Provo. Colorado: Sum-
mit, Veta Pass. Occurs from June to August.

Epicauta sanguinicollis Horn

Epicauta sanguinicollis Dejean, 1833, Cat. des Coleop. 2nd ed. :225, (nomen
nudem).

Lytta sanguinicollis Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:344. (de-
scribed from a manuscript figure so not valid).

Epicauta sanguinicollis Horn, 1873, Proc. Am. Phil. Soc. 13:103, (quotes only
Le Conte reference); 1874, Trans. Am. Ent. Soc. 5:37 (first valid descrip-
tion, from a single specimen.)

Length: 6 to 8 mm. Head and pronotum bright rufous, shiny.
Elytra black, with the margins and an oblique humeral stripe
cinereous-pubescent.

Head subquadrate, sparsely but deeply punctured, very sparsely
pubescent. Median suture distinct only on occiput. Front broadly
impressed, darker than rest of head. Mouth parts black. Eyes
prominent, broadly oval, oblique. Antennae uniform in thickness,
twice as long as an anterior tibia. First segment stout, reaching to

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the middle of the eye; second not quite half as long as first; third
almost as long as first two; rest three-fifths of third. Pronotum
broadly campanulate, punctured like head. Median suture dis-
tinct. A narrow median impression present. Basal impressed line
distinct. Line on elytra from humerus to near the inner hind angle.
Below entirely black, with scattered cinereous pubescence. Male
anterior legs not modified. Posterior tibial spurs slender, sticklike.
Type: Holotype 9 , A.N.S.P., examined.
Type locality: Florida.

Additional localities: Florida: Alachua (on sea-island cotton),
Citrus Co., Sumpter Co., Lake Poinsett.

Epicauta puncticollis (Mannerheim)

Fig. 22

Lytta -puncticollis Mannerheim, 1843, Bull. Moscou 16:288. Le Conte, 1853,

Proc. Acad. Nat. Sci. Phila. 6:336.
Epicauta puncticollis, Horn, 1873, Proc. Am. Phil. Soc. 1,3:97.

Length: 6 to 13 mm. Black, with scattered erect pubescence.
Punctures on the pronotum deep and more or less dense. Surface
shiny. Apical segments of the antennae thicker than the middle
segments. Outer division of the tarsal claws bent only near the tip.
Inner division almost as long as the outer.

Head subquadrate. Median suture fine but distinct. Antennal
calluses not prominent, punctured. Eyes rather prominent, quite
broad, transverse, excavated. Antennae longer than in oblita and
with the segments not nearly as closely articulated, reaching to the
basal fifth of the elytra. First segment moderately stout, reaching
half-way across the eye; second half as long; third almost as long
as the first two, quite slender, longer and more slender than in
oblita; rest about two-fifths as long as the third, fairly loosely
articulated, becoming thicker toward the apex. Pronotum sub-
campanuliform, one-fourth longer than broad. Median suture
very distinct. Basal impressed line not as distinct. Elytra usually
with several feebly raised lines. Male anterior legs not modified.
Posterior tibial spurs slender, sticklike.

Type: specimens probably in the Mannerheim Coll. in Moscow.
Type locality: California.
Additional localities: California to Washington, extending east

into Nevada and Montana. A single specimen in the Purdue

WERNER: NORTH AMERICAN EPICAUTA 475

University collection from Jackson Co., Indiana. This specimen

is probably a stray brought in accidentally.

The pubescence varies from very short to very long. All of the
short-haired specimens are from California. Long-haired speci-
mens make up most of the California specimens and all of the series
from the rest of the range.

Epicauta oblita (Le Conte)

Fig. 23

Lytta oblita Le Conte, 1851, Ann. Lye. Nat. Hist. N. Y. 5:162; 1853, Proc.

Acad. Nat. Sci. Phila. 6:339.
Epicauta oblita, Horn, 1873, Proc. Am. Phil. Soc. 13:97.

Length: 10 to 11 mm. Black, with short, erect pubescence.
This species is shinier than puncticollis, has the punctures on head
and pronotum fairly dense and deep. The antennae have the
apical segments slightly more slender than the middle segments.
Tarsal claws as in puncticollis.

Head subquadrate, shiny. Median impressed line fine but dis-
tinct. Antennal calluses small, punctured. Eyes rather prominent,
oval, excavated. Antennae reaching almost to the base of the
elytra. First segment stout, reaching one-third across the eye;
second more slender, half as long as first ; third slightly shorter than
first two; rest two-fifths as long as third, becoming slightly thicker
at the middle and more slender toward the apex. Pronotum sub-
quadrate, a little longer than broad. Median suture fine. Basal
impressed line distinct. Pubescence below denser than above.
Anterior legs of male not modified. Posterior tibial spurs slender,
sticklike.

Type: Lectotype 9, new designation, M. C. Z. No. 4993, exam-
ined.
Type locality: San Francisco, California.
Additional localities : Sacramento, California, Aug.

Epicauta barberi Werner

Epicauta barberi Werner, 1944, Psyche, 50:69.

Length: 12 to 13 mm. Black, fairly densely clothed with semi-
erect black pubescence. Head and pronotum shiny. The head is
much more densely and finely punctured than in puncticollis or

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oblita. The outer division of the tarsal claws is curved from the
base and the inner is not more than three-fourths as long as the
outer.

Head subquadrate, finely and densely punctured. Median suture
fine but distinct. Antennal calluses small, inconspicuous. Eyes
fairly large, moderately broad, excavated. Antennae one and three-
fourths times as long as an anterior tibia, thicker at the apex than
at the middle. First segment fairly stout, reaching one-fourth
across the eye; second small, half as long as first; third as long as
first two; fourth and following two-thirds as long as third, gradu-
ally increasing in thickness. Pronotum quadrate, slightly longer
than broad. Median suture fine but distinct, extending three-
fourths from base. Basal impressed line distinct. Punctures as
fine as on the head but not as dense. Elytra longer than in punc-
ticollis or oblita. Anterior legs of male not modified. Posterior
tibial spurs slender, the inner spiniform, the outer sticklike.
Type: Holotype o\ U.S.N.M., examined.

Type locality: La Panza, California, May 17-38, C. C. Wilson.
Additional records: 1 o71 San Luis Obispo, California, May 16-25.

C, C. Wilson. For Mr. H. S. Barber of the U. S. National

Museum.

Epicauta kansana Warner
Epicauta kansana Werner, 1944, Pyche, 50:70.

Length : 1 0 to 1 1 mm. Black, with scattered semi-erect but rather
short pubescence, resembling oblita and puncticollis. Head and
pronotum shiny, the head more densely punctured than the pro-
notum. Inner division of the tarsal claws as long as the outer, but
very slender. The outer curves near the tip. Fourth segment of
the antennae longer than the fifth.

Head subquadrate, shiny, rather densely and finely punctured,
the intervals with scattered punctulation. Median suture fine and
indistinct. Antennal calluses small, smooth. Eyes broad, exca-
vated. Antennae one and two-thirds times as long as an anterior
tibia, thicker at the apex than at the middle. First segment rather
slender, extending one-fifth across the eye; second small, half as
long as first ; third slender, as long as first two; fourth three-fourths
as long as third; rest slightly shorter than fourth. Pronotum sub-
quadrate, a little longer than broad. Median suture fine, extend-
ing three-fourths from base. Basal impressed line distinct. Surface

WERNER: NORTH AMERICAN EPICAUTA 477

as on the head but more sparsely punctured. Anterior legs of male

not modified. Posterior tibial spurs slender, the inner spiniform,

the outer sticklike. Tarsi more slender than in puncticollis, oblita,

or barberi.

Type: Holotype cf , U.S.N.M., examined.

Type locality: Sedgwick Co., Kansas. J. C. Warren, June 4,

1910.
Additional records: several more taken on June 4 and April 7.

Epicauta calcarata Werner
Epicauta calcarata Werner, 1944, Psyche 50:70.

Length: 7 to 9 mm. The mouth parts of this species would place
it in the genus Gnathos pasta, but the antennae are similar to those
of E. sericans while those of G. mimetica are very much like those
of virgulata, of the old genus Macrobasis. For this reason and
because the genitalia are of the Epicauta form, I have discarded
Gnathos pasta as a natural group. It seems rather to be a stage of
evolution within Epicauta.

Black, completely, rather densely clothed with tannish-grey
pubescence. Pronotum large and conspicuously globose. Tibial
spurs heavy and incurved.

Head triangular. Median suture fine but distinct. Antennal
calluses not denuded. Eyes moderately broad, slightly excavated.
Labrum sparsely pubescent, deeply notched but not as deeply as
in mimetica. Mandibles long and rather slender, crossing at the
tip (meeting in mimetica.) This crossing is due to an elongated tip
but the mandibles are otherwise similar in the two species. Anten-
nae short, reaching to a little beyond the base of the elytra, one
and three-fourths times as long as an anterior tibia, almost uniform
in thickness. First segment stout, reaching the middle of the eye,
clothed with cinereous pubescence; second small, half as long as
first; third as long as first; fourth to tenth subequal in length and
thickness, two-thirds as long as third; last as long as third. All but
the first are black. Pronotum subglobose, almost as wide as head.
Median suture and basal impressed line distinct. There is a trace
of a pair of midlateral impressions. Scutellum quite small. Legs
rather stout and short. Both anterior tibial spurs large, slightly
incurved, the inner slightly the stouter. Middle and posterior
tibial spurs also larger than normal, slightly incurved. Outer spur

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of hind tibiae slightly broader than inner. All spurs sticklike, blunt.

Anterior tarsi of male not modified.

Type: Holotype 9 , M. C. Z. No. 26065, examined.

Type locality: Roswell, New Mexico.

Additional localities: New Mexico: Carlsbad, Dim Lake-Pecos

Val. Texas: Culbertson Co., Ft. Stockton. Taken in August

and September. Scarce.

Epicauta heterodera Horn

Epicauta heterodera Horn, 1891, Trans. Am. Ent. Soc. 18:43.
Epicauta watsoni Blatchley, 1918, Can. Ent. 50:58.

Length: 7 to 9 mm. This species exhibits a remarkable reversal
of the ordinary. In it the female, instead of the male, departs
from the usual pattern. The pronotum is denuded, smooth in the
female, pubescent in the male. It is flattened in both sexes. Pubes-
cence pale to dark tannish-cinereous, quite dense, short.

Head triangular. Median suture distinct. Antennal calluses
small, not denuded. Eyes prominent, broadly oval to almost
circular, with an impression of very small to medium extent on
the anterior margins near the base of the antennae, none by the
mandibles. Occiput slightly notched. Antennae reaching to the
basal fifth of the elytra, two and one-fourth times as long as an
anterior tibia. First segment stout, reaching one-third across the
eye; second more slender, half as long; third as long as first two;
rest as long as the first, increasing slightly in thickness toward the
apex. Pronotum broadest at the base, three-fourths as wide as the
base of the elytra. Less than half as wide at the apex, tapering
from the base but bulging toward the middle, especially in the
male. About as long as width across base. Median suture absent.
Basal impressed line fine. A shallow median impression and a pair
of lateral impressions, connected near the base, may be present.
Pronotum of female smooth, shiny, with scattered faint punctures.
Elytra often appearing faintly margined because of the denser
pubescence on the edges. Anterior legs of male not modified.
Posterior tibial spurs slender, spiniform.

Type: Lectotype 9, new designation, A.N.S.P. No. 8093, exam-
ined.
Type locality: Florida.
of watsoni

WERNER: NORTH AMERICAN EPICAUTA 479

Type: Holotype 9 , Blatchley Coll., Purdue U., examined.

Type locality: Gainesville, Florida.

Additional localities: Florida: Kissimmee, De Funias Spgs.,

Monticello, Gainesville, Crestview, Lake Co., Alachua Co., St.

Augustine. Georgia: Thomasville, Tifton. Alabama: Kushla,

Mobile. Mississippi: Lucedale.

Epicauta californica nomen novum

Fig. 11

Lytta maura Le Conte, 1851, Ann. Lye. Nat. Hist. N. Y. 6:162; 1853, Proc.

Acad. Nat. Sci. Phila. 6:339.
Epicauta maura, Horn, 1873, Proc. Am. Phil. Soc. 13:97, nee Faldeman, 1833.

Length: 8 to 10 mm. Black, rather shiny, sparsely clothed
above with short black pubescence. Underside of head always
with some white pubescence, as in alphonsii. Femora of male with
marginal long white pubescence. Middle femora of male curved,
as in alphonsii.

Head subtriangular, sparsely and deeply punctured, finely
rugulose. Median suture distinct. Antennal calluses not strongly
elevated, less strongly rugulose than rest of head. Eyes rather
prominent, narrow, oblique, excavated. Antennae slender in the
male, reaching to the middle of the elytra, two and three-fourths
times as long as an anterior tibia, in the female reaching to the
basal third of the elytra, twice as long as an anterior tibia. Male
antennae: first segment reaching two-thirds across the eye; second
two-fifths as long as first, slender; third as long as first two; fourth
and following slender, almost uniform in thickness, three-fifths as
long as third. Antennae of female with segments of much the
same proportions as in the male but the whole shorter and more
slender. The first segment reaches one-third across the eye.
Pronotum broadly campanulate, slightly longer than broad.
Surface finely rugulose, sparsely punctured, not as deeply as on
the head. Median suture fine, indistinct. Basal impressed line
deep. Elytra duller than head and pronotum.

Below black. Male with pubescence on anterior femora and tro-
chanters mainly white. Middle and posterior tibiae of male flat-
tened and smooth behind, this surface margined with long white
pubescence; middle tibiae quite strongly curved. Anterior tarsi of

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male as in alphonsii but with the first segment narrower. Posterior

tibial spurs slender, spiniform.

Type: Lectotype cf, new designation, M. C. Z. No. 5099, exam-
ined.

Type locality : California.

Additional localities: California: Adobe Falls-San Diego Co.,
Spreckles, Riverside, Colorado Desert, Cabagon, San Ber-
nardino, Pasadena. Occurs in September and October.

Epicauta alphonsii Horn

Epicauta. alphonsii Horn, 1874, Trans. Am. Ent. Soc. 5:38; 1875, Trans. Am.
Ent. Soc. 5:153.

Length: 8 to 9 mm. Black, the elytra conspicuously margined
with white. Middle femora of male strongly curved. Close to
californica.

Head ovate in the male, subtriangular in the female, finely
rugulose, with scattered large punctures. Median suture fine.
Eyes oblique, moderately narrow, excavated. Antennae three
times as long as an anterior tibia in the male, two and one-half
times in the female. The proportions of the antennal segments
are the same in both sexes but in the male segment three and beyond
are heavier, tapering gradually. First segment reaching one-third
across the eye; second half as long as first; third two and one-half
times as long as second ; rest two-thirds as long as third. Pronotum
as broad as long, quadrate, finely rugulose and sparsely punctured.
There are some white hairs on the basal and apical margins.
Median suture feeble. Basal impressed line strong. Pubescence on
elytra sparse except on margins, where it is dense and white. Legs
of male as in californica but with a tuft of long white hairs exter-
nally at the base of the anterior tibiae, these hairs each with an
almost right-angled bend toward the apex. First segment of male
anterior tarsi much compressed, broad. Rest of segments also
compressed but narrow. Posterior tibial spurs long, slender,
spiniform.

Type: Holotype 9 , M. C. Z. No. 125, examined.
Type locality: California.
Additional localities: California: Hesperia — Mojave Desert,

Los Angeles Co. Occurs in the fall. Seems to be scarce.

WERNER: NORTH AMERICAN EPICAUTA 481

Epicauta caviceps Horn
Epicauta caviceps Horn, 1873, Proc. Am. Phil. Soc. 13:99.

Length: 6 to 9 mm. Black, moderately densely clothed with
gray to almost white pubescence. Occiput with a pair of prominent
callosities.

Head subquadrate, pubescent except for a narrow median line,
weak antennal calluses and a border around the eyes, which is
narrow except at the inner posterior margin where it is broadened
to contain a shallow depression. Occiput to front with a strong
U-shaped impression, which divides a strong bulge on the occiput,
producing a pair of prominent callosities. Eyes oblique, narrow,
especially above the antennal excavation, but not pointed, attain-
ing the occipital callosities. Antennae moderately slender, tapering
slightly toward the apex. First segment moderately stout, reaching
one-third across the eye; second half as long as first; third twice as
long as second; rest a little shorter than third, but for the last,
which equals the third. Pronotum about as broad as long, with
the sides parallel for the basal two-thirds, then converging abruptly.
Median suture and basal impressed line distinct and with a broad
oblique impression extending from just in front of the middle
almost to the anterior angles. This is usually supplemented by a
broad excavation near the posterior angles. Surface clothed with
dense pubescence, very short near the apex. Elytra with a black
scutellar spot. A raised area extends from the suture at a third
from the base, where there is a small hump, obliquely forward to
the humeral angles so that there appears to be a triangular flat
area at the base of the elytra. A narrow raised area may extend
straight back from the humeral angles. Posterior face of femora
denuded and shiny in male. Anterior tibial spurs of male short,
stout, the inner slightly incurved. Posterior tibial spurs slender,
spiniform. Third and fourth abdominal sternites sometimes with a
median posterior black spot. Fifth often with an elongate median
mark. The second to last abdominal segments have an anterior
lateral black spot near the edge, which may be concealed in
shrunken specimens.
Type: Lectotype cf, new designation, Horn Coll., A.N.S.P. No.

8094, examined.
Type locality: Arizona.
Additional localities: Arizona: Phoenix, Tucson, Sabino Canon-

Sta. Catalina Mts., Pipe Spring — Coconino Co., Peach Spg.

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Utah: Leeds, St. George. Seems to be a fall species as all of
the records are in September and October. Apparently fairly
common where it occurs as several large series have been seen
from the Phoenix region.

Epicauta impressifrons Van Dyke
Epicauta impressifrons Van Dyke, 1929, Bull. Br. Ent. Soc. 24:12.

Length: 5 to 7 mm. Black, rather sparsely clothed with long,
easily denuded, appressed cinereous pubescence. Resembles
caviceps Horn but has the occipital callosities small and lacks
velvety pubescence on the pronotum.

Head roughly triangular, sparsely but rather deeply punctured,
shiny. Median suture distinct. Occiput impressed, but not as
deeply as in caviceps Horn so that the occipital callosities are some-
what indistinct. Eyes oblique, narrow, but not as narrow as in
caviceps, surrounded by a quite broad denuded border, which
merges anteriorly with the smooth antennal calluses. A feeble
impression at inner posterior margin. Antennae slender, reaching
to the middle of the elytra, twice as long as an anterior tibia. First
segment reaching to middle of eye; second half as long as first;
third as long as first two; fourth as long as first; rest just per-
ceptibly toward apex. Pronotum bulging. Basal impressed line
distinct ? but median suture absent. There is a suboval depression
on the middle of the disc and another just before it. These vary in
depth. Elytra with black scutellar and humeral spot. Anterior
tibiae of male with two fairly stout, incurved spiniform spurs.
Posterior border of all of the femora of the male flattened, glabrous,
shiny. Middle and posterior femora with this area margined above
by long cinereous hairs. Posterior tibial spurs slender, sticklike.
Type: Holotype, Mus. Cal. Acad. Sci. No. 2592.
Type locality: Palm Springs, Riverside Co., California.
Additional localities: California: Cabazon, Indio. Specimens

taken in October, November and March.

Epicauta rileyi Horn

Epicauta Rileyi Horn, 1874, Trans. Am. Ent. Soc. 5:37.

Length: 7 to 9 mm. Black, densely clothed with yellow-cinere-
ous pubescence. A deep pit on the inner margin of the eyes.

WERNER: NORTH AMERICAN EPICAUTA 483

Head roughly triangular, moderately densely to densely pubes-
cent, except for a denuded margin around the eyes, which is very
narrow in front up to one-third the width of the eye behind. A
deep pit on the inner margin of each eye. Occiput with a denuded
median line and impression but this is not prominent. Vertex
slightly bulged. A partly denuded bulge just above the clypeus.
The eyes are very peculiar in being almost absolutely smooth and
are moderately broad, uniform, transverse, prominent. Antennae
reaching to the basal third of the elytra. First segment stout,
reaching one-third across the eye, densely pubescent, black in
front, yellow behind; second two-thirds as long as first, more
slender; third about twice as long as second; rest subequal, three-
fourths as long as third, except the last, which is equal to the third.
Pronotum a little broader than long. Basal impressed line and
median suture distinct. With a lateral pair of broad longitudinal
impressions. The pubescence may be black in these impressions,
especially anteriorly. Elytra with black scutellar spot. Suture
slightly elevated just beyond the middle. Second to fourth abdo-
minal sternites with a posterior median black spot. First to fifth
with an anterior lateral spot near the edge. The lateral spots
sometimes are lacking. Anterior tibial spurs of male shorter but
not broader than in the female, spiniform. Posterior tibial spurs
slender, spiniform. The pubescence is sparse but not completely
lacking on the posterior face of the femora of the male.
Type: Lectotype 9, new designation, Horn Coll., A.N.S.P. No.

8095, examined.
Type locality: Arizona.
Additional localities: Arizona: Phoenix, Nogales, Atascasa Mts.,

Reef, Ft. Huachuca, Baboquivari Mts., Tucson. Sonora:

Arispe. Specimens taken in October and November.

Epicauta straba Horn

Epicauta straba Horn, 1891, Trans. Am. Ent. Soc. 18:42.
Epicauta foxi Van Dyke, 1929, Bull. Br. Ent. Soc. 24:127.

Length: 5 to 8 mm. Black, clothed above with short pubescence.
Eyes oblique, strongly pointed at the inner margin. I have seen
no such eye form in any other species. Pubescence black in main
part. There is always cinereous pubescence on the femora and
trochanters. The pubescence of the underside of the body varies
from entirely black in the typical form to entirely cinereous in the

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form foxi Van Dyke, which also has the margins of the elytra
cinereous pubescent. All intergrades occur in a series from Adobe,
San Diego Co., California.

Head triangular, densely and rather deeply punctured, except
near the midline, with the intervals shiny, smooth. There is a flat
area behind each eye. This causes the middle of the occiput to
appear bulged. Median suture fine but distinct, supplemented by a
narrow but moderately deep impression on the occiput. Eyes
narrow, obliquely directed backward, reaching almost to the occi-
put. Antennae tapering gradually from the base, reaching to the
middle of the elytra. First segment stout, reaching one-fourth
across the eye; second slender, three-fifths as long as first; third
slightly longer than first two; fourth three-fourths as long as third;
rest just perceptibly shorter than fourth. Pronotum quadrate, as
broad as long. Median impressed line very distinct, but median
suture fine and indistinct. There is a shallow impression on the
middle of the disc and a pair of very shallow and indistinct im-
pressions on the sides. Femora of male flattened behind, smooth
and shiny; middle and posterior femora fringed above with long
cinereous pubescence. Anterior tibiae of male with two slender,
spiniform spurs. Posterior tibial spurs slender, spiniform.
Type: Lectotype cf, new designation, Horn Coll. A.N.S.P. No.

8096, examined.
Type locality: San Bernardino, California.
of foxi Van Dyke
Type: Holotype, Cal. Acad. Sci.

Type locality: Jacumba, San Diego, Co., California.
Additional localities: California: Pasadena, Los Angeles Co.,
Pom Mts., near Claremont, San Diego Co. Washington:
Some specimens in the Bowditch Coll. (M. C. Z.) labelled
"Wash." Many of the state labels in the Bowditch collection
have been found to be erroneous. Specimens collected in Sep-
tember and October.

Epicauta excavatifrons Maydell
Epicauta excavatifrons Maydell, 1934, Trans. Am. Ent. Soc. 60:330.

Length: 7 to 9 mm. Black, fairly densely clothed with white
pubescence. The surface is visible throughout so that the species
appear gray. Pronotum with a prominent tubercle in the middle.

Head subtriangular, with a broad depression back from each eye

WERNER: NORTH AMERICAN EPICAUTA 485

and a rather sharp depression between the eyes, back from each
pair of pits below the middle of the front. These depressions form a
short ridge back from the inner margin of each eye. Median suture
distinct. Midline, margin of eyes (broader behind) and antennal
calluses denuded, shiny. Eyes moderately narrow, broader toward
the center than outward, smooth, but not as strikingly so as in
rileyi Horn, oblique. First segment of antennae short, stout,
flattened on its inner surface as though modified to fit over the
eye; second three-fifths as long as first, also flattened on inner
surface; third as long as first two; fourth and following three-fifths
as long as third, except the last, which is three-fourths the third.
Pronotum with distinct basal impressed line. Sides straight and
slightly expanding three-fourths from the base, then converging
abruptly. Before the middle there is a prominent longitudinal
hump. Elytra without a scutellar spot, humped slightly at the
basal third. From this hump a pair of low ridges extends anteriorly
to half-way between the scutellum and the humerus. Some
specimens have a slight hump on the suture two-thirds from the
base also. Anterior tibial spurs of male shortened but not otherwise
modified, spiniform. Posterior tibial spurs slender, spiniform. All
of the femora and trochanters of the male are partly denuded
behind, with long hairs, mainly dorsally, but also on the partly
denuded area.

Type: Holotype 9, A.N.S.P. No. 8183, examined.
Type locality: Ocala, Marion Co., Florida.

Additional localities: Florida: Gainesville. Alabama: Mobile.
Mississippi: Lucedale. All of the specimens collected in Sep-
tember and October. Dr. Dietrich, who collected the Lucedale
series, says that he found the species abundant at Lucedale on
the stems of low grass.

Epicauta rehni Maydell
Epicauta rehni Maydell, 1934, Trans. Am. Ent. Soc. 60:329.

Length: 6 to 9 mm. Black, densely clothed with flavo-cinereous
pubescence, as in rileyi Horn. Easily distinguished from that
species by the lack of a pit at the inner margin of the eye and lack
of a scutellar spot on the elytra.

Head subtriangular. Vertex notched by the median suture,
which extends to the level of the eyes and is continued forward by
a narrow denuded line. A weak ridge extends from the inner margin

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of the eyes obliquely backward to near the occiput, then runs
close to the midline for a short distance. Eyes narrow, oblique,
rather small. Antennae reaching to middle of elytra, two and two-
thirds times as long as an anterior tibia, almost uniform in thick-
ness. First segment moderately stout, reaching half-way across the
eye; second more slender, three-fourths as long as first; third twice
as long as second. The first three segments have some pale pubes-
cence at the base. Fourth and following three-fourths as long as
third, diminishing just perceptibly in thickness. Pronotum
quadrate, with sides impressed slightly near the base. Median
suture obscured by the pubescence. Basal impressed line distinct.
Pubescence more or less erect, pointed in many different direc-
tions. Elytra without markings. Midline raised slightly on basal
half and there is a feeble basal elevation. Underside without spots.
Anterior tibiae of male with two stout spiniform spurs. Outer
posterior tibial spur slender, pointed, inner sticklike. Posterior
edge of male middle and posterior femora and trochanters denuded,
shiny, margined above with long pubescence.
Type: Holotype 9, A.N.S.P. No. 8184, examined.
Type locality: Schaeffer Canyon, Baboquivari Mts., Arizona.
Additional localities: Arizona: Baboquivari Mts., Tucson,
Phoenix. Occurs in September and October. Apparently scarce.

Epicauta wheeleri Horn

Epicauta wheeleri Horn, 1873, Proc. Am. Phil. Soc. 13:101, 107 (note).
Epicauta wheeleri Ulke, 1875, in U. S. Geog. and Geol. Surv. W. of 100th
Merid. 5:812, 825, PI. 61, fig 4.

Length: 6 to 10 mm. Black, finely clothed with cinereous pubes-
cence but for the dense rich rufous pubescence on the pronotum
and a black patch at the base of the elytra, which is covered when
the prothorax is raised.

Head subquadrate, shiny, sparsely but deeply punctured.
Median suture distinct, margined by a narrow denuded area. Eyes
prominent, moderately narrow, oblique. Antennae uniform in
thickness, reaching to the basal fourth of the elytra, two and one-
third times as long as an anterior tibia. Pronotum subquadrate, a
little longer than wide. Median suture and basal impressed line
distinct. With a pair of shallow impressions behind the middle.
Pubescence dense, erect, rufous, margined all around the pronotum
by cinereous pubescence. Scutellum densely clothed with cinereous

WERNER: NORTH AMERICAN EPICAUTA 487

pubescence. Base of the elytra from scutellum half-way to the

humerus with black pubescence, not visible when prothorax is

raised. Below cinereous. Black spot on midline of second to fourth

abdominal sternites. Legs cinereous-pubescent. Femora with

apical black mark, sharply margined. Tibiae also with black apices

but not so distinctly margined. Male anterior tibiae with two

slender, spiniform spurs. Inner posterior tibial spur slender, spini-

form, outer sticklike.

Type: Holotype, Horn Coll., A.N.S.P., examined.

Type locality : Arizona.

Additional localities: Arizona: Phoenix.

Epicauta diversipubescens Maydell
Epicauta diversipubescens Maydell, 1934, Proc. Am. Ent. Soc. 60:333.

Length: 6 to 9 mm. Black, densely clothed with pale tannish-
cinereous pubescence, except for black scutellar spot, extending
narrowly across the base of the elytra and black at angles of the
pronotum. Pubescence on disc of pronotum short and velvety.

Head quadrate. Median suture distinct, bordered by a narrow,
smooth denuded line. Antennal calluses small, not denuded. Eyes
fairly prominent, rather narrow, transverse. Antennae reaching
to the basal fourth of the elytra, two and one-third times as long
as an anterior tibia, almost uniform in thickness. First segment
short, stout, reaching about one-third across the eye in the male, a
little shorter in the female; second short, half as long as first; third
as long as first two. First and second segments and basal half of
third with scattered cinereous hairs. Fourth and following three-
fourths as long as third, practically equal in thickness. Pronotum
as broad as long, broadly campanulate. Median suture distinct,
supplemented by a narrow impression. Basal impressed line dis-
tinct. A broad denuded spot on the anterior angles and usually a
smaller one on the posterior angles. The denuded surface is dull.
The anterior spot is often extended toward middle by blackish
pubescence. There is always a narrow area of pale pubescence
before the anterior spots. Elytra with black scutellar spot, extended
narrowly across the base. Pale below, with an elongate spot on the
middle of the second to fifth abdominal sternites. An anterior
lateral black spot on the second to fifth sternites, usually concealed
by the elytra. Femora and tibiae with denuded tips, sharply de-

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fined. Anterior tibiae of male with two spiniform spurs, the outer
longer. Middle and posterior femora of male denuded behind,
shiny, fringed above with long hairs. Posterior tibial spurs stick-
like, the outer broader.

Type: Holotype, Leng Coll., U.S.N.M., examined.
Type locality: Albuquerque, New Mexico.
Additional localities: Texas: El Paso.

Epicauta aspera Werner
Epicauta aspera Werner, 1944, Psyche 50:70.

Length: 7 to 10 mm. Black, densely clothed with cinereous,
rarely ferrugineous, pubescence. Elytra with a scutellar spot,
extended narrowly across the base. Abdominal sternites with a
row of median black spots. Head not modified.

Head subquadrate, deeply and densely punctured and alutaceous,
except for the very small smooth antennal calluses and shallowly
impressed midline. Eyes moderately broad, normal. Antennae two
and one-third times as long as an anterior tibia, cylindrical, taper-
ing slightly. First segment short, stout, barely reaching the middle
of the eye; second half as long as first, much narrower; third two
and one-third times as long as the second, with a few cinereous
hairs; rest a little shorter than the third. Sides of pronotum sub-
parallel for the basal two-thirds, then converging abruptly. Sculp-
ture of pronotum similar to that of head. Suture distinct, discerni-
ble to one-fourth from base, but very faint. A broad, shallow de-
pression at the base. Second to fourth and sometimes fifth abdom-
inal sternites with a median black spot, sometimes transverse,
and a small spot anteriorly at the sides, usually concealed by the
elytra. One specimen lacks the median spots. Apices of femora,
tibiae, first tarsal segment and all of the rest of the tarsi black.
Male anterior tibiae with two slender spiniform spurs. Posterior
tibial spurs sticklike, rather long. Middle and posterior femora of
male denuded behind and with long pubescence fringing the
denuded area.

Type: Holotype d\ M. C. Z. No. 26063, examined.
Type locality: Salida, Colordado.
Additional localities: Colorado: Masonville, Ft. Collins, Boulder,

Morison, Boulder, B. Rotger C. R. Texas: Davis Mts. New

Mexico: Jemez Springs, Barton. Arizona: White Mts., William-

WERNER: NORTH AMERICAN EPICAUTA 489

son Valley, Copper Basin near Prescott. Occurs from August to
November.

Epicauta immaculata (Say)
Fig. 32, 33

Lytta immaculata Say, 1823, Journ. Acad. Nat. Sci. Phila. 3:304. Le Conte,

1853, Proc. Acad. Nat. Sci. Phila. 6:342.
Lytta articularis Say, 1823, Journ. Acad. Nat. Sci. Phila. 3:304.
Lytta fulvescens Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:447.
Macrobasis immaculata, Horn, 1873, Proc. Am. Phil. Soc. 13:93; Trans. Am.

Ent. Soc. 12:110.

Length : 1 2 to 20 mm. Black, rather densely clothed with f errugi-
neous to cinereous pubescence. No black markings on the elytra.

Head subquadrate. Median suture distinct but not conspicuous.
Antennal calluses rather small, denuded, shiny. Eyes only moder-
ately prominent, transverse, moderately narrow. Antennae taper-
ing from the base, heavier, shinier and more compressed in the
male than in the female. They reach to the basal third of the
elytra and are two and one-half times as long as an anterior tibia.
First segment exceeding the eye by one-fourth its length in the
male, just perceptibly shorter in the female; second two-thirds as
long as first; third and following one-fourth longer than second,
gradually decreasing in thickness. The first three segments usually
have some pale pubescence on the posterior margin. Pronotum
quadrate, slightly bulging. Median suture distinct and a narrow
median impression supplements it. Basal impressed line distinct.
Visible portion of scutellum small. Tips and often the outer edges
of femora, outer edges or all of tibiae and all of tarsi with black
pubescence. Anterior tibiae of male with two slender spiniform
spurs. Male anterior tarsi with the first segment just perceptibly
longer than in the female, not otherwise modified. Posterior tibial
spurs slender, spiniform.

Type : Say's type lost. Neotype cT and 9 neoparatypes in M. C. Z.
Type locality: "Inhabits Arkansas." Of neotype: Cambridge

Nebraska.
of fulvescens Lee.
Type: Lectotype cf, new designation, Le Conte Coll. M. C. Z.

No. 4989, examined.
Type locality: Texas.
Additional localities: Texas: Reeves Co., Pecos, Gillespie Co.,

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Rio Grande City, Davis Mts., Kingsville, Alpine, Brewster Co.,
Clarendon, Corpus Christi, Victoria. New Mexico: Las Cruces.
Oklahoma: Lawton. Colorado: Coolidge, La Junta, Las
Animas, Ft. Collins. Kansas: Belvidere, Beeler, Rossville,
Grinnelle, Ft. Hayes, Ellis, Garden City, Finney Co. Nebraska:
Indianola, Cambridge, Bartley, Lincoln, Stratton. Iowa: Lewis,
Fairfield. Missouri: Ranken. Illinois: Zeigler. Indiana:
Clarke Co. State Forest, Jackson Co., Warrick Co. Kentucky:
Berheim Forest. Occurs from June to September. Apparently
common in the western part of its range.

Epicauta segmenta (Say)
Fig. 30, 31

Lytta segmenta Say, 1823, Journ. Acad. Nat. Sci. Phila. 3:303.
Lytta segmentata Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:342.
Apterospasta valida Le Conte, 1858, Journ. Acad. Nat. Sci. Phila. (2) 4:39.
Macrobasis segmentata, Horn, 1873, Proc. Am. Phil. Soc. 13:93; 1885, Trans.
Am. Ent. Soc. 12:110. Champion, 1892, Biol. Cent.-Am. 4 pt. 2:44.

Length: 12 to 24 mm. Black, with the pubescence short, mostly
black. Pubescence always white on the posterior margin of the
pronotum and there is usually some white pubescence on the coxae,
thoracic segments, back part of head and posterior margins of the
abdominal sternites.

This species is dimorphic. In one form the wings are as long as
the elytra and in the other only one-third to one-half as long and
usually much aborted. The reduced-winged form appears heavier
because the elytra are more bulbous and they may even overlap
slightly at the suture in mounted specimens. This led Le Conte to
describe the form as a distinct species, valida, and place it in a
separate genus, Apterospasta. However, the form seems to have
no geographical distinctions from the full-winged form, nor is it
limited to one sex. Small short-winged forms have been confused
with conferta in collections but they can easily be distinguished by
the white apices of the abdominal sternites.

Head subtriangular, with a very fine median suture. Antennae
four-fifths as long as an anterior tibia, slightly heavier in the male
than in the female. First segment reaching to hind margin of
eye or slightly beyond; second half as long as first; third one and
one-half times as long as first; rest a little shorter than the third,

WERNER: NORTH AMERICAN EPICAUTA 491

decreasing gradually in thickness. Pronotum subquadrate, a little
longer than broad, with a weak median suture and a very strong
basal impressed line. Elytra always entirely black. Tibial spurs
long and sticklike on all tibiae, not modified in the male. First
segment of anterior tarsi slightly broader in the male.
Type: Say's type lost. Neotype cf and neoparatype 9 in M. C. Z.
Type locality: "Purgatory River of the Arkansa." Neotype:

Fort Hayes, Kansas.
of valida
Type: Lectotype c?, new designation, Le Conte Coll., M. C. Z.

No. 4990, examined.
Type locality : Texas.

Additional localities: Nebraska: Indianola, Cambridge, Bartley.
Kansas: Ft. Hoge, Ellis. Texas: Alpine, Marfa, Chisos Mts.,
Davis Mts.; New Mexico: Tucumcari. Arizona: Chihuahua
Mts., Fish Creek — White Mts., Palmerlee. Occurs from May to
September.

Epicauta sublineata (Le Conte)

LyUa sublineata Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:447.
Macrobasis sublineata, Horn, 1873, Proc. Am. Phil. Soc. 13:94; 1885, Trans.

Am. Ent. Soc. 12:109.
Macrobasis megacephala Champion, 1892, Biol. Cent.-Am. Coleop. 4 (pt. 2):

402.

Length: 16 to 25 mm. This is one of the most variable species
of the genus as to color pattern. The holotype of sublineata has
the head cinereous, the pronotum but for a pair of suffused dark
patches also cinereous and the elytra with brownish suffusions
between four fine lines. A specimen in the Fall Coll. from Eagle
Pass is essentially like the type. One from Devil's River in the
U.S.N.M. is similar but with the elytral suffusions very light and
the dark marks absent from the pronotum. All these have dark
suffusions on the abdominal sternites. These specimens can be
distinguished from the female of longicollis, with which Dr. Horn
synonymized the species, by their proportionately larger head,
more slender antennae and by the presence of four dark lines on
the elytra. Albida sometimes has dark lines on the elytra but there
are only three when they occur and the basal segments of the
antennae are rufous. The four remaining specimens I have seen
have the elytra black with the tip cinereous. In one, the occiput,
but for the median line, is black. This one also has the spots on the

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pronotum and the abdominal bands enlarged and dark. In the
others, these spots are weak and they have scattered cinereous
pubescence on the elytra near the suture, at the base and near the
side margins. Some specimens may possibly be confused with
segmenta but that species has the elytra entirely black and has less
pale pubescence on the pronotum.

Ground color black. Head large, suboval, with a fine suture.
Eyes moderately narrow. Antennae long and slender, tapering
from the first segment, which just exceeds the hind margin of the
eye in the male and barely attains it in the female. Last segment
only one-half the diameter of the first. (It is three-fourths the
diameter of the first in the female of longicollis.) Antennae three
times as long as an anterior tibia. (Two and one-half times in
segmenta). Palpi pale rufous. Pronotum subquadrate, about
two-thirds as wide as the head. (At least four-fifths in segmenta).
Elytra with at least a humeral and scutellar spot, a narrow sutural
line and four lines on the disc black and with at least the apex
cinereous. Abdominal sternites with at least a darker suffusion
toward the middle of some. Anterior tibiae of male with two spini-
form spurs. Posterior tibial spurs moderately stout, blunt. The
pad of the first segment of the male anterior tarsi is limited to the
apical sixth so that the segment appears narrow.
Type: Holotype 9 , Le Conte Coll. M. C. Z. No. 4979, examined.
Type locality: Eagle Pass, Texas.
of megacephala Champion
Type: presumably in the British Museum.
Type locality: Monclova, Coahuila (Mexico)

Additional localities: Texas: Devil's River, San Diego, Gold-
thwaite. Three specimens taken in May, one in September.

Epicauta lauta (Horn)
Fig. 36
Macrobasis lauta Horn, 1885, Trans. Am. Ent. Soc. 12:108.

Length: 8 to 16 mm. Tannish, finely, rather sparsely but evenly
clothed with cinereous pubescence. Elytra with dark brown
scutellar and humeral spots.

Head rounded, densely and finely punctured. Median suture dis-
tinct but not conspicuous. Antennal calluses low, smooth and shiny.
Eyes large, prominent, transverse, moderately broad. Antennae

WERNER: NORTH AMERICAN EPICAUTA 493

moderately long, slender, tapering from the basal segment, black,
rather loosely jointed, not noticeably flattened. They attain the
basal fifth of the elytra and are three times as long as an anterior
tibia. Male with first segment reaching hind margin of eye, some-
what flattened and curved to fit over the eye; second and following
three-fifths as long as first, tapering gradually. Female with first
segment not flattened and only slightly curved, reaching three-
fourths across the eye; second short, slender, half as long as the
first; third and following three-fourths as long as first. Pronotum
one-fourth longer than broad, campanuliform. Median suture and
basal impressed line distinct. There is an inconspicuous transverse
impression at the base. Apices of abdominal sternites darkened.
Tips of femora, most of tibiae and all of tarsi also dark. Male
anterior tibia somewhat curved, denuded behind, with a single
spiniform spur. First segment of anterior tarsi of male a little
shortened, slightly contorted. Posterior tibial spurs moderately
broad, flattened.
Type: Lectotype d\ new designation, Horn Coll. A.N.S.P. No.

8079, examined.
Type locality: Arizona.

Additional localities: California: Yermo, Amboina. Arizona:
Mesa, Yuma, Gila Bend, Tucson, Phoenix, Orache, Avondale
Ranch — Agua Fria R. New Mexico: Deming, Lordsburg,
Mescalera, Mesilla. Texas: Sierra Blanca, Ft. Davis. Occurs
from June to August.

Epicauta tenella (Le Conte)

Fig. 35

Lytta tenella Le Conte, 1858, Journ. Acad. Nat. Sci. Phila. (2) 4:23.
Macrobasis tenella, Horn, 1873, Proc. Am. Phil. Soc. 13:94; 1885, Trans. Am.
Ent. Soc. 12:108.

Length: 7 to 10 mm. Head, antennae, pronotum and legs pale
tan. Elytra brown to black. Underside tan to partly brown or
black. The pronotum may have darker suffusions. The whole sur-
face is sparsely closed with short cinereous pubescence. Distin-
guished from mcrkeliana by the second segment of the antennae,
which is shorter than the third.

Head subquadrate, partly denuded, sparsely punctured, the
intervals finely punctulate, dull. Median suture fine but distinct.

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Antennal calluses rather small, denuded, smooth. Last segment
of labial palpi broad, smooth. Eyes large, prominent, moderately
narrow, transverse. Antennae slender, filiform, reaching to basal
fourth of elytra. First segment slender, straight, reaching just
beyond the hind margin of the eye in the male and to just before
the hind margin in the female; second two-fifths as long as first in
male, half as long as first in female, slender; third the stoutest
segment but still slender, half again as long as second; fourth and
following four-fifths as long as third, almost uniform in thickness.
Pronotum subcampanulate, one-fourth longer than broad. Median
suture fine. Basal impressed line only slightly evident. Anterior
tibiae of male with two spiniform spurs (not one as guessed by
Horn in 1885), which are slightly heavier than in the female.
Posterior tibial spurs flattened, pointed, the inner narrow, the
outer somewhat broader.
Type: Lectotype 9 , new designation, Le Conte Coll. M. C. Z. No.

4988, examined.
Type locality: Llano Estacado (Texas or New Mexico).
Additional localities: Arizona: Safford, Tempe, Bowie, Tucson,

Phoenix, Oracle, Texas Pass — Dragoon Mts.

Epicauta merkeliana Horn
Epicauta merkeliana Horn, 1891, Trans. Am. Ent. Soc. 18:43.

Length: 7 to 10 mm. Resembles tenclla but has more brown on
the pronotum, is darker below and has the second segment of the
antennae as long as the third.

Head subquadrate, moderately sparsely and finely punctured,
shiny. Median suture distinct. Mouth parts luteous. Eyes promi-
nent, black, broadly oval. Antennae elongate, slender. First seg-
ment slender, reaching to the hind margin of the eye; second and
following subequal, a little more than half as long as the first,
decreasing gradually in thickness. Pronotum half again as long
as broad, narrow, campanulate. Median suture and basal impressed
line distinct. A basal longitudinal impression present. Scutellum
narrow, ferrugineous. Below mainly black, with some ferrugineous
areas on the thorax and near the tip of the abdomen. Male with
two slender spiniform spurs on the anterior tibiae. Posterior tibial
spurs broad, pointed.

WERNER: NORTH AMERICAN EPICAUTA 495

Type: Holotype, Horn Coll. A.N.S.P. No. 8091, examined.
Type locality: "Arizona, probably Fort Whipple."
Additional localities: Arizona: Tucson VII-30-37 by Klots.
Nevada: Las Vegas IX-2-09.

Epicauta ochrea (Le Conte)

Fig. 42

Lytta ochrea Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:342.

Macrobasis ochrea, Horn, 1873, Proc. Am. Phil. Soc. 13:91; 1885, Trans. Am.

Ent. Soc. 12:110. Champion, 1892, Biol. Cent.-Am. Coleop 4 pt. 2:401,

Tab. 18, fig. 23, 23a.
Cantharis protarsalis Duges, 1878, La Naturaleza 4:63, Tab. 2, fig. 7a-b, 8a-c.

Length: 9 to 14 mm. Ferrugineous, rather sparsely clothed with
silky ferrugineous pubescence. The moniliform antennae distin-
guish this species from all others in our fauna.

Head subtriangular, rather densely and deeply punctured.
Median suture very evident and a narrow denuded line extends
forward from it nearly to the clypeus. Antennal calluses small,
glabrous, smooth. Eyes moderately broad, transverse. Antennae
stout, moniliform, uniform in thickness. First segment in the male
reaching to just behind the eye, curved to fit over the eye and
excavated externally near the tip, in the female shorter and not as
excavated; second one-fourth as long as first in male, one-third
in female. The first two segments are shiny but not glabrous.
Third and following a little longer than the second, moniliform.
Pronotum quadrate, one-third longer than broad. Median suture
very distinct. With an apical transverse impressed line. At the
base there is a transverse impression, deep at the middle and
merging laterally into the basal impressed line. Visible portion of
scutellum small. Below darker than above. Anterior tibiae of
male with a single stout, spiniform spur. First segment of male
anterior tarsi short, smooth, contorted. Posterior tibial spurs
rather stout, sticklike, the outer shorter.

Type: Holotype d\ Le Conte Coll. M. C. Z. No. 4987, examined.
Type locality: "Texas."
of protarsalis (Duges)
Type: Presumably in Duges Coll., Mus. Nacion. Hist. Nat.,

Mexico.
Type locality : Guanajuato.

496 bulletin: museum of comparative zoology

Additional localities: Texas: Alpine, Davis Mts., Marfa, Ft.
Davis. New Mexico: Snow Coll. Arizona: Douglass, Santa
Rita Mts., Palmerlee. California: Eddy Coll.

Epicauta gissleri (Horn)
Fig. 43

Macrobasis gissleri Horn, 1878, Trans. Am. Ent. Soc. 7:58; 1885, Trans. Am.
Ent. Soc. 12:110.

Length: 9 to 12 mm. Black or very dark brown, sparsely clothed
with cinereous pubescence, which is denser on the suture and
margins of the elytra. First segment of male anterior tarsi con-
torted.

Head subquadrate. Median suture fine, supplemented by a
fairly broad denuded median line. Antennal calluses denuded,
conspicuous. Eyes rather narrow. Antennae three and one-half
times as long as an anterior tibia, with the segments strongly
separated. First segment stout, reaching a little beyond the hind
margin of the eye in the male and half-way across the eye in the
female; second short, one-third as long as first in male, one-half in
female. The first and second segments have scattered dark and
cinereous pubescence. Third segment one and one-half times as
long as second; fourth and following a little shorter than the third,
the fourth being the stoutest segment, the rest tapering gradually,
all cylindrical. Pronotum a little longer than broad, broadly
campanuliform. Basal impressed line distinct ; median suture indis-
tinct. There is a shallow midbasal impression and sometimes a
pair of small lateral impressions behind the middle. Elytra with
often indistinct scutellar and humeral dark spots. Legs slender.
All of the femora and trochanters of the male are denuded and
somewhat flattened on the posterior surface; the middle tibiae are
curved and the middle and posterior femora have fringes of long
hairs; anterior tibiae denuded behind except for a tuft of normal
cinereous pubescence at the "elbow;" anterior tibiae with a single
stout, spiniform spur; first segment of anterior tarsi partly denuded,
short, contorted, the second long, longer than the second and third
together. Posterior tibial spurs moderately slender, spiniform.
Type: Lectotype cf, new designation, Horn Coll. A.N.S.P. No.

8079, examined.
Type locality: Grant Co., New Mexico.

WERNER: NORTH AMERICAN EPICAUTA 497

Additional localities: Arizona: Williams, Pinal Mts. Rare.
Occurs in June and July.

Epicauta parkeri Werner
Epicauta parkeri Werner, 1944, Psyche 50:71.

Length: 10 to 12 mm. resembles gissleri in general appearance.
The pubescence tends to be sparser and more easily rubbed off.
The antennae are distinctly more slender, with the segments less
distinctly separated. In the form of the head it is very distinct
from gissleri. The head is elongate-triangular and the mandibles
go one-third their length beyond the labrum, have a distinct tooth,
and cross at the tips.

The lengths of the antennal segments are in the same proportion
as in gissleri. In the allotype male the segments are broader and
flat toward the apex. I believe this specimen is abnormal. The
only other male seen has the antennae broken off at the second
segment.

Type: Holotype 9 , Parker Coll., deposited in Calif. Acad. Sci.
Type locality: White Mts., Arizona, July 8, 1933.
Additional localities: Arizona: San Rita Mt., Globe, Flagstaff,

Baboquivari Mts. June and July.

Epicauta fabricii (Le Conte)
Fig. 47, 48

Lytta cinerea Fabricius, 1798, Suppl. Ent. :119, nee Meloe cinereus Forster,
1771, in Cat. An. N. Am. :62. Both belong to Epicauta.

Lytta fabricii Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:343.

Lytta debilis Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:344.

Macrobasis unicolor, Horn, 1873, Proc. Am. Phil. Soc. 13:92; 1885, Trans. Am.
Ent. Soc. 12:110 (in part).

Macrobasis fabricii, Brown, 1940, Can. Ent. 72:231.

Length : 9 to 15 mm. Black or dark brown, fairly densely clothed
with cinereous pubescence. Elytra with dark humeral and scutellar
spots. First segment of male antennae shiny, flat, reaching occiput;
second slightly more than half as long. In the female, the first
segment reaches the hind margin of the eye and the second seg-
ment is one and one-third times as long as third.

Head subtriangular, moderately densely punctured. Median

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suture distinct. Antennal calluses moderate in size, smooth, shiny.
Eyes fairly narrow, transverse. Male antennae reaching to basal
fourth to third of the elytra; first segment reaching the occiput,
curving slightly, flattened; second a little more than half as long
as first, flattened, narrower than first, straight, or curved very
slightly. The first two segments shiny, usually margined behind
with sparse cinereous pubescence. Rest of segments subequal, the
third a little shorter than the rest, tapering gradually. In the
female, the first segment slightly exceeds the hind margin of the
eye; second three-fourths as long as first; third three-fourths of
second and slightly shorter than the following, which are as in the
male. Pronotum one-fourth longer than broad, campanuliform.
Median suture and basal impressed line distinct. Elytra with
indistinct to distinct humeral and scutellar spots. Perfect speci-
mens have denser pubescence on the margins of the elytra. Ante-
rior tibiae of male with two slender, spiniform spurs. First segment
of anterior tarsi of male somewhat elongated but not otherwise
modified. Posterior tibial spurs flattened, spiniform, the outer
broader.

Type: Since fabricii is a substitute name for a homonym, the type
must be the type of Lytta cinerea Fabricius. The location of this
type is unknown to the author. If this type cannot be located,
the neotype should be the specimen labelled Fabricii in the Le
Conte collection.
Type locality: "Habitat in America Dom. Hybner."
of debilis (Lee.)

Type: Holotype 9 , Le Conte Coll. M. C. Z. No. 4984, examined.
Type locality: New York.

Additional localities: Maine, Massachusetts, Connecticut, New
York, New Jersey, Pennsylvania, Virginia, West Virginia,
Maryland, Georgia, Tennessee, Indiana, Illinois, Michigan,
Wisconsin, Iowa, North Dakota, South Dakota, Nebraska,
Kansas, Oklahoma, Texas: Tyler, Montana, Utah, Colorado,
New Mexico, Manitoba.

A common species. Feeds generally on leguminous herbs —
alfalfa, clover, Baptisia, eating the succulent growth and flowers.
It seems to choose the softest legume in the area. In eastern
Massachusetts, Baptisia tinctoria seems to be preferred, according
to Mr. Frost. In northern Illinois, Baptisia leucantha and alfalfa
are preferred. Occurs in early summer.

WERNER: NORTH AMERICAN EPICAUTA 499

Epicauta murina (Le Conte)
Fig. 49, 50

Cantharis unicolor Kirby, 1837, in Richardson's Fauna Boreali-Americana
4:241, nee Cantharis unicolor Fald., 1835, Nouv. Mem. Soc. Nat. Moscow
4:195.

Lytta murina Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:344.

Macrobasis unicolor, Horn, 1873, Proc. Am. Phil. Soc. 13:92 (in part).

Macrobasis murina ? Le Conte, 1876, U. S. Geog. Surv. W. of 100th Merid.,
Appendix J. J. :520.

Macrobasis murina, Fisher, 1919, Proc. Ent. Soc. Wash. 21:1 (in part). Chit-
tenden, 1926, Bull, Brooklyn Ent. Soc. 21:118. Brown, 1940, Can. Ent.
72:232.

Length: 7 to 11 mm. Except for the pubescence, this species
cannot be distinguished from subglabra (Fall). On the dorsal
surface, the pubescence is almost twice as dense as in subglabra,
causing a dark grey color. Some females can be confused with
females otfabricii but the second segment of the antennae is almost
as long as the first and at least one and one-half times as long as
the third. It is two-thirds as long as the first ond one and one-third
times as long as the third in fabricii. Feeds on legumes in early
summer. Two records on potatoes.

Type: Holotype d\ Le Conte Coll. M. C. Z. No. 4983, examined.
Type locality: Lake Superior.
Additional localities: New Hampshire: Mt. Washington 5000-

6000 ft. New Brunswick: Bathurst. Michigan: Eagle Harbor.

Wisconsin: Hazelhurst, Bayfield. Minnesota: Beaver R. —

Lake Co., Highland (on potatoes). Iowa: Carroll Co. North

Dakota: Mandan. South Dakota: Elmore, Black Hills.

Nebraska: Mitchell (on potatoes). Montana: Huntley, Sidney.

Wyoming: Sheridan. Colorado: Hooper.

Epicauta subglabra (Fall)

Macrobasis murina, Fisher, 1919, Proc. Ent. Soc. Wash. 21:1 (in part).
Macrobasis subglabra Fall, 1922, Can. Ent. 54:173. Brown, 1940, Can. Ent.
72:232.

Length: 6 to 10 mm. Black, sparsely clothed with short brown
hairs, which are not ordinarily visible to the unaided eye. Second
segment of male antennae two-thirds as long as first, curved.

Head subtriangular, n*inutely punctulate, dull. Median suture

500 bulletin: museum of comparative zoology

fine but distinct. Antennal calluses small, shiny. Eyes not very
prominent, quite small, oblique. Antennae of male with first
segment flattened, straight, attaining the occiput or a little beyond;
second two-thirds as long as first, as long as the next three and
one-half, flattened, curved outward; third segment the shortest,
two-sevenths as long as the second; rest slightly longer than third,
gradually tapering, slightly flattened. Female with the first seg-
ment reaching the hind margin of the eye; second four-fifths as
long; third and following two-thirds as long as second and only
slightly flattened. Pronotum one-fourth longer than broad, sub-
campanulate. Median suture and basal impressed line fine,
distinct. Surface as on head. Elytra with no sign of basal spots.
Pubescence below a little denser than above, especially on the
posterior margins of the abdominal sternites. Anterior tibiae of
male with two slender spiniform spurs. Male anterior tarsi with
first segment slightly elongated but not otherwise modified.
Posterior tibial spurs slender, spiniform, equal.
Type: Holotype <?, Fall Coll. M. C. Z. No. 24297, examined.
Type locality: Edmonton, Alberta.

Additional localities: Manitoba: Berens River, Maskado, Aweme,
Winnipeg. Minnesota: Park Point — Duluth. North Dakota:
Beach, Star Co., Mandan. South Dakota: Volga, Browns
Valley. Idaho: Paris. Arizona ?: Madera Canyon — Santa
Rita Mts., McKay's Peak— White Mts., Flagstaff. All of the
records are from June and July. Apparently limited to legumes.

Epicauta flavocinerea (Blatchley)

Fig. 51, 52

Macrobasis flavocinereus Blatchley, 1910, Cat. Coleop. Indiana: 1359.
Macrobasis flavocinerea, Brown, 1940, Can. Ent. 72:231.

Length: 9 to 11 mm. The "flavocinereous" color of the pubes-
cence cannot be relied upon to distinguish this species. All of the
specimens I have seen are flavocinerous but fabricii may be of
almost the same color. The male has the first segment of the
antennae reaching to halfway between the eye and the occiput,
not conspicuously enlarged nor flattened and has long hairs on
the hind margin of the middle and posterior femora. The female
has the first antennal segment reaching to the hind margin of the
eye, with the second segment equal to the third. The second seg-

WERNER: NORTH AMERICAN EPICAUTA 501

ment is one and one-third times as long as the third in the female
of fabricii.

The general form is very similar to that of fabricii. Only the
differences will be noted in the following description. Antennae
of male reaching to the basal sixth of the elytra, two and one-half
times as long as an anterior tibia. First segment fairly slender,
not flattened, curved to fit over the eye, reaching to half-way
between the eye and the occiput; second two-fifths as long as first,
more slender. The first and second segments have more pubescence
than in fabricii. Third and following two-thirds as second, tapering
gradually. Female with the first segment reaching to the hind
margin of the eye, second just perceptibly longer than the following.
Male anterior tarsi not modified. Posterior tibial spurs sticklike,
the outer a little broader than the inner. Posterior face of femora
of male smooth, denuded but for scattered silky hairs, which are as
long as the femur is*broad. These are scattered evenly over the
smooth surface.

Type: Holotype d71, Blatchley Coll., Purdue University, examined.
Type locality: Starke Co., Indiana.

Additional localities : Illinois: Willow Springs VI-VII. Pennsyl-
vania: "Penn." C. V. Riley. Texas: "Tex." Fall Coll. Appar-
ently everywhere rare. Not over a dozen specimens were seen.
Brown lists it from Byron and Lumington, Ontario and Aweme,
Manitoba.

Epicauta languida (Horn)
Macrobasis languida Horn, 1895, Proc. Calif. Acad. Sci. (2) 6:252.

"Form slender, general color pale luteous clothed with slightly
paler pubescence, antennae and tarsi black. Thorax longer than
wide, a feebly impressed median line. Tarsi black with a ring of
whitish pubescence at the base of each joint. Length, .40 inch;
10 mm.

Male. Antennae setaceous the first joint as long as the next
two, not sinuate at apex, second three-fourths as long and equal to
the next three joints together. Anterior tibiae with two terminal
spurs, the first joint of tarsi longer than the second. Last ventral
segment incised.

502 bulletin: museum of comparative zoology

This species resembles linearis very closely and it is possible
that the females will be difficult to separate. San Jose del Cabo."

No specimen of this species has been seen. Horn's reference to
linearis should be changed to polingi Werner, which he mistook
for linearis.

Epicauta tenuilineata (Horn)
Fig. 41
Macrobasis tenuilineata Horn, 1894, Proc. Calif. Acad. Sci. (2) 4:436.

Length: 7 to 15 mm. Ferrugineous, clothed with cinereous pub-
escence, rather sparse over the body but denser on the margins and
four narrow vittae on each elytron.

Head suboval. Median suture distinct, supplemented on the
front by a narrow denuded line. Antennal caUuses oblique, narrow,
denuded, smooth. Eyes large, bulging, quite broad. Male antennae
with the first two segments flat, sparsely punctured, shiny, with
very short hairs from the punctures. First segment reaching
three-fourths across the eye, narrow; second one-third longer than
first, more flattened and broader; third and following very slender,
the third one-third as long as second, fourth and following one-third
longer than third. First segment of female antennae reaching
two-fifths across the eye; second equal to it. Neither is flattened
and both have cinereous pubescence. The proportions of the
rest of the segments are the same as in the male but the segments
are thicker, especially toward the base. Pronotum slightly longer
than broad, broadly campanuliform. Basal impressed line dis-
tinct; median suture indistinct, replaced by a conspicuous denuded
line. Elytra with scutellar and humeral brown spot. Two of the
pubescent lines on the elytra arise from the base and two from
just behind the humerus. Femora darkened at tip. Anterior tibiae
of male with two spiniform spurs. Male anterior tarsi not modified.
Posterior tibiae with the spurs flattened and blunt, the outer
broader.
Type: Cotypes should be in the California Academy of Science

No. 153.
Type locality: Sonora, Mexico and San Jose del Cabo.
Additional localities : Arizona: Tucson, Baboquivari Mts., Tempe,

Phoenix. Occurs from May to July.

WERNER: NORTH AMERICAN EPICAUTA 503

Epicauta excors (Fall)

Macrobasis excors Fall, 1909, Can. Ent. 41:166.

Length: 14 mm. Black, quite densely clothed with cinereous

pubescence. First two segments of antennae long; second as long

■ as first, which attains the occiput in the male and the hind margin

of the eye in the female. Palpi dark. Elytra with scutellar and

humeral spots.

Head subquadrate. Median suture fine but distinct. Antennal
calluses quite large but narrow, denuded, shiny. Palpi black.
Eyes large, fairly prominent, transverse, quite broad. Male
antennae reaching two-thirds the length of the elytra. First seg-
ment shiny, flattened, slightly incurved, just exceeding the occiput;
second as long as first, shiny, slightly narrow, also incurved; third
one-third as long as second, narrow; fourth to last almost twice
as long as third, becoming very slender toward the tip. Female
with the antennae reaching one-third the length of the elytra.
First segment reaching to hind margin of eye; second just per-
ceptibly shorter; rest as in the male but a little thicker. Pronotum
campanuliform, one-fourth longer than broad. Basal impressed
line distinct. Median suture absent but a narrow denuded line
replaces it. Elytra with small scutellar and humeral black marks.
Male anterior tibiae with two slender, spiniform spurs; anterior
tarsi with first segment somewhat elongated, but not otherwise
modified. Posterior tibiae with inner spur spiniform, outer blunt,
both rather stout.

Type: Holotype &, Fall Coll. M. C. Z. No. 24296, examined.
Type locality: El Taste, Lower California.

Additional localities: Santa Rosa, Lower California Aug. (Frost
Coll.)

Epicauta tenuis (Le Conte)

Fig. 34

Lytta tenuis Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:343.
Macrobasis tenuis, Horn, 1873, Proc. Am. Phil. Soc. 13:92; 1885, Trans. Am.
Ent. Soc. 12:110.

Length: 10 to 14 mm. Slender, rather densely clothed with
cinereous to flavocinereous pubescence. Body color dark brown
to black. No basal marks on the elytra. Palpi luteous. Antennae
slender, long; reaching three-fourths the length of the elytra in the

504 bulletin: museum of comparative zoology

male, with the first two segments smooth, very long straight.
Female easily distinguished from the female of fabricii by the
second segment of the antennae, which is as long as the first.

Head subquadrate. Median suture distinct. Antennal calluses
strongly raised, denuded, shiny. Palpi luteous. Eyes moderately
prominent, narrow, transverse. Male antennae with the first
segment attaining occiput, slender, flattened^ shiny; second
slightly curved, three-fourths as long as first, also shiny; third
one-third as long as second; fourth and following one-third longer
than second third, becoming very slender toward the tip. Female
with the first segment reaching two-thirds across the eye; second
as long as first but more slender; third three-fourths as long as
second; fourth and following slightly longer than third, tapering
gradually but not as slender as in the male. Pronotum sub-
campanulate, one-fourth longer than broad. Median suture dis-
tinct, supplemented toward the middle of the disc by a narrow
denuded line. Basal impressed line distinct. Male with a single
spur on the anterior tibiae, with the first segment of the anterior
tarsi a little elongated but not otherwise modified. Posterior
tibial spurs slender, sticklike, the outer shorter.
Type: Lectotype cf, new designation, Le Conte Coll. M. C. Z.

No. 4985, examined.
Type locality : Georgia.
Additional localities: Florida: Edgewater Beach, Biscayne.

Epicauta purpurea (Horn)

Fig. 8, 44

Macrobasis -purpurea Horn, 1885, Trans. Am. Ent. Soc. 12:108. Champion,
1892, Biol. Cent.-Am.-Coleop. 4 pt. 2:397, Tab. 18, fig. 19-19a.

Length: 8 to 12 mm. Black, pubescence on body cinereous, with
a pair of black marks on head and pronotum, elytra black with
margins and a distinct vitta cinereous. Antennae of male with
first segment elongate, curved. The black pubescence may be
tinged with brown, suggesting a purplish color.

Head subtriangular, densely punctured and punctulate. Median
suture distinct. Antennal calluses moderate in size, denuded,
smooth. Eyes small, fai,rly narrow, transverse. Antennae of male
with first segment almost attaining occiput, slender, curved,
broadly excavated externally near the tip; second segment two-

WERNER: NORTH AMERICAN EPICAUTA 505

fifths as long as first, flattened on underside, forming a sort of
clasping organ with the excavation of the first. The first two seg-
ments are moderately densely punctured and punctulate. Third
three-fifths as long as second; fourth and following one and one-
half times as long as third, gradually decreasing in thickness.
Antennae of female with first segment straight, reaching to hind
margin of eye; second slender, seven-tenths as long as first; third
equal to second; fourth and following slightly longer, decreasing
just perceptibly in thickness. Pronotum quadrate. Median suture
distinct, supplemented apically by a narrow denuded area. Basal
impressed line distinct. A fairly deep midbasal impression present.
Scutellum with cinereous pubescence. Median vitta of elytra
starting at the middle of the base and ending near the apex, well
defined. Below, except for tips of femora, most of tibiae and tarsi,
with cinereous pubescence. Anterior tibiae of male with a single
straight, spiniform spur; anterior tarsi not modified. Inner
posterior tibial spur slender, spiniform, considerably longer than
outer, which is sticklike.

Type: Holotype d\ A.N.S.P. No. 8086, examined.
Type locality: Arizona.

Additional localities: Arizona: Baboquivari Mts., Sta. Rita R.,
Catalina Mts.

Epicauta polingi Werner

Fig. 37, 40

Macrobasis linearis, Horn, 1885, Trans. Am. Ent. Soc. 12:109, 110. nee

Le Conte, 1858, Journ. Acad. Nat. Sci. Phila." (2) 4:23.
Epicauta polingi Werner, 1944, Psyche 60:71.

Length: 8 to 16 mm. Elongate, slender. Head, pronotum and
scutellum black or very dark brown, elytra luteous, the whole
rather loosely and evenly clothed with cinereous to luteous pubes-
cence. First segment of male antennae elongate, flat; second one-
third as long as first.

Head subquadrate. Median suture distinct. Antennal calluses
moderate in size, denuded, smooth. Clypeus, labrum and mandi-
bles brown, the palpi luteous. Eyes moderately narrow, transverse.
First segment of male antennae exceeding the, head by one-third
its length, about one-sixth longer than an anterior tibia, not over
one-sixth as broad as long, flattened behind and with some cinere-
ous pubescence on this flat surface; second narrower, elongate —

506 bulletin: museum of compakative zoology

oval, one-third as long as first. The first two segments are smooth
and shiny. They are about equal in length to the rest of the
antenna. Third and following a little more than half the length of
the second, gradually tapering. Female with the first segment
reaching three-fourths across eye; second three-fifths as long as the
first; third and following slightly shorter than second. Pronotum
slightly longer than broad, campanuliform. Median suture dis-
tinct, bounded posteriorly by a shallow transverse basal impres-
sion. Elytra with small brown scutellar and humeral spots. Legs
ferrugineous, the rest of the underside black or dark brown, with
cinereous pubescence. Anterior tibiae of male mostly denuded,
somewhat arcuate, with a single spiniform spur. Posterior tibial
spurs stout, spiniform.

Type: Holotype d\ M. C. Z. No. 26071, examined.
Type locality: Davis Mts., Texas.

Additional localities: Texas: Alpine, Sabinol, Devil's River, Rio
Frio, Cotulla, Big Bend Park -- Brewster Co. New Mexico:
Cloudcroft, Alamagordo. Arizona: Douglass, Santa Catalina
Mts. Recorded from April to September but mainly in June
and July.

Epicauta liebecki Werner
Fig. 38
Epicauta liebecki Werner, 1944, Psyche 50:72.

Length: 9 to 14 mm. Close to poling}. Head and pronotum

black or dark brown, elytra brown, much darker than in polingi.

Pubescence cinereous, denser on the suture and margins of the

elytra.

Otherwise as in polingi except that in the male the first segment

of the antennae is one-fifth as broad as long, the third segment is

distinctly shorter than the fourth, and the first two segments are

distinctly longer than the following. Legs a little stouter. Posterior

tibial spurs quite slender.

Type: Holotype cf , M. C. Z. No. 26068, examined.

Type locality: Tucson, Arizona.

Additional localities: Arizona: Congress Junction, Wickenburg,
Columbia (on Frosopis pubesccns), Tempe, Sabina Canyon,
Santa Catalina Mts., Lowell, Globe, Bowie, Ajo, Pima Co.,
Baboquivari Mts. Sonora: Imuris (on pigweed). Taken from
mid-May to mid-July.

WERNER: NORTH AMERICAN EPICAUTA 507

- Epicauta arizonica Werner
Fig. 39
Epicauta arizonica Werner, 1944, Psyche 50:72.

Length: 9 to 13 mm. Uniformly luteous to dull brown, with
cinereous pubescence. First segment of antennae of male exceeding
the head by not more than one-sixth of its length, a little shorter
than an anterior tibia, a little less than one-fifth as broad as long.
First two segments one-fifth shorter than the rest. Eyes prominent,
much larger than in polingi or liebecki and a little broader. Other-
wise as in polingi. Averages smaller than either.
Type: Holotype d\ M. C. Z. No. 26062, examined.
Type locality: Baboquivari Mts., Arizona.
Additional localities: Arizona: Tucson, Wickenburg, Nogales,

San Bernardino Ranch — Cochise Co., Capitan Mt., Canon

Lake, Globe, Phoenix, Rice, Santa Rita Mts. Sonora: Arizpe.

Occurs from mid-July to mid-September. Commonest in

August.

Epicauta torsa (Le Conte)

Fig. 45, 46

Lytta torsa Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:343.
Macrobasis torsa, Horn, 1873, Proc. Am. Phil. Soc. 13:91; 1885, Trans. Am.
Ent. Soc. 12:110.

Length: 7 to 11 mm. Black, quite sparsely clothed with short
cinereous pubescence, as in murina. Easily distinguished from
that species by the S-shaped first antennal segment in the male and
an indication of the external apical excavation of the first segment
in the female.

Head subtriangular, finely rugulose, quite densely punctured.
Median suture distinct, supplemented by a smooth line, which
continues, narrower, to the clypeus. Antennal calluses of moderate
size, denuded, shiny. Eyes not very prominent, rather narrow,
transverse. Male with the first segment of the antennae stout,
reaching the occiput, with a 60-degree bend outward half-way and
a slight inward bend at the apex; second also stout, two-fifths as
long as first, excavated on the inner face as if to form a clasping
organ with the outer excavation of the first segment. Rest of seg-
ments slender, but not as slender as in fabricii or murina. Third

508 bulletin: museum of comparative zoology

segment two-fifths as long as second, slightly broader than long;
fourth and following a little longer than third, decreasing slightly
in thickness. Female with the first segment reaching to the hind
margin of the eye; slightly bent, with a trace of an external apical
excavation; second three-fifths as long as first; third and following
three-fourths as long as second, decreasing slightly in thickness,
not as heavy as in the male. Pronotum quadrate, as broad as long,
with the sides parallel for the basal three-fourths, then converging
abruptly. Transverse basal impression strong. Median suture
indistinct, but replaced by a fairly strong median impression from
base to middle. There is a transverse impression near the apex,
extending down the sides. Elytra with black scutellar and humeral
spots, which are indistinct because of the dark general color.
Anterior tibiae of male with a single spiniform spur. Male anterior
tarsi with the first segment flattened, slightly broadened apically.
Inner spur of posterior tibiae spiniform, the outer sticklike,
shorter.

Type: Lectotype d\ new designation, M. C. Z. No. 4982, exam-
ined.
Type locality: Texas.

Additional localities: Texas: Victoria. Alabama: Mobile. Mis-
sissippi: Poplarville. Florida: Winter Park, Little River.
North Carolina: Black Mts. Massachusetts: Wellesley,
Framingham, Randolph. Occurs in the spring. Was common in
Massachusetts in 1909, according to Mr. Frost. I have seen no
specimens taken there since that date.

Epicauta longicollis (Le Conte)

Fig. 28, 29

Lytta longicollis Le Conte, 1853, Proc. Acad. Nat. Sci. Phila. 6:343.
Macrobasis longicollis ,Horn, 1873, Proc. Am. Phil. Soc. 13:90; 1885, Trans.

Am. Ent. Soc. 12:110. Champion, 1892, Biol. Cent.-Am. Coleop. 4 pt.

2:397.

Length: 15 to 25 mm. Elongate, slender, black, densely clothed
with cinereous to tannish-cinereous pubescence. Black scutellar
and humeral spot present on the elytra. Male antennae with the
first segment elongate, cylindrical, excavated externally near the
tip. Female antennae with the second segment shorter than the
third. It is as long as the third in both sexes of immaculata (Say).

WERNER: NORTH AMERICAN EPICAUTA 509

Head subtriangular, densely and finely punctured. Median
suture visible only before the eyes. Antennal calluses small,
glabrous, smooth. Fronto-clypeal suture very strong. Eyes not
prominent, narrow. Male antennae with the first segment extend-
ing one-third beyond the occiput, stout but slender, almost cylin-
drical, only slightly flattened. There is a slight excavation extern-
ally near the apex. Second segment one-sixth as long as the first;
third and fourth slightly longer; rest a little longer than the
fourth, gradually diminishing in thickness. The first four segments
are shiny. Female antennae with the first segment exceeding the
hind margin of the eye by one-fourth, thicker toward the apex;
second one-third as long as first. First two segments rather densely
pubescent. Third and fourth one-third longer than second; rest
slightly longer than fourth, gradually decreasing in thickness.
Pronotum subquadrate, one-third longer than broad. Sides parallel
for the basal three-fourths, then converging gradually. Median
suture indistinct but there is a fairly strong median impression
from base to middle. A strong transverse basal impression con-
nects with this. There is also a transverse impression at the apex,
extending down the sides. Elytra with black humeral and scutellar
spot. Partly denuded specimens seem to have faint lines on the
elytra as in sublineata but these are due to abrasion on slightly
raised lines. Below cinereous. There are sometimes a few black
marks on the thoracic pleurites. Tips of femora, tibiae and all of
tarsi black. Tibiae of mixed black and cinereous pubescence.
Anterior tibiae of male with a single slender, spiniform spur.
Posterior tibial spurs both slender, spiniform.
Type: Lectotype d\ new designation, Le Conte Coll. M. C. Z.

No. 4978, examined.
Type locality : New Mexico, Santa Fe.

Additional localities: Texas: Alpine, Marfa. New Mexico: Fay-
wood, Albuquerque. Arizona: Chiricahua Mts., Kaibab
National Forest, Chi No Valley — Tempe, Fort Grant, Prescott,
Sulphur Spring Valley. California: Berkeley. Occurs in June
and July.

Epicauta atrivittata (Le Conte)

Fig. 24

Lytta atrivittata Le Conte, 1854, Proc. Acad. Nat. Sci. Phila. 7:224.
Macrobasis atrivittata, Horn, 1873, Proc. Am. Phil. Soc. 13:90; 1885, Trans.
Am. Ent. Soc. 12:110.

510 bulletin: MtTSEUM OF comparative zoology

Length: 20 to 25 mm. Elongate, rather slender. Black, densely
pubescent. Pubescence cinereous, with the occiput black or having
a pair of large black spots, pronotum with a pair of large black
spots, elytra with a pair of broad black vittae arising from the
scutellum and humerus and united before the apex. Below with
black marks on the thoracic pleurites and sternites and on the basal
two-thirds of the abdominal sternites. Legs mainly black. First
segment of antennae of male greatly enlarged, flattened apically,
excavated externally near the apex. A magnificent species, aver-
aging over 25 mm. long.

Head subquadrate. Median suture indistinct. Antennal calluses
large, smooth, shiny. Palpi dark brown, broad and flattened. Last
segment of labial palpi suborbicular. Eyes moderately narrow.
Male antennae with the first segment exceeding occiput by one-
third, shiny, flat, curved, greatly broadened beyond the middle,
excavated externally near the tip; second short, suborbicular,
with a short spine at the outer apex, one-fifth as long as first;
third broader than long, half as long as second; fourth as long as
second, as broad as long; fifth and sixth one-third longer than
fourth, also stout; seventh and following as long as sixth but nar-
rower, uniform in length and thickness. Female with the first seg-
ment reaching to just beyond the eye, somewhat flattened; second
one-third as long as first; third and following a little longer than
the second, tapering slightly toward the apex. Pronotum one-third
longer than broad. Sides parallel for the basal three-fourths, then
converging abruptly. Median suture absent but replaced by a
median impression, narrow before the middle and deep, fairly
broad behind. Basal impressed line deep, margined with long
pubescence. Visible portion of scutellum abnormally small, narrow.
Anterior femora with cinereous pubescence on both edges, rest on
outer edge only. Coxae and trochanters all with cinereous pubes-
cence. Rest of legs with black pubescence. Anterior tibiae of male
with a single short, spiniform spur. First segment of male anterior
tarsi a little more elongated than in the female but not otherwise
modified. Posterior tibial spurs slender, sticklike, the outer longer.
Type: Lectotype d1, new designation, Le Conte Coll. M. C. Z.

No. 4980, examined.
Type locality: "San Diego trip." Pin label "Webb b."
Additional localities: Texas: between Pecos R. and Guadaloupe

Mts., Rio Grande — Brewster Co., El Paso. New Mexico:

Roswell. Mexico: Tlahualilo. Occurs in June and July. Seems

to be quite scarce.

WERNER: NORTH AMERICAN EPICAUTA 511

Epicauta albida (Say)
Fig. 25, 27

Lytta albida Say, 1823, Journ. Acad. Nat. Sci. Phila. 3:305. Le Conte, 1853,

Proc. Acad. Nat. Sci. Phila. 6:344.
Lytta luteicornis Le Conte, 1854, Proc. Acad. Nat. Sci. Phila. 7:84.
Macrobasis albida, Horn, 1873, Proc. Am. Phil. Soc. 13:89; 1885, Trans. Am.

Ent. Soc. 12:109. Champion, 1892, Biol. Cent.-Am. Coleop. 4 pt. 2:397.

Length: 13 to 24 mm., usually 22-24. Elongate, rather slender,
black with dense cinereous to yellow-cinereous pubescence.
Oblique black markings on at least some of the abdominal sternites
and usually two longitudinal black marks on the pronotum. Often
black marks on the thoracic pleurites. First two segments of the
antennae iuteous, sparsely pubescent, the rest tan to dark brown.
Palpi pale Iuteous.

Head subquadrate, finely and densely punctured and punctulate.
Median suture distinct. Antennal calluses fairly large, denuded,
smooth. Eyes moderately narrow, transverse. Labial palpi of
male with last segment broad. Male antennae with the first seg-
ment extending slightly beyond the occiput, flattened, slightly
curved and excavated externally near the apex; second one-fourth
as long as first; third half as long as second; fourth half again as
long as third; the rest slightly longer than the fourth, decreasing
gradually in thickness. Female with the first segment of the
antennae barely reaching to the hind margin of the eye, with a
trace of the apical excavation; second half as long as first; the rest
as in the male. The first two segments have some conspicuous
dark pubescence. Pronotum subquadrate, slightly longer than
broad. Median suture absent but there is a fairly deep median
depression on the basal half. Elytra with black humeral and
scutellar spots. There is occasionally a pair of fine dark lines
from the middle of the base and the humerus, united behind. Tip
of femora, outer edge of anterior tibiae, both edges and terminal
eighth of the rest of the tibiae black. Anterior tibiae of male
slightly denuded behind. Male with two slender, spiniform spurs
on the anterior tibiae, slightly elongated first segment on the
anterior tarsi. Posterior tibial spurs slender; inner pointed, longer,
outer blunt.

Type: Say's type lost. Neotype c? and neoparatypes in M. C. Z.
Type locality : "Inhabits Arkansas. I obtained it near the Rocky

512 bulletin: museum of comparative zoology

Mountains." Neotype: La Junta, Bent Co., Colorado VI-24-25-
1885 H. F. Wickham.

of luteicornis Le Conte

Type: Lectotypecf, new designation, Le Conte Coll. No. 4977,
examined.

Type locality: Texas: "Laredo to Ringgold Barracks, Mr. Weise."

Additional localities: Kansas: Logan Co., Garden City, Reno Co.,
Wellington, Ellsworth. Colorado: Holyoke, Rocky Ford,
Pueblo, Julesburg. Oklahoma: Mangum. Texas: Victoria,
Harlingen, Dallas, Llano, Brownsville, Mason (on potatoes and
ground burnut), Cypress Mills, Canadian, Goldthwaite, Alice.
New Mexico: Roswell. Occurs from June to September.

Epicauta texana Werner
Fig. 26
Epicauta texana Werner, 1944, Psyche 50:73.

Length: 18 to 21 mm. This species is closely related to albida
(Say) but differs as following: Eyes narrower. First segment of
antennae of male elongate, flattened, extending one-third beyond
the occiput. There are no black markings on the ventral side.
Only one specimen has a slight trace of the pronotal marks. The
pubescence seems denser and natter than in albida.
Type: Holotype d", Ohio State University, examined.
Type locality: Davis Mts., Texas.
Additional localities: Texas: Alpine 4400-6000 ft., Marfa. Taken

in July and August.

Epicauta virgulata (Le Conte)
Fig. 53

Macrobasis virgulata Le Conte, 1866, Smiths. Misc. Coll. 6 no. 167, 2nd ed. :
156. Horn, 1873, Proc. Am. Phil. Soc. 13:91; 1885, Trans. Am. Ent.
Soc. 12:110.

Macrobasis hirsutipubescens Maydell, 1934, Trans. Am. Ent. Soc. 60:334.

Length: 7 to 10 mm. Black or dark brown, with rather dense
ferrugineous or cinereous pubescence. Elytra with black scutellar
and humeral spots and a line of denser pubescence from the middle
of the base to near the apex. Legs pale tan.

Head subquadrate, rather finely and densely punctulate, with

WERNER: NORTH AMERICAN EPICAUTA 513

scattered punctures. Median suture fine but distinct, accentuated
by a narrow denuded line. Antennal calluses broad, moderately
elevated, smooth, shiny. Eyes large, prominent, broad, especially
in the male. Male antennae reaching to the middle of the elytra.
First segment reaching to just behind the eye, slender, curved,
slightly excavated externally near the apex; second small, one-
sixth as long as first; third two and one-half times as long as
second; fourth the broadest segment, one-fourth longer than third;
following becoming gradually longer and more slender. Apical
segments extremely slender. Female with antennae heavier than
in the male, reaching to basal fifth of elytra. The broadest segment
is the first. First segment rather slender, curved, reaching to just
beyond the middle of the eye; second two-fifths as long as first;
third almost twice as long as second; rest becoming gradually
longer but not as markedly as in the male. Pronotum one-third
longer than broad, broadly campanulate. Basal impressed line
distinct but not denuded. Median suture distinct, supplemented
by a denuded line except at base and apex. Elytra with conspic-
uous black scutellar and humeral spots. Margin and a median line
with denser pubescence, sometimes indistinct. Legs pale tan.
Tips of femora, tibiae and tarsal segments black or dark brown.
Anterior tibiae of male with a single slender spiniform spur. First
segment of male anterior tarsi short, denuded on inside, extending
over the second segment on the outside. Posterior tibial spurs
ylender, spiniform.
Type: Lectotype cf , new designation, Le Conte Coll. M. C. Z.

No. 4981, examined.
Type locality: San Jose del Cabo, Lower California.
of hirsutipubescens

Type: Holotype cf, A.N.S.P. No. 8181, examined.
Type locality: "Texas."
Additional localities: Texas: Chisos Mts., El Paso, Nevill Spring

— Brewster Co., Ft. Stockton, Terrell Co., Davis Mts., Boquillas.

Arizona: Douglass, Nogales, Globe, Wickenburg, Orcale, Gila

Bend Mts., Texas Pass— Dragoon Mts. Sonora: Nogales.

Occurs from May to August.

Epicauta linearis (Le Conte)

Fig. 54, 55

Lytta linearis Le Conte, 1858, Journ. Acad. Nat. Sci. Phila. (2) 4:23.
Macrobasis linearis, Horn, 1873, Proc. Am. Phil. Soc. 13:94.

514 bulletin: museum of comparative zoology

Length: 8 to 9 mm. This species is not the linearis of collections.
Horn mistook poli?igi for it because of the general similarity of
color pattern. Only a single female of the true linearis, the type,
was seen by him. Apparently a very rare species. Head, pronotum
and below black; elytra and legs ferrugineous. The whole rather
densely clothed with tannish cinereous pubescence.

Head subquadrate, rather densely but finely punctured. Median
suture distinct. Antennal calluses quite large, reaching beyond
the inner margin of the eye, smooth, shiny. Eyes prominent,
moderately broad, transverse. Male antennae reaching the basal
third of the elytra, three times as long as an anterior tibia. First
segment attaining the occiput, fairly slender, curved, flattened,
deeply excavated externally near the apex, smooth, sparsely
pubescent; second cylindrical, one-fifth as long as first, small;
third half again as long as second; fourth flattened, broader than
third; one-fourth longer than second and third; following also
flattened, increasing just perceptibly in length, very narrow beyond
the sixth segment. Female with first segment reaching two-thirds
across eye; second and following as in the male but not as broad
toward the middle or as slender apically. Pronotum one-third
longer than broad, narrowly campanuliform. Basal impressed
line and median suture distinct. Elytra with a trace of a scutellar
spot but no humeral spot. Male anterior tibiae with a single stout
spiniform spur; anterior tarsi with first segment slightly shortened,
contorted. Posterior tibial spurs spiniform.

Type: Holotype 9 , Le Conte Coll. M. C. Z. No. 4986, examined.
Type locality: Llano Estacado (W. Texas or E. New Mexico)

Aug. 4, Capt. Pope.
Additional records: 1 d\ 1 9 "Tex." Eddy Coll. (M. C, Z.) 1 cT

"Tex." J. B. Smith Coll. (U.S.N.M.)

Epicauta maculifera (Maydell)
Macrobasis maculifera Maydell, 1934, Trans. Am. Ent. Soc. 60:335.

Length: 8 to 10 mm. Black or dark brown, legs ferrugineous,
clothed with rather dense cinereous pubescence, which is denuded
in small spots. Superficially resembles the macalata group because
of the spots but stands near fumosa Duges in the form of the
antennae.

Head subquadrate. Median suture distinct. Antennal calluses

WERNER: NORTH AMERICAN EPICAUTA 515

small but denuded, shiny. Eyes prominent, rather broad, trans-
verse. Antennae of male with the first segment attaining the
occiput, slender, slightly broader apically, excavated externally
near the tip; second one-fourth as long as first, flattened and
hollowed internally. First two segments shiny, sparsely pubescent.
Third and following half as long as second, decreasing in length
slightly toward the tip, loosely jointed. Female with the first
segment reaching three-fourths across the eye; second half as long
as first. First two with sparse cinereous pubescence. Third a little
longer and broader than the second; following becoming just
perceptibly shorter and somewhat narrower toward the tip.
Pronotum quadrate, as broad as long. Median suture and basal
impressed line distinct. Elytra with indistinct scutellar and
humeral dark spots. Below with denuded spots as above. Anterior
tibiae of male with a single spiniform spur. Male anterior tarsi
with first segment slightly contorted, not denuded. Posterior
tibial spurs moderately broad, flattened, pointed.
Type: Holotype cf, A.M.N.H.

Type locality: San Xavier, near Tucson, Arizona, July 24, 1916.
Additional localities: Arizona: Tempe, Phoenix.

REJECTED SPECIES

Epicauta levettei Casey

Epicauta levetti Casey, 1891, Ann. Lye. N. Y. Acad. Sci. 6:171.

Casey described this species from the Le Vette cabinet, based
on specimens with "Col." and "Colo." state labels. There are
specimens with similar state labels in the Psota collection and also
some with "Tex." labels. The type is identical with anthracina
(Erichson) from northern South America. Probably these speci-
mens are old dealers' specimens with erroneous labels. I have seen
no authentic specimens from even as far north as Panama.

516

bulletin: museum of comparative zoology

INDEX TO SPECIES

abadona 435

abandona 435

alastor 436

albida 511

albolineata 451

alphonsii 480

alpina 453

andersoni 444

anthracina 515

arizonica 507

articularis 489

aspera 488

atrata 470

atrivittata 509

balli 460

barberi 475

batesii 468

brunnea 454

calcarata 477

californica 479

callosa 468

caviceps 481

Chapmani 464

cinerea 456

cinerea 497

cinereus 456

clematides 456

conferta 448

confertus 448

conspersa 441

convolvuli 470

coracina 470

corvina 446

crassitarsis 439

debilis 497

diversipubescens 487

duplicate/, 451

elongatocalcarata 472

ensii'ormis 462

excavatif rons 484

excors 503

fabricii 497

fallax 450

ferruginea 465

fissilabris 456

flavocinerea 500

flavocinereus 500

floridensis 458

fortis 466

foxi 483

fulvescens 489

funebris 447

gissleri 496

heterodera 478

hirsutipubescens 512

immaculata 489

immerita 472

impressifrons 482

ingrata 455

insignis 437

kansana 476

languida 501

lauta 492

lemniscata 463

levettei 515

liebecki 506

linearis 513

linearis 505

longicollis 508

luteicornis 511

maculata 441

maculifera 514

magnomaculata 445

rnarginata 456

maura 479

megacephala 491

merkeliaha 494

mimetica 453

morio 447

murina 499

mutchleri 435

nigerrima 446

nigra 447

nigricornis 467

nigritarsis 438

nogales 442

normalis 442

oblita 475

occidentalis 465

ochrea 495

oregona 461

pardalis 443

parked 497

parva 449

parvulus 449

parvus 449

pedalis 440

pennsylvanica 447

pensylvanica 447

phoenix 443

piceiventris 472

polingi 505

protarsalis 495

pruinosa 471

puncticollis 474

puntipennis 459

purpurea 504

WERNER: NORTH AMERICAN EPICAUTA

517

rehni 485

rileyi 482

ruidosana 459

sanguinicollis 473

segmenta 490

segmentata 490

sericans 469

solani 457

straba . 483

strigosa 467

stuarti 462

subglabra 499

sublineata 491

techanus 448

tenella 493

tenuilineata 502

tenuis 503

texana 512

torsa 507

trichrus 470

trichura 470

unicolor 497

uniforma 452

valuta 490

ventralis 444

virgulata 512

vittata 464

watsoni 478

wheeled 486

PLATES

PLATE 1

Werner — North American Epicauta

PLATE 1

Antennae

Fig. 1 . E. floridensis cf

2. E. floridensis 9

3. E. cinerea c?

4. E. cinerea 9

5. E. Solani <?

6. E. brwnnea c?

7. E. brunnea 9

8. -B. purpurea d" (posterior tibia)

BULL. MUS. COMP. ZOOL.

Werner. North American Epicauta. Plate 1

PLATE 2

Werner — North American E picauta

PLATE 2

Antennae

Fig. 9. E. lemniscata

10. E. occidentalis

11. E. calif omica

12. E. alastor cT

13. E. alastor 9

14. E. pensylvanica d1

15. E. insignis c?

16. E. oregona d"

BULL. MUS. COMP. ZOOL. Werner. North American Epicauta. Plate 2

PLATE 3

Werner — North American Epicauta

PLATE 3

Antennae

Fig. 17.

E. ingrata cf

18.

E. atrata

19.

E. ferruginea

20.

E. fortis

21.

E. sericans

22.

E. puncticollis

23.

E. oblita

BULL. MUS. CO MP. ZOOL. Werner. North American Epicauta. Plate3

PLATE 4

Werner — North American Epicauta

PLATE 4

Antennae

Fig. 24. E. atrivittata c?

25. E. albida d>

26. E. texana d1

27. E. albida 9

28. E. longicollis cf

BULL. MUS. COMP. ZOOL.

Werner. North American Epicauta. Plate 4

PLATE 5

Werner — North American Epicauta

Fig. 29. E. longicollis 9

30. E. segmenta cT

31. E. segmenta 9

32. E. immaculata c?

33. E. immaculata 9

34. E. tenuis d>

PLATE 5

Antennae

BULL. MUS. COMP. ZOOL. Werner. North American Epicauta. Plate 5

PLATE 6

Werner — North American Epicauta

PLATE 6

Antennae

Fig. 35. E. tenella c?

36. E. lauta d"

37. E. polingi c?

38. E. liebecki <?

39. E. arizonica 6"

40. E. -polingi 9

41. E. tenuilineata d"

42. E. ochrea cf

43. E. gissleri o" -

44. E. purpurea d1

BULL. MUS. COMP. ZOOL. Werner. North American Epicauta. Plate6

PLATE 7

Werner — North American Epicauta

PLATE 7

Antennae

Fig.. 45.

E. torsa cf

46.

E. torsa 9

47.

E. fabricii cf

48.

E. fabricii 9

49.

E. murina cf

50.

E. murina 9

51.

E. flavocinerea

52.

E. flavocinerea

53.

E. virgulata cf

54.

E. linearis cf

55.

E. linearis 9

BULL. MUS. COMP. ZOOL. Werner. North American Epicauta. Plate 7

3J?

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