(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Proceedings of the California Academy of Sciences, 4th series"

PROCEEDINGS 
OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Volume 50 



Published 1997-1998 by the 

California Academy of Sciences 

Golden Gate Park 

San Francisco, California 



PROCEEDINGS 
OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Volume 50 



Published 1997-1998 by the 

California Academy of Sciences 

Golden Gate Park 

San Francisco, California 



SCIENTIFIC PUBLICATIONS DEPARTMENT 

Alan E. Leviton, Editor 
Katie Martin, Managing editor 



ISSN 0068-547X 



D California Academy of Sciences 

Golden Gate Park 
San Francisco, California 941 18 



PRINTED IN THE UNITED STATES OF AMERICA 
BY ALLEN PRESS, INC., LAWRENCE, ICANSAS 66044 



TABLE OF CONTENTS 
VOLUME 50 

1 . Brown, Walter C, Angel C. Alcala, Arvin C. Diesmos and Ely Alcala. Species of the 
Guentheri group of Platymantis (Amphibia: Ranidae) from the Philippines, with descriptions 
of four new species. (Illus.) October 8, 1997 1-20 

2. Brown, Walter C. Biogeography of amphibians in the islands of the southwest Pacific. 
(Illus.) October 8, 1997 21-38 

3. Penny, Norman D., Phillip A. Adams and Lionel A. Stange. Species catalog of the 
Neuroptera, Megaloptera, and Raphidioptera of America north of Mexico. (Illus.) 
December 9, 1997 39-1 14 

4. Zeigler, Carl V., Gordon L. Chan and Lawrence G. Barnes. A new late Miocene 
balaenopterid whale (Cetacea: Mysticeti), Parabalaenoptera baulinensis, (new genus and spe- 
cies) from the Santa Cruz mudstone, Point Reyes Peninsula, California. (Illus.) 

December 9, 1997 115-138 

5. GOSLINER, Terrence M. and David W. Behrens. Five new species of Chromodoris (Mol- 
lusca: Nudibranchia: Chromodorididae) from the tropical Indo-Pacific Ocean. (Illus.) 
February 11, 1998 139-165 

6. Brailovsky, Harry and Ernesto Barrera. A review of the Costa Rican species of Lepto- 
glossus Guerin, with descriptions of two new species (Hemiptera: Heteroptera: Coreidae: Corei- 
nae: Anisocelini). (Illus.) February 11, 1998 167-184 

7. MCCOSKER, JOHN E. A revision of the snake-eel genus Callechelys (Anguilliformes: Ophich- 
thidae) with the description of two new Indo-Pacific species and a new callenchelyin genus. 
(Illus.) April 27, 1998 185-215 

8. DANIEL, THOMAS F. Pollen morphology of Mexican Acanthaceae: diversity and systematic 
significance. (Illus.) April 27, 1998 217-256 

9. Rastegar-Pouyani, Nasrullah. A new species of Acanthodactylus (Sauria: Lacertidae) 
from Qasr-E-Shirin, Kermanshah Province, Western Iran. (Illus.) 

June 19, 1998 257-265 

10. Rastegar-Pouyani, Nasrullah and Goran Nilson. A new species of Lacerts (Sauria: Lac- 
ertidae) from the Zagros Mountains, Esfahan Province, West-Central Iran. (Illus.) 

June 19, 1998 267-277 

1 1. GOSLINER, Terrence M. and David W. Behrens. Two new discodorid nudibranchs from the 
western Pacific with a redescription of Doris luteola Kelaart, 1858. (Illus.) 

July 20, 1998 279-293 

12. Johnson, Rebecca F. and Terrence M. Gosliner. The genus Pectenodoris (Nudibranchia: 
Chromodorididae) from the Indo-Pacific, with the description of a new species. (Illus.) 

July 20, 1998 295-306 

13. Valdes, Angel and David W. Behrens. A new species of Doriopsilla (Mollusca, Nudibran- 
chia, Dendrodorididae) from the Pacific coast of North America. (Illus.) 

July 20, 1998 307-314 

14. Roberts, Tyson R. and Carl J. Ferraris. Jr. Review of South Asian sisorid catfish genera 
Gagata and Nangra, with descriptions of a new genus and five new species. (Illus.) 
September 10, 1998 315-345 

15. Gosliner, Terrence M. and Shireen J. Fahey. Description of a new species of Halgerda 
from the Indo-Pacific with a redescription of Halgerda elegans Bergh, 1905. (Illus.) 
September 10, 1998 347-359 

16. Spaulding, Sarah A. and J. Patrick Kociolek. New Gomphonema (Bacillariophyceae) 
species from Madagascar. (Illus.) November 6, 1998 361-379 



PROCEEDINGS TABLE OF CONTENTS, VOL. 50 

1 7. Alcala, Angel C, Walter C. Brown and Arvin C. Diesmos. Two new species of the 
genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. (Illus.) 

November 6, 1998 381-388 

18. VALDES, ANGEL and JESUS ORTEA. A new species ofDoriopsilla (Mollusca, Nudibranchia, 
Dendrodorididae) from Cuba. (Illus.) December 18, 1998 389-396 

19. McCosker, JOHN E. and Richard H. Rosenblatt. A revision of the eastern Pacific snake- 
eel genus Ophichthus (Anguilliformes: Ophichthidae) with the description of six new species. 
(Illus.) December 18, 1998 397-432 

Index to Volume 50. March 29, 1999 433-445 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 1, pp. 1-20, 5 figs., 4 tables. 



October 8, 1997 



SPECIES OF THE GUENTHERI GROUP OF PLATYMANTIS 

(AMPHIBIA: RANIDAE) FROM THE PHILIPPINES, WITH 

DESCRIPTIONS OF FOUR NEW SPECIES 

/ °cr* 

By *°h 

Walter C. Brown 

Department of Herpetology. California Academy of Sciences, San Francisco, California 941 18 






Angel C. Alcala 

Commission on Higher Education, DAP Building, San Miguel Ave., Ortigas Center, 

Pasig City, Philippines 



Ho, 




Arvin C. Diesmos 

College of Arts and Sciences, University of the Philippines, Los Barios, Laguna, Philippines 

and 

Ely Alcala 

Center for Tropical Conservation Studies, Silliman University, Dumaguete City 6200, Philippines 



Species of the guentheri group of Philippine Platymantis are revised. Platymantis guentheri is 
rediagnosed. Four species, previously confused with guentheri, (P. rabori, P. negrosensis, P. 
luzonensis, and P. banahao) are described. 

Received March 14, 1996. Accepted May 30, 1996. 



In a recent paper (Brown et al. 1997) three 
species-groups (assemblages) of Philippine 
Platymantis were diagnosed, based on different 
combinations of digital characters. In the same 
paper, species of the hazelae group were revised. 

For convenience those diagnoses are briefly 
restated. The hazelae Group. — This Group is 
diagnosed by the following combination of char- 
acters: ( 1 ) terminal phalanx a wide "T"; (2) disks 
of fingers, except first, broadly dilated; (3) first 
finger much shorter than second, not reaching 
beyond midpoint between subarticular tubercle 



and disk; (4) digits proximal to disks much 
broader than deep, the result of wide dermal 
flanges; (5) subarticular tubercles large, round, 
and only moderately protruding; (6) toe disks 
smaller than finger disks. 

The guentheri Group. — (1 ) terminal phalanx 
a moderate to wide "T"; (2) disks of fingers, 
except first; moderately to broadly dilated; (3) 
first finger shorter than second, reaching as far as 
base of disk for some species; (4) digits proximal 
to disks about as deep as broad, the result of 
narrow, dermal flanges; (5) subarticular tuber- 



[1] 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 



cles large and strongly protruding; (6) toe disks 
smaller than finger disks. 

The dorsalis Group. — (1) terminal phalanx 
bluntly rounded to pointed; (2) tips of fingers 
blunt, without disks, or with small to moderate 
disks; (3) first finger slightly shorter, about as 
long as, or slightly longer than second; (4) digits 
proximal to disks about as deep as broad; (5) 
subarticular tubercles large, strongly protruding 
and frequently pointed; (6) finger disks smaller 
than to slightly larger than toe disks. 

History of the Collections of Philippine 
Species of the guentheri Group 

The database for this revision of 'the Platyman- 
tis guentheri Group is the result of over a hundred 
years (1882-1996) of field sampling and obser- 
vations by numerous workers. The accumulation 
of data before 1954 was very slow. It has accel- 
erated since then. It is appropriate that readers 
understand some of the reasons for this, as well 
as the problems that limit the collection of field 
data even now. 

During the period from 1882 to 1954, the spe- 
cies of the guentheri Group, like those of the 
hazelae Group (Brown et al. 1997) were only 
rarely collected because of the sampling tech- 
niques typical of most field work in the Philip- 
pine Islands during that period. Taylor's 
extensive surveys on several islands during the 
period 1912-1 922 produced only a few examples 
of any species of these two species Groups. 

Cornufer (= Plaftmantis) guentheri was de- 
scribed on the basis of a single specimen from 
Dinagat Island (Boulenger 1882). This species 
was redescribed by Taylor (1920) based on two 
specimens from Bunawan, Agusan del Norte 
Province, Mindanao. He does not mention addi- 
tional examples of this species in his later papers 
(1922a, 1922b, 1922c, and 1923). In the 1920 
paper, however, he did recognize a second spe- 
cies of ihe guentheri Group, Cornufer worcesteri 
from Mt. Apo in Davao Province, Mindanao 
(Stejneger 1905). Taylor's surveys on several of 
the islands during the period 1912-1922 em- 
ployed sampling techniques which exploited the 
fauna inhabiting the forest floor and low shrub 
and tree habitats, but not the more arboreal habi- 
tats of the species in the guentheri and hazelae 
Groups. 

During the period from 1922 until World War 
II little field work was undertaken. Thus at the 



time of Inger's (1954) monograph on the Philip- 
pine frogs, the samples of P. guentheri were still 
very limited: Philippines 2, Dinagat 1 (type of 
guentheri), Luzon 3, Polillo 4, Leyte 1, and Min- 
danao 8 (including the type of Cornufer wor- 
cesteri). 

In 1954 the senior authors (Brown and A. 
Alcala) began their Philippine herpetofaunal 
studies that have continued to the present time. 
A population of Platymantis guentheri on Ne- 
gros Island was the first identified (Brown and 
Alcala 1957). Subsequently large samples from 
populations on both Bohol and Negros Islands 
were added to the data. This was the result of two 
factors: (1) the study of herpetofaunas of these 
two areas from a base at Silliman University for 
a period of more than a decade (1956-1972) and 
(2) the use of sampling techniques designed to 
exploit the arboreal microhabitats of amphibians, 
such as Platymantis and Philautus, as well as 
some genera of geckos and skinks. Other areas 
such as Dapitan Peak of the Malindang massif, 
Zamboanga Peninsula, and Mt. Hilong-hilong in 
northeastern Mindanao were surveyed during 
one or two expeditions to those rather remote 
regions using the same sampling techniques, but 
the samples were small (Alcala 1986). The large 
samples accumulated for Negros and Bohol did 
provide evidence of the presence of two species 
on Bohol (Brown and Alcala 1963) but only one 
species on Negros. 

It is also important for the reader to understand 
that there are difficulties which limit follow-up 
and repetition of some of these surveys. Exten- 
sive logging on both Negros and Bohol has 
greatly reduced the areas of forest normally oc- 
cupied by these frogs. A brief survey of forest 
remnants on Cuernos de Negros, southern Ne- 
gros in 1996 confirmed the continuing presence 
of a population of negrosensis in that area. 
Whether or not there is an extant population of 
either guentheri or rabori in the forest remnants 
on Bohol remains to be seen. We hope to be able 
to determine this within the year. A follow-up 
expedition to the Mt. Hilong-hilong area in 
northeastern Mindanao presents greater difficul- 
ties: ( 1 ) the lack of security in the region is a risk 
to personnel involved and (2) the survey would 
require a large commitment of men and re- 
sources. 

The recent modest samples of luzonensis and 
banahao from Mt. Banahao and Mt. Maquiling 



BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANTIS 



are the result of a base at the University of the 
Philippines at Los Bafios and the continuing 
work in the forest of the two mountains by Alcala 
and Diesmos over the period 1994-1996. Re- 
cently, populations of Platymantis guentheri on 
Cataduanes Island (Ross and Gonzales 1 992 ) and 
Panaon Island at the southern tip of Leyte 
(Gaulke 1994) and a population of P. ingeri on 
Dinagat Island (Ross and Lazell 199 1 ) have been 
reported, but the samples are limited to a few 
specimens. 

Materials and Methods 

Materials examined include: (1) holotypes of 
Platymantis guentheri, worcesteri, and ingeri; 
(2) 17 of the 19 specimens examined by Inger 
(1954); and (3) most specimens from recent col- 
lections that are referable to the guentheri group 
of species. These materials are in the custody of 
the following institutions: British Museum of 
Natural History (BMNH), California Academy 
of Sciences (CAS or CAS-SU), Field Museum of 
Natural History (FMNH), Museum of Compara- 
tive Zoology (MCZ), United States National Mu- 
seum (USNM), Delaware Museum of Natural 
History (DMNH), and Carnegie Museum (CM). 

Morphometric characters, including snout- 
vent length (SVL), head length (HL), head width 
(HW), snout length (SnL), diameter of eye (ED), 
diameter of tympanum (TD), tibia length (TiL), 
third finger length from proximal edge of basal 
tubercle (3FL), diameter of third finger disk 
(3FD), and diameter of third toe disk (3ToD) 
were measured to the nearest 0.1 mm using a 
Helios dial caliper. Other morphological charac- 
ters include: webbing of toes, shape of terminal 
phalanges, tubercles of hands and feet, skin or- 
namentation, head shape, vomerine teeth, and 
color pattern. 

Systematic Section 

Key to guentheri Group Species 
la. Dorsum smooth, without distinct ridges 
and usually with few tubercles mostly lim- 
ited to upper eyelids and dorsolateral area 

(Fig. 4A, B) 2 

lb. Dorsum with prominent ridges and scat- 
tered tubercles (Fig. 2) 3 



2a. Third toe not webbed beyond basal tuber- 
cle on outer side of third toe; brown pig- 
mentation of venter prominent on belly as 
well as head and throat; advertisement call 
(Fig. 1A) P. banahao 

2b. Third toe webbed to midpoint tubercles or 
beyond on outer side of third toe; brown 
pigmentation of venter mostly under head 
and throat; advertisement call (Fig. IB) . . 
P. luzonensis 

3a. TiL usually greater than 55% of SVL; 3FD 
less than 6% of SVL and usually less than 
100% of TD; SVL 27^10 mm for males . . 
P. guentheri 

3b. TiL usually less than 52 or 54% of SVL; 
3FD usually more than 1 15% of TD and 
7-10% of SVL; SVL 39-50 mm for males 
4 

4a. Third toe not webbed beyond basal tuber- 
cle on outer side; numerous long ridges on 
dorsum; snout with few or without tuber- 
cles; venter with scattered, small granules 
P. rabori 

4b. Third toe webbed to about midpoint be- 
tween tubercles or more distal on outer 
side; few long ridges on dorsum; snout 
with 2 or 3 rows of tubercles; venter nearly 

smooth; advertisement call (Fig. lc) 

P. negrosensis 

The Platymantis guentheri-ingeri Problem. — 
Reexamination of 17 of the 19 specimens as- 
signed to Platymantis guentheri by Inger (1954) 
indicates that the Mindanao sample includes four 
examples (CM 3424-25 and FMNH 50571, 
50573) that fit the diagnosis of P. ingeri. Reex- 
amination of the holotype of Platymantis 
guentheri along with the small series from Dina- 
gat Island reported as P. ingeri by Ross and 
Lazell (1991) shows them to be one species that 
better fits the diagnosis of P. ingeri (Brown and 
Alcala 1963) than that of P. guentheri (Brown 
and Alcala 1963, or Inger 1954). 

In view of this fact and because of the avail- 
ability of larger samples from several islands, a 
rediagnosis and redescription of Platymantis 
guentheri is necessary. Comparison of the ingeri- 
like populations of Dinagat, Mindanao, and Bili- 
ran islands shows them to be in general 
agreement with the type of guentheri, and they 
are therefore assigned to that species. Because 
the Bohol population, described as ingeri, re- 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 



veals only a few minor differences in mor- 
phometric characters, it is treated as a population 
of guentheri. Platymantis guentheri is presently 
known only from the Greater Mindanao group of 
islands. 

Platymantis guentheri (Boulenger) 

Cornufer guentheri: Boulenger, 1882:108 (Type loc.: 

Dinagat Island, Philippines; holotype in BMNH); 

(part) Taylor, 1 920:308; and (part) Inger, 1 954:362. 
Cornufer ingeri: Brown and Alcala, 1963:672 (Type 

loc.: Cantaub, Bohol Island; holotype in CAS). 
Platymantis guentheri: (part) Brown and Alcala, 

1970b: 109. 

DIAGNOSIS. — Plan-mantis guentheri differs 
from Philippine species of the hazelae and dor- 
salis Groups in the combination of digital char- 
acters used to diagnose the groups (see 
Introduction); and from other guentheri Group 
species in the following combination of charac- 
ters: ( 1 ) the longer tibia relative to S VL (TiL very 
rarely less than 57% of SVL and HW nearly 
always less than 75% of TiL); (2) the smaller 
finger disks relative to finger length (3FD 4-5% 
of SVL, rarely as great as 35% of 3FL and very 
rarely greater than 100% of TD); and (3) the 



smaller size at maturity (about 25-40 mm); see 
Tables 1, 2 and 3. 

Description. — SVL at maturity 24.4-33.2 
mm for 32 males and 27.2-39.6 mm for 45 fe- 
males; HW 99-112% of HL and 38-44% of 
SVL; snout round, pointed; upper jaw slightly 
protruding; SnL 35-45% of HL and 31^1% of 
HW; eye large; ED 71-97% of SnL, 10-13% of 
SVL, and 25-33% of HW; tympanum large; 
canthus rounded; lores concave, moderately 
oblique; vomerine teeth prominent; fingers with- 
out webs; digits proximal to disks about as wide 
as deep, with only narrow dermal flanges; first 
finger much shorter than second, reaching about 
midpoint between subarticular tubercle and disk 
of second; finger disks, except first, moderately 
large; 3FD 4-5% of SVL, 25-35% of 3FL, and 
73-108% of TD; subarticular tubercles strongly 
protruding, usually somewhat pointed; one row 
of supernumerary tubercles; inner and middle 
metacarpal tubercles large and oval, outer 
smaller and round; hind limbs long as measured 
by tibia, TiL 56-64% of SVL and HW 63-75% 
of TiL; toes webbed: to proximal edge of tubercle 
on first and second, to various levels of basal 
tubercle on third, to basal tubercle on fourth, and 
slightly distal to the basal tubercle on the fifth; 



Table 1. Snout vent length (SVL, in mm) and extent of toe webs for adults of Philippine species of the 
guentheri Group of Platymantis (N = number in sample, F = female, M = male). 



Species 



SVL 



Webbing on 3rd Toe 



P. guentheri 

F 

M 
P. rabori 

F 

M 
P. banahao 

F 

M 
P. negrosensis 

F 

M 
P. luzonensis 

F 

M 



45 


27.2-39.6 


32 


24.4-33.2 


27 


384-49.1 


22 


274-33.2 


11 


27.8-394 


15 


39.3-50.2 


25 


29.8-39.3 


8 


35.1-45.3 


12 


27.1-36.0 



not beyond basal tubercle 



not beyond basal tubercle 



not beyond basal tubercle 



midpoint between tubercles 



midpoint between tubercles 



BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANTIS 



Table 2. Comparison of body proportions (in %) of five species of the guenlheri Group (N = number of 
specimens). 



Species 



HW/TiL 



3FD/TD 



3FD/3FL 



P. guentheh 

Range 

N 
P. rabori 

Range 

N 
P. negrosensis 

Range 

N 
P. luzonensis 

Range 

N 
P. banahao 

Range 

N 



62-74 
43 

78-90 
40 

80-92 

20 

79-93 
21 

85-94 
12 



73-108 
43 


22-35 
40 


114-185 
30 


36-^4 

34 


1 30-2 1 1 
20 


35-47 
20 


116-193 
20 


30-44 
21 


100-193 
12 


31-^10 
11 



disks of toes smaller than those of fingers, 3ToD 
55-86% of 3FD; subarticular tubercles strongly 
protruding, usually somewhat pointed; inner 
metatarsal tubercle oval; outer round; no plantar 
tubercles; dorsum nearly smooth or with a few 
tubercles and frequently with a pair of short, dark 
ridges in a chevron-pattern at shoulder level; 
venter nearly smooth; posterior thighs nearly 
smooth or with small granules. 

COLOR. — In preservative dorsum light to 
dark reddish brown or a mottling of the two; 
usually an anteriorly pointed, darker chevron at 
the shoulder level; often an interorbital dark 
blotch, more rarely a broad dorsal, paler band or 
narrow, pale, dorsolateral stripes; lips dark 
barred; limbs with pale and dark, irregular trans- 
verse bands; venter pale cream, sometimes 
nearly uniform, but usually with dense, brown 
flecks beneath head and throat; belly often with 
scattered brown spots, especially anteriorly. 

Reproduction. — None of the egg clutches 
collected from any of the arboreal sites in the 
forests on Bohol Island or Mt. Hilong-hilong in 
Mindanao have been confirmed as guentheri. 
Several gravid females from these sites have 
20-24 large (2-3 mm) unpigmented, ovarian or 
oviducal eggs. These are similar to those of other 
species of Philippine Platymantis that exhibit 



direct development, and presumably guentheri 
follows that mode. 

COMPARISONS. — Platymantis guentheri dif- 
fers from rabori, negrosensis, luzonensis, and 
banahao, species with which it has long been 
confused, as indicated in the diagnosis and key. 

Inter-island Variation. — The samples of 
guentheri from Dinagat, Mindanao, and Biliran 
islands when compared to the Bohol sample, 
exhibit slightly greater SVL for females but not 
for males (Table 2). 

ECOLOGICAL Note. — As reported by Brown 
and Alcala (1963) for ingeri (= guentheri), this 
species is found in the surface stratum (among 
leaves and duff of the forest floor, occasionally 
under logs or rocks, and less frequently in the 
shrub or arboreal strata). Of a total of 91 speci- 
mens (67 Bohol and 18 northeastern Mindanao), 
67 (74%) were from the surface stratum; 4 
(4.5%) low shrubs, 5(5.5%) on trees, and 15 
( 1 6%) in arboreal ferns or leaf axils of Pandanus. 
The ferns inhabited by guentheri ranged from 4 
to 13 m above the forest floor. The range in 
elevation is from 700 m (upper dipterocarp) 
to 1900 m (montane forest) on Mt. Hilong- 
hilong in northeastern Mindanao. 

RANGE. — Known from Mindanao, Dinagat, 
Bohol, and Biliran islands. 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 



Table 3. TiL, HW, ED, TD, FD, and 3FL as a% of SVL. Species of the guentheri Group (N = number 
of specimens). 



Species 


TiL/SVL 


HW/SVL 


ED/SVL 


TD/SVL 


3FD/SVL 


3FL/SVL 


P. guentheri 














Range 


56-64 


38-43 


10-13 


4-6 


4-5 


14-17 


N 


20 


20 


20 


20 


20 


20 


P. rabori 














Range 


48-54 


40-46 


13-17 


5-7 


8-10 


20-23 


N 


20 


20 


20 


20 


20 


20 


P. negrosensis 














Range 


47-52 


38-43 


11-15 


4-6 


7-9 


19-22 


N 


20 


20 


20 


20 


20 


20 


P. luzonensis 














Range 


45-53 


39^5 


12-14 


4-6 


6-8 


17-22 


N 


20 


20 


20 


20 


20 


20 


P. banahao 














Range 


45^9 


40-44 


10-14 


4-6 


5-8 


17-21 


N 


12 


12 


11 


11 


11 


11 



Populations Confused with Platymantis 
guentheri 
Populations on Mindanao, Leyte, Bohol, Ne- 
gros, Luzon, Catanduanes, and Polillo islands 
that have erroneously been referred to guentheri 
since 1954 (see Inger 1954, Alcala and Brown 
1957, Brown and Alcala 1963, and Ross and 
Gonzales 1992) must now be reevaluated. A 
search of the literature reveals only one name that 
is possibly available, Cornufer (= Platymantis) 
worcesteri Stejneger (1905). Platymantis wor- 
cesteri is treated as a distinct species by Taylor 
( 1 920), but as a synonym off. guentheri by Inger 
(1954). However, reexamination of the unique 
holotype of worcesteri from Mt. Apo, Mindanao 
reveals the presence of an intercalary cartilage in 
the digital skeleton, placing this species in the 
Rhacophoridae. The shape of the terminal pha- 
lanx identifies it as a Philautus (Dring 1987; 
Brown and Alcala 1994). Its specific status and 
possible priority are the subject of a separate 



paper. Island populations previously confused 
with guentheri are therefore all unnamed. 

Recent collections (about 35 specimens) from 
Mt. Maquiling and Mt. Banahao in southern 
Luzon provide evidence that two, cryptic species 
are sympatric on Luzon, as is the case for Min- 
danao and Bohol islands. Comparison of these 
Luzon samples with the large samples (more than 
50 specimens from each of the Greater Mindanao 
and Negros areas) indicates that these four popu- 
lations differ in several morphological and mor- 
phometric characters. These differences are 
sufficiently great that we treat the populations 
from the Greater Mindanao, Greater Negros, and 
the two from Greater Luzon areas as distinct 
species. The advertisement calls for the two 
populations on Luzon and the population on Ne- 
gros also differ from one another (Fig. 1). 



-> 



Figure 1. Advertisement calls of: A. Platymantis banahao (CAS 201532), B. P. luzonensis (PNM5495), C. P. negrosensis 
(PNM 5493). 



BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANTIS 



fliH-'tf * ' j 



o ss^iLlliraias^iM^ 



II 1 V! 



| 



rsi 

x 



>- 
(J 

z 

LU 

a 



m 



B 



Q ■• •■ i-l p -,t |i!^ 1, ■ ■ ii;. >fil ,i-|i|»n.|j n-ri =' M.||, ri |fh"ir r i! |»"lr,W ;i-'ii|.|«|i 




1 TIME (sec) 2 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 



Platymantis rabori new species 
(Fig. 2) 

Cornufer guentheri (part): Brown and Alcala, 

1963:674. 
Platymantis guentheri (part): Brown and Alcala, 

1970b: 109. 

The diagnosis and description of this species 
are based on a large sample (230 specimens) 
from Bohol Island. Small samples of five or six 
specimens from Leyte and Mindanao islands are 
assigned to this species but are not designated as 
paratypes. 

Holotype. — CAS 136889, a gravid female, 
collected at Cantaub, Sierra Bullones, Bohol Is- 
land, February 13, 1972, by L. Alcala and party. 

Paratypes. — Bohol Island (same general lo- 
cality as holotype): CAS-SU 17284, 21721, 
21723,21733,21738-39,21741,21748,21756, 
21775-81, 23148-64, 23166-68, 23170-82, 
23184-212, 23222-23, 23425; CAS 136888, 
136890-137006, 138169-70, 139019-21, 
145697, 145699, 186065; FMNH 134987-88; 
MCZ 39052-53; PNM 5282. 

Description of holotype. — A gravid female 
(measurement in mm) SVL = 48.9, HL = 18.5, 
HW = 20.7, TiL = 23.8, SnL = 7.8, ED = 6.6, TiL 
= 2.5, 3FL = 10.2, 3FD - 3.5, 3ToD = 1.8; 
dorsum with tubercles and some ridges; toe-webs 
to basal tubercle or beyond except for fourth toe. 

DIAGNOSIS. — Platymantis rabori differs 
from species of the hazelae and dorsalis Groups 
in the combination of characters diagnosing 
those groups (see Introduction). It differs from 
other species of the guentheri Group in these 
characters: (1) snout without tubercles or only a 
few; (2) dorsum with some tubercles as well as 
short and long ridges; (3) venter posterior to 
forelimbs with scattered, small granules; (4) toe 
webs reach about midpoint of tubercle on first 
and second toes, midpoint or distal edge of basal 
tubercle of third toe, and does not reach basal 
tubercle or only its basal edge on fourth toe. 

Description. — SVL 27.4-35.3 mm for 22 
males and 38.4-49.1 mm for 27 females; HW 
107-125% of HL and 40-46% of SVL; snout 
rounded, upper jaw only slightly protruding; SnL 
36^12% of HL and 30-40% of HW; ED 77-98% 
of SnL, 24-33% of HW, 13-17% of SVL; tym- 
panum exposed; TD 3 1-45% of ED and 5-7% of 
SVL; canthus rounded; lores moderately 



oblique, concave; fingers without webs and with 
very narrow, lateral, dermal flanges; first finger 
much shorter than second, reaching about mid- 
way between tubercle and disk or slightly be- 
yond; disks of fingers (except first) broadly 
dilated, truncate; 3FD 33^13% of 3FL and 
120-180% of TD; subarticular tubercles large, 
strongly protruding and somewhat pointed; one 
row of supernumary tubercles; inner metacarpal 
tubercle prominent, large oval; middle and outer 
large, low; hind limbs long, TiL 48-54% of SVL 
and HW 76-88% of TiL; toes webbed, first toe 
to base or midpoint of subarticular tubercle, sec- 
ond to about midpoint of tubercle, third to mid- 
point or distal edge of tubercle, fourth short of 
basal tubercle or to its proximal edge, fifth be- 
yond basal tubercle or midpoint between tuber- 
cles (Fig. 3a); subarticular tubercles like those of 
fingers; inner metatarsal tubercle prominent, 
elongate, outer small, round to oval; disks of toes 
smaller than of fingers, 3 ToD 41-51% of 3FD; 
plantar surface smooth; snout smooth or with a 
few tubercles; a white-tipped tubercle on poste- 
rior upper eyelid; dorsum posterior to eyes with 
some tubercles as well as short and intermediate 
length ridges; throat smooth; venter posterior to 
forelimbs with small, scattered granules; poste- 
rior thighs with moderate granules. 

COLOR. — In preservative background color 
of dorsum variable, tannish brown to light choco- 
late brown, dorsal ridges with narrow, blackish 
borders; a few specimens with a narrow, median 
pale stripe or a pair dorso-laterally; venter pale, 
with chocolate brown flecks and pale spots, groin 
and anterior thighs similarly flecked, but without 
prominent, pale creamy spots. 

In life the color is variable as in preservative 
but the dark background phase appears darker, 
almost blackish brown. The short dorsal ridges 
are outlined in black. The vertebral or median 
pale stripes, when present, are tannish brown to 
almost grayish. 

Reproduction. — Gravid females examined 
for eggs have large (about 2 mm) eggs in the 
ovaries or oviducts. Egg masses collected from 
arboreal ferns (Asplenium) or leaf axils of Pan- 
danus, habitats most frequently occupied by 
adults of this species as well as Platymantis 
guentheri and Philautus surdus, all give evidence 
of a direct development mode, but the only ones 
including sufficiently late stages to assign to 
genus and species are examples of Philautus 



BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANT1S 




Figure 2. Platvmantis rabori from Mindanao Island. 



swdus. None of the clutches have been positively 
identified with this species. 

Etymology. — Named for Dioscoro S. 
Rabor, Filipino ornithologist and mammalogist, 
who added greatly to our knowledge of the 
ranges of Philippine amphibians and reptiles. 

COMPARISONS. — Platvmantis rabori differs 
from guentheri, negrosensis, luzonensis, and ba- 
nahao as indicated in the diagnosis and key. 

ECOLOGICAL NOTE. — Preferred microhabitat 
is arboreal ferns or leaf axils of Pandanus. Of 56 
examples collected in 1962 in original forest near 
Cantaub, Siena Bullones, Bohol Island 50 were 
from arboreal ferns, 4 from leaf axils of Pan- 
danus and 2 from duff on the forest floor. The 
elevation is between 500 and 800 m (Brown and 
Alcala 1963). This contrasts with Platvmantis 
guentheri with 41 of 57 specimens from the 



forest floor and only 9 from arboreal ferns. In a 
later (1972) study of the arboreal fern microhabi- 
tat in the same forest area on Bohol, 88 specimens 
were P. rabori and 20 were P. guentheri. The 
height range of the ferns for P. rabori is 0.5 to 
12.5 m. 

RANGE. — Populations are recorded from Bo- 
hol, Leyte, and Mindanao islands. 

Platymantis negrosensis new species 

Cornufer guentheri: Alcala and Brown, 1957:182; 

Brown and Alcala, 1961:632, 1964:600. 
Platvmantis guentheri (part): Brown and Alcala, 

1970b: 109. 

This new species, like the sympatric Platyman- 
tis hazelae, has been extensively studied in the 



10 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 













?* i 



;■'-'■-'. V 



FiGURE 3. Undersurface of foot, showing webbing and tuberculation: A. Platymantis rabori (illustrative of the webbing- 
pattern of rabori, guentheri, and banahao); B. P. negrosensis (illustrative of the webbing-pattern of negrosensis and 

hizonensis). 



BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANT1S 






m 



il 



1 



■ ! 



Ik: 


" ^ ^'U : : 


£;■" :•'* 
?.*'•" 


' i ".-'*$S?zr, 


%. 




k }: 




/|: 


a 


It 


§j 



■J 






-hi 



:J: : ? 
















. ); - . f ?«" 










?.5j 







f 



B 



w 



3 

■8# 



n'. zawojsw -J>ec. 62 . 



12 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 



field over the past 35 years, and its biology is thus 
better known than that of most species of the 
genus. 

Holotype. — CAS 137416, an adult female, 
collected in original forest at Lake Baling- 
sasayao, Negros Oriental Province, Negros Is- 
land, on September 19, 1972, by L. Alcala. 

Paratypes. — Negros I. (same locality as holo- 
type): CAS 137386-415, 137417-28; Cuernos 
de Negros: CAS-SU 17996-18002, 18434-35, 
18487, CAS 89804; other localities of Negros I. 
CAS-SU 19527-30, 23703, CAS 128903-12 
128914, 128919, 133863, 133899-905, 133911 
133955-56, 133988, 134319, 134325-26 
137993-138000, 138151-60, 138213, 138215 
138232-33, 138236-39, 138245—46, 139281 
139283, 139286, 145611-20, 145807-08 
145851-52, 145892, 145973-74, 147332 
147335-43, 185651, 200202-07; PNM 5283 
USNM 512300. 

Description of holotype. — An adult female 
(measurement in mm): SVL = 50.2, HL = 17.7, 
HW = 120.9, SnL = 6.7, ED 5.7, TD = 2.5, TiL = 
24.4, 3FL = 10.3, 3FD = 4.6, 3 ToD= 1.9; dorsum 
with tubercles and short ridges; toe-webs beyond 
basal tubercle except for fourth toe. 

DIAGNOSIS. — Platymantis negrosensis dif- 
fers from the hazelae and dorsalis Groups of 
species in those characters that diagnose the 
groups (see Introduction); and from other species 
of the guentheri Group in: ( 1 ) two or three rows 
of tubercles on snout; (2) usually only short 
ridges and some tubercles on dorsum; (3) toe 
webs usually reach distal edge of tubercle or 
beyond for first and second toes, midway be- 
tween tubercles on third toe, and distal edge of 
basal tubercle or beyond on fourth; (4) venter 
nearly smooth; (5) advertisement call. 

Description. — SVL 29.8-39.3 mm for 25 
males and 39.3-50.2 mm for 15 females; HW 
105-125% of HL and 38^43% of SVL; snout 
rounded to round-pointed, SnL 35—42% of HL 
and 30-36% of HW; ED 86-104% of SnL, 
29-36% of HW, and 1 1-15% of SVL; tympa- 
num exposed, TD 25^13% of ED and 4-6% of 
SVL; canthus rounded; lores moderately 
oblique, concave; fingers without webs; First fin- 
ger much shorter than second, reaching beyond 
basal tubercle to midway between tubercles of 
second finger; disks of fingers (except first) 
broadly dilated, truncate and with narrow, der- 
mal flanges proximal to disks; 3FD 35^17% of 



3FL, 130-21 1% of TD, and 7-9% of SVL; 3FL 
19-22% of SVL; subarticular tubercles large, 
strongly protruding; one row of supernumary 
tubercles; inner metacarpal tubercle elongate to 
oval; middle and outer low, usually partially 
fused; hind limb moderately long, TiL 46-53% 
of SVL and HW 80-92% of TiL (only three of 
20 below 83%); toes webbed: to midpoint or 
distal edge of subarticular tubercle on first toe, 
distal edge or slightly beyond tubercle on second 
toe, about midway between tubercles or slightly 
beyond on third toe; basal tubercle or slightly 
beyond on fourth toe, and usually base of distal 
tubercle on fifth toe; disks of toes smaller than 
those of fingers, 3ToD 38-54% of 3FD; subar- 
ticular tubercles large, protruding; plantar area 
smooth; inner metatarsal tubercle elongate; outer 
round; plantar area smooth (Fig. 3b); usually two 
or three rows of low tubercles on snout; few to 
numerous short ridges and usually some tuber- 
cles on dorsum; a few including one pale-tipped 
tubercle on upper eyelids; venter usually smooth; 
posterior thighs with small granules. 

COLOR. — In preservative background color 
of dorsum and upper lateral surfaces somewhat 
reddish brown to chocolate brown; dorsal ridges 
with nauow, blackish borders; venter with dense 
brown flecks or numerous brown spots; groin 
area and adjacent surface of thighs creamy-white 
with brown intrusions. 

In life the dorsal background color is variable 
as it is for preserved specimens, but the dark 
specimens appear darker in life. The vertebral or 
dorsolateral stripes are grayish to tan. 

REPRODUCTION. — Platymantis negrosensis, 
like other species of Platymantis for which the 
life cycle is known, produce large, unpigmented 
eggs which undergo direct development within 
the egg capsule. Clutch size was reported as 8^47 
(Alcala 1962) but a couple of gravid females 
(SVL about 45 mm) indicate a clutch size of 
± 40. The clutch of eight from the leaf axil of an 
arboreal fern may have been that of Platymantis 
hazelae (clutch size 5-9). The developmental 
period to hatching is reported as 39+ days. For 
more detail on development see Alcala (1962) 
and Brown and Alcala (1983). 

Advertisement Call. — The call of ne- 
grosensis sounds like "kwek-kwek-kwek." The 
first, brief part of each note is between 1,000 and 
about 2,000 Khz. The second, longer part of the 
note is between 1,200 and about 2,300 Khz. The 



BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANTIS 



13 



Table 4. Some advertisement call parameters for Platymantis negrosensis, P. luzonensis, and P. 
banahao (n = number of voice records, s = seconds). 



Species 


n 


Sound to human 
ear 


Notes in a 
series 


Frequency 
(Khz) 


Length of note 
(s) 


P. negrosensis 


2 


Kwek-Kwek 


1 or 2-5 in 
succession 


1000-2300 


0. 1 06-0. 1 22 


P. luzonensis 


3 


Kwenk-Kwenk 


1 or 2-6 in 
succession 


1 800-3300 


0.106-0.116 


P. banahao 


4 


Tut-Tut-Tut 


6-24 in 
succession 


1 200-2400 


0.0593-0. 1 1 25 
(usually < 0.100) 



duration of each note is about 0.1060-0.1220 
seconds, and the time between notes ranges from 
about 0.15 to 0.54 seconds. There are usually 
three to four notes in quick succession following 
an initial note that is separated by a longer inter- 
val (see Fig. 1C and Table 4). 

ETYMOLOGY. — The name is derived from 
Negros, the name of the island where this species 
occurs. 

COMPARISONS. — Platymantis negrosensis 
differs from other species of the guentheri Group 
as indicated in the diagnosis and key. 

Ecological Note. — Of 36 specimens col- 
lected in the mountains of southern Negros dur- 
ing field work 1956-58, 28 are from arboreal 
ferns, 5 from leaf axils of Pandanus, 2 from leaf 
axils of gabi plants, and Ion the forest floor. This 
species occurs in the upper dipterocarp and sub- 
montane forest zones at elevations from about 
500 to 1050 m. During field surveys in the Lake 
Balinsasayo area in 1970 and 1972, this species 
was found in arboreal ferns ranging from about 
2-24 m above the forest floor. 

Range. — Known only from Negros Island, 
primarily from populations in the southern 
mountains. Only one specimen was recorded 
from Mt. Canlaon in northern Negros during a 
field survey in 1962. 

Platymantis luzonensis new species 
(Fig. 4) 

Platymantis guentheri (part): Inger, 1954:362; Brown 
andAlcala, 1970b: 109. 



Recent samples from populations on various 
mountains in southern Luzon Island and on 
Catanduanes Island have added to our knowl- 
edge of the variability of this species, but, except 
for the populations on Mt. Banahao and Mt. 
Maquiling, the samples are, at this time, too small 
to determine if they should be treated as subspe- 
cies or new species. We therefore limit the holo- 
type and paratypes to the Banahao-Maquiling 
samples. 

Holotype. — CAS 196368, a female, collected 
in forest at about 600 m elevation, Mt. Maquil- 
ing, Laguna Province, Luzon Island, Philippines, 
July 1 1, 1994, by Angel C. Alcala. 

Paratypes. — CAS 196364, 196369-70, 
200404-08, 201544-45, 201538-39 Mt. 
Maquiling; CAS 201218-21 Mt. Banahao at 
Lucban. 

Description of holotype. — An adult female 
(measurement in mm): SVL = 44.9, HL = 15.6, 
HW = 18.8, TiL = 20.3, SnL = 6.2, ED = 5.2, TD 
= 1.95, 3FL = 8.3, 3FD = 3.5, 3ToD = 1.7, TiL = 
20.3. A few low tubercles on dorsum, posteri- 
orly; toes about half webbed. 

DIAGNOSIS. — Differs from species of Platy- 
mantis of the hazelae and dorsalis Groups in 
those characters that diagnose the groups. Platy- 
mantis luzonensis differs from species of the 
guentheri Group, other than banahao, in the skin 
of snout and dorsum being smooth or nearly so. 
It differs from all species but negrosensis in the 
greater webbing which reaches the distal edge of 
the basal tubercle on first and second and midway 
between tubercles on fourth. It differs from 



14 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 




BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANTIS 



15 



banahao in the lighter pigmentation of the ven- 
ter. It also differs from negrosensis and banahao 
in its advertisement call. 

DESCRIPTION. — SVL 27.1-36.0 mm for 12 
mature males and 35.1^5.3 mm for 8 females; 
HW 106-123% of HL and 39^15% of SVL; 
snout round-pointed; upper jaw moderately pro- 
truding and 30-35% of HW: ED 83-100% of 
SnL, 26-34% of HW, and 12-14% of SVL; 
tympanum exposed; TD 28^45% of ED and 
4-6% of SVL; canthus rounded; lores moder- 
ately oblique, concave; fingers without webs, 
only slightly broader than deep throughout 
length proximal to disk; lateral, dermal flanges 
very narrow; first finger much shorter than sec- 
ond, reaching midway or slightly more between 
tubercle and disk of second finger; disks of fin- 
gers (except first) broadly dilated, truncate; 3FD 
30^13% of 3FL, 1 16-193% of TD, and 6-8% of 
SVL; 3FL 17-22% of SVL; subarticular tuber- 
cles large, strongly protruding; one row of super- 
numary tubercles; inner metacarpal tubercle 
large, oval, prominent large; outer small, more 
round; hind limbs long, TiL 44-54% of SVL and 
HW 79-93% of TiL; toes webbed: to about distal 
edge of basal tubercle on inside of first and 
second toes, nearly halfway between tubercles on 
third, and halfway between to near basal edge of 
distal tubercle on fifth toe (Polillo specimens 
exhibit similar webbing); disks of toes smaller 
than those of finger 3ToD 33^19% of 3FD; 
subarticular tubercles prominent, usually slightly 
pointed; plantar area smooth; inner metatarsal 
tubercle elongate; outer small round; dorsum and 
upper lateral surfaces smooth with no distinct 
ridges and at most only a few vague tubercles; 
venter smooth; posterior thighs with some small, 
low granules. Polillo specimens exhibit more 
tubercles and short ridges on dorsum and upper 
lateral surfaces. 

COLOR. — In preservative background color 
of dorsum tannish to pale brown with variable 
darker blotches; sometimes with pale dorsolat- 
eral streaks or a pale vertebral line; venter 
densely flecked with light brown under head and 
throat and scattered brown spots on belly. 

In life dorsal ground color variable from rather 
uniformly tannish including the lateral surfaces 
or the lateral surfaces darker. Some specimens 
are more rusty brown or dark brown bordered by 
narrow or broad, pale dorsolateral lines on the 
dorsum. There is usually a very dark interorbital 



bar, and lips and limbs with alternating pale and 
dark transverse bands. Brown flecks on venter 
are mostly under head and throat. 

Reproduction. — Four clutches of eggs and 
embryos still in their capsules were collected at 
Mt. Banahao de Lucban from arboreal ferns and 
leaf axils of Pandanus at altitudes of 
800-1,500 m on April 21 and September 17. 
1996. These have been identified as Platymantis 
based on Alcala (1962), but the species identifi- 
cation is still unknown. 

Advertisement Call. — The call of 
luzonensis sounds like "kwenk-kwenk-kwenk." 
Each note is in two parts. The first brief part is 
between 1,800 and 2,200 Khz; the second be- 
tween 2,200 and 3,100 or 3.200 Khz. The dura- 
tion of each note is about 0.1060 to 0.1156 
seconds, and the time between notes ranges from 
0.1437 to about 0.1 563 seconds (see Fig. IB and 
Table 4). 

COMPARISONS. — Platymantis luzonensis dif- 
fers from other species of ihe guentheri Group as 
indicated in the diagnoses and key. 

Ecological Note. — Twelve specimens 
from Mt. Maquiling were from various, arboreal 
sites (leaf axils of Pandanus and Araceae, leaves 
of shrubs and trees, tree trunks) 1-3 m above 
ground, in forest at 550-1200 m elevation. Six of 
the seven specimens from Mt. Banahao were 
found in sites 1-3 m above ground at elevations 
of 650-1, 250 m in the upper lowland to montane 
forests on Mt. Banahao; 600-1,160 m on Mt. 
Maquiling. 

RANGE. — Mt. Maquiling and Mt. Banahao, 
southern Luzon Island. 

Platymantis banahao new species 
(Fig. 5) 

This species is known thus far only from for- 
ests on the upper slopes of the Mt. Banahao 
massif. 

Holotype. — CAS 201208, an adult male, col- 
lected in original forest at 1100m on the NE 
slope of Mt. Banahao at Lucban, Quezon Prov- 
ince, Luzon Island, by R. Quiver and R. Reso, 
April 20, 1996. 

Paratypes. — CAS 201003-07, 201209-10, 
201231, 20153 1-32, 201544 Mt. Banahao (same 
general locality as holotype); CAS 201015 Mt. 
San Cristobal. Quezon Province, Luzon Island. 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. I 



Description of holotype. — An adult male 
(measurement in mm): SVL = 33.3, HL = 13.0, 
HW = 14.8, SnL = 4.8, ED = 3.9, TD = 2.0, TiL 
= 15.9, 3FL = 6.4, 3FD = 2.15; 3ToD = 1.1; 
dorsum free of tubercles or ridges; toe-webs 
reach basal tubercle except for fourth toe. 

Diagnosis. — Platymantis banahao differs 
from species of the hazelae and dorsalis Groups 
in those characters that diagnose these groups 
(see Introduction). It differs from other species 
of the guentheri Group in the following combi- 
nation of characters: (1) webs of toes not reach- 
ing distal end of basal tubercle or beyond on any 
toe except fifth; (2) heavily pigmented venter; (3) 
smooth dorsum, except for luzonensis; (4) a dis- 
tinctive advertisement call when compared with 
negrosensis and luzonensis. 

Description. — SVL 27.8-39.4 mm for 1 1 
males; HW 111-123% of HL and 40.44% of 
SVL; snout rounded; upper jaw only slightly 
protruding; SnL 36-44% of HL and 30-39% of 
HW; eye moderate, ED 73-92% of SnL, 26-30% 
of HW, and 10-14% of SVL; tympanum with 
thin skin; TD 44-54% of ED and 4-6% of SVL; 
canthus sharply rounded; lores moderately 
oblique, concave; fingers without webs; first fin- 
ger much shorter than second reaching about 
midway between tubercle and disk of second; 
fingers (except first) with broad, truncate disks; 
proximal to disks not much broader than deep 
and with narrow dermal flanges; 3FD 29-44% of 
3FL, 100-192% of TD, and 5-8% of SVL; 3FL 
17-21% of SVL; subarticular tubercles moder- 
ately protruding, broadly oval; inner metacarpal 
tubercle moderate, oval; middle one large round 
and outer small, round; a row of moderate to 
prominent supernumary tubercles; hind limb 
moderately long, TiL 45^9% of SVL and HW 
85-94% of TiL; toes with short webs: to tubercle 
on first and second toes, to proximal tubercle on 
third, not reaching proximal tubercle on fourth, 
and slightly beyond tubercle on fifth; disks of 
toes much smaller than those of fingers, 3ToD 
41-60%of3FD; subarticular tubercles like those 
of fingers; inner metatarsal tubercle elongate- 
oval; outer small round; plantar area smooth; 
dorsal surfaces smooth except for a tubercle pos- 
teriorly on the upper eyelid or occasionally a few 
scattered tubercles; upper lateral surfaces smooth 
or with a few, scattered, low tubercles; venter 
smooth; posterior thighs nearly smooth or with a 
few, vague tubercles. 



COLOR. — In preservative background color 
light to dark chocolate brown, with a few darker 
spots and small blotches; usually a paler band 
between the anterior half of the eyes; sometimes 
paler snout and upper lateral surfaces; lips nearly 
uniformly dark or with a couple of distinct, dark 
bars; hind limbs with vague, dark, transverse 
bars; venter heavily flecked with brown, slightly 
more dense on head and throat. 

In life the dorsal background color is nearly 
uniformly light tan to lemon yellow including the 
head as well as the body. Even an interorbital 
dark bar is absent. For a few specimens there are 
some dark flecks. The lateral surfaces for these 
specimens are chocolate brown. Other specimens 
are rusty chocolate on both dorsal and lateral 
surfaces. The venter is densely covered with 
brown flecks and spots. 

REPRODUCTION. — See note under Platyman- 
tis luzonensis. 

ADVERTISEMENT Call. — The call of bana- 
hao sounds like "tuk-tuk-tuk-tuk," usually re- 
peated rapidly 6-20 or more times like a machine 
gun. Each note is between 1,100 or 1,200 and 
2,100 to 2,400 Khz. The duration of each note is 
0.0593 to 0.1 125 seconds (usually less than 0.1 
second), and the time between notes ranges from 
0.1219 to 0.3562 seconds (see Fig. 1A and Table 
4). 

ETYMOLOGY. — The name is that of the 
mountain massif to which it appears to be en- 
demic. 

COMPARISONS. — Platymantis banahao dif- 
fers from other species of the guentheri Group as 
indicated in the diagnoses and key. 

Ecological Note. — One specimen was 
found on the forest floor and four in leaf axils of 
Pandanus. Field notes for the other six simply 
state arboreal. The sites are in original montane 
forest at elevations of 800 to 1400 m on Mt. 
Banahao at Lucban and Mt. San Cristobal in the 
Banahao massif. 

RANGE. — Mt. Banahao massif, southern 
Luzon Island. 

Summary 

Factors Limiting Sample Size 

In this revision of the Platymantis guentheri 

Group, we recognize five species: P. guentheri, 

P. rabori. P. negrosensis, P. luzonensis, and P. 

banahao. In the introduction we noted the distri- 



BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANTIS 



17 




Figure 5. Map of the Philippines showing present islands (light stippled areas) and late Pleistocene islands (dark stippled 
areas). 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 



bution patterns of the species, the reasons for the 
large samples from Bohol and Negros and rela- 
tively small samples from other localities, the 
recent reduction of suitable habitat for some 
populations, and the difficulties which limit fur- 
ther sampling in areas other than Cuernos de 
Negros (Negros Island) and Mt. Maquiling and 
Mt. Banahao (Luzon Island). 

Factors Limiting Recording of Advertisement 

Calls 
The advertisement calls reported here are the 
result of the recent aquisition of suitable record- 
ing equipment and the availability of bases at the 
University of the Philippines, Los Banos, Luzon 
Island and Silliman University, Negros Island. 
These bases made possible the numerous field 
trips necessary (1994-96) to secure suitable re- 
cordings of the voices of the species of Platyman- 
tis in those areas. We can continue this study in 
these localities when environmental conditions 
are suitable. The possibility of obtaining voice- 
recordings for the populations of P. guentheri 
and P. rabori on Bohol or northeastern Mindanao 
Islands are, however, subject to the same limita- 
tions as attempts to increase the size of the sam- 
ples from those populations (see Introduction). 

Voice Records of Platymantis on Mt. 
Maquiling and Mt. Banahao 

Recent field work on Mt. Maquiling and Mt. 
Banahao has not only provided voice records 
reported here for P. luzonensis and P. banahao, 
but also records of the advertisement calls of 
other species of Platymantis. During some field 
trips, calls of three species {P. dorsalis, P. mimu- 
lus, and P. luzonensis) were heard and recorded 
on Mt. Maquiling and six species (P. dorsalis, P. 
mimulus, P. banahao, P. luzonensis, P. mon- 
tanus, and P. undescribed [dorsalis Group]) on 
Mt. Banahao. Frequently all six species were 
calling at the same time. 

The advertisement calls reported here for P. 
luzonensis and P. banahao are very different (to 
the human ear), as were the calls for P. dorsalis 
and P. mimulus reported earlier (Brown et al. 
1997). The analysis of their voice records con- 
tributed to more complete diagnoses of P. ne- 
grosensis, P. luzonensis, and P. banahao. We 
hope that similar voice records can be obtained 
for P. guentheri and P. rabori when conditions 



permit. The voice of P. luzonensis sounds like 
"kwenk-kwenk-kwenk". The notes are repeated 
three to five or six times in one or two seconds, 
or the notes may be more isolated and widely 
spaced. The voice of P. banahao sounds like 
"tuk-tuk-tuk-tuk" repeated six to 20 or more 
times at a rate of four to five times a second. It 
can be described as the sound of a machine gun. 
The voice of P. negrosensis is more like that of 
P. luzonensis but differs in dominant frequency 
and other characteristics (see Fig. 1A-C and 
Table 4). Advertisement calls such as those re- 
corded for P. luzonensis and P. banahao are 
generally accepted as probably the primary 
means of species identification (see Blackwell 
and Passmore 1991). 

Voice Records of Other Species of Platymantis 
The only other voice record for a Philippine 
population of Platymantis is that of/ 3 , dorsalis 
from Cuernos de Negros, Negros Island (Kura- 
moto 1 997). Our voice record for this population 
of P. dorsalis differs in the lower kilohertz, closer 
to 3,000 than 4,000, and time intervals separating 
calls (or notes?). Our records have not yet been 
published. 

Records of voices for several non-Philippine 
species were reported much earlier: that of P. 
papuensis from New Guinea (Zweifel 1969) and 
those of P. schmidti, P. solomonis, P. magnus, 
and P. neckeri from the Bismarcks and Solomons 
(Menzies 1982). These reports provide varying 
amounts of information in terms of sonograms, 
their analyses, and inteipretations. Certainly es- 
tablishment of uniform terminology as well as 
reanalysis of these published sonograms which 
would permit direct comparisons of both Philip- 
pine and non-Philippine species are desirable 
projects, but are not the purpose of this study. 

Discussion 

The five species of the P. guentheri Group 
recognized at this time are based on large sam- 
ples (over 100 specimens each for P. guentheri, 
P. rabori, and P. negrosensis) and smaller sam- 
ples of 12-20 specimens for P. banahao and P. 
luzonensis. Differences in their advertisement 
calls were important clues to our diagnosing the 
latter two species. Other populations from iso- 
lated mountains such as Mt. Isarog on Luzon, or 



BROWN, A. ALCALA, DIESMOS AND E. ALCALA: PLATYMANTIS 



19 



islands such as Cataduanes, may be recognized 
as additional species of the P. guentheri Group 
when larger samples and audiosterograms of 
their advertisement calls become available. 



Platymantis luzonensis. — - See list of holo- 
type and paratypes for that species. 

Platymantis banahao. — See list of holotype 
and paratypes for that species. 



Acknowledgments 

Early field work by E. H. Taylor, A. C. Alcala, 
and W. C. Brown provided most of the collec- 
tions and much of the data on the ecology and 
zoogeography of this species-Group. Recent 
field work by A. C. Alcala, A. C. Diesmos, A. 
Ross, E. L. Alcala. and numerous people who 
assisted them have provided data on populations 
occurring on southern Luzon and associated 
small islands. We are indebted to R. C. Drewes 
(CAS), J. E. Cadle (MCZ), R. F. Inger (FMNH), 
P. C. Gonzales (PNM), and E. N. Arnold and 
B. T. Clarke (BMNH) for permission to examine 
material in their collections. We also wish to 
thank A. E. Leviton, R. C. Drewes, and R. F. 
Inger for their critiques during the prepara- 
tion of this paper. Drawings were prepared 
by C. Sudekum, California Academy of Sci- 
ences. Photographs were provided by A. C. Al- 
cala and E. L. Alcala. 

Appendix A 
Specimens Examined 

Platymantis guentheri. — Dinagat I., Philip- 
pines: BMNH 1947.2.31.34 (holotype), Esper- 
anza: DMNH 151-153, Mt. Magkono: DMNH 
135; Mindanao I., Davao Prov.: FMNH 50571, 
50573; Zamboanga Prov.. Dapitan Peak: CAS- 
SU 19987, 20121, 20125; Agusan del Norte 
Prov.: CM 3424-29, Mt. Hilong-hilong: CAS 
133148-49, 133287, 133307, 133313, 133332- 
33, 133531, 133540. 133548, 133651-53, 
133780-81, 186066-67, 186124-26, 196378-79; 
Biliran I.: FMNH 318381-83; Bohol I., Sierra 
Bullones area (within radius of 1 5 km): CAS-SU 
21214 (holotype of Cornufer ingeri), CAS-SU 
21 196-210, 21212. 21216-17, 21219, 21709-16, 
21719-20, 21724-32, 21734, 21736, 21740, 
21745, 21747, 21749-55, 21757, MCZ 38097- 
98, FMNH 134985-86 (paratypes of Cornufer 
ingeri). 

Platymantis rabori. — See list of holotype 
and paratypes for that species. 

Platymantis negrosensis. — See list of holo- 
type and paratypes for that species. 



Literature Cited 

Alcala, A. C. 1963. Breeding behavior and early 
development of frogs of Negros, Philippine Islands. 
Copeia 1963:679-726. 



. 1986. Guide to Philippine flora and fauna: 

Amphibians and Reptiles. Univ. Philippines 
10:1-195. 

Alcala, A. C. and W. C. Brown. 1957. Discovery 
of the frog Cornufer guentheri on Negros Island. 
Philippines, with observation on its life history. 
Herpetologica 13:182-184. 

Boulenger, G. A. 1882. Catalogue of the Batrachia, 
Salientia, and Ecaudata in the Collection of the 
British Museum. Taylor and Francis, ed. 2: 1-503. 

Blackwell, P. R. Y. and N. T. Passmore. 1991. 
Advertisement calls and female phonotaxis in the 
Natal dwarf Afri.xalus (Anura: Hyperolidae). J. Afr. 
Zool. 105:275-280. 

Brown, W. C. and A. C. Alcala. 1961. Populations 
of amphibians and reptiles in the submontane and 
montane forests of Cuemos de Negros, Philippine 
Islands. Ecology 42:628-636. 

— . 1963. A new frog of the genus Cornufer 
(Ranidae) with notes on other amphibians known 
from Bohol Island, Philippines. Copeia 
1963:672-675. 

. 1 970a. A new species of the genus Platyman- 
tis (Ranidae) with a list of amphibians known from 
South Gigante Island, Philippines. Occas. Pap. 
Calif. Acad. Sci. 84:1-8. 

— . 1 970b. The zoogeography of the herpeto- 



fauna of the Philippine Islands, a fringing archipel- 
ago. Proc. Calif. Acad. Sci. 38(6): 105-1 30. 

. 1983. Modes of reproduction of Philippines 



anurans. Pp. 4 1 5^428 in Advances in Herpetology 
and Evolutionary Biology, A. G. J. Rhodin and K. 
Mujata, eds. Harvard Univ. Press, Cambridge. 

Brown, W. C, A. C. Alcala., and A. C. Diesmos. 
1997. A new species of Platymantis (Amphibia: 



20 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. I 



Ranidae) from Luzon Island, Philippines. Proe. 
Biol. Soc. Washington 110:18-23. 

Brown, W. C, R. M. Brown, and A. C. Alcala. 
1997. Speeies of the hazelae group of Platymantis 
(Amphibia: Ranidae) from the Philippines, with de- 
scriptions o\' two new speeies. Proe. Calif. Aead. 
Sci.49(ll):405-421. 

GAULKE, M. 1994. Notes on the herpetofauna of 
Panaon and Samar, east Visayans, Philippines. 
Hamadryad 19:1-10. 

Hi aney, L. and E. A. Rickart. 1990. Correlations of 
elades and elines: geographic, elevational, and phy- 
logenetic distribution patterns among Philippine 
mammals. Pp. 321-332 in Vertebrates in the Trop- 
ics. G. Peters and R. Hutterer, eds. Zoologisches 
Forschungsinstitut und Museum Alexander 
Koenig, Bonn. 424 pp. 

[NGER, R. F. 1954. Systematics and zoogeography of 
Philippine amphibia. Fieldiana: Zool. 33:181-531. 

KURAMOTO, M. 1997. Relationships of the Palau frog, 
Platymantis pelewensis (Anura: Ranidae): morpho- 
logical, karyological, and acoustic evidences. 
Copeia 1 997(1): 183-1 87. 

Mi n/iis. J. 1. 1982. The voices of some male Platy- 
mantis species of the New Guinea region. Brit. J. 
Herpetol. 6:241-245. 



mals along an elevational transect in southeastern 
Luzon, Philippines. .1. Mamm. 72(3):458-469. 

Ross, C. A. and P. C. Gonzales. 1992. Amphibians 
and reptiles of Catanduanes Island, Philippines. 
Zool. Pap. Nat. Mus., Manila 2(2):50-76. 

Ross, C. A. and J. D. Lazell. 1991. Amphibians and 
reptiles of Dinagat and Siargao Islands, Philippines. 
Philip. J. Sci. 119(3):257-280. 

Stejneger, L. 1905. Three new frogs and one new 
gecko from the Philippine Islands. Proe. U. S. Nat. 
Mus. 28:343-348. 

TAYLOR, E. H. 1920. Philippine Amphibia. Philip. J. 
Sci. 16:213-359. 

— . 1922a. Additions to the herpetologieal fauna 
o\' the Philippine Islands, I. Philip. J. Sci. 
21:161-206. 

— . 1922b. Additions to the herpetologieal fauna 
o\~ the Philippine Islands, II. Philip. J. Sci. 
21:257-303. 



. 1922c. Herpetologieal fauna oi' Mount Makil- 

ing. Philip. Agriculturist 1 1:127-140. 

— . 1923. Additions to the herpetologieal fauna 
of the Philippine Islands, 111. Philip. J. Sci. 

22:515-557 



RICKART, E. A.. L. R. HEANEY, AND R. C. B. LJTZUR- 
RUM. 1991. Distribution and ecology of small mam- 



ZWEIFEL, R. G. 1969. Frogs of the genus Platymantis 
(Ranidae) in New Guinea, with description of a new 
species. Am. Mus. Nov. 2374:1-19. 



CALIFORNIA ACADEMY OF SCIENCES. 1997 
Golden Gate Park 
San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 2, pp. 21-38, 3 figs., 2 tables. October 8, 1997 



BIOGEOGRAPHY OF AMPHIBIANS IN THE ISLANDS OF THE 

SOUTHWEST PACIFIC 

/ -v W C ln 

Walter C. Brown U ' 2 ]$Qy 

Department ofHerpetology, California Academy of Sciences, 



San Francisco, CA 941 18 






Present patterns of distribution and endemism of frogs in the islands of the southwest Pacific are 
examined. These patterns are analyzed in terms of the generally accepted geological history and 
sea-level changes for the region. 

The evidence indicates that the island anuran fauna includes three components. One is an Asian 
component. Representation of this component on the Sunda Shelf islands includes almost all the 
mainland genera and four on Borneo that are not known on the mainland. There is a great 
diversity and high endemism for species. The Philippine fauna includes less than half of the genera 
of the Sunda Shelf islands and a greatly reduced number of species. The islands of western 
Wallacea have about one-fourth of the Sundaland genera and about one-seventh as many species. 
Only two of the ranid genera (Rana and Limnonectes) of the Asiatic component occur in the islands 
east of western Wallacea; and there are only two or three species on any of these islands except 
for New Guinea. There the genus Rana has several endemic species. 

The second component is one derived from the Australian anuran fauna. In New Guinea (the 
primary Sahul Shelf island), two of the three Australian genera of Hylidae (Pelodryadidae of 
some) occur, with a large number of species, most of them endemic. One of these (Litoria) dispersed 
to the Melanesian Arcs and the islands of eastern Wallacea, the second (Nyctimystes) only to the 
latter. Only 5 of 20 genera of the Myobatrachidae occur in New Guinea, and they have not 
dispersed beyond the Sahul Shelf. 

The third component includes 16 genera in 2 endemic subfamilies of microhylids (centered in 
New Guinea) and 4 genera in an endemic subfamily of ranids. Three of these genera are centered 
in the Melanesian arcs, and one in Melanesia and the Philippines. Relationships of these 
subfamilies to other microhylid and ranid lineages are not clear at this time. Also one genus 
(Batracliylodes) in the subfamily Raninae is known only from the Solomons. It is not clearly related 
to any of the Asiatic ranid genera. 

Received February 28, 1995. Accepted May 19, 1997. 



The numerous islands of the southwestern and forms of life on these islands have been the basis 

central regions of the Pacific Ocean, extending for numerous biogeographic hypotheses, 
from the coast of southeast Asia and the northern From the time of Wallace's "Island Life" 

coast of Australia to the Hawaiian, Line, and (1880) to Darlington's "Zoogeography: The 

Tuamota islands in the Central Pacific (Fig. 1), Geographical Distribution of Animals" (1957), 

provide isolated land areas of various ages. The zoogeographers explained the presence of ani- 

[21] 



22 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 



mals on islands in terms of the animals' use of 
known or imagined previous land bridges, their 
abilities to cross water barriers, or their possible 
transport by man. These explanations were based 
on the geological concept that the earth's crust 
was a surface covering of ocean and land areas 
occupying relatively fixed positions. Since the 
1960s, the geological concept of continental drift 
has provided for very different explanations of 
island biogeography. 

Recently, there has been renewed interest and 
a number of papers on the geological history and 
biogeography of the islands of the southwest 
Pacific, with emphasis on those islands between 
the Sunda and Sahul Shelves, the region often 
referred to as Wallacea (Fig. 2). Many of these 
papers were published in 3 volumes: "Wallace's 
Line and Plate Tectonics," 1981, "Biogeography 
of the Tropical Pacific," 1984, and "Biogeo- 
graphical Evolution of the Malay Archipelago," 
1987, Oxford Monogr. Biogeography, Claren- 
don Press, Oxford. As to organisms: angio- 
sperms receive the most attention, with emphasis 
on palms in the plant kingdom; birds, mammals, 
and Lepidoptera receive most attention in the 
animal kingdom. Amphibians and other terres- 
trial, vertebrate fauna are dealt with in the paper 
by Cranbrook (1981). Shore-fish distribution is 
discussed in a separate paper (Springer 1982). 
Numerous other papers, limited to the geology of 
the region, have also been published in various 
geological journals or special publications dur- 
ing this same period. Other islands of the region 
including the Greater Sundas, Philippines, New 
Guinea and its satellites, Bismarcks, Solomons, 
and Fiji, are considered in many of these papers. 
Several papers have summarized the biogeogra- 
phy of the heipetofaunas on some major groups 
of islands (Allison 1996; Brown and Alcala 
1970; Inger 1954, 1966). The present paper is 
limited to the amphibian fauna of these islands. 

Methods 

In this study, distributional data for anuran 
species in the southwest Pacific islands is primar- 
ily based on island localities cited in Frost 
(1985), Duellman (1993), Allison (1996), 
Menzies ( 1982), Zweifel and Tyler ( 1 982), Inger 
(pers. comm.), and Zweifel (pers. comm.). Deter- 
mining accuracy of species' assignments in these 
sources, or by preceding authors, was not a part 



of this study. These data are converted to dis- 
tributional patterns of genera and species as- 
signed to families or subfamilies native to 
southeast Asia and Australia, or subfamilies in 
Microhylidae or Ranidae that are centered in the 
islands. 

These distributional patterns are evaluated in 
relation to: (1) the geological events that pro- 
duced the islands and established their current 
spatial relationships, (2) past sea-level changes, 
(3) climatological history, (4) their dispersal 
abilities, routes, and opportunities, and (5) evo- 
lution and extinction events. Biogeographical 
hypotheses concerning these island anurans are 
reevaluated. 

Geological History of the Islands 

The islands in the area under consideration can 
be assigned to one of five groups: ( 1 ) those on 
the shallow Sunda Shelf off the Asian coast, (2) 
those on the Sahul Shelf off northern Australia, 
(3) the clusters of islands between these two 
shelves, usually termed Wallacea (Fig. 2), (4) the 
Philippines, and (5) the Melanesian arcs: Admi- 
ralties, Bismarcks, Solomons, and Fiji. 

The geological history of this region is com- 
plex, and many uncertainties still exist. But re- 
cent studies have changed interpretations of the 
origins and history of southeast Asia and the 
islands as well as their probable ages. Geological 
processes are measured in terms of megayears; 
some of the important sea-level changes and 
climatological events in terms of thousands or 
hundreds of thousands of years. 

All of southeast continental Asia from the Hi- 
malayan-Tibet region and southern China, as 
well as the Greater Sunda Islands, is believed to 
be the result of accretion of terranes over millions 
of years. The origins of these terranes and the 
times of their collisions are fairly well estab- 
lished for some but poorly understood for others, 
especially the older ones. 

At least five such terranes have been identified 
in the Himalayan-Tibet region to the north of 
India. The origin of the oldest (most northerly) is 
thought not to be Gondwanan (Nishiwaki and 
Uyeda 1987), but at least the more recent of these 
terranes (south China and part of Indochina-Ma- 
lay-Sundaland) are rift-blocks from eastern 
Gondwana along the north rim of the Australian 
region (Murphy 1987; Nishiwaki and Uyeda 



BROWN: BIOGEOGRAPHY OF AMPHIBIANS 



23 



1987; Audley-Charles 1987, 1988). This rift pe- 
riod may have extended from the late Carbonif- 
erous to the mid-Jurassic (150-300 MaBP) and 
was followed by long periods of northward drift- 
ing across the changing Tethys Ocean. The ear- 
liest terranes may have accreted to southern Asia 
before the end of the Jurassic (Audley-Charles 
1987). Data indicate that India rifted from Gond- 
wana about 140 MaBP and collided with Asia 
about 50+ MaBP (Butler 1995; Hallam 1981). 

Greater Sunda Islands. — The Greater Sunda 
Islands include Sumatra. Java, Borneo, Palawan 
(western Philippines), and several (isolated or 
clustered) small islands, all situated on the Sunda 
Shelf off the coast of southeast Asia (Fig. 2). At 
times, these have been united as a subaerial ex- 
tension of the Indochina-Malay region, most re- 
cently about 20.000 years BP, a result of lower 
sea levels during the last ice age. The origin of 
Sundaland has been open to several interpreta- 
tions. Recent geological research indicates that 
it, like the Malay region, is the result of bringing 
together fragments of eastern Gondwana, Pacific 
arcs, and Seamounts. These are the most recent 
terranes to be added to the southeast margin of 
Asia, beginning about 50 MaBP (Audley-Char- 
les 1987). However, the main geological proc- 
esses which resulted in the present shape of 
Sundaland were completed about 15 MaBP (Oi- 
lier 1985). 

Palawan, at the north end of the Sunda Shelf, 
is a composite island. The southern part is asso- 
ciated with Borneo and its development. The 
northern part and the Calamian Group are from 
a segment of a continental terrene that rifted from 
China, probably in the Cretaceous, and collided 
with the western edge of the Philippine plate and 
south Palawan in the mid- to late Miocene 
(McCabe and Cole 1987). 

Sahul Shelf islands. — These islands, with the 
exception of the northern half of New Guinea, are 
part of the Australian continent that rifted from 
Antarctica and drifted northward between 45 
MaBP and 15 MaBP. At that time the collision 
with the Pacific, Asian, and minor border plates 
occurred. The northern part of New Guinea and 
small satellites along the north coast and Vo- 
gelkop Peninsula are the result of the accretion 
of Tethys Ocean marginal arcs onto the New 
Guinea portion of the Gondwanan Australia- 
New Guinea Plate about mid-Miocene. There is 
evidence that the central region of New Guinea, 



along the central mountain range, became 
subaerial in early Miocene (Hamilton 1979; 
Audley-Charles 1981). These islands, like those 
of the Sunda Shelf, have been connected to Aus- 
tralia by land bridges during periods of lowered 
sea levels. 

Wallacea. — The islands of Wallacea, be- 
tween the shelves (Fig. 2), are partly Asian and 
partly Australian, and vary in age and type. Their 
present positions are the result of compression 
between the converging plates over the past 40 
million years. The Asiatic (western) part includes 
two groups. In the south are the Lesser Sunda 
Islands, a volcanic arc extending from Bali 
through Flores and continuing as the Inner Banda 
Arc through Wetar Island. This arc dates from at 
least 20 MaBP (Audley-Charles 1987). In the 
north is Sulawesi, the western part of which is 
Asian in origin and was joined to Borneo until 
the Eocene (Audley-Charles 1981, 1987; 
McCabe and Cole 1987). The northern and east- 
ern parts, derived from the margin of the advanc- 
ing Australia-New Guinea Plate, probably began 
to emerge as an island area about 5-10 MaBP at 
about the same time as it joined with western 
Sulawesi (McCabe and Cole 1987). 

The remaining islands of Wallacea on the 
south and east originated from the advancing 
front of the Australia-New Guinea Plate. Those 
of the Outer Banda Arc from Sumba through 
Timor, Tanimbar, Kaie, and Ceram and Buru in 
the southern Moluccas (Fig. 2), are not volcanic, 
but the result of recent, probably Pliocene ( 1 .5-5 
MaBP), upthrust along the margin of the Austra- 
lian Plate (Milsom and Audley-Charles 1986; 
McCabe and Cole 1987; Nunn 1994). The Bang- 
gai-Sula island group, according to one interpre- 
tation, became subaerial by late Miocene. 3 
MaBP (Audley-Charles 1981, 1987). The north- 
ern Moluccan Islands, centered about the large 
island of Halmahera, are volcanic. Some of the 
volcanoes on Halmahera are Miocene, but most 
of the small islands are young, Quaternary vol- 
canoes (Audley-Charles 1987; McCabe and Cole 
1987). Areas of Wallacean islands are not cur- 
rently as great as during Pleistocene periods of 
lower sea levels. At such times, some of them 
may have been united but not joined to the ex- 
posed areas of the Sunda or Sahul shelves. 

Philippine Islands. — The Philippine Archi- 
pelago is a region with a very complex geological 
history. The main islands of Luzon and Min- 



24 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 




BROWN: BIOGEOGRAPHY OF AMPHIBIANS 



25 




_n — £ 

— o « 

!»o.£ 

— r- u 

^ J § 

I E S. 

•a 2 Si 



5 



u 



<u 3 ca 
_c — u 

TO — — - 

3 •> 

-. D £ 

■§ jj-2 

o g S 

— w "3 

£ § i 

X 5 u 
a) o -2 

3J "Z § 

£c£g 

^ c ° 

.2 £ "3 



3 ul <U 
(3 <U C 

g a.s 

.5 8 in 

n S< 
O _c u 

£ <U <D 

U ,3 -3 

Z -a £ 

° § 8 





as 


<D tn 




<u 


-3 a> 






O ? 






J= c 




o 


U 1) 














o 


2-S 




C 


"2. £ 


z 


.2 




< 


<2 c 
= 2 


o 


u 


(U O 


O 


.3 


\3 <D 






3 _S 


Z 


o 


O — 
O en 


M 




3 b 


Q 

Z 




<u 2 

.a e 


IH 




"3 'c3 




Ph 


ca | 






(U 




<u 


■£ - 05 




£ 


J= <u 




j= 


.3 3 




3 


£- 




O 


"3^ 




<u 


3 F- 




-5 


'O rj 




14-1 









"2 ° 

5 




•a 


* L/~l 




3 


T3 1 




3 


<u | 








=' ' T ~'"~ ■ 


™ 


a 

3..S 




ri 


X — 






u "3 




s 


.3 " 




C 


2-§ 




£ 





26 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 



danao have large pre-Miocene areas (McCabe 
and Cole 1987;Heaney 1991). There is evidence 
to support the view that some of the volcanoes of 
these islands were subaerial as long as 70-140 
MaBP in the Cretaceous (Audley-Charles 1981; 
McCabe and Cole 1987). Other islands are gen- 
erally viewed as more recent arcs: Mindoro 8-10 
Ma, Negros-Panay-Cebu 1-4 Ma, and some of 
the smaller islands such as Camiguin and 
Sibuyan 0.1-1.0 Ma (Heaney 1991). One view 
also holds that the Philippine Archipelago 
achieved its present position by a northward drift 
from the Eocene to present (Jarrand and Sasajima 
1980; Oilier 1985; McCabe and Cole 1987). 
Although there is general agreement that most of 
the Philippines are oceanic in origin, the North 
Palawan continental terrane was a part of south 
China at least into the Cretaceous. Following the 
rift from south China, the North Palawan block 
began a southward drift in the Oligocene, collid- 
ing with and contributing to the formation of 
northern Palawan, western Mindoro, and western 
Panay. Other areas of Mindoro and Panay are 
derived from the Mindoro-Panay disrupted ter- 
rane and the Central Philippine Arc terrane 
(McCabe and Cole 1987; McCabe et al. 1985). 

There is also evidence that sea-level changes 
caused temporary land connections with north- 
eastern Borneo at various times. The lowering of 
sea level during the late Pleistocene glaciation, 
18,000-20,000 years ago would have created the 
Greater Islands shown in Figure 3 and joined 
Palawan to Borneo, but would not have closed 
the water barriers between Borneo and Greater 
Sulu, Greater Sulu and Greater Mindanao, or 
Greater Mindanao and Greater Luzon. It has been 
suggested (Morley and Flenley 1987) that in 
mid-Pleistocene (perhaps 100,000 years ago or 
more) even lower sea levels may have closed 
these gaps and also those between Greater Luzon 
and Greater Negros as well as Greater Luzon and 
Greater Mindoro (Figs. 2 and 3). These island 
connections would have required a sea-level 
lowering of 300 m or more, unless the sea floor 
was higher at that time. 

Melanesian Arcs. — The island arcs which 
contributed to the formation of northern New 
Guinea at the collision of the Australia-New 
Guinea and Pacific Plates, as well as the more 
eastern Admiralty, Bismarck, Solomon, Fiji 
Arcs, and the Palau Arc are also outside of Wal- 
lacea. Based on recent geological evidence, one 



hypothesis concerning the history of the Melane- 
sian Arcs is that they had their origin from a 
double-arc system (Tethys Arcs) on separate sub- 
plates that began to break off from northeastern 
and eastern Gondwanan- Australia between 
100-60 MaBP, at about the same time that New 
Zealand to the south was rifted from the north- 
eastern Antarctica segment. These arc-blocks be- 
came widely separated from Australia by sea 
floor spreading while drifting north and north- 
westward during the subsequent 30^0 million 
years (Hamilton 1979; Coleman 1980;Halloway 
1984). 

Following compression between the Pacific 
and Australia-New Guinea Plates, 15-20 MaBP, 
the central portion of these arcs gradually fused 
to the advance edge (southern New Guinea) of 
the Australian Plate. Part of the central mountain 
region is derived from islands of the Inner Arc 
and the Vogelkop and northcoast mountain 
ranges from the Outer Arc (Hamilton 1979; 
Audley-Charles 1981; Halloway 1984). Other 
parts of the Outer Arc persist as an alignment of 
archipelagos to the east, Bismarcks, Solomons, 
Vanuatu, and Fiji (Coleman 1980; Halloway 
1984). The Vanuatu Arc underwent a southward 
rotation beginning 6-8 MaBP that brought it to 
its present position just east of New Caledonia 
(Coleman 1980; Kroenke 1984). 

Just as lower sea levels caused by Pleistocene 
glaciation events resulted in land connections 
between Borneo and Palawan and possibly be- 
tween Borneo and Greater Sulu at times in the 
recent past, they created similar land connections 
between Australia and New Guinea. Although 
the Bismarcks and Solomons were not connected 
to New Guinea during these periods, the water 
channels separating them were narrowed. Also 
within the Solomons and Bismarck archipelagos, 
the Pleistocene lower sea-level periods resulted 
in the uniting of existing islands into much larger 
islands for varying periods of time (see Diamond 
andMayr 1976). 

Anuran Families in the Islands 

Components I, II and III 

The anuran fauna of the southwest Pacific Is- 
lands can be assigned to one of three compo- 
nents: (1) Component I, those resulting from 
colonization by present-day Asian stocks, (2) 
Component II, those resulting from colonization 



BROWN: BIOGEOGRAPHY OF AMPHIBIANS 



27 



Batan Islands • 



#122° 



KEY 



Babuan Islands 



Greater Luzon 
Pleistocene Island 



1 . Lubang 

2. Burias 

3. Tablas-Romblon 

4. Sibuyan 

5. Camiguin 



ss than 120 meters 
rent water depth 

200 



Mindoro Island 



/ <& . 

Greater Palawan / o \ j 

Pleistocene Island^. , 4 ; <_ 



•■A 




18°- 



Samar Island 



y-V^l "'!tt\ 



Mlfe 



i>'^ 



10° 



si 



X^/ Great* 



£*v \ 3**r Leyte 



- 6 ° 1a 



Borneo 



Negros-Panay 
Pleistocene Island 







«^ tl_ • v 7^' Greater 

(^—5^, ^^ Mindanao 

<# Pleistocene Island 

IP 
— Greater Sulu 

Pleistocene Island 

122° 



Figure 3. Philippines: present islands (pale stippled areas) and late Pleistocene, about 20,000 years BP (dark stippled areas) 
based on presumed lower sea levels of about 1 20 m (after Brown and Alcala 1 994). 



28 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 



by present-day Australian stocks, and (3) Com- 
ponent III, those genera in distinct subfamilies 
that are centered in the islands. Adler, Austin and 
Dudley ( 1995) divided the island skinks that they 
were studying into comparable categories for 
their analysis of distribution patterns. 

The first two components are easy to identify 
based on the genus and family (sometimes sub- 
family) to which the species are assigned. Six of 
the eight families represented in the island fauna 
are Asian (Component I) and nearly all of the 
genera are mainland genera (Table 1, Appendix 
A). The other two families are Australian (Com- 
ponent II), and the genera are mainland genera 
(Table 1, Appendix A). Component III includes 
two distinct subfamilies (Asterophryinae and 
Genyophryninae) of the family Microhylidae, 
with most of the genera restricted to New Guinea, 
and a distinct subfamily (Platymantinae) of the 
fami ly Ranidae with most of the genera restricted 
to the Solomons and Bismarcks but with one 
genus (Plat\'mantis) also centered in the Philip- 
pines (Table 2, Appendix C). 

History of the Data Base 

Data on the species of anurans in the islands of 
the southwest Pacific have been acquired over 
the past century and a half. Only a few species 
had been recorded prior to publication of "Island 
Life' 1 (Wallace 1880), most notably the several 
species described from New Guinea (Peters and 
Doria 1878) and several from the Philippines 
(Boulenger 1882). 

During the last couple of decades of the I9th 
century and the first couple of the 20th, collecting 
in the Solomons, New Guinea, and the Philip- 
pines greatly increased. Of particular interest was 
the discovery of the genera that we now classify 
as the platymantine ranids in the Solomons 
(Boulenger 1886). One of the genera (Platyman- 
tis) was also known from the Philippines. Two 
monographs (Kampen van 1923 and Taylor 
1920) summarized the data on the frogs of the 
Indo-Australian Archipelago and the Philippines 
respectively. 

A renewed interest in the herpetofauna of these 
islands began during World War II, and contin- 
ues to intensify even today. The resulting litera- 
ture includes numerous short papers describing 
species. Major publications during this period 
that summarize data on systematics and distribu- 
tion of anurans worldwide include: Frost 1985; 



Ford and Cantella 1993; and Duellman 1993. 
Several other major papers are limited to the 
southwest Pacific Islands (Brown 1953; Inger 
1 954, 1966; Brown and Tyler 1968; Zweifel and 
Tyler 1982; Allison 1996). 

Results 

Because this paper is concerned with the bio- 
geography of the anurans of the southwest Pa- 
cific Islands, but not those of mainland Asia and 
Australia, only the number of genera and species 
in Australia and southeast Asia are indicated in 
the tables. For the island areas, not only the total 
number but the number that are endemic is also 
given. The number of genera and species of the 
Asian and Australian anurans (Components I and 
II) and their distribution patterns are summarized 
in Table l and are listed by name in Appendix B. 

Component I. — For the families Bufonidae, 
Megophryidae. and Rhacophoridae and the 
Asian lineages of the Microhylidae and Ranidae, 
27 of 34 (82%) of the genera occur in the Greater 
Sundas (Appendix B). This, along with 67 of 
southeast Asian species on the larger Sunda Is- 
lands, suggests good dispersal opportunities in 
the relatively recent past. 

This is consistent with the evidence that the 
Sunda Shelf islands have been connected to each 
other and to mainland southeast Asia at various 
times in the Pleistocene as a result of lower sea 
levels, most recently about 18,000-20,000 years 
ago. The presence of one endemic genus for both 
the Megophryidae and the Ranidae in Borneo 
and another Staurois in both Borneo and the 
Philippines may be the result of an extinction of 
those generic lineages on the mainland, their 
evolution in situ, or a sampling weakness for 
some areas of southeast Asia. I treat Barbourula 
in the Bombinatoridae (Ford and Cantella 1993) 
known from one species in Borneo and one in 
Palawan, as a relict genus of that Asian family 
represented by the genus Bombina in China but 
not in southeast Asia. The Eurasian lineage of the 
family Hylidae, with one species of the genus 
Hyla recorded from Indochina and Thailand, is 
not known from the Pacific Islands. 

Fifteen (48%) of the Bornean genera, exclu- 
sive of the relict Barbourula, are known from the 
Philippines. Eighteen species of these genera are 
shared with Borneo and 30 species are endemic. 
Six of the seven families in southeastern Asia and 



BROWN: BIOGEOGRAPHY OF AMPHIBIANS 



29 



=2 ^ 

b * 

ra O 



^ GO 

^5 00 

O .2 

oo o 

.5 o- 

00 

CD ~D 

C C 

3 3 



o 

cs 



co 

c « 

CO <y 

■3 c 

CO Cl> 

e db 

< 2"S 
co S "o 

S 01 o 

co -= C 

< H „ 



"> "« co 
.52 fc -3 



>> 



00 ■ — 

3 ^ 



ca 



00 






.5 -n -C 



-a E ^c 

« to g 
►— a> ° 

CO £ CO 

c •£ "" 

a> 3 -a 

C o S 

O to CO 

0- - 1= 
C - — >.S 

§ .2 S 

U >» V 

1- w c 

,2n o 



a. E 



_c 


> 


£ 




00 




c 


D 


3 
-J 


ca 

Ph 


— 


CO 


c 


w 


T3 


~ 






■j 


< 


Xi 


S- 


5 


B 



U £ 



>> 

I 



o 

JS 
D. 
O 



oi 



CQ 



b 



^■vO o 


O /— 
— CN 


c-> CN 


m 

co 


cn ^0" m c~- — "3" 
O co O oo O on 


OOO h (N 
CN — 

a oo a on 


"* zz ^ ** 
a oo O oo 


On r~- 

o 

a oo 



cn "3" — — ^r o 
cn t-~ 

O oo O oo O oo 



IT) t-~- 

O oo 



a oo 



CN — t~~ 

— iri CN fN CNO — CN 

OO 

O oo O oo O oo 



O oo 



a oo 



NO — v~t rn -rf v~t CN ^ — — " — — 

r*l "0" — 

OooOooOoo OooOoo Ooo 



NO CN — — OO, 

^dTj-oo rs ^ (N n r-iO „_ cn — — — — 
OooOooOoo OooOoo Ooo Ooo Ooo OooOooOoo 



OooOooOoo OooOoo 



so o no no m 

CN 



OooOooOoo OooOoo 



_ — CN 



O oo O oo O oo 



< 5S 



^ 8 

E £ 

00 3 

^ oo 



^ CQ 

c a3 B 

a a < 

00 (~\ 

LQ 







TJ 


00 










a 


T3 










a 


C 


- 1 




oo 

ca 






3 


< 




u 

3 


is 
"3 


3 

'3 


O 


C 
3 




G 

•> 


oo 


O 


~ 


00 

U 

3 


3 

E 


z 


3 

■§ 

OO 


Z 


3 
00 


_3 
'J 


CQ 



30 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 



southern China are represented. This data is con- 
sistent with the concept that very narrow water 
channels or direct land connections may have 
existed between northwestern Borneo and 
Palawan and northeastern Borneo and Mindanao 
as recently as the lower sea-level event 
(18,000-20,000 years ago), and during earlier 
periods when sea levels were even lower (Fig. 3), 
mid-Pleistocene (100,000-500,000 years ago) 
and at other times back to the late Tertiary (Mor- 
ley and Flenley 1987). The effect of sea-level 
changes on the distribution patterns of rhaco- 
phorid frogs within the Philippine Archipelago is 
discussed by Brown and Alcala (1994). There is 
no direct evidence as to how the composition of 
the current faunas have been affected by extinc- 
tion or dispersal from the Philippines to Borneo. 

Western Wallacea. — The sharp reduction in 
the number of Asiatic genera and species in Su- 
lawesi and the Lesser Sundas (Table 1 , Appendix 
B) indicates much more limited dispersal oppor- 
tunities between eastern Sundaland and the is- 
lands of western Wallacea than existed between 
Sundaland and the mainland or the Philippines 
and Borneo. Eight (30%) of the Asia-Sundaland 
genera occur in Sulawesi and seven (26%) in the 
lesser Sundas. But at the species level, the 
number recorded from Sulawesi is almost twice 
that of the Lesser Sundas and endemism is five 
times greater. This is consistent with the lack of 
evidence of land connections between Borneo 
and Sulawesi or between Java and the Lesser 
Sundas at low sea-level times during the Pleisto- 
cene. The narrow, but deep Makassar Strait sepa- 
rates Sulawesi from Borneo and the Lesser 
Sundas from Java. Also, Sulawesi is geologically 
much more complex than the Lesser Sundas, 
with western Sulawesi older than the eastern part 
or the Lesser Sundas (Audley-Charles 1987; 
McCabe and Cole 1987). 

Only three ( 1 0%) of the genera of the Greater 
Sundas (Table 1 , Appendix B) are recorded from 
eastern Wallacea. Of the five species of ranids, 
one species Limnonectes verruculosa is treated 
as endemic, and L. modesta is otherwise known 
only from Sulawesi. These should be validated. 
Two species are conspecific or closely related to 
a group of species of Rana that evolved in New 
Guinea. 

Only two (6%) Sundaic genera have success- 
fully colonized the Sahul Shelf island of New 
Guinea. Limnonectes has only one species, the 



widespread L. grunniens. Rana has a group of 
nine species, seven closely related and restricted 
to New Guinea, and two less clearly related. The 
seven species are very similar morphologically 
and cytologically (Menzies 1987). None of these 
species are conspecific with species in the 
Greater Sundas or western Wallacea. 

Only one genus (3%) of the Sundaland genera 
is known from the Melanesian Arcs, with two 
species (one endemic) in the Bismarcks and one 
endemic species in the Solomons. These are the 
result of secondary colonization from the New 
Guinea radiation. One of the widespread species 
in New Guinea, Rana daemeli, is also known 
from a population on Cape York, Australia. 

Component II. — For the two families (Hyli- 
dae and Myobatrachidae) of Australia, repre- 
sentation in the Sahul Shelf islands of New 
Guinea and its satellites differs in several re- 
spects from that of the representation of the 
southeast Asian fauna in the Greater Sunda Is- 
lands. Of the 23 genera in Australia, only seven 
(30%) are presently known from New Guinea 
(Table 1, Appendix A). However, the two fami- 
lies are very different. Two (67%) of the three 
genera of the Hylidae are in New Guinea, 
whereas only five (25%) of the 20 genera of the 
Myobatrachidae are in New Guinea. 

At the species level they also differ greatly. For 
the Myobatrachidae there are only 7 species in 
the 5 genera, and none are endemic. For the 
Hylidae there are 80 species, 71 (89%) endemic. 
The number of species is actually greater than the 
69 recorded from Australia (Table 1). For one of 
the genera, Nyctimystes, there is only one species 
known from Australia but 22 in New Guinea. 
These differences in colonization and radiation 
success of the two families suggest an earlier 
colonization for Hylidae than for Myobatrachi- 
dae. 

Because land bridges between Australia and 
New Guinea would have been created by the 
same lower sea-level events as were the Sun- 
daland bridges, the dispersal opportunities for 
anurans during the Pleistocene were probably 
similar between Australia and New Guinea to 
those proposed between southeast Asia and the 
Sunda Shelf islands. Two factors are most impor- 
tant: the large land mass of Australia compared 
to that of southeast Asia adjacent to the Sunda 
Shelf and variation in climate for different re- 
gions of Australia. A comparison limiting the 



BROWN: BIOGEOGRAPHY OF AMPHIBIANS 



31 



Table 2. Distribution patterns for anurans of Component III (subfamilies Platymantinae in Ranidae and 
Asterophryinae and Genyophryninae in Microhylidae) in the southwestern Pacific Islands and Australia. 
The number of genera (G) and species (S) is followed by the number (in parentheses) that are endemic. 



Microhylidae 



Ranidae 



Philippines 


G 


1 




S 


2 (2) 


Western Wallacea 






Sulawesi 


G 


1 




S 


3 (3) 


Lesser Sundas 


G 


1 




S 


2 (2) 


Eastern Wallacea 






Outer Banda Arc 


G 

S 


1 

1 


Northern Moluccas 


G 


3 




S 


5 (5) 


Sahul Shelf 






New Guinea and satellite islands 


G 


16 (13) 




S 


101 (99) 


Melanesian Arcs 


G 


2 




S 


2 (1) 


Fiji Islands 






Palau Islands 






Australia 


G 


2 



G 1 

S 15 (15) 



G 1 

S 4 (3) 

G 4 

S 23 (23) 

G 1 

S 2 (2) 

G 1 

S 1 (1) 



source fauna of Australia to that of the northern 
subtropical and tropical regions might well re- 
veal colonization success similar to that found 
between southeast Asia and the Greater Sunda 
Islands. The radiation of the hylid genera in New 
Guinea is independent of dispersal opportunities. 
It is doubtless related to the availability, at the 
time, of unoccupied anuran niches in the diverse 
communities of the large, mountainous, tropical 
island of New Guinea. 

Both genera of Hylidae (Litoria and Nyctomys- 
tes) have colonized islands of eastern Wallacea, 
but Myobatrachidae is limited to New Guinea. 
Litoria has two species (one endemic) in the 
outer Banda Arc and four (two endemic) in the 
northern Moluccas along with an endemic spe- 
cies of Nyctomystes (Table 1, Appendix B). Li- 
toria also occurs in the Bismarcks and the 
Solomons with two non-endemic species (Table 
1, Appendix B). 



Component III. — For the subfamilies Astero- 
phryninae and Genyophryninae (Microhylidae), 
centered in New Guinea and its satellite islands, 
all 16 recognized genera occur there, with 13 
(81%) endemic. There are 101 species, 99 (98%) 
endemic. One to three genera (6 to 19%), and 
from two to five species in each genus are known 
from several islands of Wallacea as well as the 
Philippines and Melanesian Arcs. Two genera 
with 15 species are recorded from northern Aus- 
tralia (Table 2, Appendix C). 

For the subfamily Platymantinae (Ranidae), 
the four recognized genera are in the Solomons, 
two (50%) are known from the Bismarcks, and 
one (25%) is known from New Guinea, Fiji, 
Palau, and the Philippines. The number of spe- 
cies of Platymantis is 23 in the Melanesian Arcs 
(all endemic), one endemic in the Palaus, two 
endemic in Fiji, and 15 endemic in the Philip- 
pines (Table 2, Appendix C). In addition, there is 



32 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 



one genus (Batrachylodes) in the subfamily 
Raninae (Ranidae) with eight endemic species in 
the Solomons. Its relationship to other ranine 
genera is unknown. 

In summary, the Asiatic families (Component 
I) have all been highly successful in colonizing 
the Greater Sunda Islands, moderately so for the 
Philippines, but in western Wallacea only a few 
genera and species have succeeded. These differ- 
ences correlate well with the geological histories 
of the three areas. The successful colonization of 
New Guinea and a few islands in eastern Wal- 
lacea may be better explained as probably as- 
sisted by man at one or more points in time since 
his migrations began, perhaps as early as 50,000 
years ago as well as recently. There is no evi- 
dence that these two genera are more successful 
in dispersal across marine barriers by means 
other than human assistance. 

The two Australian families (Component II) 
are very uneven in their colonization of the Sahul 
Shelf island of New Guinea. The Hylidae have 
been highly successful and have undergone an 
extensive radiation in the diverse habitats pro- 
vided by this large, mountainous island. The 
Myobatrachidae are represented by only a few 
genera and species. 

The boundary between the predominantly 
Asian fauna and predominantly Australian fauna 
(Components I and II), as originally proposed by 
Wallace (1863) for amphibians, was along the 
western edge of Wallacea (the Makassar Strait) 
and extended north between Borneo and the Phil- 
ippines. He later modified this to include the 
Philippines on the Asian side. A distribution 
pattern of predominantly Australian or Asian 
components holds for all classes of land verte- 
brates, based on the large data bases available 
today (Cranbrook 198 1). The line for the anuran 
fauna, based on the data currently available, is 
close to the midpoint of Wallacea, between the 
Outer Banda Arc and the Lesser Sundas in the 
south and between Sulawesi and northern 
Moluccas (Fig. 2). 

The Problem of the Island-Centered Microhylids 
and Ranids (Component III) 

Although some genera of the microhylids have 
been recognized as unique to New Guinea for a 
century (Mehely 1898) and some of the ranid 
genera unique to the Solomons (Boulenger 
1886), both families were known to be in Asia, 



which was thought to be the immediate source 
area. The geological concept of stable land and 
ocean areas also supported this view until the late 
1960-early 1970 period (e.g. Boulenger 1920; 
Noble 1931; Darlington 1957). However, the 
systematic treatment of these (Component III) 
subfamilies has differed since the time of their 
discovery. I, therefore, summarize separately the 
treatments accorded them. 

The New Guinea subfamilies (Asterophryinae 
and Genyophryninae) of microhylids have been 
regarded as restricted to the southwest Pacific 
Islands and secondarily, Australia by nearly all 
systematists and biogeographers (e.g. Kampen 
van 1923; Parker 1934; Tyler 1979; Zweifel and 
Tyler 1982; Allison 1996). Savage (1973) dif- 
fered in including the Asian genus Calluella in 
the Papuan subfamily Asterophryinae. 

The ranid subfamily Platymantinae (Cornufer- 
inae of Noble) is diagnosed by early authors as 
including various southeast Asian genera 
(Boulenger 1920; Noble 1931; Savage 1973). To 
the best of my knowledge the first diagnosis, 
limiting the genera to those restricted to the is- 
lands, was that of Tyler (1979) and Zweifel and 
Tyler (1982). In this conclusion they were fol- 
lowed by Duellman and Trueb (1985). The latter 
were presumably unaware of descriptions of 
three species (Cornufer [= Platymantis] xi- 
zangensis Hu, Platymantis liui Yang, and Platy- 
mantis reticulatus Zhao and Li) found in western 
China and Tibet between 1977 and 1984. Dubois 
(1987) further confused the issue by erecting the 
genus Ingerana to include these three species and 
several Asian Rana and Micrixalus. He included 
Ingerana in the same subfamily as the island 
platymantine frogs. Zhao and Adler (1993) 
adopted a conservative view and referred all spe- 
cies of Ingerana to Micrixalus. I follow the last 
classification. 

Discussion 

Whether we are considering hypotheses pro- 
posed during the period advocating fixed land 
areas or continental drift, some elements of the 
island fauna are unquestionably relatively recent 
colonizers. These are genera common to the is- 
lands and mainland faunas (Components I and 
II). The island-centered genera and subfamilies 
of the Microhylidae and Ranidae (Component 
III) are the source of many differences in bio- 



BROWN: BIOGEOGRAPHY OF AMPHIBIANS 



33 



geographic hypotheses. My starting point is the 
close of the fixed-land area period and the views 
of Darlington (1957). 

Darlington regarded the old-world tropics (Af- 
rica and Asia) as the source area for amphibians. 
Successive waves dispersed from that center as 
the more recent families evolved and older ones 
became extinct except in peripheral areas (e.g., 
New Zealand). He thought the southwest Pacific 
Island anurans to be the result of relatively recent 
dispersal through southeast Asia and the fauna of 
Australia to be somewhat older but by way of the 
same conduit (Darlington 1957). 

Most biogeographic papers since 1957 that 
have included the New Guinea-centered micro- 
hylids have continued to cite southeast Asia as 
the immediate source area (Duellman and Trueb 
1985; Zweifel and Tyler 1982; Allison 1996). 
The most radical dissent from this view is that of 
Savage ( 1 973). He would derive the New Guinea 
microhylids from an Australian stock that be- 
came extinct in that continent during the Ceno- 
zoic. Because he included the southeast Asian 
genus Calluella in the Asterophrynae, he viewed 
that family as dispersing from New Guinea to 
southeast Asia. He viewed other southern Asian 
genera as derived from a stock that reached Asia 
by way of the Indian subcontinent. 

Recent biogeographic papers that included the 
platymantine frogs have also treated southeast 
Asia as the source area, although presumably at 
an early date. As previously noted, many of these 
authors included south Asian genera in the Platy- 
mantinae (Savage 1973; Dubois 1987, 1992). 
Other papers (Tyler 1979; Zweifel and Tyler 
1982; Duellman and Trueb 1985; Allison 1996) 
limit the platymantine frogs to the island genera, 
but as having evolved from Asian stocks that 
colonized the islands very early. A satisfactory 
explanation of how they were able to disperse 
through the islands as far as Fiji has not yet been 
proposed. Inger (pers. comm.) refers to the Phil- 
ippine Platymantis as Papuan elements and sug- 
gests they may have colonized the Philippines in 
the pre-Oligocene period when the Philippines 
had a more southern position. 

One answer to the question of how and when 
the ancestral platymantine and microhylid stocks 
arrived in Melanesia is more attainable today 
than it was in 1982. If the hypothesis that the 
Inner and Outer Tethys Arcs were derived from 
terranes rifted as ridges from eastern Australia 



prior to the separation of that continent from the 
rest of Gondwana (p. 26) is correct, the ancestors 
to the endemic platymantines and microhylids 
could presumably have been transported on those 
rift blocks such as the Solomon Ridge. This 
assumes that parts of the ridge were subaerial 
throughout the time, and either prior to or sub- 
sequently, these stocks became extinct in Austra- 
lia. The presence of leiopelmatid anurans in New 
Zealand and their probable presence in Australia 
is analogous, although there is no fossil evidence 
to support this. 

Patterns of endemism and distribution within 
islands beyond the Sunda Shelf for the genera 
Metroxyton and Pigafetta in the palm family 
(Arecaceae, subfamily Calamoideae) are similar 
to the patterns of the platymantines and micro- 
hylids; and a Gondwanan origin is also hypothe- 
sized (Dransfield 1987). 

Similarly, the carphodactyline geckos and 
some of the skinks of New Caledonia are be- 
lieved to be the result of vicariant events of stocks 
isolated following rifting from the eastern part of 
Gondwanan Australia in the Mesozoic (Bauer 
1988, 1990; Bauer and Vindum 1990). New 
Caledonia is east of Australia at the southern tip 
of islands that were part of the Inner and Outer 
Arcs. 

In this study I have isolated recent and very old 
anuran colonizers of the Pacific Islands and ana- 
lyzed their distribution patterns independently. 
Phylogenetic studies at the levels of species, 
genera, and subfamilies are much needed for 
both the ranids and microhylids. Such studies 
should help either to confirm, modify, or deny 
the interpretation presented here. Also a better, 
future understanding of the geological history, 
especially of the Inner and Outer Tethys Arcs, 
may provide a more adequate explanation of the 
current island distributions of these endemic ra- 
nid and microhylid frogs. 

ACKNOWLEDGMENTS 

I thank my colleagues in the Department, A. E. 
Leviton and R. C. Drewes, for helpful discus- 
sions and assistance, R. F. Inger, L. R. Heaney, 
R. G. Zweifel, and G. Zug for their valuable 
comments on early drafts of this paper, J. S. 
Brown for typing and editing the manuscript, and 
C. Sudekum for illustrations. 



34 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 



Literature Cited 

Adler, G. H., C. C. Austin, and R. Dudley. 1995. 
Dispersal and speciation of skinks among archipel- 
agos in the tropical Pacific Ocean. Evol. Ecol. 
9:529-541. 

Allison, A. 1993. Biodiversity and conservation of 
the fishes, amphibians, and reptiles of Papua New 
Guinea. Pp. 1 57-225 in Papua New Guinea Conser- 
vation Needs Assessment, Vol. 4, B. M. Beehler, 
ed. Biodiversity Support Program, Washington, 
DC. 

. 1996. Zoogeography of Amphibians and 



Reptiles of New Guinea and the Pacific Region. Pp. 
407-436 in The Origin and Evolution of Pacific 
Island Biotas, New Guinea to Eastern Polynesia: 
Patterns and Processes, A. Kaost and S. E. Miller, 
eds. Academic Publishing, Amsterdam. 

Audlly-Charles, M. G. 1 98 1 . Geological history of 
the region of Wallace's line. Pp. 24-35 in Wallace's 
Line and Plate Tectonics, Oxford Monog. Biogeog. 
2, T. C. Whitmore, ed. Clarendon Press, Oxford. 



— . 1987. Dispersal of Gondwanaland: relevance 
to evolution of the angiosperms. Pp. 5-25 in Bio- 
geographical Evolution of the Malay Archipelago, 
Oxford Monogr. Biogeog. 4, T. C. Whitmore, ed. 
Clarendon Press, Oxford. 

— . 1988. Evolution of the southern margin of 



Tethys (north Australian region) from early Per- 
mian to late Cretaceous. Pp. 79-100 /'// Gondwana 
and Tethys, M. G. Audley-Charles and A. Hal lam, 
eds. Oxford University Press, Oxford. 

Bauer, A. M. 1988. Reptiles and the biogeographic 
interpretation of New Caledonia. Tuatara 30:39-50. 



. 1990. Phylogenetic systematica and bio- 
geography of the Carphodactilini (Reptilia: 
Gekkonidae). Bonn. Zool. Monogr. 30:65-88. 

Bauer, A. M. and J. V. Vindum. 1990. A checklist 
and key to the herpetofauna of New Caledonia, with 
remarks on biogeography. Proc. Calif. Acad. Sci. 
47(2): 1 7-45. 

Boulenger, G. A. 1882. Catalogue of the Batrachia 
Salientia Ecaudata in the Collection of the British 
Museum, London. 1 :503 pp. 

— . 1 886. On reptiles and batrachians of the Solo- 



— . 1920. A monograph of the south Asian, Pap- 
uan, Melanesian, and Australian frogs of the genus 
Rana. Rec. Ind. Mus. 20:1-226. 

Brown, W. C. 1 953. The amphibians of the Solomon 
Islands. Bull. Mus. Comp. Zool. 1 07( 1 ): 1 -64. 

Brown, W. C. and M. J. Tyler. 1968. Frogs of the 
genus Platymantis (Ranidae) from New Britain with 
descriptions of new species. Proc. Biol. Soc. Wash- 
ington. 81:69-86. 

Brown, W. C. and A. C. Alcala. 1970. The zooge- 
ography of the herpetofauna of the Philippine Is- 
lands, A fringing archipelago. Proc. Calif. Acad. 
Sci. 38(6)405-130. 

. 1994. Philippine frogs of the family Rhaco- 

phoridae. Proc. Calif. Acad. Sci. 48(10): 1 85-220. 

Butler, R. 1995. When did India hit Asia? Nature 
373:20-21. 

COGGER, H. G. 1992. Reptiles and Amphibians of 
Australia, A. H. and A. W. Reed, eds. Sydney, Aus- 
tralia. 775 pp. 

Coleman, P. J. 1980. Plate tectonics background to 
biogeographic development in the southwest Pa- 
cific over the last 100 million years. Palaeogeogr. 
Palaeoclimat. Palaeoecol. 31 : 105-121. 

Coulson, F. I. and J. G. Vedder. 1986. Geology of 
the central and western Solomon Islands, Pp. 59-86 
in Geology and Offshore Resources of Pacific Is- 
land Arcs — Central and Western Solomon Islands, 
Circum-Pacific Council for Energy and Mineral 
Resources, Earth Sci. Ser. V. 4, J. G. Vedder, K. S. 
Pound, and S. Q. Boundy, eds. Houston, Texas. 

Cranbrook, (Earl of) 1981. The vertebrate fauna. 
Pp. 57-69 in Wallace's Line and Plate Tectonics. 
Oxford Monog. Biogeog. 2, T. C. Whitmore, ed. 
Clarendon Press, Oxford. 

Darlington, P.J. 1957. Zoogeography: The Geo- 
graphical Distribution of Animals. Wiley and Sons, 
New York. 675 pp. 

Diamond, J. 1984. Biogeographic mosaics in the Pa- 
cific. Pp. 1-14 in Biogeography of the Tropical 
Pacific, Spec. Pub. No. 72, F. J. Radovsky, P. H. 
Raven, and S. H. Sohmer, eds. Association of Sys- 
tematica Collections and Bemiee P. Bishop Mu- 
seum. 



mon Islands. Trans. Zool. Soc. London 12:35-62. 



BROWN: BIOGEOGRAPHY OF AMPHIBIANS 



35 



Diamond, J. M. and E. Mayr. 1976. Species-area 
relation for birds of the Solomon Archipelago. Proc. 
Nat. Acad. Sci. USA. 73:262-266. 

DranSFIELD, J. 1987. Bicentric distribution in Male- 
sia as exemplified by palms. Pp. 60-72 in Bio- 
geographical Evolution of the Malay Archipelago, 
Oxford Monogr. Biogeog. 4, T. C.Whitmore, ed. 
Clarendon Press, Oxford. 

Dubois, A. 1987. Miscellanea taxinomica ba- 
trachologica(I). Alytes, 5:7-95. 



— . 1992. Notes sur la classification des Ranidae 
(Amphibiens Anoures). Bull. Mens. Soc. Linn. 
Lyon 61:305-352. 

Duellman, W. E. 1993. Amphibian Species of the 
World: Additions and Corrections. Univ. Kansas 
Mus. Nat. Hist. Spec. Pub. 21:1-372. 

Duellman, W. E. and L. Trulb. 1985. Biology of 
Amphibians. McGraw-Hill Book Co., New York. 
670 pp. 

Erwin, T. L. 1 98 1 . Taxon pulses, vicariance, and dis- 
persal: an evolutionary synthesis illustrated by 
carabid beetles. Pp. 159-183 in Vicariance Bio- 
geography a Critique, G. Nelson and D. E. Rosen, 
eds. Columbia University Press, New York. 

Ford, L. S. and D. C. Cantella. 1993. The major 
clades of frogs. Herp. Monogr. 7:94-1 1 7. 

FROST, D. R., ed. 1985. Amphibian Species of the 
World. Allen Press and Association of Systematic 
Collections, Lawrence, Kansas. 



ciation of Systematics Collections and Bernice P. 
Bishop Museum. 

Inger, R. F. 1954. Systematics and zoogeography of 
Philippine Amphibia. Fieldiana: Zool. 33: 1 8 1-53 1 . 

. 1966. The Amphibia of Borneo. Fieldiana: 



Zool. 52:1-402. 

Jarrand, R. D. and S. Sasajima. 1980. Palaeomag- 
netic synthesis for Southeast Asia: constraints on 
plate movements. Pp. 293-3 1 6 in The Tectonic and 
Geologic Evolution of Southeast Asian Seas and 
Islands, D. E. Hayes, ed. American Geophysical 
Union, Geophys. Monogr. 23. 

Kampen van, P. N. 1923. The Amphibia of the Indo- 
Australian Archipelago. E. J. Brill, Leiden, The 
Netherlands. 304 pp. 

Kroenke, L. W. 1984. Cenozoic tectonic develop- 
ment of the southwest Pacific. United Nations Eco- 
nomic and Social Commission, Committee for 
Coordination of Joint Prospecting for Mineral Re- 
sources in the South Pacific Offshore Area. Tech. 
Bull. 6:1-122. 

LlEM, S. -S. 1970. The morphology, systematics, and 
evolution of the Old World treefrogs (Rhacophori- 
dae and Hyperolidae). Fieldiana: Zool. 52:1-145. 

McCabe, R., J. N. Almasco, and G. Yumul. 1985. 
Terranes of the central Philippines. Pp. 421^36 in 
Tectonostratigraphic Terranes of Circum-Pacific 
Region, Circum-Pacific Energy and Mineral Re- 
sources, Earth Science Series Vol. 1 , Howell, D. H., 
ed. 



Hamilton, W. 1979. Tectonics of the Indonesian 
region. U. S. Geol. Surv. Prof. Pap. 1078:1-345. 

Heaney, L. R. 1991. An analysis of patterns of distri- 
bution and species richness among Philippine fruit 
bats (Pheropodidae). Bull. Amer. Mus. Nat. Hist. 
206:145-167. 

Hallam, A. 1 98 1 . Relative importance of plate move- 
ments, eustacy, and climate in controlling major 
biogeographical changes since the early Mesozoic. 
Pp. 303-330 in Vicariance Biogeography a Cri- 
tique, G. Nelson and D. E. Rosen, eds. Columbia 
University Press, New York. 

Halloway, J. D. 1984. Lepidoptera and the Melane- 
sian Arcs. Pp. 129-163 in Biogeography of the 
Tropical Pacific, Spec. Pub. No. 72, F. J. Ra- 
dovsky, P. H. Raven, and S. H. Sohmer, eds. Asso- 



McCabe, R. and J. Cole 1987. Speculations on the 
late mesozoic and cenozoic evolution of the south- 
east Asian margin. Pp. 375-394 in Transactions of 
the Fourth Circum-Pacific Energy and Mineral Re- 
sources Conference, M. K. Horn, ed. Circum-Pa- 
cific Council for Energy and Mineral Resources, 
Tulsa, Oklahoma. 

Mehley van, L. 1 898. An account of the reptiles and 
batrachians collected by Mr. Lewis Biro in New 
Guinea. Tennesz. Fiiz. 21:1 65-1 78. 

Menzies, J. I. 1982. Systematics of Platymantis 
papuensis (Amphibia: Ranidae) and related species 
of the New Guinea region. British J. Herp. 
6:236-240. 

. 1987. A taxonomic revision of Papuan Rana 



(Amphibia: Ranidae). Aust. J. Zool. 35:373^118. 



36 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 



Metcalfe, I. 1988. Origin and assembly of south-east 
Asian continental terranes. Pp. 101-118 in Gond- 
wana and Tethys, M. G. Audley-Charles and A. 
Hallam, eds. Oxford University Press, Oxford. 

Milsom, J. S. and Audley-Charles, M. G. 1986. 
Post-collision isostatic readjustment in the southern 
Banda arc. Pp. 353-364 in Collision Tectonics, M. 
P. Coward and A. Ries, eds. Geological Society of 
London. 

MORLEY, R. J. AND J. R. Flenley. 1987. Late Caino- 
zoic vegetational and environmental changes in the 
Malay Archipelago. Pp. 50-59 in Biogeographical 
Evolution of the Malay Archipelago, Oxford 
Monogr. Biogeog. 4, T. C. Whitmore, ed. Claren- 
don Press, Oxford. 

Murphy, R. W. 1987. Southeast Asia: A Tectonic 
Triptich. Pp. 395^400 in Transactions of the Fourth 
Circum-Pacific Energy and Materials Resources 
Conference, M. K. Horn, ed. Circum-Pacific Coun- 
cil for Energy and Mineral Resources. Tulsa, Okla- 
homa. 

Nishiwaki, C. and S. Uyeda 1987. Mode of subduc- 
tion, collision, and types of metallogenesis. Pp. 
455-464 in Transactions of the Fourth Circum-Pa- 
cific Energy and Mineral Resources Conference, 
M. K. Horn, ed. Circum-Pacific Council for Energy 
and Mineral Resources, Tulsa, Oklahoma. 

Noble, G. K. 1931. Biology of the Amphibia. 
McGraw-Hill, New York. 577 pp. 

Nunn, P. D. 1994. Oceanic Islands. Blackwell, Ox- 
ford, England. 413 pp. 

Ollier, C. D. 1985. The geological background to 
prehistory in Southeast Asia. Pp. 25-42 in Modern 
Quaternary Research in Southeast Asia, J. Bartstra 
and W. A. Casparje, eds. Biologisch Archaeolo- 
gisch Institute, Rotterdam. 

Parker, H. W. 1 934. A monograph of the frogs of the 
family Microhylidae. British Museum (Natural His- 
tory), London. 208 pp. 

Peters, W. and G. Doria. 1 878. Catologo der rettili 
e dei batraci raciolti da O. Beccari, L. M. D'Alber- 
tise, e A. A. Bruijnnela Salto-Regione Austro- 
Malese. Ann. Mus. Civ. Stor. Nat. Genova 
13:323-450. 

Savage, J. M. 1973. The geographic distribution of 
frogs: patterns and predictions. Pp. 35 1-445 in Evo- 



lutionary Biology of Anurans, J. L. Vial, ed. Univ. 
Missouri Press, Columbia. 

Springer, V. G. 1982. Pacific plate biogeography 
with special reference to shore fishes. Smithson. 
Contrib. Zool. 367:1-182. 

TAYLOR, E. H. 1920. Philippine Amphibia. Phillip. J. 
Sci. 16:213-359. 

Tyler, M.J. 1979. Herpetological relationships of 
South America and Australia. Monogr. Mus. Nat. 
Hist. Univ. Kansas 7:73-106. 



Wallace, A. R. l: 
London. 526 pp. 



Island Life, Macmillan Co. 



Whitmore, T. C. 1981. Introduction. Pp. 1-2 in Wal- 
lace's Line and Plate Tectonics, Oxford Monogr. 
Biogeog. 2, Whitmore, T. C, ed. Clarendon Press, 
Oxford. 

Zhao, Er-mi and K. Adler. 1993. Herpetology of 
China. Society for the Study of Amphibians and 
Reptiles, in cooperation with the Chinese Society 
for the Study of Amphibians and Reptiles, 1 52 pp. 

Zweifel, R. G. AND M. J. Tyler. 1 982. Amphibia of 
New Guinea. Monogr. Biol. 42:759-801. 

Appendix A 
Southeast Asian Anuran Genera, (B) means 
also in Borneo. 

Megophryidae 

Leptobrachium (B) 

Leptolalax (B) 

Megophrys (B) 
Hylidae 

Hyla 
Bufonidae 

Ansonia (B) 

Bufo (B) 

Leptophryne (B) 

Pedostibes (B) 

Pelophryne (B) 

Pseudobufo (B) 
Microhylidae 

Calluella (B) 

Chaperina (B) 

Glyphoglossus 

Kalophiynus (B) 

Kaloula(B) 

Metaphtynella (B) 



BROWN: BIOGEOGRAPHY OF AMPHIBIANS 



37 



Microhyla (B) 

Phrynella (B) 
Ranidae 

Amalops 

Elachyglossa 

Hoplobatrachus (B) 

Huia (B) 

Limnonectes (B) 

Micrixalus 

Occidozyga (B) 

Paa 

Phiynoglossus (B) 

Rana (B) 
Rhacophoridae 

Chihxalus 

Nyctixalus (B) 

Philautus (B) 

Polypedates (B) 

Rhacophorus (B) 

Theloderma (B) 

Australian Anuran Genera, (NG) means also 
in New Guinea. 

Hylidae 

I//o/7fl (NG) 

Nyctimystes (NG) 

Cyclorana 
Myobatrachidae 

Arenophryne 

Assa 

Crinia (NG) 

Geoc/7/7/V/ 

Heleioporus 

Lechriodus (NG) 

Limnodynastes (NG) 

Metac/7/7/a 

Mixophyhyes (NG) 

Myobatrachus 

Neobatrachus 

Not a den 

Paracrinia 

Philoria 

Pseudoplvyne 

Rheobatrachus 

Taudachlus 

Uperoleia (NG) 



Appendix B 
Asiatic and Australian Families and Genera of 
Anurans in the Islands of Wallacea, New Guinea, 
and the Melanesian Arcs. 

Sulawesi and Sula-Banggai 

Bufonidae. — Bufo celebensis 

Microhylinae (Microhylidae). — Kaloula 
baleata, K. pulchra 

Ranidae. — Limnonectes cancrivorus, L. 
grunniens, L. heinrichi, L. kuhli, L. mod- 
esta, Phiynoglossus celebensis, P. laevis, 
P. semipalmata, Rana arathooni, R. cele- 
bensis, R. chalconta, R. eiythraea, R. 
macrops 

Rhacophoridae. — Rhacophorus edentulus, 
R. georgi, R. monticola, and Polypedates 
leucomystax 
Lesser Sundas (Lomboc, Sumbawa, Flores, 
Wetar) 

Bufonidae. — Bufo biporcatus 

Microhylinae (Microhylidae). — Kaloula 
baleata 

Ranidae. — Limnonectes cancrivorus, L. 
limnocharis, L. macrodon, L. verrucu- 
losa, Phiynoglossus florensis, P. laevis, 
Occidozyga lima, Rana florensis, R. el- 
berti, R. daemeli 

Rhacophoridae. — Polypedates leucomystax 

Outer Banda Arc (Sumba, Timor, Tanimbar, 
Seram, Ambon. Bum) 

Hylidae. — Litoria amboinensis, L. capitula, 

L. everetti, L. infrafrenata, L. vagahunda 

Ranidae. — Limnonectes grunniens, L. mod- 

esta, L. verruculosa, Rana elberti, R. 

grisea 

Rhacophoridae. — Polypedates leucomystax 

Northern Moluccas (Halmahera, Morotai. Obi. 

Bacan. Ternate) 

Hylidae. — Litoria infrafrenata, Nyctimyec- 

tes rueppelli 
Ranidae. — Limnonectes modesta, Rana 
moluccana 
New Guinea 

Ranidae. — Limnonectes grunniens, Rana 
arfaki, R. daemeli, R. garritor, R. grisea, 
R.jimiensis, R. novaeguinae, R.papua, R. 
semevella, R. supragrisea 



38 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 2 



Melanesian Arcs 
Bismarcks (New Britain, New Ireland) 
Ranidae. — Rana kreffti, R. daemeli 
Hylidae. — Litoria infrafrenata, L. the- 
saurensis 
Solomon Islands 
Ranidae. — Rana kreffti 
Hylidae. — Litoria infrafrenata, L. the- 
saurensis 
Australia 

Ranidae. — Rana daemeli 

Appendix C 
Non-Asiatic and Non-Australian Subfamilies 
of Anurans in the Islands of Wallacea, Philip- 
pines, New Guinea, and Melanesian Arcs. 

Western Wallacea 

Sulewesi and Sula-Banggai 

Genyophryninae (Microhylidae). — Oreo- 
phryne celebensis, O. variabilis, O. zim- 
meri 
Lesser Sundas 

Genyophryninae (Microhylidae). — Oreo- 
phiyne jeffersoniana, O. monticola 
Eastern Wallacea 
Outer Banda Arc 
Asterophryinae (Microhylidae). — Callu- 
lopsfuscus 
Northern Moluccas (Halmahera) 
Asterophryinae (Microhylidae). — Callu- 

lops boettgeri, C. dubia 
Genyophryninae (Microhylidae). — Cophix- 
alus montanus, Oreophiyne frontifas- 
ciata, O. moluccensis 
Philippines 

Platymantinae (Ranidae). — Platymantis 
corrugatus, dorsalis, insulatus, guen- 
theri, hazelae, ingeri, lawtoni, levigatus, 
mimulus, montanus, polilloensis, pan- 
ayensis, reticulums, spaeleus, subterres- 
stris 

Genyophryninae (Microhylidae). — Oreo- 
phiyne annulata, O. nana 



Sahul Shelf 

New Guinea and satellite islands 

Platymantinae (Ranidae). — Platymantis 
cheesmanae, P. batantae, P. punctata, P. 
papuensis 
Astereophryinae and Genyophryninae (Mi- 
crohylidae). — The 16 genera and 101 spe- 
cies (99 endemic) are not listed here but 
may be found in Frost (1985) and Duell- 
man(1993). 
Melanesian Arcs 

Bismarcks (New Britain, New Ireland, New 
Hanover) 
Platymantinae (Ranidae). — Platymantis 
akari thymus, P. boulengeri, P. gilliardi, 
P. macroceles, P. magnus, P. mimicus, P. 
nexipus, P. schmidti, Discodeles guppyi 
Geneophryne (Microhylidae). — Oreo- 

phiyne brachypus, Sphenophryne me- 
helyi 
Admiralty Islands 

Platymantinae (Ranidae). — Discodeles ven- 
tricosus, Platymantis gilliardi, P. species 
Solomon Islands 

Platymantinae (Ranidae). — Platymantis 
acrochordus, P. aculeodactylus, P. gup- 
pyi, P. myersi, P. neckeri, P. parkeri, P. 
solomonis, P. weberi, Discodeles bufoni- 
formis, D. guppyi, D. malukuna, D. opis- 
thodon, Ceratobatrachus guentheri, 
Palmatorappia solomonis 
Raninae (Ranidae). — Batrachylodes ele- 
gans, B. gigas, B. mediodiscus, B. minu- 
tus, B. montanus, B. trossulus, B. 
vertebralis, B. wolfi 
Fiji 

Platymantinae (Ranidae). — Platymantis vi- 
tianus, P. vitiensis 
Palau Islands 

Platymantinae (Ranidae). — Platymantis pe- 
lewensis 



© CALIFORNIA ACADEMY OF SCIENCES, 1997 

Golden Gate Park 

San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 3, pp. 39-114. 



December 9, 1997 



SPECIES CATALOG OF THE NEUROPTERA, MEGALOPTERA, 
AND RAPHIDIOPTERA OF AMERICA NORTH OF MEXICO 

1 WOOds L/Jf D ' 'OQ/caM T — 

By / Libjy raphlc lns[ lt ut l0 . 

Norman D. Penny 

Department of Entomology, California Academy of Sciences, 
San Francisco, CA 941 IS 

Phillip A. Adams 

California State University, Fullerton, CA 92634 



and 



Lionel A. Stange 

Florida Department of Agriculture, Gainesville, FL 32602 



The 399 currently recognized valid species of the orders Neuroptera, Megaloptera, and 
Raphidioptera that are known to occur in America north of Mexico are listed and full synonymies 
given. Geographical distributions are listed by states and provinces. Complete bibliographic 
references are given for all names and nomenclatural acts. Included are two new junior 
homonyms indicated, seven new taxonomic combinations, two new changes of rank, fourteen new 
synonymies, three new lectotype designations, and one new name. 



Received March 20, 1996. Accepted June 3, 1997. 



The recent publication of Nomina Insecta 
Nearctica, A Check List of the Insects of North 
America (Poole 1996) has given us a listing of 
North American Neuropterida (Neuroptera + 
Megaloptera + Raphidioptera) species for the 
first time in more than a century. However, for 
anyone trying to identify these species, the litera- 
ture is scattered and obscure. Only with the great- 
est difficulty is a non-specialist able to keep up 
with current useage of names. With this lack of 
reference materials in mind, the present catalog 
has been developed to allow the non-specialist 
and specialist alike to grasp the limits and diver- 
sity of these groups. The primaiy literature has 



been consulted whenever possible, as well as 
Zoological Record, and appropriate mono- 
graphic revisions published up to 1 January 1997. 
A number of taxonomic changes are incorpo- 
rated into this catalog: there are two new junior 
homonyms indicated, seven new taxonomic 
combinations, two new changes of rank, fourteen 
new synonymies, three new lectotype designa- 
tions, and one new name. However, the classifi- 
cation of neuropteroid insects is constantly being 
modified and enhanced as we understand the 
group better. Thus, the authors know that the 
genus "Mantispa"' is currently being redefined by 
Kevin Hoffman, and the genus "Brachynemu- 



[39] 



40 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



ms" was recently redefined by Stange (1994). 
Such studies enhance our knowledge of these 
groups and are a normal part of taxonomic devel- 
opment and maturity, but at the same time they 
require that this list, as with any catalog, be used 
in conjunction with other recent taxonomic lit- 
erature to ensure the most up-to-date treatment 
of any particular taxon. To help maintain cur- 
rency, we have listed most recent treatments of 
included taxa at the end of each family. 

The geographical coverage of this catalog in- 
cludes continental United States (including 
Alaska with its offshore islands), and Canada. 
Greenland has been excluded. The southern lim- 
its are the political boundaries of the United 
States, excluding the Caribbean and all offshore 
islands, such as the Bahamas, not belonging to 
the United States. The Hawaiian Islands have 
been excluded because they belong to another 
faunal region, as well as having a current, on-line 
faunal listing through the Hawaiian Biological 
Survey URL address (http: //www. bishop. 
hawaii.org/bishop/HBS/). 

This portion of the Nearctic Region, excluding 
Mexico, currently contains 399 valid species, 
distributed in the following families: 
Family Genera Species 

Neuroptera 
Ascalaphidae 3 8 

Berothidae 1 10 

Chrysopidae 15 81 

Coniopterygidae 8 55 

Dilaridae 1 2 

Hemerobiidae 6 61 

Ithonidae 1 1 

Mantispidae 4 15 

Myrmeleontidae 17 94 

Polystoechotidae 2 2 

Sisyridae 2 6 

Raphidioptera 
Inocellidae 1 3 

Raphidiidae 2 18 

Megaloptera 
Corydalidae 7 19 

Sialidae 1 24 



Total 



71 



399 



Distributional information is credited to an 
author only if a state or province is specifically 
mentioned in the referenced work. Thus, dis- 
tributional records associated with some species 



are sparse, even though earlier authors may have 
mentioned such regions as "Atlantic Coastal 
States" or "Maritime Provinces." New distribu- 
tional data introduced in this catalog are credited 
to the contributing author. 

Some of the published records must be viewed 
with caution, particularly among older literature. 
Where we know records are wrong, such as with 
Chrysoperla externa in Canada, we have indi- 
cated this. However, other records which may 
have been correct are listed without comments 
under each species treatment. 

Type label information has come from three 
sources. Where possible, information has been 
transcribed from the type specimens. Much label 
information has also been cited from the original 
literature. A valuable third source has been the 
electronic type catalog at Harvard University's 
Museum of Comparative Zoology (gopher: 
//huh. harvard.edu: 179/7). which houses the 
entensive Banks collection. 

There are a surprisingly large number of North 
American types in the collection at the Museum 
of Comparative Zoology, Harvard University. In 
addition to the expected types of Hagen, Banks, 
and Caipenter, there are also primary types de- 
scribed by Schneider, Burmeister, Smith and 
Navas, among others. Information about the 
types at this museum are now available as a 
listing on internet. 

Within the main body of the catalog the orders 
are arranged: Neuroptera, Raphidioptera, and fi- 
nally Megaloptera. Families are arranged alpha- 
betically. The included species are arranged 
within a taxonomic infrastructure of subfamilies 
and tribes in the larger families, with included 
genera and species organized alphabetically. 
Each valid species includes in sequential order: 
reference to the original description; original ge- 
neric placement; label data of type specimens and 
type depository; synonyms, their label data and 
type depository; taxonomic status changes with 
appropriate references; and finally geographical 
distribution by state or province within the de- 
fined area. Acronyms for states/provinces are 
alphabetized and followed by a superscript 
number that keys to the citation at the end of the 
entry, e.g., AZVSay, 1824:305). 

Generic synonymies were published by 
Oswald and Penny (1991) and are not repeated 
here. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



41 



Abbreviations 

The state and province abbreviations used 
throughout this paper are the two letter codes 
used by the U. S. Postal Service for the United 
States and Canada. Collection abbreviations are 
as follows: 

AMNH = American Museum of Natural His- 
tory, New York, NY 

ANSP = Academy of Natural Sciences, Phila- 
delphia, PA 

BMNH = The Natural History Museum, Lon- 
don, England 

CAS = California Academy of Sciences, San 
Francisco, CA 

CIC = Cawthron Institute Collection, New 
Zealand 

CMNH = Carnegie Museum of Natural His- 
tory, Pittsburgh, PA 

CNC = Canadian National Collection, Ottawa, 
Canada 

CORN = Cornell University, Ithaca, NY 

DEIC = Institut fur Pflantzenschutzforschung, 
Eberswalde, Germany 

HMB = Museum fur Naturkunde der Hum- 
boldt Universitat, Berlin, Germany 

INHS = Illinois State Natural History Sur- 
vey, Champaign, IL 

IRSNB = Institut Royal des Sciences 
Naturelles de Belgique, Brussels, Belgium 

LACM = Los Angeles County Natural History 
Museum, Los Angeles, CA 

MACN = Museo Argentino de Ciencias Natu- 
rales, Buenos Aires, Argentina 

MCZ = Museum of Comparative Zoology, 
Harvard University, Cambridge, MA 

MLP = Museo de La Plata, La Plata, Argentina 

MNHN = Museum National d'Histoire 
Naturelle, Paris, France 

MZB = Museu de Zoologia, Barcelona, Spain 

MZH = Universitetets Zoologiska Museum, 
Helsingfors, Finland 

NMV = National Museum of Victoria, Mel- 
bourne, Australia 

NRS = Naturhistoriska Riksmuseum, Stock- 
holm, Sweden 

OHSTU = Ohio State University Insect Col- 
lection, Columbus, OH 

PSU = Pennsylvania State University Insect 
Collection, State College, PA 



ROM = Royal Ontario Museum, Ontario, 
Canada 

SDNHM = San Diego Natural History Mu- 
seum, San Diego, CA 

SEM = Snow Entomological Museum, 
Lawrence, KS 

USNM = National Museum of Natural His- 
tory, Smithsonian Institution, Washington, DC 

UCB = University of California, Berkeley, CA 

UCD = University of California, Davis, CA 

YPM = Yale University Peabody Museum, 
New Haven, CT 

ZIL = Zoological Institute, Lund, Sweden 

ZMC = Zoologiske Museum, Copenhagen, 
Denmark 

ZMH = Zoologisches Museum, Hamburg, 
Germany 

ZMUO = Zoological Museum, University of 
Oslo, Norway 

ZSBS = Zoologische Sammlungen des Bayer- 
ischen Staates, Munich, Germany 

ASCALAPHIDAE 
Ascalaphidae, or owlflies, as adults are large 
crepuscular aerial predators. For some species 
the daily flight period is very short, less than a 
half hour before total darkness in the evening 
(MacNeill 1962). Eggs are laid on the tips of 
twigs (Henry 1977), often near streams. The lar- 
vae live on the soil surface or on leaves, where 
they await passing prey. Adults are strong fliers 
and superficially resemble dragonflies, except 
for their distinctive long antennae. Males and 
females are often dimorphic in wing coloration, 
wing shape, and even sometimes antennal length. 
The world fauna was monographed by Weele 
( 1 909a). Shetlar's ( 1 977) unpublished Ph.D. the- 
sis is the most recent treatment of North Ameri- 
can species. Only three genera are known from 
North America. 

Haplogleniinae 

Ascalobyas Penny, 1982:395 

Ascalobyas albistigma (Walker), 1853:452 [As- 
calaphus]. 

Holotype (sex unknown): Honduras (BMNH). 

Taxonomy. — To Haploglenius (see 
McLachlan, 1871:236); to Byas (see Weele, 
1908:30); to Ascalobyas (see Penny, 1982:395). 



42 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Distribution. 
1990:158). 



USA: TX (Vogtsberger, 



Ascaloptynx Banks, 1915:350 

Ascaloptynx appendiculatus (Fabricius), 
1793:96 [Ascalaphus]. 

Holotype female: USA, "Carolinas" (BMNH). 

= Ptynx juvenilis McLachlan, 1871:239. 
Type(s) (sex unknown): USA, Texas, Belfrage 
(BMNH). NEW SYNONYMY. 

= Ptynx furciger McLachlan, 1891:509. Five 
syntypes (3 males, 2 females): USA, Arizona, H. 
K. Morrison (BMNH). NEW SYNONYMY. 

TAXONOMY. — To Haploglenius (see Ram- 
bur, 1842:363); to Ptynx (see McLachlan, 
1871:239); to Neuroptynx (see Banks, 
1907c:32); to Ascaloptynx (see Banks, 
1 9 1 5a:350); {P. juvenilis = A. appendiculatus, P. 
furciger = A. appendiculatus (see Shetlar, 
1977:99). 

Distribution. — USA: AL 5 , AR 5 , AZ 2 , FL 3 
GA 3 , MO 4 , MS 6 , NC 3 , OK 5 , SC 5 , TN 5 , TX 1 
VA 5 ('McLachlan, 1871:239, as P. juvenilis 
2 McLachlan, 1891:509; 3 Weele, 1908:58, 59 
4 Froeschner, 1947:129; 5 Shetlar, 1977:102 
6 Lago& Testa, 1989:12). 

Remarks. — The synonymy of Ptynx ju- 
venilis and Ptynx furciger with Ascaloptynx ap- 
pendiculatus was first suggested by Shetlar in 
1977. However, his Ph.D. thesis was never pub- 
lished and therefore cannot be used for nomen- 
clatorial purposes. The present publication is the 
first official record of these synonymies. 

ASCALAPHINAE 

Ululodes Currie in Smith, [1900] 1899:57 

Ululodes arizonensis Banks, 1907a:275. Three 
syntypes (sex unknown): USA, Arizona, Palmer- 
lee, IX, C. R. Biedrman; same data, except VII 
(two specimens) (MCZ). NEW STATUS. 

TAXONOMY. — To U. quadrimaculata ari- 
zonensis (see Weele, 1908: 1 10); to U. arizonen- 
sis (see Shetlar, 1977: 142). 

DISTRIBUTION. — USA: AZ 1 , CA 2 NM 2 , 
NV 3 , TX 2 , UT 2 ('Banks, 1907a:275; "Shetlar, 
1977:147, 148); 3 Penny, NEW STATE RE- 
CORDS). 



Remarks. — The elevation of Ululodes ari- 
zonensis to species rank was first suggested by 
Shetlar in 1977. However, his Ph.D. thesis was 
never published and therefore cannot be used for 
nomenclatorial purposes. The present publica- 
tion is the first official record of this status 
change. 
Ululodes bicolor (Banks), 1895a:521 [Ulula]. 

Three syntypes (sex unknown): Mexico, Baja 
California Sur, San Jose del Cabo; Sierra El 
Taste, IX (3 specimens) (MCZ). 

TAXONOMY. — To Ululodes (see Weele, 
1908:105). 

Distribution. — USA: AZ 1 , CA 1 , LA 2 , TX 1 
('Shetlar, 1977: 138; 2 Penny, NEW STATE RE- 
CORD). 

Ululodes floridana (Banks), 1906a:99 [in Suh- 
palacsa as "Suhpalasca"] 

Holotype male: USA, southern Florida 
(MCZ). REVISED STATUS. 

TAXONOMY. — To subspecies of Ululodes 
quadrimaculata (see Weele, 1908:1 1 1); to spe- 
cies status (see Shetlar, 1977:139). 

REMARKS. — The elevation of Ululodes flori- 
dana to species rank was first suggested by Shet- 
lar in 1977. However, his Ph.D. thesis was never 
published and therefore cannot be used for no- 
menclatorial purposes. The present publication is 
the first official record of this status change. 

Distribution. — USA: FL 1 , GA 2 , SC 2 
('Banks, 1906a:99; 2 Shetlar, 1977:141). 
Ululodes macleayana (Guilding), 1825:140 
[Ascalaphus]. 

Holotype male: Antilles, Saint-Vincent, 
VI. 1824, Guilding (BMNH). 

= Ascalaphus senex Burmeister, 1839:1001. 
Syntype(s) unknown: locality unknown (collec- 
tion unknown). 

= Ulula microcephala Rambur, 1842:359. 

= Ascalaphus avunculus Hagen, 1861:238. 
Syntype(s) unknown: Cuba, Poey (USNM). 

= Ululodes macleavana hageni Weele, 
1908:101. 

TAXONOMY. — To Ulula (see McLachlan, 
1 87 1 :247); to Ululodes as synonym oft/, hyalina 
(see Banks, 1907c:32); as valid species (see 
Weele, 1908:101). 

Distribution. — USA: AL 4 ,AR 4 FL 1 GA 4 , 
KS 2 , LA 4 , MO 3 , MS 4 , NC 5 , NJ ' , OK 4 , SC 4 , TX ' 
(' Weele, 1908:99 as subspecies U. m. hageni; 
"Smith, 1925:170; 3 Froeschner, 1947:129; 



PENNY, ADAMS AND STANGE: SPECIES CATALOG 



43 



4 Shetlar, 1977: 1 16-120; 5 Penny, NEW STATE 
RECORD) 

Ululodes nigripes Banks, 1943:79. 

Holotype (sex unknown): USA, Texas, Davis 
Mountains, D. J. & J. N. Knull (OHSTU). 

Distribution. — USA: TX (Banks, 
1943:79). 

Ululodes quadripunctatus (Burmeister), 
1839:1001 [Ascalaphus]. NEW STATUS 

Type(s) (female): USA, New York (MCZ). 

= Ascalaphus quadrimaculata (Say), 
1824:305. Ajunior primary homonym of Asca- 
laphus quadrimaculatus Lichtenstein, 1 796: 192. 
Holotype female: USA, Pennsylvania (MCZ). 

= Colobopterus excisus Hagen, 1887:193. 
Holotype male: USA, Florida, 1862, Uhler 
(MCZ). 

= Ulula albifrons Banks, 1901c: 172. Twosyn- 
type females: USA, Arizona, Phoenix, 20. IX, 
Kunze (MCZ). 

TAXONOMY. — To Ulula (see McLachlan. 
1 87 1:247); to Ululodes (see Currie, 1900:57); {A. 
quadripunctatus = U quadrimaculata; U. albi- 
frons = U quadrimaculata albifrons) (see 
Weele, 1908:109); (C. excisus = U. quadrimacu- 
lata) (see Beatty & Runner, 1970,: 153). 

Distribution. — CAN: ON 13 , USA: AL 12 , 
AR 14 . AZ 6 , DC 4 . DE 12 ,FL 3 , IA 12 , IL 12 , IN*, 
KS 7 , KY 12 , LA 1 ", MD", MI 12 , MO 10 , MS 12 



,2 ,pa\sc 



NC^NE ,4 ,NJ 5 ,NY-, OH 1 
TN 12 , TX 12 , VA 12 , WS fl , WV H ('Say, 
1824:305; 2 Hagen, 1861:239, as U. quadripunc- 
tata; Hagen, 1887:193, as C. excisus; Banks, 
1892:361; 5 Smith 1900:57; 6 Banks, 1901c:172; 

7 8 

Smith, 1925:170, as C. excisus; Montgomery 
& Trippel, 1933:260, as C. excisus; Brimley, 
1938:32; 10 Froeschner, 1947:129; "Throne, 
1972:121; 12 Shetlar, 1977:124-127; 13 Garland 
& Marshall, 1980:637; 14 Penny, NEW STATE 
RECORDS) 



Berothidae 
Berothidae, or beaded lacewings, are so called 
because of the development of secretion en- 
crusted setae on wings of females. Adults of most 
North American species are immediately recog- 
nizable by the falcate shape of the forewing. 
Females lay stalked eggs on wood surfaces, often 
near termite nests. Larvae are inquilines in dry- 
wood termite nests, where they feed by injecting 
a paralyzing chemical into the termite, and then 
sucking out the body fluids of the immobile prey. 



The only genus of berothids known from North 
American is Lomamyia Banks 1904. The species 
were reviewed and a key provided by Carpenter 
(1940). Several additional undescribed species 
are known from North America. 

Lomamyia Banks, 1904c:209 

Lomamyia banksi Carpenter, 1940:260. 

Holotype male: USA, South Carolina, Clem- 
son College, 30.IV. 1932, D. Duncan (MCZ, 
specimen not found). 

DISTRIBUTION. — USA: AL 1 , AZ 1 , DC ' , FL 1 , 
IL 1 IN 2 , KS 1 , LA 3 , MI 1 , MO 1 , MS 1 , NC 1 , NY 1 , 
SC, VA 1 , ('Carpenter, 1940; 2 Lawson & 
McCafferty, 1984:130; J Penny, NEW STATE 
RECORD). 

Lomamyia flavicornis (Walker), 1853:278 [He- 
merobius]. 

Lectotype (without abdomen): North America 
(BMNH). 

= Isoscelipteron pennsylvanicum Brauer, 
1 864:898. Type(s) sex(es) unknown: USA, Penn- 
sylvania (Vienna). 

= Lomamyia nearctica Navas, 19 13b: 19. 
Holotype male: USA, New York, Japhank [Long 
Island], 23.IX.191 1, De la Torre Bueno (MZB). 

Taxonomy. — To Micromus (see Hagen, 
1861:198); to Lomamyia (see Banks, 1905a:27); 
(L. nearctica = L. flavicornis; I. pennsylvanicum 
= L. flavicornis) (see Navas, 1929a:26); lecto- 
type designated (see Carpenter 1940:260). 

Distribution. — USA: AL 8 , DC 2 , FL , GA 1 , 
IN 9 KS 4 , KY 8 LA 8 , MS 10 , NC 7 , NJ 3 , NY 5 , 
PA , VA 6 ( f Walker, 1853:278; 2 Hagen, 
1861:199; 3 Smith, 1900:56; 4 Smith, 1925:167; 
5 Leonard, 1928:40; 6 Navas, 1929:26; 7 Brimley, 
1938:30; 8 Carpenter, 1940:260; 9 Lawson & 
McCafferty, 1984:130; 10 Penny, NEW STATE 
RECORD). 
Lomamyia fulva Carpenter, 1940:264. 

Holotype female: USA, California, Riverside 
Co., San Jacinto Mts., 21.VII.1929, R. H. 
Beamer (MCZ, specimen not found). 

Distribution. — USA: CA (Carpenter, 
1940:264). 

Lomamyia hamata (Walker), 1853:278 [He- 
merobius]. 

Holotype male: North America (BMNH). 

= Lomamyia hubbardi Banks, 1924:430. 
Type(s) sex(es) unknown: USA, Florida, Cedar 
Keys, 5.IV. 1878, Hubbard and Schwarz (MCZ). 



44 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



TAXONOMY. — To Micromus (see Hagen, 
1861:199); to Lomamyia (see Banks, 1905a:48); 
(H. hamatus = L. flavicornis) (see Navas, 
1929a:26); (L. hamatus not equal to L. flavi- 
cornis, L. hubbardi = L. hamata) (see Carpenter, 
1940:265). 

Distribution. — USA: FL 1 , SC 2 ('carpen- 
ter, 1940:265; 2 Brushwein, 1987a:671). 
Lomamyia latipennis Carpenter, 1940:262. 

Holotype male: USA, California, Marin Co., 
Phoenix Lake, 4.VII.1927, H. H. Kiefer (CAS). 

Distribution. — AZ, CA (Carpenter, 
1940:262). 

Lomamyia longicollis (Walker), 1853:281 [He- 
merobius]. 

Two syntype males: USA, Georgia, Abbot 
(BMNH). 

TAXONOMY. — To Lomamyia (see Caipenter, 
1940:267). 

Distribution. — GA 1 , MS 2 , NC 2 , SC 3 
('Walker, 1853:281; 2 Carpenter, 1940:267; 
3 Brushwein, 1987b: 151). 
Lomamyia occidentalis (Banks), 1905a:89 
[Berotha]. 

Lectotype female: USA, Nevada, Ormsby Co., 
6.VII, Baker (MCZ). 

TAXONOMY. — To Lomamyia (see Banks, 
1905c:48); considered a synonym of L. texana 
(see Banks, 1907c:23); considered a valid species 
(see Caipenter, 1940:263); lectotype designated 
(see Carpenter, 1940:263). 

Distribution. — CAN: BC 2 , USA: AZ 1 , 
CA 2 , NV 1 ('Banks, 1905a: 89; 2 Carpenter, 
1940:263). 
Lomamyia squamosa Carpenter, 1940:266. 

Holotype male: USA, Texas, Brownsville, 
1 1 16.VI. 1933, P. J. Darlington, Jr. (MCZ). 

Distribution. — USA: TX (Carpenter, 
1940:266). 
Lomamyia tenuis Carpenter, 1940:261. 

Holotype male: USA, New Mexico, Torrance 
Co., VII. 1925, C. H. Martin (MCZ). 

Distribution. — USA: NM (Carpenter, 
1940:262). 
Lomamyia texana (Banks), 1897:24 [Berotha]. 

Lectotype male: USA, Texas (MCZ). 

TAXONOMY. — To Lomamyia (see Banks, 
1905a:27); lectotype designated (see Carpenter, 
1940:265). 

Distribution. — USA: AZ 2 , TX 1 , UT 2 
('Banks, 1897:24; 2 Carpenter, 1940:265). 



Chrysopidae 
Most chrysopid adults are green with golden 
eyes, but members of the genera Nothochrysa, 
Pimachiysa, Eremochrysa, and Yumachiysa are 
usually predominately brown or black. Most spe- 
cies have characteristic red or dark markings, 
which often fade within a few months after death. 
Eggs are laid on long stalks, usually singly. Lar- 
vae are voracious predators, those of some spe- 
cies carrying packets of trash or corpses of their 
prey on their backs (Smith 1922). Most species 
will feed upon a wide variety of slow-moving 
insects, although some have more specific food 
requirements. Larvae of Belonopterygini 
(Abachrysa and Nacarina in our region) live in 
ant nests, presumably feeding upon the ants. 
Adult chrysopids are very conservative morpho- 
logically, with many of the most taxonomically 
significant characteristics found in the male ter- 
minalic structures. Thus, to determine the generic 
placement of an unfamiliar male specimen, the 
abdomen must be cleared (in 10% KOH), and 
subsequently may be stained with chlorazol 
black-E. A key to subfamilies and genera of 
Nothochrysinae was provided by Adams (1967). 
A worldwide generic revision of the family con- 
tains helpful descriptions and illustrations of 
wings and genitalic structures, as well as a key to 
genera (Brooks and Barnard 1990). The species 
of Chrysopidae of Canada were included in a key 
by Garland (1985). All other literature is too old 
to be of much help with identification. There are 
14 genera of Chrysopidae known from North 
America; several new species are in collections, 
awaiting description. 

Nothochrysinae 

Nothochrysa McLachlan, 1868:195 

Nothochrysa californica Banks, 1892:373. 

Holotype male: USA, California, Los Angeles 
(MCZ). 

Distribution. — CAN: BC", USA: CA 1 , 
OR 3 , WA 3 ("Banks, 1892:373; 2 Smith, 
1932:582; 3 Adams, 1967:224). 

Pimachrysa Adams, 1957a:67 
Pimachrysa albicostales Adams, 1967:226. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



45 



Holotype male: Mexico, Baja California, 22 
miles north of Punta Prieta, 9.XII.1958, H. B. 
Leech (CAS). 

Distribution. — USA: AZ 1 , CA" ('Adams, 
1967:227; 2 Penny, NEW STATE RECORD) 
Pimachrysa fusca Adams, 1967:227. 

Holotype male: USA, California, Riverside 
Co.. L. Covington Flat, Joshua Tree National 
Monument. 19.111.1961, E. L. Sleeper (CAS). 

Distribution. — USA: CA (Adams, 
1967:227). 
Pimachrysa grata Adams, 1957a:68. 

Holotype female: USA, Arizona, Santa Rita 
Mountains, Madera Canyon, 26. VIII. 1 949, P. A. 
Adams (MCZ). 

Distribution. — USA: AZ (Adams, 
1957a:67). 
Pimachrysa intermedia Adams, 1967:228. 

Holotype female: USA, California, Riverside 
Co., Whitewater. Snow Creek, 1500 ft., 
8. III. 1955, W. R. M. Mason (CNC). 

Distribution. — USA: CA (Adams, 
1967:228). 
Pimachrysa nigra Adams, 1967:228. 

Holotype male: USA, California, Riverside 
Co.. Gavilan, 19.111. 1936, P. Timberlake (CAS). 

Distribution. — USA: CA (Adams, 
1967:228). 

Chrysopinae 
Belonopterygini 

Abachrysa Banks, 1938a:75 

Abachrysa eureka (Banks), 1931:174 
[Chrysopa]. 

Holotype (sex unknown): USA, Arkansas, 
Hope, Knobel (MCZ). 

Taxonomy. — To Abachrysa (see Banks, 
1938a:75). 

Distribution. — USA: AR 1 , FL 5 GA 3 , 
MS 2 , TX 4 , ('Banks, 1931:174; "Banks, 
1938a:75; 3 Bickley & MacLeod, 1956: 197; 4 Ag- 
new et al., 1981:4; 5 Brooks & Barnard, 
1990:165). 

Nacarina Navas, 19 15c: 133 

Nacarina robusta (Banks), 1905b:5 
[Chrysopa]. 



Type(s) (sex unknown): USA, North Carolina, 
Tyron (MCZ). 

TAXONOMY. — To Nacarina by Adams 
(NEW COMBINATION). 

DISTRIBUTION. — USA: FL 2 , NC 1 ('Banks 
1905b:5; 2 Stange, NEW STATE RECORD). 

LEUCOCHRYSINI 

Leucochrysa (Leucochrysa) McLachlan, 
1868:208 

Leucochrysa (Leucochrysa) ampla (Walker), 
1853:268 [Chrysopa]. 

Lectotype female: no locality data (BMNH). 

Taxonomy. — Listed as possible synonym of 
C. lineaticornis (see Banks 1907c:27); to Leuco- 
chrysa (Leucochrysa) as valid species (see 
Brooks & Barnard, 1990:276); lectotype desig- 
nated (see Kimmins, 1940:444). 

Distribution. — USA: GA (Walker, 
1853:268). 

Leucochrysa (Leucochrysa) arizonica 
(Banks), 1906a:98 [Allochrysa]. 

Holotype male: USA, Arizona, Palmerlee, VII 
(MCZ). 

TAXONOMY. — To Leucochrysa (see Adams, 
1977:95). 

Distribution. — USA: AZ (Banks, 
1906a:98). 

Leucochrysa (Leucochrysa) insularis 
(Walker), 1853:269 [Chiysopa]. 

Holotype male: Jamaica (BMNH). 

= Chrysopa virginica Fitch, 1855:795. Holo- 
type (sex unknown): USA, Virginian Carters- 
ville, T. A. Culbertson (type probably lost). 

= Nothochrysa phantasma MacGillivray, 
1894:170. Lectotype male: USA, Massachusetts, 
West Chop, 8.VIII. 1 893 (MCZ). 

= Leucochrysa cerverai Navas, 1924a:325. 
Type(s) (sex unknown): Cuba, Habana, Santiago 
de las Vegas, 18.IX.1923, F. Z. Cervera (MCZ). 

= Leucochrysa joannisi Navas, 1925d: 13. No 
type specimen designated: Cuba, Habana, Santi- 
ago de las Vegas, 17. VII and 20.VIII.1924, F. 
Cervera (no types located). 

= Allochrysa virginica ocala Banks, 
1938b: 122. Holotype female: USA, Florida, Jef- 
ferson Co., Lloyd Sink, 9.VIII. 1935, G. Fairchild 
(MCZ). 



46 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Taxonomy. — To Protochrysopa (see 
Kolbe, 1888:74); to Leucochrysa (see Adams, 
1977:97); N. phantasma = C. virginica (see 
Banks, \%95b:?>\5)\ L.joannisi = L. insularis (see 
Alayo, 1968:57); L. virginica = L. insularis, L. 
cerverai = L. insularis, L. virginica ocala = L. 
insularis (see Adams, 1977:97). It appears that 
no formal description of L. joannisi was ever 
published; referred to by Alayo, 1968:57. 

DISTRIBUTION. — USA: AL 8 , AR 8 , DC 7 , FL 4 , 
IA 9 , MA 2 , MO 8 , MS 8 , NC 5 , NJ 3 , TN 6 , VA 1 , 
WV 8 ("Fitch, 1855:795, 2 MacGillivry, 
1894:170, as Nothochtrsa phantasma; Smith, 
1900:55; 4 Banks, 1938b: 122; 5 Brimley, 
1938:30; 6 Bickley, 1941:187; 7 Bickley & 
MacLeod, 1956:184; 8 Adams, 1977:98; 9 Penny, 
NEW STATE RECORD). 

Leucochrysa (Nodita) Navas, 19 16a:2 1 

Leucochrysa (Nodita) americana Banks, 
1897b:175. 

Holotype female: USA, Alabama, Auburn, 
1938 (MCZ). 

TAXONOMY. — To Nodita (see Navas, 
1917b:280); to Leucochrysa (Nodita) (see 
Brooks & Barnard, 1990:277). 

Distribution. — USA: AL 1 , KS 3 , TX 2 
('Banks, 1897b: 175; 2 Banks, 1907c:26; 3 Bick- 
ley& MacLeod, 1956:188). 
Leucochrysa (Nodita) antennata Banks, 
1905b:5. 

Type (sex unknown): Mexico, Tuxpan (MCZ). 

Taxonomy. — To Nodita (see Banks, 
1939:2); to Leucochrysa (Nodita) (see Brooks & 
Barnard, 1990:277).' 

NOTE. — A male specimen in MCZ from 
Texas, Brownsville, 13.VIII.1909, H. S. Barber 
is erroneously labelled as type. 

DISTRIBUTION. — USA: TX (Banks, 
1907c:26). 

Leucochrysa (Nodita) callota Banks, 
1915b:626. 

Holotype female: USA, Texas, Austin, 
3.V.1901 (MCZ). 

Taxonomy. — To Nodita (see Banks, 
1939:2); returned to Leucochrysa (Leucochrysa) 
(see Brooks & Barnard, 1990:276); to Leuco- 
chnsa (Nodita) by Adams, NEW COMBINA- 
TION 



DISTRIBUTION. — USA: CA 2 , FL 2 , LA 2 , TX 1 
('Banks, 1915b:626; 2 Adams, NEW STATE 
RECORDS). 

Leucochrysa (Nodita) explorata (Hagen), 
1861:217 [Chrysopa]. 

Type(s) (sex unknown): Mexico, Cordova, 
Saussure (MCZ, specimen not found). 

TAXONOMY. — To Leucochrysa (Nodita) (see 
Brooks and Barnard, 1990:277). 

DISTRIBUTION. — USA: NM (Hagen, 
1875:921). 

Leucochrysa (Nodita) floridana Banks, 
1897c: 184. 

Holotype female: USA, Florida, Lake Worth 
(MCZ). 

Taxonomy. — To Nodita (see Navas, 
1917b:280); to Leucochysa (Nodita) (see 
Brooks & Barnard, 1990:277). 

DISTRIBUTION. — USA: FL 1 , MS 2 , NC 3 , TX 4 
('Banks, 1897c:184; 2 Banks, 1903a: 144; ^rim- 
ley, 1938:30; 4 Agnewetal., 1981:14). 
Leucochrysa (Nodita) nigrinervis (Banks), 
1939:1 [Nodita]. 

Type(s) (sex unknown): USA, New Mexico, 
McKinley Co., Satan Pass (MCZ, specimen not 
found). 

TAXONOMY. — To Leucochysa (Nodita) (see 
Brooks & Barnard, 1990:277).' 

DISTRIBUTION. — USA: AZ 3 , CA 3 , CO 3 , 
NM 1 , TX 2 ('Banks, 1939:1; 2 Bickley & 
MacLeod, 1956:188; 3 Adams, NEW STATE 
RECORDS). 

Leucochrysa (Nodita) pavida (Hagen), 
1861:216 [Chrysopa]. 

Syntype series (sex unknown): Mexico, Cor- 
dova, Saussure, Deppe (NHB); USA, South 
Carolina, Zimmerman (female) (MCZ). 

Taxonomy. — To Nodita (see Banks, 
1939:2); to Leucochysa (Nodita) (see Brooks & 
Barnard, 1990:277).' 

DISTRIBUTION. — USA: FL 2 , IN 4 , NC 2 , SC 1 , 
TX 3 ('Hagen, 1861:216; 2 Bickley & MacLeod, 
1956:188; 3 Agnew et al., 1981:14; 4 Lawson & 
McCafferty, 1984:130). 

Leucochrysa (Nodita) rufina (Banks), 1950:54 
[Eremochrysa]. 

Holotype male: USA, Arizona, Coconino Co., 
Grand Canyon, 24.VII.1934 (MCZ). 

TAXONOMY. — To Leucochysa (Nodita) by 
Adams, NEW COMBINATION 

Distribution. — USA: AZ (Banks, 
1950:55). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



47 



Leucochrysa (Nodita) texana (Banks), 1939:3 
[Nodita]. 

Holotype male: USA, Texas, Travis Co. 
(MCZ). 

TAXONOMY. — To Leucochrysa (Nodita) (see 
Brooks & Barnard, 1990:278). 

Distribution. — USA: TX (Banks, 1939:3). 

Chrysopini 
Ceraeochrysa Adams, 1982a:70 

Ceraeochrysa cincta (Schneider), 1851:86 
[Chrysopa]. 

Syntype series (sex unknown): Brazil, RJ, Rio 
de Janeiro (HMB and IRSNB). 

= Chrysopa bUineata Navas, 1914e:91. Holo- 
type female: Guatemala, Vulcan de Attilan, 
2500-3500 ft., Champion (BMNH). 

= Chrysopa lafonei Navas, 1914a:222. Holo- 
type male: Argentina (MLP - Bruch collection). 

= Chrysopa bnichi Navas, 1914a:222. Lecto- 
type male: Argentina, Huasan, 25.11.1912 
(MACN) (Adams, NEW SYNONYMY). 

= Chrysopa incalis Banks, 1915b:627. Syn- 
types (sex unknown): Peru, Matucana, 7780 ft., 
14. VI, Parish (male); Chosica, 2800 ft., 10.VI, 
Parish (MCZ). 

= Chrysopa bicarnea Banks, 1920:338. Holo- 
type male: USA, Florida, Miami, G. F. Moznette 
(MCZ). 

= Chrysopa advena Navas, 1922a:51. Holo- 
type male: Coast of Uruguay, 1920, Renato Mar- 
tin (MNHN). 

= Chrysopa habana Navas, 1922a:52. Holo- 
type (sex unknown): Cuba, Havana, Fermin Z. 
Cervera (MZB). 

= Chrysopa bessona Navas, 1924d:361. Holo- 
type female: Argentina, La Plata, 20.XI.1920, 
Bruch (MLP). 

= Chrysopa mestizo Navas, 1924a:330. Lecto- 
type female: Cuba, Santiago de las Vegas, 
IX. 1923; Havana, Rio Almendares, 19.VII.1923, 
Fermin Z. Cervera (MCZ). NEW LECTO- 
TYPE DESIGNATION. 

= Chrysopa villosula Navas, 1924b:337. Holo- 
type (missing abdomen): Cuba, Havana, Santi- 
ago de las Vegas, 4.XII.1923 (MCZ). 



= Cintameva bina Navas, 1924a:331. Holo- 
type male: Cuba, Havana, Rio Almendares, 
9.VIII. 1923 (MCZ). 

= Chrysopa cornuta Navas, 1925a:65. Holo- 
type female: Brazil, RJ, Niteroi, 21. XL 1924 
(MNHN). 

= Chrysopa alternans Navas, 1933a:306. 
Holotype male: Peru, Lima, 4.X.1933 (HMB). 

= Chrysopa wollebaeki Esben-Petersen, 
1934:291. Seven syntypes (six males, one fe- 
male): Ecuador, Galapagos Islands, Santa Maria, 
Post Office Bay, 1-10.X.1925, one male; 
29.X. 1925, four males, one female; 48.XI.1925, 
one male, all collected by Wollebaek (ZMUO). 

= Chrysopa iona Banks, 1944:12. Holotype 
male: Surinam, Paramaribo. 7.XII.1938, Gei- 
jskes (MCZ). 

= Chrysopodes sal lei Banks, 1946:171. Holo- 
type male: Mexico, Salle (MCZ). 

TAXONOMY. — C. mestiza = C habana (see 
Alayo, 1968:30); C. advena, C alternans, C 
bessona, C. bicarnea, C. bilineata, C. bina, C 
caligata, C. cornuta, C. habana, C. incalis, C 
iona, C lafonei, C mestiza, Chrysopodes sallei, 
C villosula, C wollebaeki = C cincta (Adams, 
1982:72); C. caligata not = C cincta (Adams & 
Penny, 1987:442). 

Distribution. — USA: FL (Banks, 
1920:338). 

Ceraeochrysa cubana (Hagen), 1861:215 
[Chrysopa], 

Neotype male: Cuba, Soledad, near Cienfue- 
gos, 6-20. VIII, N. Banks (MCZ). 

= Chrysopa tolteca Banks, 1901a;364. Holo- 
type (female): Mexico, Oaxaca, Tomellin, VI 
(MCZ). 

= Chrysopa venularis Navas, 1913b:20. Holo- 
type female: Jamaica (ZSBS). 

= Chrysopa albatala Banks, 19 13a: 139. Holo- 
type (abdomen missing): Guyana, Bartica, XII, 
Parish (MCZ). 

= Chrysopa imbecilla Navas, 1914e: 107. 
Holotype male: Barbados (BMNH). 

= Chrysopa epheba Navas, 1924a:328. Holo- 
type (sex unknown): Cuba, Calabazas, 
28.VIII. 1923 (MZB). 

= Chrysopa seminole Banks, 1924:432. Holo- 
type female: USA, Florida, Marco, 24. VII, Mott 
(MCZ). 

= Chrysopa freemani Smith, 1931:81 1. Holo- 
type female: Haiti, Port-au-Prince, 500 ft., 
16.XI. 1929 (MCZ). 



48 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



= Chrysopafischerina Navas, 1 933 : 1 96. Holo- 
type female: Brazil, Central. Mang., 13.X.1913, 
R. Fischer (DEIC) (Adams, NEW SYNON- 
YMY). 

= Chtysopa damiensis var. jamaicensis Banks, 
1941a:394. 

Syntype series (male, female): Jamaica, King- 
ston, 27— 29.VIII, P. J. Darlington; Trelawny, 
Perkins (MCZ). 

TAXONOMY. — To Ceraeochrysa (see 
Adams, 1982:72); C. epheba = C cubana, C. 
jaeobaea = C. cubana (see Alayo, 1968:28); 
Adams, 1982:72 (C. jaeobaea = C giindlachi, C. 
tolteca = C. cubana, C. albatala = C. cubana, C. 
venularis = C. cubana, C. imbecilla = C. cubana, 
C. epheba = C. cubana, C. seminole = C. cubana, 
C.jreemani=C. cubana, C damiensis jamaicen- 
sis = C. cubana, C. scapularis = C. cubana (see 
Adams, 1982:72); C. scapularis not= C. cubana 
(see Adams & Penny, 1987:448); neotype desig- 
nated (see Adams & Penny, 1987:446). 

Distribution. — USA: FL 2 , NC 3 , VA 1 
('Hagen, 1861:215; 2 Banks, 1924:432, as C. 
seminole; 3 Banks, 1938b: 121). 
Ceraeochrysa lineaticornis (Fitch), 1855:795 
[Chtysopa]. 

Holotype (sex unknown): USA, New York 
(MCZ). 

= Chrysopa puncticornis Fitch, 1855:796. 
Type(s) (sex unknown): USA, New York (MCZ, 
specimen not found). 

= Allochysa parvula Banks, 1903a: 143. Holo- 
type (male): USA, Florida, Runnymede (MCZ). 

= Chrysopa columbiana Banks, 1903a: 150. 
Holotype male: USA, D. C, Washington 
(MCZ). 

= Chrysopa stichoptera Navas, 19 14c:61. 
Holotype (sex unknown): USA, New York, Long 
Island, Yaphank, 1 0. VII. 1 9 1 3, J. R. de la Torre- 
Bueno (MZB). 

TAXONOMY. — To Ceraeochrysa (see 
Adams, 1982:73) C. puncticornis = C. lineati- 
cornis (see Banks, 1907c:27); C. stichoptera = 
C. lineaticornis (see Bickley & MacLeod, 
1956: 190); C. columbiana = C. lineaticornis (see 
Br am & Bickley, 1963: J 6); A. parvula- C. linea- 
ticornis (see Adams, 1982:73). 

DISTRIBUTION. — CAN: BC 3 PQ 3 USA: 
DC 2 , FL 2 GA 10 , IN 9 , KS 10 , MA" MD" ME 10 , 
MI 2 NC 4 , NH 2 , NY 1 , PA 10 , TN 5 , TX 8 , VA 6 , 
WS 7 ('Fitch, 1855:795; 2 Banks, 1903a:143, 150, 
151 as A. parvula, C lineaticornis and C. colum- 



biana; Smith, 1932:585,as C. columbiana and 
C. lineaticornis; Brimley, 1938:30; Bickley, 
1941:197; 6 Bickley & MacLeod, 1956:191; 
7 Throne, 1971a:74; 8 Agnew et al., 1981:12; 
9 Lawson & McCafferty, 1984:1 30; 10 Adams, 
NEW STATE RECORDS) 
Ceraeochrysa placita (Banks), 1908a:259 
[Chtysopa]. 

Male syntypes: USA, Colorado, Chimney 
Gulch, 20.1.1907; Clear Creek (MCZ). 

= Chrysopa intacta Navas, 19 12a: 199. Holo- 
type (sex unknown): Canada, Quebec, Toronto, 
1. VI. 1908 (MZB). 

= Chrysopa forreri Navas, 1914e:97. Type(s) 
(sex unknown): Mexico, Mexico City, 8100 ft., 
Forrer (BMNH). 

TAXONOMY. — To Ceraeochn'sa (see 
Adams, 1982:73);C. forreri = C. placita (see 
Adams, 1982:73); C intacta = C placita (see 
Brooks & Barnard, 1990:269). 

Distribution. — CAN: PQ 2 , USA: CA 4 , 
CO 1 , MO 3 , NH 4 , OR 4 ('Banks, 1908a:259; 
2 Navas, 1912a:200; 3 Froeschner, 1947:134, as 
C. intacta; 4 Adams, NEW STATE RE- 
CORDS) 

Ceraeochrysa smithi (Navas), 19 14e: 105 
[Chrysopa]. 

Two male syntypes: Grenadines, Union Island, 
W. J. H. H. Smith; West Indies, Mustiques 
(BMNH). 

= Chtysopa poeyi Navas, 1924a:330. Holotype 
(male): Cuba, Havana, Rio Almendares, 
20.VII.1923, Cervera (MCZ). 

= Chtysopa neotropica Navas, 1929b:3 17. 
Syntype series (sex unknown): Honduras, Ama- 
pala, 9.VII.1908, R. Paessler; El Salvador, La 
Union, 3. XII. 1 907, R. Paessler; Mexico, Jalisco, 
La Garita, 1912, W. Fritsche; Colombia, Behn 
verid., 3.XII.1900 (Hamburg, specimen lost). 

TAXONOMY. — To Ceraeochrysa (see 
Adams, 1982:73); C. neotropica = C. smithi, C 
poeyi = C smithi (see Adams, 1982:73). 

Distribution. — USA: FL (Adams, NEW 
NATIONAL RECORD) 

Remarks. — This species has been men- 
tioned before from Florida by Banks under the 
names C. claveri and C. cubana sanchezi. Sev- 
eral specimens have been identified by other 
chrysopid specialists, but this distribution has not 
been reported in the taxonomic literature. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



49 



Ceraeochrysa valida (Banks), 1895:517 
[Chrysopa]. 

Two syntypes (sex unknown): Mexico, Baja 
California Sur, Serra El Taste, IX; San Jose del 
Cabo, IX (MCZ). 

= Chrysopa bimaculata McClendon, 
1901:215. Lectotype female: USA, Texas, 
Laredo, VIII. 1900 (MCZ) NEW LECTOTYPE 
DESIGNATION 

= Chiysopa limitata Navas, 1913c:84. Holo- 
type (sex unknown): Brazil, Rio de Janeiro, Nova 
Friburgo, XII. 19 1 1 , Ramos (MZB). 

= Chrysopa longicella Navas, 19 14e:97. Holo- 
type male: Guatemala, Zapote, G. C. Champion 
(BMNH). 

= Chrysopa breviata Banks, 19 15b: 628. One 
male syntype, one syntype of unknown sex. 
Ecuador, Guayaquil, 8. VI, Parish; Quevedo 
(MCZ). 

= Chiysopa lioni Navas, 1927a:54. Holotype 
female: Haiti, Port-au-Prince, 1910, G Lion 
(MNHN). 

= Clvysopa damiensis Smith, 193 1 :803. Holo- 
type male: Haiti, Hatte Lathan, 18.1.1929, R. C. 
Smith (MCZ). 

= Chiysopa wolcotti Smith, 1931:810. Holo- 
type male: Haiti, Damien (MCZ). 

TAXONOMY. — To Ceraeochrysa (see 
Adams, 1982:73); C bimaculata = C. tolteca, C 
longicella = C tolteca, C tolteca (?) = C. valida 
(see Banks, 1946:148); C. bimaculata = C. 
valida, C limitata = C. valida, C longicella = C 
valida, C. breviata = C. valida, C lioni = C 
valida, C. damiensis = C valida, C. wolcotti - C 
valida, C camposana = C valida, C. seminole = 
C. valida, C. tolteca = C cubana (see Adams, 
1982:73); C. seminole = C. cubana; C cam- 
posana = Clvysopa incertae sedis (see Brooks & 
Barnard, 1990:268,279). 

Distribution. — USA: FL 2 , TX 1 ('McClen- 
don, 1901:215, as C. bimaculata; "Banks, 
1903a:153). 

Chrysopa Leach in Brewster, 1815:138 

Chrysopa chi Fitch, 1855:791. 

Holotype female: USA, New York (MCZ). = 
Chrysopa upsilon Fitch, 1855:791. Holotype fe- 
male: no label data (MCZ). 

= Chiysopa upsilon var. haematica Navas, 
1918a:354. (as Chrysopa hypsilon [sic]) Holo- 



type (sex unknown): USA, New York, White 
Plains. 3 1 V. 1 9 1 5, de la Torre Bueno (MZB). 

Taxonomy. — C. upsilon = C. chi (see Smith, 
1922:1352); C. hypsilon var. haematica - C. chi 
(see Smith, 1932:592). 

Distribution. — CAN: AB 5 ^ BC 5 MB 5 , 
NS 5 ON 5 PQ 5 , SK 5 , USA: DC", MD . ME 7 
MN 8 , NJ 3 , NH 4 , NY 1 , TN 6 , WA 11 , WS 10 
('Fitch, 1855.791; 2 Hagen, 1861:214; 3 Smith, 
1900:55; 4 Banks, 1903a: 148; 5 Smith, 1932:592; 
6 Bickley, 1941:188; 7 Procter, 1946:43; 8 Parfin, 
1952:424; 9 Bram & Bickley, 1963:8; 10 Throne, 
1971a:70; ' 'Penny, NEW STATE RECORD). 
Chrysopa coloradensis Banks, 1895b:314. 

Holotype male: USA, Colorado, Fort Collins 
(MCZ). 

Distribution. — CAN: BC 3 , USA: AZ 2 , 
C A 2 , CO ' , ID 3 ^ NM 2 , OR 2 , UT 4 , WA 2 (' Banks, 
1895b:315; "Banks, 1903a:151; 3 Smith, 
1932:585; 4 Bickley & MacLeod, 1956:190). 
Chrysopa excepta Banks, 1911:340. 

Holotype (sex unknown): USA, New Mexico, 
Fort Wingate, 25.VII, Woodgate (MCZ). 

DISTRIBUTION. — CAN: AB 2 , BC", USA: 
NM 1 , UT 3 WY 3 ('Banks, 1911:340; 2 Smith, 
1932:585; 3 Bickley & MacLeod, 1956:190). 
Chrysopa incompleta Banks, 1911:340. 

Syntypes (sex unknown): USA: North Caro- 
lina, Beaufort, 15.V.1906, Woglum (male); 
Raleigh, 15.VII.1902, Sherman (MCZ). 

DISTRIBUTION. — USA: FL 6 , GA 2 IN 5 , MA 2 . 
MD 2 , NC 1 , NJ 2 , TX 4 , VA 2 , WS 3 ('Banks, 



i 



1911:341; "Bram & Bickley, 1963:7; "Throne, 

1971a:72; 4 Agnew et al,, 1981:12; 5 Lawson & 

McCafferty, 1984:130; 6 Adams, NEW STATE 

RECORD). 

Chrysopa nigricornis Burmeister, 1839:980. 

Holotype male: North America (Berlin). 

= Chrysopa colon Fitch, 1855:792. Type (sex 
unknown): (MCZ). 

= Chiysopa eiythrocephala Banks, 1898:201 
not C. eiythrocephala (Rambur) 1842:428 [He- 
merobius]. Holotype female: USA, California, 
San Francisco, VII. 1897, Morse (MCZ). 

= Chrysopa majuscula Banks, 1906a:98. Re- 
placement name for Chrysopa erythrocephala 
Banks. 

= Chiysopa vegeta Navas, 1917:6. Type(s) 
(sex unknown): USA, New Mexico, Jemez 
Springs, 2 l.VI. 19 16, John Woodgate (MZB). 



50 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



= Chn'sopa crotchi Banks, 193 8a: 76. Holo- 
type (female): Canada, British Columbia, Van- 
couver Island, Victoria, VII, Crotch (MCZ). 

Taxonomy. — C. colon = C nigricornis (see 
Hagen, 1875:920); C. majuscula, new name for 
C. erythrocephala Banks (see Banks, 1906a:98); 
C. majuscula = C. nigricornis (see Bram & Bick- 
ley, 1963:10); C. crotchi = C. nigricornis; C. 
vegeta = C. nigricornis (Adams, NEW SYNON- 
YMY). 

Distribution. — CAN: AB 5 , BC 5 , ON 5 
PQ 4 USA: CA 3 , CO 2 , DC 4 , IN 6 , KS 4 , MA 4 
MD, ME 9 , MN 11 , MO 10 , NC 7 , NM 2 , NY'l 
OH 4 , RI 4 TN 8 , TX 14 , VA 4 , WS 13 ('Fitch 
1855:792; "Hagen, 1875:920; 3 Banks, 1898:201 
4 Banks, 1903a: 150; 5 Smith, 1932:584 
6 Montgomery & Trippel, 1933:260; Brimley 
1938:30; 8 Bickley, 1941:188- 9 Procter, 1946:43 
10 Froeschner, 1947:133; "Parfin, 1952:424 
12 Bram&Bickley, 1963:9; 13 Throne, 1971a:69 
14 Agnewetal., 1981:12). 
Chrysopa oculata Say, 1839:45. 

Type(s) (sex unknown): USA (depository un- 
known). 

= Chrysopa chlorophana Burmeister, 
1839:979. Type(s) (sex unknown): North Amer- 
ica (depository unknown). 

= Chrysopa euryptera Burmeister, 1839:980. 
Type(s) (sex unknown): North America (?) 
(Halle). 

= Chrysopa latipennis Schneider, 1851:118. 
Holotype (sex unknown): North America (HMB 
or Frankfurt). 

= Chrysopa albicornis Fitch, 1855:788. Holo- 
type (sex unknown): USA, Mississippi (MCZ). 

= Chn'sopa illepida Fitch, 1 855:788. Holotype 
(female): USA, New York; Illinois (MCZ). 

= Chn-sopa omikron Fitch, 1855:789. Holo- 
type (female): USA, New York (MCZ). 

= Chrysopa xanthoccphala Fitch, 1855:789. 
Holotype (sex unknown): USA, New York; 
Michigan (MCZ). 

= Chrysopa fulvihucca Fitch, 1855:790. 
Type(s) (sex unknown): USA, New York (Type 
not found). 

= Chrysopa mississippiensis Fitch, 1855:790. 
Holotype female: USA, Mississippi, Jackson, 
IV, Fitch (MCZ). 

= Chrysopa hipunctata Fitch, 1855:791. Holo- 
type female: USA, New York (MCZ). 

= Chrysopa transmarina Hagen, 1861:213. 
Syntypes (sex unknown): Canada, Ontario, La 



Chine, near Montreal, Barnston; Trenton 
Falls,New York; Newfoundland, St. John; Nova 
Scotia (Types should be in BMNH). 

= Nothochrysa annulata MacGillivray, 
1894:169. Holotype female: USA, Massachu- 
setts, Wellesley, 22.VI.1892 (CORN). 

= Chrysopa assimilis Banks, 1898:202. One 
male syntype, one syntype of unknown sex: 
USA, Oregon, Ashland (male) and Hood River 
(MCZ). 

= Chrysopa mexicana Banks, 1901:364. Six 
syntypes (sex unknown): Mexico, Veracruz, 
Thalpan, VII (MCZ) Adams, NEW SYNON- 
YMY. 

= Chn'sopa separata Banks, 1911:341. Syn- 
types (sex unknown): USA, Colorado, Golden, 
Chimney Gulch, Oslar; New Mexico, Pecos, 
23.VI, Cockerell (MCZ). 

= Chrysopa rubicunda Navas, 1913b:20. 
Type(s) (sex unknown): USA, New York, Long 
Island, Yapbank, 23.IX.1911, Torre Bueno 
(MZB). 

= Cintameva conspersa Navas, 1929:318. 
Adams, NEW SYNONYMY. Holotype male: 
Colombia (?), Behn vend., 3.XII.1900 (Ham- 
burg, specimen destroyed). 

TAXONOMY. — C. euryptera = C oculata, C 
illepida = C oculata, C omikron = C oculata, 
C. fulvihucca = C oculata, C mississippiensis = 
C oculata, C. latipennis = C. chlorophana, C 
xanthocephala = C chlorophana, C hipunctata 
= C chlorophana, C transmarina = C. chloro- 
phana (see Banks, 1903a: 161); C. chlorophana 
and earlier synonymies = C oculata, C albi- 
cornis = C oculata, Nothochrysa annulata (?) 
= C. oculata, C separata = C oculata, C ru- 
bicunda = C oculata (see Bickley & MacLeod, 
1956: 19 1); C. assimilis = C oculata, C. separata 
is valid species, C. chlorophana is valid species 
with associated earlier synonymies (see Brooks 
& Barnard, 1990:270). We consider that the syn- 
onymy of C chlorophana and C. oculata by 
Bickley and MacLeod is a correct decision. 

Distribution. — CAN: AB 8 , BC 8 , MB 8 , 
NF 6 , NS 6 ON 8 PQ 6 , SK 8 , USA: AZ 6 , CO 6 , 
DC 2 , GA 2 IA 16 , IL 2 , IN 9 , KS 7 , LA 2 MD 2 , 
ME 11 , MI 3 , MN 13 , MO 12 MS 2 , NC 10 , NJ^, 
NM 6 NY 1 , OH 6 , OR 4 , PA", TN 2 , TX 15 , VA 2 , 
WS 14 ('Fitch, 1855:788; 2 Hagen, 1861:212; 
3 Banks, 1 892:358; 4 Banks, 1 898:202, as C. chlo- 
rophana; 5 Smith, 1900:55; 6 Banks, 1903a: 148, 
as C. chlorophana; 7 Smith, 1925:168; 8 Smith, 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



51 



1932:587; ^Montgomery & Trippel, 1933:260 
lu Brimley, 1938:30; "Procter, 1946:43 
12 Froeschner, 1947:133; 13 Parfin, 1952:424 
l4 Throne, 1971a:69; ,5 Agnew et al., 1981:12 
16 Penny, NEW STATE RECORD) 

REMARKS. — Carpenter (1992:538) has indi- 
cated that Burmeister's Handbuch der Entolo- 
mologie was apparently published in two parts. 
The first 400 pages were published in 1839, 
while the remainder, including all Neuroptera, 
were published in 1838. If this is found to be 
correct, it could lead to serious nomenclatorial 
instability. The synonymy of Chrysopa oculata 
and Chiysopa chlorophana would be reversed. 
Almost certainly the International Commission 
on Zoological Nomenclature would be petitioned 
to declare, for nomenclatorial purposes, that the 
entire book was published in 1839. 
Chrysopa pleuralis Banks, 191 1:341. 

One male syntype, one syntype of unknown 
sex: USA, Colorado, Boulder Co., North Boulder 
Creek, 21.VIII.1907, S. A. Rohwer; Steamboat 
Springs, 27.V, Cockerell (male) {MCZ). 

DISTRIBUTION. — CAN: AB", BC", USA: 
CO 1 , MT 3 , UT 3 , WY 3 ('Banks, 1911:342; 
2 Smith, 1932:591; 3 Adams, NEW STATE RE- 
CORDS) 

Chrysopa quadripunctata Burmeister, 
1839:980. 

Type(s) (sex unknown): North America 
(Halle). 

= Chiysopa sulphured Fitch, 1855:793. 
Type(s) (sex unknown): USA, New Jersey, IX 
(MCZ, specimen not found). 

= Chrysopa sichelii Fitch, 1855:793. Holotype 
male: USA, New York (MCZ). 

TAXONOMY. — C sulphurea = C. quadri- 
punctata, C sichelii = C. quadripunctata (see 
Banks, 1903a: 162). 



CAN: BC 



USA: DC" 

10 



Distribution. 
IL 6 , IN 7 , KS 5 .^MD 12 , MN", MO iu , NC 8 , NJ"\ 
NY 1 , PA 2 , SC 2 , TN 9 , TX 4 , VA 4 , WS 13 ('Fitch, 
1855:793; 2 Banks, 1892:359; 3 Smith, 1900:55; 
4 Banks, 1903a: 153; 5 Smith, 1925:168; 6 Smith, 
1932:597; Montgomery & Trippel, 1933:260; 
8 Brimley, 1938:30; 9 Bickley, 1941:189; 
10 Froeschner, 1947:134; "Parfin, 1952:425; 
l2 Bram & Bickley, 1963:8; 13 Throne, 
1971a:70). 
Chrysopa slossonae Banks, 1924:432. 

Syntypes females: USA, North Carolina, Hen- 
dersonville, F. Sherman; New Hampshire, Fran- 



conia, Mrs. A. T. Slosson; Virginia, Great Falls 
(MCZ). 

Distribution. — USA: NC ' , NH ' , NY 2 VA ' 
('Banks, 1924:432; 2 Eisner et al., 1978:790). 



Chrysoperla Steinmann. 1964:260 

Chrysoperla adamsi Henry, Wells and Pupedis. 
1993:9. 

Holotype male: USA, Oregon, Benton Co., 
Philomath (Mary's Peak), 28.IX.1989, C. S. 
Henry & M. M. Wells (YPM). 

DISTRIBUTION. — USA: CA, ID, OR, WA 
(Henry, Wells & Pupedis, 1993:9). 
Chrysoperla comanche (Banks), 1938b: 119 
[Chrysopa]. 

Holotype female: USA, Texas, Laredo (MCZ). 

= Chn'sopa sperryae Banks, 1943:74. Lecto- 
type female: USA, California, Riverside, 
7.IX.1940, Sperry(MCZ). 

TAXONOMY. — To Chrysoperla (see Brooks 
& Barnard, 1990:271); C. spenyae = C. co- 
manche (see Adams, 1958:45); lectotype desig- 
nated for C. spenyae (see Adams, 1958:45). 

Distribution. — CAN: BC 2 , USA: AZ 1 , 
CO 2 , NM 1 , TX 1 ('Banks, 1938b: 119; 
Bickley & MacLeod, 1956: 194, as C. spenyae). 
Chrysoperla downesi (Smith). 1932:594 
[Chrysopa]. 

Holotype female: Canada, British Columbia, 
Kelowna, W. Downes (CNC). 

= Chrysopa mohave Banks, 1938b: 120. Syn- 
types (sex unknown): USA, California, Clare- 
mont; Stanford University campus; Arizona, 
Chiricahua Mountains (MCZ). 

TAXONOMY. — To Chrysoperla (see Henry, 
1985:965); C. mohave synonymized with C. 
carnea (see Tauber & Tauber, 1973:1153); C. 
downesi synonymized with C. carnea (see Gar- 
land, 1985:753); C. downesi and C mohave con- 
sidered synonyms of C. carnea (see Brooks & 
Barnard, 1990:271); C. downesi considered valid 
species with C. mohave as synonym (see Henry, 
1993:23). 

DISTRIBUTION. — CAN: BC 1 , SK',_USA: 



CA 



AZ Z , CA", ID 3 , MA 3 , MT 3 , NH 3 , NY 3 , VT 3 

('Smith, 1932:595; 2 Banks, 1938b: 120, as C. 

mohave; 3 Henry, 1985:967). 

Chrysoperla externa (Hagen), 1861:221 

[Chrysopa]. 

Neotype female: USA, Winter Park, Florida, 
3.IX, E. M. Davis (MCZ). 



52 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



= Chrysopa lanata Banks, 1 9 1 0: 1 54. Holotype 
male: Argentina, Mendoza, Jensen-Haarup Ex- 
pedition (MCZ). 

= Chrysopa graciana Navas, 19 19b:30 1 . 
Holotype (sex unknown): Argentina, Alta Gra- 
cia, 11.1918, C. Bruch (MLP). 

Taxonomy. — C. lanata = C. externa 
(Adams, 1963:222); C. graciana = C externa 
(Adams & Penny, 1987:42 1). 

DISTRIBUTION. — USA: FL 1 , SC 1 , TX 2 
('Adams, 1963:222; 2 Agnew et al,, 1981:9). 

Note. — The specimens listed by Hagen, 
1875:921 and Smith, 1932:596 are misidentifi- 
cations. 

Chrysoperla harrisii (Fitch), 1855:794 
[Chrysopa]. 

Holotype male: USA, New York (MCZ). 

= Chrysopa stenostigma Navas, 1914c:61. 
Holotype (abdomen missing): USA, New York, 
Long Island, Yaphank, 25.VII.1913 (MZB). 

TAXONOMY. — C. harrisii is a replacement 
name for the C. perla of Harris, 1841:197. To 
Chrysoperla (see Garland, 1985:755); C. 
stenostigma = C harrisii (Bickley & MacLeod, 
1956:193). 

DISTRIBUTION. — CAN: BC 5 ON 5 USA: 
AK 1 ', DC 3 , IN 6 , KS 4 , MD 12 , MI 3 , MN 10 MO 9 , 
NC 7 , NH 3 , NJ 2 , NY 1 , TN 8 , TX 14 , WS 13 (Titch, 
1855:794; 2 Smith, 1900:55; 3 Banks, 1903a: 155; 
4 Smith, 1925:169; 5 Smith, 1932:595; 
Montgomery & Trippel, 1933:260; 7 Brimley, 
1938:30; 8 Bickley, 1941:188; 9 Froeschner, 
1947:134; 10 Parfin, ^1952:425; "Bickley & 
MacLeod, 1956:193; 12 Bram& Bickley, 1963:7; 
13 Throne, 1971a:73; 14 Agnew et al., 1981:9). 
Chrysoperla johnsoni Henry, Wells and Pu- 
pedis, 1993:10. 

Holotype male: USA, Oregon, Benton Co., 
Philomath (Mary's Peak), 28. IX. 1989, C. S. 
Henry & M. M. Wells (YPM). 

DISTRIBUTION. — USA: AZ, CA, ID, OR, 
WA (Henry et al., 1993:10). 
Chrysoperla plorabunda (Fitch), 1855:792 
[Chrysopa]. 

One male syntype, one syntype of unknown 
sex: USA, Illinois (no head or abdomen) 
(USNM); female (no label data, except #8592) 
(MCZ). 

= Chrysopa robertsonii Fitch, 1 855:792. Holo- 
type female: USA, Arkansas, V, W. S. Robertson 
(MCZ). 



= Chrysopa pseudographa Fitch, 1855:793. 
Lectotype male: USA, Illinois, X (MCZ) NEW 
LECTOTYPE DESIGNATION 

= Chrysopa illinoiensis Shimer, 1865:208. 
Type(s) (sex unknown): USA, Illinois (deposi- 
tory unknown). 

= Chrysopa californica Coquillett, 1890:288. 
Holotype female: USA, California, Los Angeles, 
Coquillett (USNM). 

= Chrysopa simplex Navas, 1908:402. Type(s) 
(sex unknown): USA, Rocky Mountains, R. P. 
Joannis (MZB). 

TAXONOMY. — To Chrysoperla (see Brooks 
& Barnard, 1990:271); C. robertsonii = C. 
plorabunda, C pseudographa = C plorabunda, 
C illinoiensis = C plorabunda (see Banks, 
1903a: 162); C. californica = C. plorabunda (see 
Smith, 1932:594); C. californica = subspecies of 
C plorabunda (see Adams, 1956:45); C 
plorabunda = C carnea (see Tjeder, 1960: 148); 
C. plorabunda not = C. carnea (see Henry, 
1983:293); C. simplex = C plorabunda (Adams, 
NEW SYNONYMY). 

DISTRIBUTION. — CAN: AB 5 , BC 5 , MB 5 , 

SK 5 , USA: AK 10 , AZ 14 , CO 3 , CT 13 , DC 14 , FL 14 , 

IA 3 , ID 13 FL 1 , IN 6 , KS 4 , MD 11 ME 14 , MI 3 , 

MN 3 , MO 9 , MS 3 MT 13 , NC 7 , NJ", NM 14 , NY 1 

OH 3 , OK 1 , OR 14 , TN 8 , TX 3 , WA 14 WS 1 " 

( 'Fitch, 1855:792; 2 Smith, 1900:55; 3 Banks, 

1903a:155; 4 Smith, 1925:168; 5 Smith, 

1932:593; Montgomery & Trippel, 1933:260; 

7 Brimley, 1938:30; 8 Bickley, 1941:188; 

9 Froeschner, 1947:134; l0 Bickley & MacLeod, 

1956:193; n Bram & Bickley, 1963:4, as C. 

12 
carnea; ^Throne, 197 la:72, as C. carnea; 

13 Henry, 1985:967; 14 Henry et al., 1993:9). 

REMARKS. — Henry ( 1985:967) indicated that 
the holotype of C. plorabunda is at MCZ. In fact, 
this is one of two syntypes, and no lectotype has 
yet been designated. Henry, Wells and Pupedis 
(1993:9) reported that C. plorabunda possibly 
does not occur in California, so that previous 
records from this state probably pertain to C. 
adamsi or C. johnsoni, which are morphologi- 
cally indistinguishable from C. plorabunda. 
Chrysoperla rufilabris (Burmeister), 1839:979 
[Chrysopa]. 

Syntypes (sex unknown): USA and Mexico 
(ZMH, specimen destroyed). 

= Chrysopa interrupta Schneider, 1851:76. 
Holotype male: USA, Pennsylvania (HMB). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



53 



= Chrysopa attenuata Walker, 1853:242. Lec- 
totype male: USA, east Florida, St. John's Bluff, 
E. Doubleday (BMNH). 

= Chrysopa repleta Walker, 1853:244. Holo- 
type male: USA, Georgia, Abbot (BMNH). 

= Chn'sopa novaeboracensis Fitch, 1855:794. 
Holotype female: USA, New York (MCZ). 

= Chiysopa tabida Fitch, 1855:796. Type(s) 
(sex unknown): USA (MCZ, specimen not 
found). 

= Chnsopa citri Ashmead, 1 880: 12. Holotype 
male: USA, Florida, W. H. Ashmead & S. Hen- 
shaw (MCZ). 

= Chrysopa medialis Banks, 1 903a: 1 54. Holo- 
type female: USA, Maryland, High Island, IX 
(MCZ). 

TAXONOMY. — To Chrysoperia (see Garland, 
1985:756); Hagen, 1866:395, 398 (C. novaebo- 
racensis = C rufilabris, C. tabida = C. rufilabris) 
(see Hagen, 1866:395, 398); C. repleta = C. 
rufilabris, C. attenuata = C. rufilabris (see 
Banks, 1903a: 161); C. citri = C. rufilabris (see 
Bickley & MacLeod, 1956:194); C. medialis = 
C. rufilabris, C interrupta = C rufilabris (see 
Bram & Bickley. 1963:6); lectotype designated 
for C. attenuata (see Kimmins, 1940:447). 

Distribution. — CAN: NB 4 ON 6 , USA 
DC 4 , FL 2 , GA 2 , IN 7 , KS 5 , LA 4 , MD 12 , MI 4 
MN 11 , MO 10 , NC 8 , NJ 3 , NY 3 , OH 4 , PA 1 , TN 9 
TX 14 , VA 4 , WS 13 ('Schneider, 1851:79 
2 Walker, 1853:242, 244; 3 Smith, 1900:55 
4 Banks, 1903a: 152; 5 Smith, 1925:168; 6 Smith 
1932:597; Montgomery & Trippel, 1933:260 
8 Brimley, 1938:30; 9 Bickley, 1941:189 
10 Froeschner, 1947:134; ' 'Parfin, 1952:425 
l2 Bram& Bickley. 1963:5; 13 Throne, 1971a:74 
14 Agnew et al., 1981:5). 

Chrysopodes (Neosuarius) Adams and 

Penny, 1987:435 

Chrysopodes (Neosuarius) collaris 

(Schneider), 1 85 1 :80 [Chrysopa]. 

Holotype (abdomen missing): Puerto Rico 
(HMB). 

= Chrysopa thoracica Walker, 1853:243. 
Holotype (abdomen missing): Dominican Re- 
public, Santo Domingo, M. A. Pierret (BMNH). 

= Chrysopa bugii Kolbe, 1888:173. Type(s) 
(sex unknown): Puerto Rico (HMB). 



= Chrysopa signatalis Banks, 191 1:342. Syn- 
type females: USA, Texas, Brownsville, VI, 
Schaeffer (MCZ, USNM). 

= Chnsopa rufolinea Banks, 1914a:24. Holo- 
type male: Colombia, Cali, 1000m, Fassl (MCZ). 

= Chrysopa acolhua Banks, 1948:156. Holo- 
type male: Mexico. Guerrero, Pungarabato. 1260 
m, 22.VIII.1930, Jose Paner (MCZ). 

TAXONOMY. — C. thoracica = C. collaris, C 
b-ugii = C. collaris, C rufolinea = C collaris, C 
signatalis = C. collaris, C acolhua = C. collaris 
(see Adams & Penny, 1987:436). 

DISTRIBUTION. — USA: FL 2 , TX 1 ('Banks, 
191 1:342; 2 Banks, 1938b: 121, as C thoracica). 

Eremochrysa (Chrysopiella) Banks. 191 1:344 

Eremochrysa (Chrysopiella) brevisetosa 

(Adams & Garland), 1981:1 [Chrysopiella]. 

Holotype male: USA, Utah, Uintah Co., 
22.V. 1954, M. Cazier (AMNH). 

TAXONOMY. — To Eremochrysa (Chryso- 
piella) (see Brooks & Barnard, 1990:272). 

Distribution. — CAN: AB, USA: ID, NV, 
OR, UT, WY (Adams & Garland, 1981:3). 
Eremochrysa (Chrysopiella) minora (Banks), 
1935:55 [Chrysopiella]. 

Holotype (sex unknown): USA, Oregon, Uma- 
tilla, 24.VI.1882, S. Henshaw (MCZ). 

TAXONOMY. — To Eremochrysa (Chryso- 
piella) (see Brooks & Barnard, 1990:272). 

Distribution. — USA: CA 2 , CO 2 , OR 1 
('Banks, 1935:55; 2 Adams, NEW STATE RE- 
CORDS) 

Eremochrysa (Chrysopiella) pallida (Banks), 
1911:345 [Chrysopiella]. 

Holotype male: USA, New Mexico, Rincon, 
3LVIII,Cockerell(MCZ). 

TAXONOMY. — To Eremochrysa (Chryso- 
piella) (see Brooks & Barnard, 1990:272). 

Distribution. — AZ 2 , CA 3 , NM 1 , OR-, 
UT 2 , ('Banks, 191 1:345; 2 Bickley & MacLeod, 
1956: 198; 3 Adams, NEW STATE RECORD). 
Eremochrysa (Chrysopiella) sabulosa 
(Banks), 1897b: 174 [Chrysopa]. 

Holotype female: USA, Colorado, Fort Col- 
lins, 1916 (MCZ). 

TAXONOMY. — To Chrysopiella (see Banks, 
1911:344); to Eremochrysa (Chrysopiella) (see 
Brooks & Barnard, 1990:272). 

DISTRIBUTION. — CO 1 , KS 3 , NM 2 , UT 4 , 
WY 4 ('Banks, 1897b: 174: 2 Banks, 1903a: 151; 



54 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



J Smith, 1925:169; 4 Bickley & MacLeod, 
1956:198). 

Remarks. — Bickley and MacLeod 
(1956:198) erroneously reported that Banks 
(1948) listed specimens from Texas. 

Eremochrysa (Eremochrysa) Banks, 
1903a: 158 

Eremochrysa (Eremochrysa) altilis Banks, 
1950:56. 

Syntype male and females: USA, Arizona, 
Graham Co., Pinaleno Mtns., Stockton Pass, 
5440 ft., 4.VIII.1948, W. Nutting and F. Werner 
(MCZ). 

Distribution. — AZ (Banks, 1950:57). 
Eremochrysa (Eremochrysa) californica 
Banks, 1905b:6. 

Holotype male: USA, California, Santa Clara 
Co. (MCZ). 

Distribution. — AZ 2 , CA 1 ('Banks, 
1905b:6; 2 Banks, 1950:63). 
Eremochrysa (Eremochrysa) canadensis 
(Banks), 1911:339 [Chrysopa]. 

Holotype female: Canada, Ontario, Lake 
Huron, Go Home Bay, 12.VII.1907 (MCZ). 

TAXONOMY. — To Eremochn'sa (see Banks, 
1950:64). 

Distribution. — CAN: ON 1 , USA^ MA 2 , 
ME 2 , NH 2 , WS 3 ('Banks, 1911:339; 2 Banks, 
1950:66; 3 Throne, 1971a: 76). 
Eremochrysa (Eremochrysa) fraterna 
(Banks), 1897b: 174 [Chrysopa]. 

Holotype female: USA, Colorado (MCZ). 

TAXONOMY. — To Eremochrysa (see Banks, 
1903a: 159). 

Distribution. — CAN: BC 4 , USA: CO 1 , 
KS 3 , NV 2 ('Banks, 1897b: 174; 2 Banks, 
1903a:159; 3 Smith, 1925:169; 4 Smith, 
1932:582). 

Eremochrysa (Eremochrysa) hageni (Banks), 
1903a: 158 [Chrysopa]. 

Holotype female: USA, Texas, Austin, 
20. V. 1900 (MCZ). 

Distribution. — USA: AZ 2 , NM 2 , TX 1 , 
UT 2 ('Banks, 1903a: 159; 2 Banks, 1950:60). 
Procter (1946:44) recorded a specimen of E. 
ha-geni from Maine, but E. canadensis is the 
only species of Eremochrysa which is found in 
the northeastern U. S. 

Eremochrysa (Eremochrysa) pima Banks, 
1950:61. 



Two syntypes (one female, one missing abdo- 
men): USA, Arizona, White Mtns., South Fork 
Camp, 22.VI. 1 947, G. H. and J. L. Sperry (miss- 
ing abdomen); New Mexico, Santa Fe, Cockerell 
(female) (MCZ). Note that MCZ records only 
indicate presence of paratype. 

Distribution. — USA: AZ, NM (Banks, 
1950:62). 

Eremochrysa (Eremochrysa) pumilis Banks, 
1950:58. 

Holotype male: USA, Colorado, Garland, VII, 
Yarrow (MCZ). 

Distribution. — USA: CA 2 , CO 1 , TX 1 , UT' 
('Banks, 1950:58; 2 Bickley & MacLeod, 
1956:200). 

Eremochrysa (Eremochrysa) punctinervis 
(McLachlan), 1869:24 [Chrysopa]. 

Holotype female: USA, Texas, Bosque Co. 
(BMNH). 

TAXONOMY. — To Eremochrysa (see Banks, 
1903a:159). 

Distribution. — CAN:^ BC 4 , USA: AZ 2 , 
CA 2 , CO 2 , FL 5 , KS 3 , NM 2 , TX 1 ('McLachlan, 
1869:24; 2 Banks, 1903a: 159; 3 Smith, 1925:169; 
4 Smith. 1932:583; 5 Banks, 1950:53). 
Eremochrysa (Eremochrysa) rufifrons Banks, 
1950:57. 

Syntype series (both sexes): USA, Arizona, 
Globe, 9 .VIII. 1933, Parker; Arizona, Globe, 
19. VII, F. Werner and W. Nutting (MCZ). 

Distribution. — USA: AZ (Banks, 
1950:58). 

Eremochrysa (Eremochrysa) spilota Banks, 
1950:61. 

Five syntype females: USA, California, Fort 
Yuma, 6-13.VI.1948, Andahl; California, 
Brawley, 8. IV, A. L. Melander (MCZ). Note that 
MCZ records only indicate presence of paratype. 
Distribution. — USA: CA (Banks, 1950:61). 
Eremochrysa (Eremochrysa) tibialis Banks, 
1950:55. 

Syntype male: USA, Arizona, Florence Junc- 
tion, 18.IV.1935, F. H. Parker; Utah, Waston 
22.VII, F. M. Carpenter; California, Vidal, 9.IV, 
Sperry (MCZ). 

DISTRIBUTION. — USA: AZ'XA'^V-^T 1 
('Banks, 1950:56; 2 Bickley & MacLeod, 
1956:199). 

Eremochrysa (Eremochrysa) yosemite Banks, 
1950:63. 

Holotype male: USA, California, Yosemite, 
12.VI.1931,Essig(MCZ). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



55 



Distribution. 
1950:64). 



USA: CA (Banks, 



Meleoma Fitch. 1855:786 

Meleoma arizonensis (Banks), 1903a: 155 
[Chrysopa], 

Holotype female: USA, Arizona, Prescott 
(MCZ). 

TAXONOMY. — To Meleoma (see Tauber, 
1969:18). 

Distribution. — USA: AZ 1 , CA~, Nivr, 
TX 3 , UT 3 ('Banks, 1903a: 155; 2 Bickley & 
MacLeod, 1956:195; 3 Tauber, 1969:18). 

Meleoma beardi Tauber, 1969:20. 

Holotype male: USA, Arizona, Chiricahua 
Mountains, Rustler's Park. 8,400 ft., 
1 1 .VI. 1956, H. & A. Howden (CNC). 

Distribution. — USA: AZ (Tauber, 
1969:21). 

Meleoma dolicharthra (Navas), 1914e:96 
[Chrysopa]. 

Holotype male: Guatemala: Capetillo, 
XI. 19 14, G. C. Champion (BMNH). 

= Meleoma cavifrons Banks, 1950:46. Holo- 
type male: USA, California, Tuolumne Co., 
Pinecrest, 10.VII.1948.P. H. Arnaud, Jr. (MCZ). 

TAXONOMY. — To Meleoma (see Kimmins, 
1940:448); M. cavifrons = M. dolicharthra (see 
Adams, 1969:6). 

Distribution. — CAN: BC 2 , USA: AZ 2 , 
CA 1 , CO 2 , OR 2 , UT 2 , WA 2 ('Banks, 1950:46; 
"Tauber, 1969:36). 

Meleoma emuncta (Fitch). 1855:792 
[Chrysopa]. 

Holotype (sex unknown): USA, New York 
(MCZ). 

= Meleoma slossonae Banks, 1896:95. Holo- 
type male: USA, New Hampshire, Mount Wash- 
ington (MCZ). 

= Meleoma verticalis Banks. 1908a:259. Holo- 
type male: USA, Colorado. Golden, Chimney 
Gulch, 3.VIII. 1907, Oslar (MCZ). 

— Meleoma comata Banks, 1950:45. Holotype 
male: USA, California, San Bernardino Co., Up- 
per Santa Ana River, 15.VII.1948, G. H. & J. L. 
Sperry (MCZ). 

Taxonomy. — To Meleoma (see Banks, 
1924:432); M. slossonae = M. emuncta (see 
Banks. 1924:432); M. verticalis = M. emuncta, 
M. comata = M. emuncta (see Tauber, 1969:23). 



Distribution. — CAN: BC 5 , MB 8 , NB 8 , 
NF 8 NS 8 , ON 5 , PQ 5 , USA: AZ 7 , CA 4 , CO 3 , ID 8 , 
MA 8 , ME 6 . MI 8 NC 8 , NH 2 . NM 7 , NV 8 NY 1 , 
OR 8 , UT 8 ^ VA , VT 8 , WA 8 . WS 8 (Fitch, 
1855:792; "Banks, 1896:95; 3 Banks, 1908a:259; 
4 Banks, 1950:45; 5 Smith, 1932:583; 6 Procter, 
1946:44; 7 Bickley & MacLeod, 1956:185, 186; 
8 Tauber, 1969:23). 

Meleoma furcata (Banks), 1911:342 
[C/uysopa]. 

Holotype female: USA, New Mexico, Fort 
Wingate, 19. VII (MCZ). 

= Meleoma delicata Banks, 1 950:48. Holotype 
male: USA. New Mexico, Fort Wingate, 
14.VIII.1908, J. Woodgate (MCZ). 

Taxonomy. — To Meleoma (see Tauber, 
1969:40); M. delicata = M. furcata (see Tauber, 
1969:40). 

Distribution. — USA^ AZ 2 , CA 2 , NM 1 . 
UT 3 ('Banks, 1911:342; 2 Bickley & MacLeod. 
1956:186, 196; 3 Tauber, 1969:41). 
Meleoma hageni Banks, 1948:170. 

Holotype male: Mexico. Distrito Federal, 
Mexico City, Lomas de Chapultepec, 
25.VII.1939, A. Dampf (MCZ). 

Distribution. — USA: AZ, NM (Tauber, 
1969:38). 
Meleoma kennethi Tauber, 1969:42. 

Holotype male: USA, Nevada, Clark Co., 38 
miles northwest of Las Vegas, Lee Canyon, 
6,900 ft., 29.VII.1966. F., P., and M. Rindge 
(AMNH). 

Distribution. — USA: CA, NV (Tauber, 
1969:42). 
Meleoma pallida Banks, 1908a:260. 

Holotype male: USA, Arizona, Huachuca 
Mountains, 8,000 ft., 20.VIII.1905. Oslar 
(MCZ). 

Distribution. — USA: AZ 1 , CA 2 ('Banks, 
1908a:260; 2 Bickley & MacLeod, 1956:186). 
Meleoma pinalena (Banks), 1950:49 
[Clvysopa]. 

Holotype (abdomen missing): USA, Arizona. 
Pinals-Globe, 18.VII.1948,F.Werner&W. Nut- 
ting (MCZ). 

TAXONOMY. — To Meleoma (see Tauber, 
1969:13). 7 

Distribution. — USA: AZ , NM - ( Banks, 
1950:49; 2 Tauber, 1969:13). 
Meleoma schwarzi (Banks), 1903a: 146 
[Chrysopa]. 



56 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Holotype (abdomen missing): USA, New 
Mexico, Las Vegas Hot Springs, 5. VIII, Barber 
& Schwarz (USNM). 

TAXONOMY. — To Meleoma (see Adams, 
1962:178). 

DISTRIBUTION. — USA: AZ 2 , CA 2 , CO 2 , 
NM 1 , NV 2 , OR 2 , UT 2 ('Banks, 1903a: 147; 
2 Tauber, 1969:14). 
Meleoma signoretii Fitch, 1855:786. 

Type(s) (sex unknown): USA, Vermont, Ru- 
pert (depository unknown) 

Distribution. — CAN: BC 3 MB 9 , NS 9 , 
ON 3 , PQ 3 , USA: DC 8 , IL 9 , IN 10 MA 9 MD 8 , 
ME 6 , MI 9 , MN 7 , NC 4 , NH 8 , NJ 9 , NY 2 , PA 9 , 
TN 5 , VA 8 , VT 1 , WS 9 , WV 9 , ('Fitch, 1855:786; 
2 Leonard 1928:40; 3 Smith, 1932:584; 4 Brimley, 
1938:31; Bickley, 1941: 187; 6 Procter, 1946:44; 
7 Parfin, 1952:424; 8 Bickley & MacLeod, 
1956:185; 9 Tauber, 1969:33; l0 Lawson & 
McCafferty, 1984:130). 

Meleoma tezcucana (Banks), 1948:157 
[Chrysopa]. 

Holotype female: Mexico, Distrito Federal, 
Lomas de Chapaltepec, Mexico City, 
8.VIII.1940, A. Dampf (MCZ). 

Taxonomy. — To Meleoma (see Adams, 
1969:8). 

Distribution. — USA: AZ (Adams, 
1969:8). 

Nineta Navas, 1912b:98 

Nineta gravida (Banks), 191 1:343 [Chrysopa]. 

Type male: USA, California, Yosemite 
(MCZ). 

Taxonomy. — to Nineta (see Brooks & Bar- 
nard, 1990:275). 

Distribution. — CAN: BC 2 , USA: CA 1 
('Banks, 191 1:343; 2 Smith, 1932:597). 
Nineta nanina (Banks), 191 1:344 [Chtysopa]. 

Type(s) (sex unknown): USA, Arizona, Pal- 
merlee (MCZ). 

Taxonomy. — To Nineta (see Brooks & Bar- 
nard, 1990:275). 

Distribution. — USA: AZ 1 , UT 2 ('Banks, 
191 1:340; 2 Adams, NEW STATE RECORD). 



Holotype (female): Brazil (HMB). 

= Chtysopa cubanaNavas, 1921:120. A junior 
primary homonym of Chn'sopa cubana Hagen, 
1861:215. 

= Chtysopa antillana Navas, 1922c:392. A 
replacement name for Chrysopa cubana Navas. 
Lectotype male: Cuba, Havana, F. Cervera 
(MCZ). 

= Chrysopa bouvi eri Navas, 1924c: 1 12. Holo- 
type female: Costa Rica, Paul Serre (MNHN). 

= Chtysopa rata Lacroix, 1926:68. Syntype 
series (one male, one female, one without abdo- 
men): Brazil, Minas Gerais, Uberaba (MNHN). 

= Chtysopa uribei Navas, 1927b:5. Holotype 
male: Colombia, Cundinamarca, San Antonio de 
Tena, San Pedro Claver (MZB, specimen not 
found). 

= Chtysopa scalaris Navas, 1929c:26. Holo- 
type male: Colombia, Behn vend., 3.XII.1900 
(Hamburg, specimen destroyed). 

= Chtysopa yucatanensis Navas, 1929c:28. 
Holotype male: Mexico, Yucatan, Progreso, 
5.V.1905, W. Schwinghammer (Hamburg, 
specimen destroyed). 

= Chrysopa divergens Navas, 1931:82. Holo- 
type (sex unknown): Costa Rica, San Jose, 8 km 
wsw San Jose, Farm La Caja, V. 1925 (Hamburg, 
specimen destroyed). 

TAXONOMY. — To Plesiochtysa; C bouvieri 
= C. brasiliensis, C antillana = C brasiliensis, 
C. rata = C brasiliensis, C uribei = C brasilien- 
sis, C scalaris = C brasiliensis, C yucatanensis 
= C brasiliensis, C divergens = C brasiliensis 
(see Adams, 1982:29). 

Distribution. — USA: FL (Adams, NEW 
NATIONAL RECORD) 

Pseudomallada Tsukaguchi, 1995:67 

Pseudomallada luctuosus (Banks), 1911:343 
[Chrysopa]. 

Holotype male: USA, New Mexico, Fort Win- 
gate, 26.VI (MCZ). 

TAXONOMY. — To Mallada (see Adams, 
1975:172); to Pseudomallada by Adams, NEW 
COMBINATION 



Plesiochrysa Adams, 1982b:28 

Plesiochrysa brasiliensis (Schneider), 1851: 83 
[Chrysopa]. 



Distribution. 



USA: AZ 2 , CO 2 , IN 3 , NE 2 , 



NM' ('Banks, 1911:343; "Adams & Garland, 
1982:246; 3 Lawson & McCafferty, 1984: 130). 
Pseudomallada macleodi (Adams & Garland), 
1982:240 [Mallada]. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



57 



Holotype male: USA, Texas, Erath Co., 
Stephenville. 20.IV. 1 98 1 , C. W. Agnew (MCZ). 

TAXONOMY. — To Pseudomallada by 
Adams, NEW COMBINATION. 

Distribution. — CAN: ON, USA: AZ, KS, 
MD. TX (Adams & Garland, 1982:242). 
Pseudomallada perfectus (Banks). 1895a:516 
[Chrysopa]. 

Holotype female: Mexico. Baja California Sur, 
Serra El Taste (MCZ). 

= Chrysopa cockerelli Banks, 1903a: 154. 
Holotype female: USA, New Mexico, East Las 
Vegas (MCZ). 

= Clvysopa injusta Banks. 1906a:98. Holo- 
type female: USA, California, mountains near 
Claremont, Baker (MCZ). 

= Clvysopa marginalis Banks, 1905b:5 not 
Navas 1905:122. Holotype (sex unknown): 
USA, California, mountains near Claremont, 
Baker (MCZ, specimen not found). 

Taxonomy. — To Mallada (see Adams, 
1975:172): C. cockerelli = M. perfectus, C in- 
justa = M. perfectus (see Adams & Garland, 
1982:244); to Pseudomallada by Adams, NEW 
COMBINATION 

Distribution. — CAN: BC 4 , USA: AZ 5 
CA 2 , CO 5 , KS 3 , NM 1 , OR 5 , UT 5 , WA 5 , WY 5 
('Banks, 1903a: 154; 2 Banks, 1906a:98; 3 Smith, 
1925:168, as C. cockerelli; 4 Smith, 1932:597, as 
C. cockerelli; 5 Adams & Garland, 1982:244). 

REMARKS. — The Navas name of Clvysopa 
marginalis was published in June. 1905 while the 
Clvysopa marginalis of Banks was published in 
November, 1905. 

Pseudomallada sierra (Banks), 1924:431 
[Chrysopa]. 

Holotype female: USA, California, Los Ange- 
les Co., San Gabriel Mtns., Sister Elsie Peak, 
10.VI, F. Grinnell (MCZ). 

Taxonomy. — To Mallada (see Adams & 
Garland, 1982:245); to Pseudomallada by 
Adams, NEW COMBINATION 

Distribution. — USA: ^AZ 2 , CA 1 , OR 2 , 
WA 2 ('Banks, 1924:431; 2 Adams & Garland, 
1982:245). 

Yumachrysa Banks, 1950:51 

Yumachrysa apache (Banks), 1938b: 120 
[Chrysopa]. 

Holotype female: USA, Arizona, Globe, 
27.VI.1933, Parker (MCZ). 



TAXONOMY. — To Clvysopa (Yumachrysa) 
(see Bickley & MacLeod, 1956:189); to Suarius 
(Prochrysopa) (see Tauber, 1975:696), to Yu- 
machrysa (see Brooks & Barnard, 1990:276). 

Distribution. — USA: AZ 1 , CA 3 , TX 2 
('Banks, 1938b: 121; 2 Bickley & MacLeod, 
1956:189; 3 Tauber, 1975:697). 
Yumachrysa clarivena (Banks). 1950:50 
[ Clin 'sopa (Yi imachrysa) ] . 

Holotype female: USA, Arizona, Yuma Co.. 
Ehrenberg, 11. VII. 1948, W. Nutting and F. 
Werner (MCZ). 

Taxonomy. — Yumachrysa elevated to ge- 
neric status (see Brooks & Barnard, 1990:241). 

Distribution. — USA: AZ (Banks, 
1950:50). 

Yumachrysa yuma (Banks), 1950:49 
[Clvysopa (Yumachrysa)]. 

Two syntypes (male and female): USA, Cali- 
fornia, Fort Yuma, 6. VI and 13. VI, Andahl 
(MCZ). 

TAXONOMY. — To Suarius (Proclvysa) (see 
Tauber, 1975:697); to Yumachrysa (see Brooks 
& Barnard, 1990:241). 

Distribution. — USA: AZ 2 , CA 1 ('Banks, 
1950:50; 2 Tauber, 1975:698). 

NOTE. — Leucochrysa californica Navas, 
1928a:35 was described from California, and the 
type specimen is present in the Stockholm Mu- 
seum. Adams, 1977:94 has subsequently shown 
that this species is a synonym of Leucochrysa 
Colombia Banks, 1910:150, known only from 
Colombia. The locality label associated with the 
type specimen of L. californica dates from the 
last century and may have originally been noted 
as "Calf or "Col." 

CONIOPTERYGIDAE 
Coniopterygidae, or dusky-wings, comprise 
the smallest and some of the most highly modi- 
fied lacewings. Adults are generally only 
2-3 mm long, with reduced wing venation and a 
white powder covering the wings and body. Be- 
cause of this white powder, they often are mis- 
taken for white-flies (Aleyrodidae) in flight. 
They are most active near dawn and dusk, flut- 
tering slowly between plants, where they lay 
eggs and feed on soft-bodied insects. Eggs are 
usually laid singly on bark or leaves. Withy- 
combe (1923) indicated that a larval dusky-wing 
will consume between 150 and 300 aphid eggs 



58 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



or young nymphs before metamorphosing to a 
pupa. Larvae and adults can feed on a wide 
variety of sluggish, soft-bodied insects, like 
aphids, white-flies, mealy-bugs, scale insects, 
etc. This makes them one of our more important 
biological control agents, although because of 
their size, they are often overlooked in the field. 
Although a wide variety of insects are consumed 
by the family in general, many species are closely 
associated with a specific species of tree or bush, 
indicating a close association with a specific type 
of host insect. The taxonomy of Coniopterygidae 
has been covered in a series of monographs and 
papers by Meinander (1972, 1974, 1975, 1990) 
and Johnson (1976, 1981a, 1981b). 

ALEUROPTERYGINAE 

Aleuropteryx Low, 1885:79 

Aleuropteryx arceuthobii Meinander, 1975:28. 

Holotype male: USA, Colorado (USNM). 

DISTRIBUTION. — USA: CO 1 , TX 2 ( ' Meinan- 
der, 1975:28; 2 Johnson, 1981a:272). 
Aleuropteryx arizonica Johnson, 1981a:272. 

Holotype male: USA, Arizona, Pinal Co., near 
Superior, Boyce Thompson Southwestern Arbo- 
retum, 1 1. VII. 1949, B. W. Benson (INHS). 

Distribution. — USA: AZ (Johnson, 
1981a:274). 
Aleuropteryx cupressi Meinander, 1974:218. 

Holotype male: USA, California, Marin Co., 
emerged ex Cupressus goveniana, 2.V.1961, J. 
Powell (CAS). 

Distribution. — USA: CA (Meinander, 
1974:218). 
Aleuropteryx dragoonica Johnson, 198 la:275. 

Holotype male: USA, Arizona, Cochise 
Stronghold, Dragoon Mountains, 

15-18.VIII.1969, R. J. Shaw (USNM). 

Distribution. — USA: AZ (Johnson, 
1981a:275). 
Aleuropteryx juniperi Ohm, 1968:14. 

Holotype male: Germany, Treisbach (Ohm 
Collection). 

Distribution. — USA: PA 1 , VA 2 ('Henry, 
1974:643; 2 Flint, 1974:703). 
Aleuropteryx knovvltoni Johnson, 198 la:278. 

Holotype male: USA, Utah, Logan, 
29.VI.1950, G. F. Knowlton (USNM). 



Distribution. — USA: UT (Johnson, 
1981a:279). 

Aleuropteryx longipennis Meinander, 
1974:218. 

Holotype male: Mexico, Baja California 
Norte, 1 mile north of Meling Ranch, 2100 ft., 
17.111. 1972, J. Doyen & J. Powell (CAS). 

Distribution. — USA: CA 1 , NM 1 , UT 2 
('Meinander, 1974:219; 2 Johnson, 1981a:281). 
Aleuropteryx maculipennis Meinander, 
1972:45. 

Holotype male: USA, California, Kern Co., 19 
miles north northeast of Mojave, 14. VI. 1962, C. 
MacNeill, D. Rentz & R. Brown (CAS). 

Distribution. — USA: AZ 2 , CA 1 , NM 3 , 
TX 2 ('Meinander, 1972:45; 2 Johnson, 
1981a:282; 3 Meinander, 1990:7). 
Aleuropteryx megacornis Johnson, 198 la:284. 

Holotype male: USA, Arizona, Cochise 
Stronghold, Dragoon Mountains, 9-12.V.1970, 
R. J. Shaw (USNM). 

Distribution. — USA: AZ, NM, TX 
(Johnson, 1981a:284). 
Aleuropteryx punctata Meinander, 1974:220. 

Holotype male: Mexico, Sinaloa, 20 mi W 
Rosario, 30.1.1964, E. SchlingerfCAS). 

Distribution. — AZ 1 , NM" ('Meinander, 
1974:220; 2 Johnson, 1981a:287). 
Aleuropteryx simillima Meinander, 1972:46. 

Holotype male: Mexico, Baja California, San 
Felipe, 7.III.1963, P. H. Arnaud (CAS). 

Distribution. — USA: AZ 2 , TX ' ( 'wheeler, 
1980:51; 2 Johnson, 1981a:289). 
Aleuropteryx unicolor Meinander, 1972:48. 

Holotype male: USA, California, Riverside 
Co., Palm Desert, 4 miles south of Boyd Desert 
Research Center, 11.IV.1963, W. A. Steffan 
(CAS). 

Distribution. — USA: AZ 2 , CA 1 ('Meinan- 
der, 1972:49; 2 Johnson, 1981a:290). 
Aleuropteryx vulgaris Meinander, 1972:51. 

Holotype male: USA, Texas, Kerrville, 
VII. 1954, L. J. Bottimer (USNM). 

Distribution. — USA: AZ, CA, NM, TX, 
UT (Meinander, 1972:51). 
Aleuropteryx werneri Johnson, 1981a:294. 

Holotype male: USA, New Mexico, Las Cru- 
ces, 28.VII-3.VIII. 1975, W. P. Morrison 
(USNM). 

Distribution. — USA: NM (Johnson, 
1981a:294). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



59 



Bidesmida Johnson, 1976:192 

Bidesmida morrisoni Johnson, 1976:193. 

Holotype male: USA. New Mexico, Las Cru- 
ces, 30.XI.1975, W. P. Morrison (USNM). 

Distribution. — USA: NM (Johnson, 
1976:294). 

Helicoconis Enderlein, 1905b:226 

Helicoconis (Helicoconis) californica Meinan- 
der, 1972:119. 

Holotype male: USA, California, Mono Co., 1 
mile southwest of Tom's Place, 10.VIII.1963, 
M. J. Tauber & C. A. Toschi (CAS). 

DISTRIBUTION. — USA: CA (Meinander, 
1972:119). 

Helicoconis (Helicoconis) lutea (Wallengren), 
1871:55 [Coniopteiyx]. 

Type lost: Sweden, Gotland (NRS). 

Taxonomy. — To Aleuropteryx (see 
Klapalek, 1894: 12 1 ); to Helicoconis (see Ender- 
lein, 1905b:226). 

Distribution. — USA: PA (Meinander, 
1972:124). 

Helicoconis (Helicoconis) similis Meinander, 
1972:124. 

Holotype male: Canada, British Columbia, 
Top of Moyle Mountains.. 6868 ft.. East 
Kootenaya, H. B. Leech (CAS). 

Distribution. — CAN: BC (Meinander, 
1972:124). 

Helicoconis (Helicoconis) walshi (Banks), 
1906b: 83 [Aleuropteryx]. 

Holotype male: USA, Michigan, Michigan 
Agricultural College, 3.VII.1896 (MCZ). 

TAXONOMY. — To Helicoconis (see Ender- 
lein, 1908:16): to Cryptoscenea (see Enderlein, 
1930: 113); returned to Helicoconis (see Meinan- 
der, 1972:125). 

Distribution 
WS 2 ('Banks, 1906:83; 2 Meinander, 1972:125; 
3 Meinander, 1974:223; 4 Lawson & McCafferty, 
1984:130). 

Neoconis Enderlein, 1930:112 

Neoconis bifurcata Meinander, 1974:223. 

Holotype male: USA, Arizona, Cochise Co., 
Stewart Camp, 1 mile south of Portal, 
12-15.VIII.1971, J. Doyen (CAS). 



USA: CA 3 , IN 4 , ME 2 , MI 1 



DISTRIBUTION. — USA: AZ (Meinander, 
1974:223). 
Neoconis inexpectata Meinander. 1972:155. 

Holotype male: USA, Arizona, Cochise Co.. 
Southwest Research Station, 5400 ft., Chiraca- 
hua Mts.. 58.VII.1964, D. R. Davis (MZH). 

DISTRIBUTION. — USA: AZ (Meinander, 
1972:155). 
Neoconis marginata Meinander, 1972:156. 

Holotype male: USA, Texas, Kerrville, 
13.VIII.1954, L. J. Bottimer (USNM). 

Distribution. — USA: AZ, CA, TX (Mei- 
nander, 1972:157). 

CONIOPTERYGINAE 

Coniopteryx (Coniopteryx) Curtis, 1834: 
unpaginated letterpress to plate 528 

Coniopteryx (Coniopteryx) californica Mei- 
nander. 1974:225. 

Holotype male: USA, California, San Luis 
Obispo Co.. Oro Haco Lake, 5.VIII.1962, E. 
Schlinger(CAS). 

Distribution. — USA: CA 1 . TX" ('Meinan- 
der, 1974:226; 2 Meinander, 1990:61). 
Coniopteryx (Coniopteryx) dorsicornis 
Johnson, 1981b:183. 

Holotype male: USA, Florida, Highlands Co.. 
Archbold Biological Station, 

30.III-5.IV.1967, S. W. Frost (PSU). 

Distribution. — USA: FL (Johnson, 
1981b:183). 

Coniopteryx (Coniopteryx) fitchi Banks. 
1895b:315! 

Type lost: USA, Colorado (MCZ). 

= Malacomyza farinosa Banks, 1906b:85. 
Type lost: USA, California, San Mateo Co. 
(MCZ). 

Taxonomy. — To Malacomyza (see Banks, 
1906b: 84); to Semidalis (see Enderlein, 
1907:12); returned to Coniopteiyx (see Meinan- 
der, 1972:256); M. farinosa = C. fitchi (see Me- 
inander, 1972:256). 

Distribution. — USA: AZ, CA, NV, TX 
(Meinander, 1972:257). 

Coniopteryx (Coniopteryx) forcipata Johnson, 
1981b:185. 

Holotype male: USA, Arizona, Cochise 
Stronghold, Dragoon Mountains, 

20-24.VI.1970, R. J. Shaw (USNM). 



60 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Distribution. — USA: AZ (Johnson, 
1981b:185). 

Coniopteryx (Coniopteryx) latipalpis Meinan- 
der, 1972:257. 

Holotype male: USA, Colorado, Junction 
Creek Road, 10,000 ft., 12-17.VII.1968, E. C. 
Becker (CNC). 

Distribution. — USA: AZ, CA, CO (Mei- 
nander, 1972:259). 

Coniopteryx (Coniopteryx) mexicana Mei- 
nander, 1974:226. 

Holotype male: Mexico, Sinaloa, 20 miles east 
of Villa Union, 31.1.1964, E. Schlinger & M. 
Irwin (CAS). 

Distribution. — USA: TX (Meinander, 
1990:62). 

Coniopteryx (Coniopteryx) mi nut a Meinan- 
der, 1972:259. 

Holotype male: USA, California, San Luis 
Obispo Co., Morro Bay 8 miles, 2.V. 1962, C. A. 
Toschi (CAS). 

Distribution. — USA: CA (Meinander, 
1972:260). 

Coniopteryx (Coniopteryx) pal pal is Meinan- 
der, 1972:260. 

Holotype male: Mexico, Mexico, Route 15, 
km 152 west of Bosencheve, 20. VII. 1966, Flint 
& Ortiz (USNM). 

DISTRIBUTION. — USA: CA (Meinander, 
1990:62). 

Coniopteryx (Coniopteryx) quadricephaia 
Johnson, 198 lb: 188. 

Holotype male: USA, Utah, Cache Co., Green 
Canyon, 25-30.VII.1968, W.J. Hanson 
(USNM) 

Distribution. — USA: UT (Johnson, 
1981b:188). 

Coniopteryx (Coniopteryx) simplex Meinan- 
der, 1974:228. 

Holotype male: USA, California, El Dorado 
Co., Pino Grande, southwest of Lake Edson, 
8. VII. 1 967, W. J. Turner (CAS). 

Distribution. — USA: CA (Meinander, 
1974:228). 

Coniopteryx (Coniopteryx) simplicior Mei- 
nander, 1972:261. 

Holotype male: USA, Texas, Kerr Co., Ker- 
rville, VII. 1953, L. J. Bottimer (USNM). 

Distribution. — USA: AR 2 , CA~, FL 1 , 
G A 1 , IN 3 , MD 1 ^, NY 1 , TN^TX^V A ' (' Meinan- 
der, 1972:262; 2 Meinander, 1974:64; 3 Lawson& 
McCafferty, 1984:130). 



Coniopteryx (Coniopteryx) tineiformis Curtis, 
1834: text to plate 528. 

Lectotype male: England (NMV). 

= Aleyrodes dubia Stephens, 1829:367(nomen 
nudum). 

= Malacomyza lactea Wesmael, 1836:166. 
Type (sex unknown): (locality unknown) 
(IRSNB). 

= Sciodus lacteus Zetterstedt, 1840:1051. 
Type (sex unknown): Sweden, Bohuslan (NRS). 

= Sciodus fuscus Zetterstedt, 1 840: 1051. Type 
(sex unknown): Sweden. Gotland (NRS). 

= Coniopteryx tineiformis var. xaveriana 
Navas, 1918:20. Holotype female: Spain, Javier 
(MZB). 

TAXONOMY. — Originally described from a 
mixed syntype series of seven specimens, with 
the text describing one species, and associated 
plate describing another. Meinander, 1972:254 
has corrected this problem by designating a lec- 
totype. Aleyrodes dubia is a nomen nudum, with 
no description and no type designation. A. dubia 
= C. tineiformis, M. lactea = C. tineiformis (see 
Enderlein, 1905b: 197); S. lacteus = C. tineifor- 
mis, S. fuscus = C. tineiformis, S. albus = C. 
tineiformis, C. t. xaverina = C. tineiformis (see 
Meinander, 1972:256). 

Distribution. — CAN: BC 3 , MB 4 , NF 1 . 
pqVyu^usa^k'xa'jd'jn^mi^tn 1 , 

"Lawson & 



1972:254; 
I 984: 130; 3 Meinander, 1990:61 



VA ( Meinander, 

McCafferty, 

4 Penny, NEW STATE RECORD). 

Coniopteryx (Coniopteryx) westwoodii 

(Fitch), 1855:802 [Aleuronia]. 

Type lost: USA, New York 

= Malacomyza ventralis Navas, 1912a: 198. 
Type (very deteriorated): USA, New York, 
White Plains (MZB). 

= (?) Coniopteiyx gidae Procter, 1938:44 (no- 
men nudum). 

TAXONOMY. — No description or type were 
designated for C. gidae. M. ventralis = C. west- 
woodii (see Meinander, 1972:263). 



MB",. 



USA: FL' 



NY', TN C 



Distribution. — CAN: 
IN 7 ME 3 , MI 6 , MN 5 , MO 4 , 
TX , VA 6 WS 6 ('Fitch, 1855:98; 2 Smith, 
1900:54; Procter, 1946:44, as C. gidae; 
4 Froeschner, 1947:135; 5 Parfin, 1952:426; Me- 
inander, 1972:263; 7 Lawson & McCafferty, 
1984:130; 8 Penny, NEW STATE RECORD). 



PENNY, ADAMS AND STANGE: SPECIES CATALOG 



Coniopteryx (Xeroconiopteryx) Meinander, 
1972:203 

Coniopteryx (Xeroconiopteryx) canadensis 

Meinander, 1972:211. 

Holotype male: Canada, Saskatuwan, Elbow, 
10.VI. 1960, A. R. Brooks (CNC). 

DISTRIBUTION. — CAN: SA 1 , USA: AK 2 , 
WS 1 ('Meinander, 1972:212; 2 Meinander, 
1990:40). 

Coniopteryx (Xeroconiopteryx) diversicornis 
Meinander! 1972:213. 

Holotype male: USA, Texas, Kerr Co., Kerr- 
ville, 13.X.1953, L. J. Bottimer (USNM). 

DISTRIBUTION. — USA: AZ, CA, FL, TX 
(Meinander, 1972:214). 

Coniopteryx (Xeroconiopteryx) meinanderi 
Johnson, 1981b: 186. 

Holotype male: USA, Arizona, Cochise 
Stronghold, Dragoon Mountains, R. J. Shaw 
(USNM). 

Distribution. — USA: AZ 1 , CA" ( Johnson, 
1981b:186; 2 Meinander, 1990:40). 
Coniopteryx (Xeroconiopteryx) texana Mei- 
nander, 1972:214. 

Holotype male: USA, Texas, Kerr Co., Ker- 
rville, 30.VI.1953, L. J. Bottimer (USNM). 

Distribution. — USA: AZ 1 , CA 2 , TX 1 
('Meinander, 1972:215; 2 Meinander, 1990:40). 

Conwentzia Enderlein, 1905a: 10 

Conwentzia barretti (Banks), 1898:202 
[Coniopteryx]. 

Holotype female: Mexico, Amecameca 
(MCZ). 

= Parasemidalis flaviceps Banks, 1906b:81. 
Lectotype female: USA, California, Los Angeles 
(MCZ). 

Taxonomy. — To Semidalis (see Enderlein, 
1907:12); to Conwentzia (see Meinander, 
1972:295); P. flaviceps = C barretti (see Mei- 
nander, 1972:295). 

Distribution. — USA: CA (Fleschner & 
Ricker, 1953:458, as Parasemidalis flaviceps). 
Conwentzia californica Meinander, 1972:297. 

Holotype male: USA, California, Mono Co., 
Lee Vining Campground, west of Mono Lake, 
VII.1961,D. C. Rentz(CAS). 

Distribution. — CAN: BC 2 USA: AZ 1 , 
CA 1 , CO 1 , NM 3 , OR 3 , UT 1 , WA 1 , WY 1 ("Mei- 



nander, 1972:298; "Meinander, 1990:72; 
3 Penny, NEW STATE RECORDS). 
Conwentzia pineticola Enderlein, 1905a:40. 

Lectotype male: Berlin, Germany (HMB). 

= Conwentzia pineticola var. furcilla Ender- 
lein, 1906a: 194. Two syntype males: Berlin, 
Germany (probably lost). 

= Conwentzia pineticola var. tetensi Enderlein, 
1906a: 195. Lectotype female: Finland, Jakob- 
stad (MZH). 

= Conwentzia hageni Banks, 1906b:82. Lecto- 
type female: USA, Virginia (MCZ). 

= Coniopteryx reticulata Tullgren, 1906:14. 
Syntypes male and female: Sweden, Stockholm 
(depository unknown). 

= Conwentzia angulata Navas, 19 14b: 16. 
Holotype female: USA, New York (MZB). 

= Conwentzia axillata Navas, 1 9 1 4b: 1 7. Holo- 
type male: USA, New York (MZB). 

= Conwentzia cnptoneuris Bagnall, 1915: 192. 
Syntype male, 2 females: England, Newcastle- 
upon-Tyne (depository unknown). 

TAXONOMY. — C. p. war. furcilla = C pineti- 
cola, C. p. var. tetensi = C. pineticola, C. hageni 
= C pineticola, C. angulata = C pineticola, C 
axillata = C pineticola; lectotypes designated for 
C. pineticola and C hageni (see Meinander, 
1972:301). 

Distribution. — CAN: MB 9 , NF 6 , NS 6 , 

ON 6 , USA: AR 7 , DC 6 , FL 6 , LA 9 IN 8 , MA 6 , 

MD 6 , ME 3 , MI 6 , MN 5 , MO 4 , NC 6 , NH 6 , NJ 6 , 

NY 2 , OH 6 , PA 6 , VA 1 , WS 6 ('Banks, 1906b:82; 
2 3 

"Navas, 19 14b: 16, as C. angulata; "Procter, 

1946:44, as C. hageni; Froeschner, 1947: 135, as 
C. hageni; ~ Parfin, 1 952:426, as C. hageni; Me- 
inander, 1972:303; 7 Throne, 1972:126; 8 Lawson 
& McCafferty, 1984:130; 9 Penny, NEW 
STATE RECORDS). 

Conwentzia psociformis (Curtis), 1834: plate 
528 [Coniopteryx]. 

Type deposition unknown: England, specimen 
presumed lost. 

= Coniopteryx aphidiformis Rambur, 
1842:316. Type depository unknown. 

TAXONOMY. — To Coniortes (see Walker, 
1853:299); to Conwentzia (see Enderlein, 
1905:10); C aphidiformis = C psociformis (see 
Meinander, 1972:305). 

Distribution. — CAN: BC, USA: NJ, OH 
(Meinander, 1972:306). 



62 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Parasemidalis Enderlein, 1905c: 197 

Parasemidalis fuscipennis (Reuter), 1894:13 
[Coniopteryx]. 

Holotype male: Finland (MZH). 

= Parasemidalis annae Enderlein, 1905c: 198. 
Holotype female: Germany. Berlin (HMB). 

TAXONOMY. — To Parasemidalis (see Ender- 
lein, 1906:219); P. annae = P. fuscipennis (see 
Meinander, 1972:281). 

Distribution. — USA: AZ, CA, FL, MI 
(Meinander, 1972:283). 

Semidalis Enderlein, 1905c: 197 

Semidalis angusta (Banks), 1906b:81 [Coniop- 
teryx]. 

Lectotype male: USA, Arizona, Williams, 
29.V, Barber & Schwarz (USNM). 

TAXONOMY. — To Semidalis (see Enderlein, 
1908:1 1); lectotype designated (see Meinander, 
1972:327). 

Distribution. — USA: AR, AZ, CA, MT, 
TX (Meinander, 1972:327). 
Semidalis arnaudi Meinander, 1972:328. 

Holotype male: Mexico, Sonora, Alamos, 
27.11.1963, P. H. Arnaud (CAS). 

DISTRIBUTION. — USA: AZ (Meinander, 
1990:78). 
Semidalis bituberculata Meinander, 1990:79 

Holotype male: USA, California, Riverside 
Co., hills on west side of Menifee Valley, 
27.VII.1980, J. D. Pinto (CAS). 

Distribution. — USA: CA (Meinander, 
1990:79). 
Semidalis deserta Meinander, 1974:230. 

Holotype male: USA, California, Riverside 
Co., Boyd Desert Research Center, 3 miles south 
of Palm Desert, 24-26.VI.1969, S. Frommer & 
B. Worley (CAS). 

Distribution. — USA: CA (Meinander, 
1974:231). 
Semidalis flinti Meinander, 1972:335. 

Holotype male: USA, Texas, Ken - Co., Ker- 
rville, 17.VIII.1954, L. J. Bottimer (USNM). 

Distribution. — USA: AZ 1 , CA 2 , TX 1 
('Meinander, 1972:336; 2 Meinander, 1974:231). 
Semidalis frommeri Meinander, 1974:231. 

Holotype male: USA, California, Riverside 
Co., Deep Canyon, 22.VII.1964, M. Irwin 
(CAS). 



Distribution. — USA: CA (Meinander, 
1974:232). 
Semidalis inconspicua Meinander, 1972:336. 

Holotype male: USA, Virginia, Falls Church, 
Holmes Run, 6.VI.1961, W. W^ Wirth (USNM). 

Distribution. — USA: AR 2 , AZ 1 , CA 1 , IN 3 , 
MD 1 , Ok' v TX 1 , VA 1 , WS 1 ('Meinander, 
1972:336; 2 Meinander, 1974:232; 3 Lawson & 
McCafferty, 1984:130). 
Semidalis tricornis Johnson, 1981b: 190. 

Holotype male: USA, Arizona, Cochise 
Stronghold, Dragoon Mountains, R. J. Shaw 
(USNM). 

Distribution. — USA: AZ (Johnson, 
1981b:191). 

Semidalis vicina (Hagen), 1861:197 [Coniop- 
teryx]. 

Holotype female: USA, Washington, D. C. 
(MCZ). 

= Parasemidalis nigriceps Navas, 1918a:355. 
Holotype female: USA, New York, White 
Plains, 18.VII.1915(MZB). 

= Semidalis ribesi Ohm, 1973:237. Holotype 
male: France, Pyrenees orientales, above the 
Bains du Boulou, 27.V.1963 (Ohm Collection). 

Taxonomy. — To Semidalis (see Enderlein, 
1906:215); to Niphetia (see Enderlein, 
1930:106); returned to Semidalis (see Procter, 
1938:44); P. nigriceps = S. vicina, S. ribesi = S. 
vicina (Meinander, 1972:330). 

Distribution. — CAN: AB 8 , PQ 8 USA: 
CT 8 DC 1 FL 8 , GA 8 IN 9 KS"\ ME 5 MO^ 
MN , MA 8 , ME 8 , MI 8 ,MS , NC 4 , NH 8 , NY 2 , 
OH 8 , PA 8 ,^SC 8 , TN 8 , VA 8 , WS 8 ('Hagen, 
1861:197; 2 Navas, 1918:19, as P. nigriceps; 
3 Smith, 1925:170; 4 Brimley, 1938:32; Procter, 
1946:44; 6 Froeschner, 1947:135, 7 Parfin, 
1952:425; 8 Meinander, 1972:331; Xawson & 
McCafferty, 1984:130). 
Semidalis wallacei Meinander, 1972:337. 

Holotype male: USA, Pennsylvania, Pitts- 
burgh, 15.VII.1932, H. Kahl (CMNH). 

DISTRIBUTION. — USA: PA (Meinander, 
1972:338). 
Semidalis xerophila Meinander, 1990:84. 

Holotype male: USA, California, Riverside 
Co., south of Palm Desert, Boyd Desert Research 
Center, 19-21. VI. 1978 (CAS). 

DISTRIBUTION. — USA: CA (Meinander, 
1990:84). 



PENNY, ADAMS AND STANGE: SPECIES CATALOG 



63 



DILAR1DAE 

The dilarids, or pleasing lacewings, comprise 
a small family of tropical, moth-like lacewings 
which are rarely seen in North America. This 
family has broad, darkly-banded wings with con- 
siderable long pilosity on the wings, giving them 
a "hairy" appearance. Males have distinctive 
pectinate antennae. However, unlike most moths, 
the eyes are very large in proportion to the rest of 
the head. Females bear ovipositors which are 
recurved over the abdomen. Larvae are very 
elongate, and live under the bark of dead trees, 
where they are presumed to feed on beetle adults 
or larvae (Gurney, 1947). Tropical American 
species appear to be most abundant in the forest 
canopy as adults, and seem to emerge most fre- 
quently during the driest part of the year (Penny 
and Arias, 1981). Only two species are known 
from the U. S., both in the genus Nallachius 
Navas. 1909a, and although one species is rather 
widely distributed over eastern North America, 
it is seldom collected. A key to New World 
species can be found in Adams (1970). 

Nallachius Navas, 1909a:666 

Nallachius americanus (McLachlan), 1881:55 
[Dilar]. 

I lolotype female: USA, Kentucky, Bee Spring, 
VI. 1874, Sanborn (MCZ). 

TAXONOMY. — To Nallachius (see Navas, 
1909a:669). 

DISTRIBUTION. — USA: GA 5 , IN 4 , KY 1 , 
MD 2 , SC 5 , TX 3 . VA 2 ('McLachlan, 1881:55; 
2 Carpenter, 1940:274; 3 Stange, 1961b: 144; 
4 Lawson & McCafferty. 1984:130; 5 Hoffman, 
1990:155). 

Nallachius pulchellus (Banks), 1938c:289 [Di- 
lar]. 

Male and female syntypes: Cuba, Soledad 
(Cienfuegos), 4.V.1930, P.J. Darlington, Jr. 
(MCZ). 

Taxonomy. — To Nallachius (see Carpenter, 
1940:274). 

Distribution. — USA: AZ (Carpenter, 
1940:275). 

HEMEROBIIDAE 
Hemerobiidae, or brown lacewings, comprise 
one of the more commonly encountered families 
of Neuroptera. Adults are predominantly brown 



or black in coloration. They can be characterized 
as having the Rs and MA fused with R in the 
forewing to give the appearance of having mul- 
tiple radial sectors. The number of these apparent 
radial sectors is important in recognizing individ- 
ual genera. They are rather small insects, and 
individuals of the genus Sympherobius can have 
forewing length 2—3 mm. Adults are most abun- 
dant on trees and bushes, although some groups, 
most notably Micromus, can be found on grasses. 
Both adults and larvae feed on soft-bodied in- 
sects. The genus Sympherobius was revised by 
Oswald (1988), and he has dealt with the family 
level classification (Oswald, 1993). The Cana- 
dian species have been redescribed by Kevan and 
Klimaszewski (1986, 1987) and Klimaszewski 
and Kevan (1985, 1987a, 1987b. 1988a, 1988b, 
1989. 1990a, 1990b). 

Hemerobiinae 

Hemerobius Linnaeus, 1758:549 

Hemerobius alpestris Banks, 1908a:261. 

Holotype male: USA, Colorado, Sugar Loaf 
Mountain, 13.V, S. A. Rohwer (MCZ). 

Distribution. — USA: AZ", CO r ('Banks, 
1908a:261; 2 Carpenter, 1940:208). 
Hemerobius bistrigatus Currie, 1904a:79. 

Holotype female: USA, California. Humboldt 
Co., Little River, 31. V. 1903 (USNM). 

Distribution. — CAN: BC 2 , USA: CA 1 , 
ID 2 , OR 2 (' Currie, 1904a:79; 2 Carpenter. 
1940:208). 
Hemerobius conjunctus Fitch, 1855:798. 

Holotype male: Kelson's Cabin, 7.VII (MCZ). 

= Hemerobius citrinus Hagen, 1861:204. 
Holotype (male): America Septentrionalis. 
Knoch (HMB). 

= Hemerobius venustus Banks, 1897a:25. 
Holotype female: USA, New Hampshire, Mount 
Washington, N. Banks (MCZ). 

= Hemerobius cockerelli Banks, 1901b:286. 
Holotype male: USA, New Mexico, Las Vegas 
Range, 1 1000 ft., Cockerell (MCZ). 

= Hemerobius caudelli Currie, 1904b:87. 
Holotype female: Canada, British Columbia, 
London Hill Mine, Bear Lake, 7000 ft., 
29.VII. 1903 (USNM). 

= Hemerobius glacialis Currie, 1904b:88. 
Holotype female: Canada, British Columbia. 



64 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Kokanee Mountain, 9000 ft., 10.VIII.1903 
(USNM). 

Taxonomy. — H. venustus = H. conjunctus, 
H. caudelli = H. cockerelli (see Banks, 
1905c:49); H. citrinus = H. conjunctus, H. 
glacial is = H. conjunctus, H. cockerelli = H. 
conjunctus (see Carpenter, 1940:209). Carpen- 
ter, 1940:209 considered the single Fitch speci- 
men of//, conjunctus at MCZ to be the holotype. 
Klimaszewski andKevan (1985:40) rejected this 
point of view and designated this specimen as 
lectotype. As there is no evidence of additional 
specimens in the original type series, we agree 
with Carpenter and herein list it as holotvpe. 

Distribution. — CAN: AB 6 , BC, LB 6 , 
MB 10 , NF 6 , NS 6 , ON 6 , PE 6 , PQ 6 SK 6 YT 10 , 
USA: AK 6 , CO 6 , IN 9 , ME 6 , MN, MT 6 , NC 5 , 
NH ', NM 2 , NY 4 , TN 6 , VT 6 , WS 8 , WY 6 ( 'Banks, 
1897a:25,as//. venustus; 2 Banks, 1901b:286,as 
H cockerelli; 3 Currie 1904b: 87, as H. caudelli; 
4 Leonard, 1928:39- 5 Brimley, 1938:29; Car- 
penter, 1940:209; Parfin, 1952:423; 8 Throne, 
1971b:80; 9 Lawson & McCafferty, 1984:130; 
10 Klimaszewski & Kevan, 1985:39). 
Hemerobius costalis Carpenter, 1940:2 13. 

Holotype male: USA, Maine, Eastport, 
14.VII.1909, C. W. Johnson (MCZ). 

DISTRIBUTION. — CAN: AB 1 , BC 1 NS 1 , 
NT 1 , ON 1 PQ 1 , SK 3 , YT 2 , USA: AK 1 , CO 1 , 
MA 1 , ME 1 , NH 1 , VT 1 (' Carpenter, 1940:213; 
2 Klimaszewski & Kevan, 1985:69; kli- 
maszewski & Kevan, 1989:205). 
Hemerobius discretus Navas, 19 1 7a:5. 

Lectotype female: USA, New Mexico, Jemez 
Springs, 16.VI.1916 (MZB). 

= Hemerobius neglectus Hagen, 1861:206. 
Type(s) unknown: Mexico, Ehrenberg (ZMB). 

= Hemerobius mexicanus Navas, 1 92 1 : 1 23. 

= Hemerobius pallidulus Kimmins, 1928:366. 

= Hemerobius neadelphus Gurney, 1948:214. 
Holotype male: Canada, British Columbia, 
Kaslo, 4.VII. 1903, R. P. Currie (USNM). 

Taxonomy. — H. discretus = H. pacificus 
(see Carpenter, 1940:204).//. discretus not equal 
H. pacificus; H. mexicanus = H. discretus, H 
neglectus = H discretus, H. pallidulus = H 
discretus (see Monserrat, 1996:416-417). //. 
neadelphus = H. discretus (Penny, NEW SYN- 
ONYMY). 

Distribution. — CAN: BC 1 , USA: AZ 3 
CA 2 , CO 3 , LA 3 , NM 3 , OK 3 , OR 1 , TX 3 , UT 2 



( Gurney 1948:214; Klimaszewski & Kevan, 
1985:25; 3 Monserrat, 1996:421). 
Hemerobius dorsatus Banks, 1904a:61. 

Lectotype male: USA, Colorado, Veta Pass, 
1. VII (MCZ). 

Taxonomy. — Lectotype designated (see 
Carpenter, 1940:211). 

DISTRIBUTION. — CAN: AB 2 , BC 2 , NT 2 
ON 3 , PQ 3 , SA 2 , YT 3 , USA: AK 2 , CO 1 , UT" 
(' Banks, 1904a:61; 2 Carpenter, 1940:212; kli- 
maszewski & Kevan, 1985:66). 
Hemerobius humulinus Linnaeus, 1758:550. 

Type lost. 

= Hemerobius crispus Stephens, 1836:112. 
Type lost: England. 

= Hemerobius maculatus Wesmael, 1 84 1:215. 
Type(s) (unknown): Belgium, Brussels 
(IRSCB). 

= Hemerobius obliteratus Walker, 1853:289. 
Holotype male: USA, Georgia (BMNH). 

= Hemerobius castaneae Fitch, 1855:798. 
Holotype female: USA (MCZ). 

= Hemerobius matrix Fitch, 1855:798. Holo- 
type male: USA (MCZ). 

= Hemerobius gossypii Ash mead, 1895:27. 
Lectotype female: USA, Georgia (BMNH). 

= Hemerobius algonquinus Banks, 1924:429. 
Lectotype female: USA, New Hampshire, 
Hampton, 6.III.1904, S. A. Shaw (MCZ). 

= Hemerobius obtusus Nakahara, 1954:42. 
Holotype male: Japan, Mejiro, Tokyo (Nakahara 
Collection). 

= Hemerobius shikotanus Kuwayama, 
1956:77. Holotype female: Kurile Islands, Shik- 
otan Island, Shakotan, 23-27.VI.1935, Y. Sugi- 
hara (Kuwayama Collection). 

Taxonomy. — H. humuli = H. humulinus, H. 
maculatus = H. humulinus, H. castaneae = //. 
humulinus, H. tutatrix = H. humulinus, H. 
gossypii = H. humulinus, H. algonquinus = //. 
humulinus, H. obliteratus = H. humulinus (see 
Carpenter, 1940:201); H. shikotanus = H. hu- 
mulinus (see Makarkin, 1985:105). 

Distribution. — CAN: AB 1 ;, BC 6 , MB 6 , 
NF 6 , NS 6 ON 6 , PQ 6 SK 6 YT 12 , USA: AK 10 , 
CO 6 , DC 3 , FL 6 GA 1 IN 4 , KS 3 , MD 13 , ME 7 , 
MI 3 , MN 9 , MO B , NC 5 ND 6 ,^NH 3 , NJ 2 , NY 2 , 
PA 3 , SD 6 , TN 6 , TX 6 , VA 2 , WA 10 , WS 11 
( ' Walker, 1 853:289; 2 Smith 1 900:56, as H cas- 
tanea and //. tutatrix; Banks, 1905c:32; 
Montgomery & TrippeL 1933:259; Brimley, 
1938:29, as H. humuli; Xarpenter, 1940:202; 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



65 



7 Procter, 1946:43; 8 Froeschner, 1947:131; 9 Par- 
fin, 1952:423; 10 Nakahara, 1966:212; "Throne, 
1971b:80; 12 Klimaszewski & Kevan, 1985:56; 
13 Monserrat, 1996:429). 
Hemerobius kokaneeanus Currie, 1904b:86. 

Holotype female: Canada, British Columbia, 
Kokanee Mountain, 9000 ft., 10.VIII.1903 
(USNM). 

= Hemerobius hesperus Banks, 1924:429. 
Lectotype female (abdomen missing): USA, 
California, Tahoe, Angora Pk., 10.VII.1915, E. 
P. Van Duzee (MCZ). 

Taxonomy. — H. hesperus = H. kokanee- 
anus (see Carpenter, 1940:206); lectotype desig- 
nated for H. hesperus (see Klimaszewski & 
Kevan, 1985:30). 

Distribution. — CAN: ^BC'^MB 4 ?q\ 
YT 3 , USA: AK 5 , CA 2 , CO 2 , MT 2 , NV 5 , WA" 
('Currie, 1904b:85; 2 Carpenter, 1940:206; kli- 
maszewski & Kevan, 1985:32; Klimaszewski & 
Kevan, 1989:205; 5 Penny, NEW STATE RE- 
CORDS) 
Hemerobius nigrans Carpenter, 1940:207. 

Holotype male: Canada, British Columbia, 
Kamloops, Mount Lobo, 2.VI.1938, J. K. Jacob 
(CNC). 

Distribution. — CAN: BC 1 , YT 2 , USA: 
CA 1 , CO 1 , WY 1 ('Carpenter, 1940:207; kli- 
maszewski & Kevan, 1985:36). 
Hemerobius ovalis Carpenter, 1940:205. 

Holotype male: USA, California, Claremont, 
C.F.Baker (MCZ). 

Distribution. — CAN: AB 1 , BC 1 , NF 2 , 
PQ 2 YT 2 , USA: AK 1 , CA 1 OR 1 , UT 1 , WA 1 , 
WY 1 , ('Carpenter, 1940:205; 2 Klimaszewski & 
Kevan, 1985:28). 
Hemerobius pacificus Banks, 1897a:24. 

Lectotype male: USA, Washington, Olympia, 
T. Kincaid (MCZ). 

= Hemerobius pallescens Currie, 1904a: 80. 
Holotype female: USA, California, Humboldt 
Co., Fieldbrook, 30.V.1903 (USNM). 

TAXONOMY. — H. pallescens = H. pacificus 
(see Banks, 1905c:49). Lectotype designated for 
H. pacificus (see Carpenter, 1940:204). 

Distribution. — CAN: AB 4 , BC 4 , MB 5 , 
SK 4 USA: AK 6 , AZ 4 , CA 2 , CO 4 , ID 4 IN 5 , MT 8 , 
NM 3 , NV 7 y OR 4 , TX 4 , UT 8 WA 1 ('Banks, 
1897a:24; 2 Currie, 1904a:80; 3 Navas, 1917a:5; 
4 Carpenter, 1940:204; 5 Lawson & McCafferty 
1984:130; 6 Klimaszewski & Kevan, 1985:23; 



Monserrat, 1996:415; 8 Penny, NEW STATE 
RECORDS). 

REMARKS. — The Indiana record is probably 
in error. However, one of us (NDP) has seen a 
series of specimens of the closely related H. 
neadelphus labelled from northwestern Minne- 
sota. 
Hemerobius pinidumus Fitch, 1855:799. 

Holotype male: no locality data (MCZ). 

= Hemerobius hyalinatus Fitch, 1855:799. 
Holotype male: no locality data (MCZ). 

= Hemerobius canadensis Banks, 1897a:26. 
Holotype female: Canada, Quebec, Sherbrooke, 
October, L'Abbe, P. A. Begin (MCZ). 

= Hemerobius kootcnarensis Currie, 
1904b:88. 

Holotype female: Canada, British Columbia, 
Kaslo, 17. VI. 1903 (USNM). 

TAXONOMY. — H. canadensis = H. hyali- 
natus) (see Banks, 1905c:34); H. kootenayensis 
= H. conjunctus (see Carpenter, 1940:209); H. 
hyalinatus = H. conjunctus var. pinidumus) (see 
Carpenter, 1940:211); H. kootenayensis = H. 
pinidumus) (see Klimaszewski & Kevan, 
1985:41); H. pinidumus variety of//, conjunctus 
(see Carpenter, 1940:209); H. pinidumus is valid 
species (see Throne, 1971 b:81). 

DISTRIBUTION. — CAN: AB 6 , BC 3 , PQ 1 , 
USA: CO 4 , IN 5 ,MA 4 , ME 4 , MN 4 , NH 4 , NJ 2 , 
NY 2 . WS 4 ('Banks, 1897a:26, as H canadensis; 
"Smith 1900:56, as H. pinidumus and H. hyali- 
natus; Currie, 1904b:88, as H. kootenayensis; 
4 Carpenter 1940:21 1; 5 Lawson & McCafferty, 
1984:130; 6 Klimaszewski & Kevan, 1985:42). 
Hemerobius simulans Walker, 1853:285. 

Holotype male: Canada, Hudson's Bay, St. 
Martin's Falls, Albany River (BMNH). 

= Hemerobius orohpus Wallengren, 
1870:155. Syntypes (number unknown): Swe- 
den, regions of Dalarne, Norbotten and Vastbot- 
ten; one syntype in BMNH, others may be in ZIL 
andNRS. 

= Hemerobius nevadensis Banks, 1904a:61. 
Holotype female: USA, Nevada, Ormsby Co.. 
3. VII. 1903, Baker (MCZ). 

= Hemerobius placidus Banks, 1908a:260. 
Lectotype male: USA, New York, Lake Placid, 
12.VIII.1904, E. C. Van Duzee (MCZ). 

= Hemerobius piceus Navas, 1925e:2. Three 
male, four female syntypes: Russia, Kamchatka 
Peninsula (NRS). 



66 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Taxonomy. — H. orotypus = H. simulans 
(see Kimmins, 1932:88); H.placidus = H. simu- 
lans, H. nevadensis=H. simulans (see Carpenter, 
1940:212); H. piceus = H. simulans (see Tjeder, 
1960:148); lectotype designated for H. placidus 
(see Klimaszewski & Kevan, 1985:60). 

Distribution. — CAN: n AB 3 , BC 3 MB 5 , 
NB 3 , NF 3 , NS 5 ON 3 PQ 3 SK 5 YT 5 , USA: 
AK 3 , CO 4 , MA 3 , ME 3 , MI 3 , NH 3 , NV 1 , NY 2 
('Banks, 1904a:61; 2 Banks, 1908a:260; Car- 
penter, 1940:212; ^akahara, 1966:213; kli- 
maszewski & Kevan, 1985:63). 
Hemerobius stigma Stephens, 1836:112. 

Lectotype (without abdomen): England, near 
Ripley (BMNH). 

= Hemerobius irroratus Stephens, 1836:111. 
Lectotype (without abdomen): England, London 
metropolitan district (BMNH). 

= Hemerobius strigosus Zetterstedt, 
1840:1049. Type(s) (unknown): Lapponia-Up- 
land-Ostrogoth-cania (ZIL). 

= Hemerobius limbatellus Zetterstedt, 
1840:1050. Type(s) (unknown): Lapponia 
meridional (ZIL). 

= Hemerobius limbatus Wesmael, 1841:215. 
Type(s) (unknown): Belgium, Brussels 
(IRSNB). 

= Hemerobius crispus Stephens, 1836:112. 
Single remaining syntype male: North America 
(BMNH). 

= Hemerobius stigmaterus Fitch, 1855:797. 
Lectotype male: no locality data (MCZ). 

= Hemerobius moestus Banks, 1897a:25. 
Holotype female: USA, Washington, Olympia, 
T. Kincaid (MCZ). 

= Hemerobius dyari Currie, 1904b:85. Holo- 
type female: Canada, British Columbia, Kaslo, 
17.VII.1903(USNM). 

= Hemerobius simplex Banks, 1905c:32. Lec- 
totype male: USA, Arizona, Prescott, 1 8. V. 1902, 
Oslar (MCZ). 

= Hemerobius buyssoni Navas, 1909b:217. 
Type(s) (unknown): France, La Bouboule (Puy- 
de-D6me) (MNHN). 

= Hemerobius periphericus Navas, 1 9 1 3d: 8 1 . 
Type(s) (unknown): Spain, Escorial (depository 
of type unknown). 

TAXONOMY. — H. crispus = H. stigmaterus, 
H. moestus = H. stigmaterus, H. dyari = H. 
stigmaterus, H. simplex = H. stigmaterus) (see 
Carpenter, 1940:202); H. irroratus = H. stigma, 
H. strigosus = H. stigma, H limbatellus = H 



stigma, H. limbatus = H. stigma, H. stigmaterus 
= H. stigma, H. buyssoni = H. stigma, H. 
periphericus = H. stigma (see Klimaszewski & 
Kevan, 1985:44); lectotype designated for H 
stigmaterus (see Carpenter, 1940:202); lectotype 
designated for H. simplex (see Klimaszewski & 
Kevan, 1985:46). 

Distribution. — CAN: AB 11 , BC 3 , LB 6 , 
MB 11 , NF 11 , NS 11 , ON 11 , PQ 11 , SK 11 YT 1 ', 
USA: AK 12 , AZ 3 , CA 3 . CO 3 , DC 3 , FL, IA 3 , 
ID 3 IN 4 KS 3 ^MD 6 ME 7 ^ MI 3 MN 3 MO 3 
NC 5 , NJ-, NM 3 , NV, NY-, TX , WA, WS g 
('Banks, 1897a:25; 2 Smith, 1900:56, as H. stig- 
materus; Banks, 1905c:49; Montgomery & 
Trippel, 1933:259; 5 Brimley, 1938:29, as//, stig- 
materus; Carpenter, 1940:203; Procter, 
1946:43; 8 Throne, 1971b:79; Vgnew et al., 
1981:16; l0 MacLeod & Stange, 1981:1; n Kli- 
maszewski & Kevan, 1985:49; Klimaszewski 
& Kevan, 1989:205). 

REMARKS. — Hemerobius californicus 

Banks, 1905a:90 (nomen nudum). 



Wesmaelius Kriiger, 1922:170 

Wesmaelius brunneus (Banks), 1920:333 
[Boriomyia]. 
Holotype female: USA, Montana, Midvale, C. 

E. Brown (MCZ). 

TAXONOMY. — To Kimminsia (see Carpenter, 
1940:221); to Wesmaelius (see Kevan & Kli- 
maszewski, 1986:19). 

Distribution. — CAN: AB 2 , BC 4 YT 4 , 
USA: AK 3 , CA 2 , CO 2 , MT 1 , UT 3 , WA", WY 2 
('Banks, 1920:333. 2 Carpenter, 1940:222; 3 Par- 
fin, 1956:208; 4 Klimaszewski & Kevan, 
1987b:256). 

Wesmaelius coloradensis (Banks), 1897a:26 
[Hemerobius]. 

Lectotype male: USA, Colorado, Fort Collins, 

F. C. Baker (MCZ). 

TAXONOMY. — To Boriomyia (see Banks, 
1905c:38); to Kimminsia (see Carpenter. 
1940:217); to Wesmaelius (see Kevan & Kli- 
maszewski, 1986:19); lectotype designated (see 
Carpenter, 1940:217). 



Distribution. — CAN: AB~^ BC% YT. 
USA: CA 2 , CO 1 , NV 3 , OR 2 , UT", WA J , WY" 
(' Banks, 1897a:26; 2 Carpenter, 1940:217; 3 Par- 



fin, 1956:208; 
1987b:252). 



Klimaszewski & Kevan, 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



67 



Wesmaelius constrictus (Parfln), 1956:203 
[Kimminsia], 

Holotype male: USA, Alaska, 10-15 miles 
below Gulkana Lake along Gulkana River, 
27.VI-20. VII. 1955, G. O. Schumann (USNM). 

= Kimminsia olympica Nakahara, 1966:216. 
Holotype male: USA, Washington, Deer Park. 
5,400 ft., 17.VII.1948, C. P. Alexander 
(USNM). 

TAXONOMY. — To Wesmaelius: K. olympica 
= W. constrictus (see Kevan & Klimaszewski, 
1986:19). 

DISTRIBUTION. — USA: AK 1 , WA 2 ('Parfin, 
1956:203; 2 Nakahara, 1966:216, as A', olym- 
pica). 

Wesmaelius fumatus (Carpenter), 1940:225 
[Kimminsia]. 

Holotype male: USA, Colorado, Boulder Co., 
Long's Peak Inn, 9000 ft., 13.VII.1926, E. C. 
Van Dyke (CAS). 

TAXONOMY. — To Wesmaelius (see Kevan & 
Klimaszewski, 1986:19). 

DISTRIBUTION. — CAN: ^AB 2 , USA: CO 1 
('Carpenter, 1940:225; 2 Klimaszewski & 
Kevan, 1987b:258). 

Wesmaelius furcatus (Banks), 1935:55 
[Boriomyia]. 

Holotype male: USA. Colorado. Argentine 
Pass, T. D. A. Cockerell (MCZ). 

TAXONOMY. — To Kimminsia (see Caipenter, 
1940:219); to Wesmaelius (see Kevan & Kli- 
maszewski, 1986:19). 

Distribution. — CAN: AB 2 , BC 3 MB 3 , 
NT 3 , YT 3 , USA: AK 3 , CA 3 , CO 1 ('Banks, 
1935:55; 2 Carpenter, 1940:219; 3 Klimaszewski 
& Kevan, 1987b:254). 

Wesmaelius involutus (Carpenter), 1940:21 
[Kimminsia]. 

Holotype male: Canada, Alberta, Banff, Run- 
die Mountain, 5000-7000 ft., 25.VI. 1925, Owen 
Bryant (MCZ). 

TAXONOMY. — To Wesmaelius (see Kevan & 
Klimaszewski, 1986:19). 

DISTRIBUTION. — CAN: AB 1 , BC 1 , NF 3 , 
NS 3 , NT 3 , PQ 3 , YU 1 , USA: AK 2 , CO 1 , ID 1 . 
UT 2 , WY 1 ('Carpenter, 1940:220; 2 Parfin, 
1956:209; 3 Klimaszewski & Kevan, 1987a: 175). 
Wesmaelius longifrons (Walker), 1853:291 
[Hemerobius]. 

Holotype female: Canada. Ontario, Albany 
River, St. Martin's Falls (BMNH). 



= Hemerobius alternatus Fitch, 1855:797. 
Holotype female: USA, New York (MCZ). 

= Hemerobius transvcrsus Banks, 1904a:61. 
Holotype male: USA, Colorado, Denver (MCZ). 

= Allotomyia borealis Banks, 1935:56. Holo- 
type male: USA, New Hampshire, Hillsboro, 
11. VI (MCZ). 

TAXONOMY. — To Boriomyia (see Banks. 
1905c:37); to Wesmaelius (see Caipenter, 
1940:226). 

Distribution. — CAN: AB\ BC 3 MB 3 , 
NB 5 , NF 5 , NS 3 , ON 1 , PQ 3 , SK J YT 5 , USA: 
AK 3 , AZ 3 , CO 3 , ME 3 , MI 3 , MN 4 NH J , NY 2 , 
OR 3 . UT 3 , VT 3 , WA 3 , WY 3 ('W alker < 
1853:291; 2 Smith, 1900:55; Carpen- 
ter, 1940:226; 4 Parfin, 1952:423; 5 Klimaszewski 
& Kevan, 1987b:266). 

Wesmaelius longipennis (Banks), 1920:333 
[Boriomyia]. 

Holotype female: USA, California, Alameda 
Co.. Berkeley. 1 .IV. 19 15 (MCZ). 

TAXONOMY. — To Kimminsia (see Carpenter, 
1940:221); to Wesmaelius (see Klimaszewski & 
Kevan, 1987a: 163). 

Distribution. — USA: CA (Banks, 
1920:333). 

Wesmaelius nervosus (Fabricius). 1793:85 
[Hemerobius]. 

Two syntypes of unknown sex: France (Kiel). 

= Hemerobius betulinus Strom, 1788:387 (no- 
men dubium). Type(s) lost. 

= Hemerobius nebulosus Stephens, 1836:107. 
Type(s) (sex unknown): England, presumably 
near London (depository unknown). 

= Hemerobius conspersus Burmeister, 
1839:974. Type(s) (sex unknown): Germany, 
Halle (Halle). 

= Mucropalpus distinctus Rambur. 1842:421. 
Type(s) (sex unknown): Spain, Andalusia (de- 
pository unknown). 

= Hemerobius disjunctus Banks, 1897:25. 
Lectotype female: USA, New Hampshire, Mount 
Washington (MCZ). 

= Hemerobius frostinus Navas. 1933b: 109. 
Type(s) (sex unknown): USA, Massachusetts, 
Framingham (depository unknown). 

= Kimminsia alexanderi Nakahara, 1966:217. 
Holotype male: USA, Alaska, Haines Highway. 
M. P., 5.VII. 1952, C. P. Alexander (USNM). 

= Kimminsia melaleuca Nakahara, 1966:219. 
Holotype female: Canada, Yukon Territory, 



68 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Alaska Highway, M. P. 320, 2.VII.1952, C. P. 
Alexander (USNM). 

Taxonomy. — To Boriomyia (see Banks, 
1905c:29); to Kimminsia (see Hale Carpenter, 
1938:532); to Wesmaelius (see Tjeder, 1967:5); 
H. conspersus = H. nervosus (see Wallengren, 
1 87 1 :37); H. nebulosus = B. betulina, M. distinc- 
tus = B. betulina (see Killington, 1937:5); H. 
disjunctus = W. nervosus, H. frostinus = W. ner- 
vosus, K. alexanderi = W. nervosus, K. melaleuca 
= W. nervosus) (Kevan & Klimaszewski, 
1986: 1 1 ); lectotype designated for H. disjunctus 
(see Klimaszewski & Kevan, 1987a: 169). 

DISTRIBUTION. — CAN: BC 5 , MB 4 , NB 8 , 
NF 8 , NS 4 , NT 8 . ON 4 , PQ 4 , SK 4 , USA: AK 4 , 
CO 4 , ID 4 , IN 7 MA 3 , ME 4 . MI 5 , MT 4 , NC 4 , 
NH 1 , NY 2 , OR 5 , RT 5 , UT 5 , WS 6 , WY 4 ('Banks, 
1897:25; 2 Leonard, 1928:40; 3 Navas, 
1933b: 109; 4 Carpenter, 1940:216; 5 Parfin, 
1956:208, as K. disjuncta; 6 Throne, 197 lb: 84, as 
K. disjuncta; 7 Lawson & McCafferty, 1984: 130, 
as K. disjuncta; Klimaszewski & Kevan, 
1987a: 172). 

Wesmaelius posticatus (Banks), 1905c:39 
[Boriomyia]. 

Holotype female: USA, south Utah, VII. 1900 
(MCZ). 

Taxonomy. — To Kimminsia (see Carpenter, 
1940:218); to Wesmaelius (see Kevan & Kli- 
maszewski, 1986:19). 

DISTRIBUTION. — CAN: AB 2 , YT 3 , USA: 
AZ 2 , CA 3 , CO 2 , NM 2 , UT 1 , ('Banks, 1905c:39; 
2 Carpenter, 1940:219; 3 Nakahara, 1966:217). 
Wesmaelius pretiosus (Banks), 1908a:260 
[Boriomyia]. 

Lectotype male: USA, Colorado, Golden, 
22.VII. 1907, Oslar (MCZ). 

TAXONOMY. — To Kimminsia (see Carpenter, 
1940:223); to Wesmaelius (see Kevan & Kli- 
maszewski, 1986:19; lectotype designated (see 
Carpenter, 1940:223). 

Distribution. — CAN: BC 5 , PO 5 , USA: 
AZ 2 , CO 1 , NE 2 , NV 6 OR 2 , UT 3 , WS 4 ('Banks, 
1908a:260; 2 Carpenter, 1940:223; 3 Parfin, 
1956:209; 4 Throne, 1971b:84; 5 Klimaszewski & 
Kevan, 1987b:259; 6 Penny, NEW STATE RE- 
CORD). 

Wesmaelius schwarzi (Banks), 1903b:24 [He- 
merobius]. 

Holotype female: USA, Arizona, Williams, 
23.VII, H. S. Barber (USNM). 



USA: AZ',CA 2 , CO 2 , NM 2 



= Boriomyia ultima Banks, 1930a:223 Holo- 
type female: USA, California, Angora Park, Ta- 
hoe, E. P. Van Duzee (MCZ). 

TAXONOMY. — To Boriomyia (see Banks, 
1905c:38); to Kimminsia (see Carpenter, 
1940:223); to Wesmaelius (see Kevan & Kli- 
maszewski, 1986:19); B. ultima = K. schwarzi 
(see Carpenter, 1940:223) 

Distribution 
('Banks, 1903b:241; Carpenter, 1940:224). 
Wesmaelius subnebulosus (Stephens), 
1836:107 [Hemerobius]. 

Type(s) (sex unknown): England, presumably 
near London (depository unknown). 

= Hemerobius fuscus Stephens, 1836:107. 
Type(s) (sex unknown): England, presumably 
near London (depository unknown). 

= Boriomyia maorica Tillyard, 1923:221. 
Holotype female: New Zealand, Dunedin. 
George Street (CIC). 

TAXONOMY. — To Boriomyia (see Banks, 
1905c:29); to Kimminsia (see Killington, 
1937:255); to Wesmaelius (see Tjeder, 1967:5); 
H. fuscus = H. subnebulosus, H. nebulosus = H. 
subnebulosus (see Wallengren, 1871:36); B. 
maorica = W. subnebulosus (see Wise, 
1973:181,182). 

Distribution. — CAN: NS 3 , ON J , PQ 3 , 
USA: CT 1 , IN 2 , NY 1 ('Parfin, 1956:206; 
2 Lawson & McCafferty, 1984:130; kli- 
maszewski & Kevan, 1987b:248). 
Wesmaelius yukonensis Klimaszewski & 
Kevan, 1987b:262. 

Holotype male: Canada, Yukon Territory, km 
1,683 Alaska Highway, 7.VI.1979 (ROM). 

DISTRIBUTION. — CAN: YT (Klimaszewski 
& Kevan, 1987b:263. 

Megalominae 

Megalomus Rambur, 1842:418 

Megalomus angulatus Carpenter, 1940:242. 

Holotype male: USA, New York, Ithaca, 
3.IX.1939, J. S. Franclemont (Cornell). 

Distribution. — CAN: ON, USA: AZ, ME, 
NH, NY (Carpenter, 1940:242). 
Megalomus carpenteri Penny, Adams & 
Stange, NEW NAME. 

= Hemerobius speciosus (Banks), 1904a:62. A 
junior primary homonym of Hemerobius specio- 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



69 



sus Linnaeus, 1758:551 (now Palpares specio- 
sus). Holotype female: USA, Maryland, Plum- 
mer's Island, 9.IX (MCZ). 

Taxonomy. — To Boriomyia (see Banks, 
1904c:209); to Allotomyia (see Banks, 
1930a:224); returned to Boriomyia (see Carpen- 
ter, 1940:245); to Megalomus (see Oswald, 
1993:241). 

Distribution. — USA: FL 2 , MD 1 , VA 2 
("Banks, 1904a:62; 2 Carpenter, 1940:245). 
Megalomus fidelis (Banks), 1897:27 [Hemero- 
bius]. 

Holotype female: USA, New York, Gowanda, 
25.VI, E. P. Van Duzee (MCZ). 

TAXONOMY. — To Boriomyia (see Banks, 
1904c:209): to Allotomyia (see Banks, 
1930a:224); returned to Boriomyia (see Carpen- 
ter, 1940:244); to Megalomus (see Oswald, 
1993:239). 

DISTRIBUTION. — CAN: BC 4 , ON 4 USA: 
DC 4 , FL 6 , IL 4 , IN 7 , MA 4 , MD 4 . MN" MS 4 
NC 3 , NH 4 , N J 2 , NY ' , PA 4 , SC 4 , TX 4 , V A 4 , W V 4 
('Banks, 1897:27; 2 Smith, 1900:56; 3 Brimley, 
1938:29; 4 Carpenter, 1940:244; 5 Parfin, 
1952:423, as Boriomyia: MacLeod & Stange, 
1981:1, as Boriomyia; Lawson & McCafferty, 
1984:130). 
Megalomus minor Banks, 1905a:90. 

Lectotype male: USA, Nevada, Ormsby Co., 
G.F.Baker (MCZ). 

TAXONOMY. — Lectotype designated by Car- 
penter, 1940:242. 

Distribution. — USA: AL 2 , CA 2 , MS 2 , 
NV 1 ("Banks. 1905a:90; 2 Penny, NEW STATE 
RECORDS) 
Megalomus moestus Banks, 1895b:3 14. 

Holotype (missing abdomen): USA, New 
Mexico, Santa Fe, VII, T. D. A. Cockerell 
(MCZ). 

= Megalomus latus Banks, 1903b:240. 

Holotype female: USA, Arizona, Williams 
(USNM). 

Taxonomy. — M. latus = M. moestus (see 
Carpenter, 1940:241). 



Distribution. 
NM 1 , TX 2 . UT 



USA: AZ~, CA~, CO", 
("Banks, 1895b:314; 



WY 
2 Carpenter, 1940:241). 
Megalomus parvulus Kimmins, 1935:614. 

Holotype male: West Indies, St. Vincent Is- 
land, windward side. H. H. Smith (BMNH). 

Distribution. — USA: FL (Stange. 1995:7). 



Megalomus uniformis Banks, 1935:56. 

Holotype female: USA, Texas, Brownsville, 
VI, F. H. Snow (MCZ). 

Distribution. — USA: TX (Banks, 
1935:56). 

MlCROMINAE 

Micromus Rambur, 1842:416 

Micromus angulatus (Stephens). 1836:106 
[Hemerobius]. 

Holotype female: no locality data (BMNH). 

= Hemerobius villosus Zetterstedt. 1840:1050 
(nomen nudum). Type(s) (sex unknown): (local- 
ity unknown) (Lund). 

= Hemerobius intricatus Wesmael, 1841:214. 
Type(s) (sex unknown): (locality unknown) 
(IRSNB). 

= Micromus tendinosus Rambur, 1842:417. 
Type(s) (sex unknown): Sardinia (MNHN). 

= Hemerobius lineatus Goszy, 1852:345. 
Type(s) (sex unknown): no locality data (prob- 
ably Vienna). 

= Micromus jonas Needham, 1 905: 15. Type(s) 
(sex unknown): USA, New York (depositoiy 
unknown). 

TAXONOMY. — To Micromus; H. villosus = 
M. angulatus, H. intricatus = M. angulatus, M. 
tendinosus = M. angulatus, H. lineatus = M. 
angulatus, M. aphidivorus = M. angulatus (see 
Hagen, 1889:280); M. jonas = M. angulatus (see 
Carpenter, 1940:247). 

DISTRIBUTION. — CAN: AB 2 BC 2 , MB 2 , 
NF 3 , NS 3 NT 3 7 , ON 2 PQ 2 SK", YT". USA: 
CO 2 , ME 2 , MI 2 , MN~, NH", NY 1 , SD 2 , VT 2 , 
WS 2 (" Leonard, 1928:40; 2 Carpenter, 1940:247; 
Klimaszewski & Kevan, 1990b:65). 
Micromus borealis Klimaszewski & Kevan, 
1988:49. 

Distribution. — CAN: AB, BC, YT, USA: 
AK (Klimaszewski & Kevan, 1990b:65). 
Micromus montanus Hagen, 1886:279. 

Lectotype male: USA, Massachusetts, Natick 
(MCZ). 

Distribution. — CAN: AB 4 , BC 4 , NS 4 , 
PQ 4 SK 7 USA: AZ 4 , CA 4 , CO 4 , CT 4 , ID 4 , 
MA 1 , ME 5 , NC 3 , ND 4 , NH 7 , NV 4 , NY 2 , TN 4 , 
UT 4 , VT 7 , WA 4 , WS 6 ("Hagen, 1889:279; 
2 Leonard, 1928:40; 3 Brimley, 1938:30; 4 Car- 



70 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



penter, 1940:249; 5 Procter, 1946:43; 6 Throne, 
1971b:83; 7 Klimaszewski & Kevan, 1990b:64). 
Micromus posticus (Walker), 1853:283 [He- 
merobius]. 

Male and female syntypes: USA, Georgia 
(BMNH). 

= Micromus insipidus Hagen, 1861:199. Syn- 
types (sex unknown): USA, New York; Pennsyl- 
vania, Philadelphia (MCZ). 

= Micromus sobrius Hagen, 1861:199. Holo- 
type (sex unknown): USA, Illinois, Chicago, Os- 
ten Sacken (MCZ). 

Taxonomy. — McLachlan, 1867:271 (M. in- 
sipidus = M. posticus, M. sobrius = M. posticus 
(see McLachlan, 1867:271). 

DISTRIBUTION. — CAN: BC ",0-N 7 PQ 11 
YT 11 , USA: AZ 7 , CO 7 , CT 1 ', FL 7 , GA 1 , IA 12 
IL 2 IN 6 , KS 5 , MA 3 , ME 8 MN 7 , MO 9 , NC 3 
ND 7 , NE 7 , NJ 4 , NY 2 , OH 1 ", PA 2 , TX 7 ^ VA 1 ' 
WA 11 , WS 10 ('Walker, 1853:283; "Hagen 
1861:199 200; 3 Hagen, 1889:286, as M. in- 
sipidus; 4 Smith, 1900:56; 5 Smith, 1925:167 
6 Montgomery & Trippel, 1933:259- Carpenter 
1940:248;.. Procter, 1946:43; Troeschner 



,11 



ON 1 



10^ 



1947:131; ,u Throne, 1971b:82; "Klimaszewski 

& Kevan, 1990b:65; l2 Penny, NEW STATE 

RECORDS). 

Micromus remiformis Oswald, 1987:1. 

Holotype male: USA, California, Madera Co., 
4.75 miles east southeast of Fish Camp, 6400 ft., 
9. VIII. 197 1, H. B. Leech (CAS). 

Distribution. — USA: CA, ID, NV, OR, UT 
(Oswald, 1987:3). 

Micromus subanticus (Walker), 1853:282 [He- 
merobius]. 

Holotype male: USA, East Florida, St. John's 
Bluff (BMNH). 

= Micromus angustus Hagen, 1 889:287. Seven 
syntypes: USA, Florida and North Carolina 
(MCZ and CNC). 

= Micromus nesoticus Navas, 19 14b: 16. Holo- 
type (sex unknown): USA. New York, Long 
Island, Yaphank (MZB). 

= Micromus haitiensis Smith, 193 1 :800. Holo- 
type female: Haiti, Petionville near Port-au- 
Prince, 17.11.1930, H. L. Dozier (MCZ). 

TAXONOMY. — To Micromus (see Banks, 
1905c:46); Carpenter, 1940:250, 251 (M. angus- 
tus = M. subanticus, M. nesoticus = M. suban- 
ticus (see Caipenter, 1940:250, 251); M. 
haitiensis = M. subanticus (see Monserrat, 
1993:511). 



Distribution. — CAN: BC", MB 1 
PQ 11 , USA: AZ 5 , CA 5 , FL 1 IA 1 ', IL 1 ', IN 1( ;, 
KS 1 '. LA 2 , MA 5 , MN 7 , MO , NC 4 , NV 5 . NY J , 
OH 12 , SC 12 . TX 9 , VA 2 , WS 8 ('Walker, 
1853:282; "Banks, 1905c:46; 3 Leonard, 
1928:40; 4 Brimley, 1938:30; 5 Carpenter, 
1940:250, 251; 6 Froeschner, 1947:131: 7 Parfin, 
1952:423; 8 Throne, 1971b:82; 9 Agnew et al., 
1981:16; 10 Lawson & McCafferty, 1984:130; 
1 'Klimaszewski & Kevan, 1990b:65; '"Monser- 
rat, 1993:511). 

Micromus variegatus (Fabricius, 1793:85) [He- 
merobius]. 

Type(s): location unknown. 

DISTRIBUTION. — CAN: BC (Klimaszewski 
& Kevan, 1990b:66). 

NOTE. — This species appears to be estab- 
lished on Galiano Island, off the west coast of 
British Columbia, Canada. Klimaszewski and 
Kevan suspect this to be an introduction from 
Japan. 
Micromus variolosus Hagen, 1886:284. 

Holotvpe female: USA, Colorado, Denver 

(MCZ) -' 1 6 6 

Distribution. — CAN: BC 3 SK°, YT b . 
USA: AZ 3 , CA 3 , CO 1 , ID 3 , IN 5 , KS 2 , MN 4 , 
NM 3 , NV 3 , OR 3 , UT 3 , WA 3 , WY 3 ('Hagen, 
1886:284; 2 Smith, 1925:167; 3 Carpenter, 
1940:251; 4 Parfin. 1952:423; 5 Lawson & 
McCafferty, 1984:130; 6 Klimaszewski & 
Kevan, 1990b:65). 



NOTIOBIELLINAE 
Psectra Hagen, 1866:376,458 

Psectra diptera (Burmeister), 1839:973 [He- 
merobius]. 

Holotype male: Germany. Saxony, near 
Leipzig (Halle). 

= Hemerobius delicatulus Fitch, 1855:800. 
Two syntypes: USA, Illinois (MCZ). 

= Psectra buenoiNavas, 1909b:2 18. Five syn- 
types (1 male, 4 females): USA, New Jersey, 
Westfield, 5.VIII.1904; New York, Putnam Co., 
5.IX. 1903 (MZB). 

TAXONOMY. — To Psectra (see Hagen, 
1866:376); H. delicatulus = P. diptera, P. buenoi 
= P. diptera (see Carpenter, 1940:252). 

DISTRIBUTION. — CAN: NF 7 , ON J , USA: 
IA 8 , IL 1 , IN 6 , MI 3 , MN 4 , NY 2 , VA 3 , WS 5 , WV 3 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



71 



('Fitch, 1855:800, "Leonard, 1928:39; ^Carpen- 
ter, 1940:252; 4 Parfin, 1952:423; 5 Throne, 
1971b:84; 6 Lawson & McCafferty, 1984:130; 
Kevan and Klimaszewski, 
NEW STATE RECORD) 



,986:9; "Penny, 



. ON 4 , PQ 8 , 



Sympherobiinae 
Sympherobius Banks, 1904c:209 

Sympherobius amiculus (Fitch), 1855:799 [He- 
merobius]. 

Lectotype female: USA, New York, Washing- 
ton Co., Salem, 20. VIII. 1852. A. Fitch (MCZ). 

= Sympherobius buenoi Navas. 19 12a: 198 
Holotype (sex unknown): USA, New York, 
White Plains (MZB). 

TAXONOMY. — To Sympherobius (see Banks, 
1904c:209); S. buenoi = S. amiculus (see Carpen- 
ter, 1940:230). See note (Oswald. 1988:451) for 
clarification of type status. 

Distribution. — CAN: 
USA: AL 8 , AR 4 AZ 8 , CO 8 , CT 8 , DC 8 , FL 4 , 
GA 4 , IA 8 , IL 4 , IN 7 , KS 2 . LA 4 MA 4 , MD 8 , ME 5 . 
MI 4 , MN 4 , MO 6 . MS 8 , NC, NE 4 NH 4 , NJ 8 , 
NY ' , OH 4 . OK 4 , PA 8 , SC 8 , TN 8 TX 4 , V A 4 , VT 4 , 
WS 4 ,WV 4 ( 'Fitch. 1855:799; 2 Smith. 1925:167; 
3 Brimley. 1938:29; 4 Carpenter. 1940:229: 
5 Procter, 1946:42; 6 Froeschner. 1947:130; 
7 Lawson & McCafferty, 1984:130; 8 Oswald, 
1988:433). 

Sympherobius angustus (Banks), 1904b: 102 
[Hemerobius]. 

Lectotype female: USA. New Mexico, 
Mesilla. A. P. Morse (MCZ). 

= Sympherobius tristis Navas, 19 14b: 15. 
Type(s) (sex unknown): USA, New Mexico, Pe- 
cos (MZB). 

= Sympherobius stangei Nakahara, 1960:16. 
Type (sex unknown): USA, California, San Ber- 
nardino Co.. Barton Flats (probably in Nakahara 
Collection, Japan). 

= Sympherobius brunneus Nakahara, 
1966:207. Holotype male: USA, California, 
Mariposa Co., Miami Ranger Station (CAS). 

TAXONOMY. — To Sympherobius (see Banks, 
1905c:4 1 ); S. tristis = S. angustus (see Carpenter, 
1940:233); S. stangei = S. angustus, S. brunneus 
= S angustus (see Oswald, 1988:418). 

Distribution. — CAN: BC 2 , USA: AZ 3 ^ 
CA 3 , CO 2 , ID 3 , NM 1 , NV 3 , OR 3 , SD 3 , UT~ 



(' Banks, 1904b: 102; "Carpenter. 1940:233; 

3 Oswald, 1988:421). 

Sympherobius arizonicus Banks. 191 1:346. 

Holotype female: USA, Arizona, Prescott. 
Oslar(MCZ). 

Distribution. — USA: AZ 1 , CA 2 ('Banks, 
1911:346; 2 0swald, 1988:408). 
Sympherobius barberi (Banks). 1903b:241 
[Hemerobius]. 

Two female syntypes: USA, Arizona, Wil- 
liams, 20,2 1 .VII, H. S. Barber and E. A. Schwarz 
(USNM). 

TAXONOMY. — To Sympherobius (see Banks, 
1905c:42). 

Distribution. — USA: AL 8 , AR 3 Az' 
CA 3 CO 3 , FL 3 , IA 8 , IL 8 , IN 7 , KS 2 , LA 8 . MD 8 . 
MN" , MO 4 , MS 8 , NC 3 , NJ 8 , NM 3 , OK 3 , OR J , 
PA 3 , SC 3 , TX 3 , UT 3 , VA 3 , WS^ ('Banks, 
1903b:241; 2 Smith, 1925:166; 3 Carpenter, 
1940:236; 4 Froeschner, 1947:130; 5 Parfin, 
1952:423; 6 Throne, 1971b:83; 7 Lawson & 
McCafferty, 1984:130; 8 Oswald, 1988:429). 
Sympherobius beameri Gurney, 1948:220. 

Holotype male: USA, California, Lake Tahoe, 
11. VIII. 1940, R. H. Beamer(SEM). 

DISTRIBUTION. — USA: AZ 2 , CA 1 , UT~ 
('Gurney, 1948:220; 2 Oswald, 1988:406). 
Sympherobius bifasciatus Banks, 191 1:347. 

Holotype male: USA, Colorado, G. F. Baker 
(MCZ). 

Distribution. — CAN: AB-, USA: CO 1 , 
AZ 3 , CA 3 , UT 3 ('Banks, 191 1:347; 2 Carpenter, 
1940:233; 3 Oswald, 1988:423). 
Sympherobius californicus Banks. 191 1:346. 

Lectotype male: USA, California, Pasadena 
(MCZ). 

Taxonomy. — Lectotype designated by Car- 
penter, 1940:232. 

DISTRIBUTION. — USA: CA 1 , WA" ('Carpen- 
ter. 1940:232; 2 Oswald, 1988:425). 
Sympherobius constrictus Oswald. 1988:425. 

Holotype male: USA, California, San Fran- 
cisco, Lobos Creek, 16.11. 1967, J. Powell (CAS). 

Distribution. — USA: CA (Oswald, 
1988:427). 
Sympherobius distinctus Carpenter, 1940:238. 

Holotype male: USA, Colorado, Pingree Park, 
18.VIII.1926, R. C. Smith (MCZ). 

Distribution. — USA: AZ", CO ( Carpen- 
ter, 1940:238; 2 Oswald, 1988:418). 
Sympherobius killingtoni Carpenter, 

1940:238. 



72 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Holotype male: USA, Utah, Eureka, 13. VIII, 
T. Spaulding (MCZ). 

= Sympherobius texanus Nakahara, 1966:209. 
Holotype male: USA, Texas, Kerr Co., Kerrville, 
VI. 1954, L. J. Bottimer (USNM). 

Taxonomy. — S. texanus = S. killingtoni (see 
Oswald, 1988:408). 

DISTRIBUTION. — USA: AZ 1 , CA 1 , CO", ID 1 , 
NM 1 , NV 2 , OR 2 , TX 1 , UT 1 , WA 2 ('Carpenter, 
1940:239; "Oswald, 1988:410). 
Sympherobius limbus Carpenter, 1940:236. 

Holotype male: USA, Texas, Alpine, 
ll.VII.1928,R.H. Beamer (SEM). 

DISTRIBUTION. — USA: AZ 2 , NM 2 , TX 1 
('Carpenter, 1940:236; 2 Oswald, 1988:416). 
Sympherobius occidentalis (Fitch), 1855:799 
[Hemerobius]. 

Holotype (sex unknown): USA, Illinois, Hen- 
derson River, 2.X.1854, A. Fitch (lost). 

TAXONOMY. — To Sympherobius (see Banks, 
1905c:40). 

DISTRIBUTION. — USA: AL 10 AR 4 DC 4 , 
FL 9 GA 10 IL 1 , IN 8 , KS 2 , LA 10 , MO 5 NC 3 , 
NY, OH f0 PA 10 , TX 7 , VA 10 , WS 6 (Titch, 
1855:799; 2 Banks, 1905c:40; 3 Brimley, 
1938:29; 4 Carpenter, 1940:231, 5 Froeschner, 
1947:130; 6 Throne, 1971b:83; 7 Agnew et al., 
1981:16; 8 Lawson & McCafferty, 1984:130; 
9 MacLeod & Stange, 1981:2; 10 Oswald, 
1988:415). 

Sympherobius perparvus (McLachlan), 
1869:22 [Hemerobius]. 

Lectotype male: USA, Texas, Bosque, G. W. 
Belfrage (BMNH). 

= Sympherobius sparsus Banks, 1911:346. 
Holotype female: USA, Kansas (MCZ). 

TAXONOMY. — To Sympherobius (see Banks, 
1905c:41); S. sparsus = S. perparvus (see Car- 
penter, 1940:238); lectotype designated by 
Oswald, 1988:403. 

Distribution. - 



CA 3 , CO 3 , NM 



-CAN: SK 3 , USA: AZ 3 , 
OK 3 , IA 4 , KS 2 , MN 4 , MT 4 
NE 4 , NV 4 , OR 4 , SD 4 , TX 1 , UT 4 , WY 4 
('McLachlan, 1 869:22; 2 Banks, 191 1:346; Car- 
penter, 1940:238; 4 Oswald, 1988:405). 
Sympherobius pictus (Banks), 1904a:62 [He- 
merobius]. 

Lectotype female: USA, southwestern Colo- 
rado, Oslar (MCZ). 

Taxonomy. — To Sympherobius (see Banks, 
1905c:41); lectotype designated by Carpenter, 
1940:235. 



Distribution. — USA: AZ~, CA;, CO 1 , 
NM 2 , NV 2 ('Banks, 1904a:62; 2 Oswald, 
1988:413). 

Sympherobius quadricuspis Oswald, 
1988:421. 

Holotype male: USA, Arizona, Cochise Co., 
Southwest Research Station, 22-30.IV. 1963, 
Vincent Roth (AMNH). 

Distribution. — USA: AZ (Oswald, 
1988:422). 
Sympherobius similis Carpenter, 1940:236. 

Holotype male: USA, Arizona, Santa Rita 
Mountains, 17.VII.1932, R. H. Beamer (SEM). 

DISTRIBUTION. — USA: AZ (Carpenter, 
1940:236). 

Sympherobius umbratus (Banks), 1903b:242 
[Hemerobius]. 

Holotype male: USA, Arizona, Williams, 
lO.VI(USNM). 

= Sympherobius gracilis Carpenter. 1 940:23 1 . 
Holotype male: USA, North Carolina, Raleigh, 
1 1. VI. 1934, R. R. Leiby (MCZ). 

TAXONOMY. — To Sympherobius (see Banks, 
1 905c:4 1 ); S. gracilis = S. umbratus (see Oswald, 
1988:430). 

Distribution. — USA: AR 5 , AZ ' , FL 4 , GA" , 
MO 3 , NC 2 , NM 2 , OK 5 , PA 5 . SC 5 , VA 5 ('Banks, 
1903b:242; 2 Carpenter 1940:230, 231; 
3 Froeschner, 1947:130; 4 MacLeod & Stange, 
1981:2, as S gracilis; 5 Oswald, 1988:431). 

ITHONIDAE 
Ithonidae, or moth-like lacewings, are known 
from only one species in the genus Oliarces 
Banks 1908 in the United States, and are prob- 
ably encountered less than any other family of 
Neuroptera. The original type locality is now 
flooded by the Salton Sea, and only one specimen 
(a male) was known until 1949. Its known distri- 
bution includes the desert of southern California 
and Arizona, being seen most often between the 
Gila Mtns. of Arizona and the Salton Sea. Adults 
are known to have mass emergences, and the 
larvae appear to feed on the roots of creosote 
bush (Faulkner, 1990a: 18). Another species was 
recently described from Honduras (Penny, 
1996). All other known species of this family are 
from Australia, although there are other unde- 
scribed species known from Mexico, and it is 
thought that the Mexican species Narodona 
mexicana Navas is actually an ithonid, rather 
than an osmylid, as originally indicated by Navas 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



73 



(Adams, 1969). The adult behavior of Oliarces 
clara was described by Faulkner ( 1 990a, 1 990b). 

Oliarces Banks, 1908b:203 

Oliarces clara Banks, 1908b:203. 

Holotype male: USA, California, Imperial 
Co.,Walter's Station, IV, J. B. Smith (MCZ). 

Distribution. — USA: AZ 2 , CA 1 NV 3 
('Banks, 1908b:203; 2 Belkin, 1954:67; 3 Penny, 
NEW STATE RECORD). 

Mantispidae 
Mantispidae. or mantid-flies, superficially re- 
semble preying mantids, or in the case of Clima- 
ciella, Polistes wasps. All species have raptorial 
forelegs as adults. Larvae are hypermetamor- 
phic, being very active triungulins in the first 
instar, and becoming inactive scarabaeiform lar- 
vae in subsequent stages. Primitive members of 
the family are known to feed on noctuid pupae, 
solitary bee larvae, and polybiine social wasp 
larvae. Other species, in the more advanced Man- 
tispinae, are spider egg sac predators, having a 
number of strategies for seeking out and entering 
the spider egg sacs (Redborg and MacLeod, 
1985). In this family, adult size is dependent on 
larval food availability, and striking differences 
in size are common. The higher classification of 
Mantispidae has been discussed by Penny 
(1982b) and Lambkin (1986). 

Symphrasinae 

Plega Navas, 1928b:326 

Plega banksi Rehn, 1939:248. 

Holotype male: USA, Arizona, Pima Co., Kits 
Peak Rincon. Balboquivari Mts., 4050', 
14.VIII.1916, Lutz & Rehn (AMNH). 

DISTRIBUTION. — USA: AZ (Rehn, 

1939:250). 
Plega dactylota Rehn. 1939:250. 

Holotype male: USA, Arizona, Mohave Co., 
Kingman, 20.VII.1920, O. C. Poling (ANSP). 

= Plega dactylota lipanica Rehn, 1939:254. 
Holotype male: USA, Texas, Brewster Co., hills 
west of Ord Mountains, 1-15.VI.1926, O. C. 
Poling (ANSP). 

Taxonomy. — Two subspecies synonymized 
(see Rice, 1987:342). 



USA: 



NV 



AZ'^ CA" 

UT' ('Rehn, 1939:253; "Penny, NEW 



Distribution 
TX 
STATE RECORD) 

Plega fratercula Rehn, 1939:247. 

Holotype male: USA, Arizona, Gila Co., Capi- 
tan Mountain, 8.VIII. 1933, R. Anderson (MCZ). 

Distribution. — USA: AZ (Rehn, 
1939:248). 
Plega signata (Hagen). 1877:208 [Symphrasis]. 

Holotype female: USA, California, Kern Co., 
Fort Tejon. 1879, F. Brown (MCZ). 

Taxonomy. — To Plega (see Navas, 
1928b:326). 

DISTRIBUTION. — USA: AZ 2 , CA 1 (' Hagen, 
1877:208; 2 Rehn, 1939:246). 

Calomantispinae 

Nolima Navas, 1914f:100 

Nolima dine Rehn, 1939:261. 

Holotype male: USA, Arizona, Pima Co., 
Santa Catalina Mts., Pepper Sauce Canyon, 
16.VIII.1924, J. O. Martin (CAS). 

Distribution. — USA: AZ (Rehn, 
1939:262). 
Nolima kantsi Rehn, 1939:260. 

Holotype male: USA, Texas, Brewster Co.. 
Chisos Mts., 16.VII.192I, C. D. Duncan, (CAS). 

Distribution. — USA: TX (Rehn, 
1939:261). 
Nolima pinal Rehn, 1939:257. 

Holotype male: USA, Arizona, Gila Co., base 
of Pinal Mts., IX, D. K. Duncan (MCZ). 

Distribution. — USA: AZ (Rehn, 
1939:259). 

Mantispinae 

Climaciella Enderlein, 1910:360 

Climaciella brunnea (Say), 1824:309 [Mon- 
ti spa]. 

Syntype male: USA, Minnesota, St. Peter's 
River (depository unknown). 

= Mantispa denarius Taylor, 1862:494 No 
types designated. 

= Mantispa burquei Provancher, 1875:247. 
Type(s) (sex unknown): Canada, Quebec (de- 
pository unknown). 



74 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



= Mantispa brunnea var. occidentis Banks, 
1911:348. Syntype males: USA, Colorado, 
Golden, Chimney Gulch, Oslar; Nevada, 
Ormsby Co., Baker; Washington, Wawawai, 
Mann (MCZ). 

= Climaciella rubescens Stitz, 1913:37. Holo- 
type (sex unknown): Mexico, Tampico (HMB). 

= Climaciella rubescens var. itnicolor Stitz, 
1913:39. Holotype (sex unknown): Mexico, Si- 
erra Mixteca (HMB). 

= Climaciella rubescens var. laciniata Stitz, 
1913:39. Holotype (sex unknown): Mexico, 
Mexico (HMB). 

TAXONOMY. — To Climaciella (see Ender- 
lein, 1910:360); M. burquei = M. brunnea (see 
Provancher, 1877:174); C. rubescens and varie- 
ties unicolor and laciniata = C. brunnea (see 
Handschin, 1960:546547); M. denarius = C. 
brunnea (see Kevan, 1989:8); C. brunnea var. 
occidentis = C. brunnea (see Welch & Kondra- 
tieff, 1991:70). 

DISTRIBUTION. — CAN: BC 10 , ON 12 , PQ 4 , 
USA: AZ 14 , CA 16 , CO 15 , FL 2 , GA 2 , ID 15 , IL 3 , 
IN 8 , KS 6 , MN 1 , MO 11 NC 9 NJ 5 , NV 15 , NY 7 , 
PA 3 ,TX ,4 ,UT 15 ,WA .WS f2 ('Say, 1824:309; 
2 Walker, 1853:214; 3 Hagen, 1861:208; 
4 Provancher, 1875:247. 5 Smith, 1900:54; 
6 Smith, 1925:166; Xeonard, 1928:39; 
Montgomery & Trippel, 1933:259; 9 Brimley, 
1938:29; 10 Spencer, 1942:24; n Froeschner, 
1947:124; ' "Robert, 1949:10; l3 Throne, 
1972:119; l4 Redborg & MacLeod, 1983:63; 
15 Welch&Kondratieff 1991:71; l6 Penny,NEW 
STATE RECORD) 

Mantispa Illiger in Kugleann, 1798:499 

Mantispa floridana Banks, 1897a:23. 

Holotype (sex unknown): USA, Florida, Lake 
Worth, A. T. Slosson (MCZ). 

Distribution. — USA: FL (Banks, 
1897a:23). 

Mantispa interrupta Say, 1825: unpaginated 
text to plate 25. 

Type(s) (sex unknown): USA, Pennsylvania, 
Philadelphia (specimen probably lost). 

= Mantispa aureus Taylor, 1 862:494. No types 
designated. 

= Mantispa cincticornis Banks, 1911:347. 
Holotype female: USA, Texas, near Browns- 
ville, Snow (MCZ). 



TAXONOMY. — M. aureus = M. interrupta 
(see Kevan, 1989:8); M. cincticornis = M. inter- 
rupta (see Welch & Kondratieff 1991:72). 

CAN: PQ 6 , USA: CO 10 , 
5 „ TT 2 Cr,,4 tw 1 o^ll 



Distribution. 

t9 „n3 WXT 7 



IN , KS J , MN', MO , NJ", NY\ PA'^ SC 
TX, VA 1 , WS 8 ('Banks, 1892:358; "Smith, 
1900:54; 3 Smith, 1925:166; 4 Leonard, 1928:39; 
5 Froeschner, 1947:124; 6 Robert, 1949:12; 7 Par- 
fin, 1952:423; 8 Throne, 1972:120; 9 Lawson & 
McCafferty, 1984:130; 10 Welch & Kondratieff, 
1991:72; ' 1 Hoffman & Brushwein, 1992:161). 
Mantispa moesta Hagen, 1861:210. 

Type(s) (sex unknown): USA. Tennessee 
(MCZ). 

Distribution. — USA: TN (Hagen, 
1861:210). 

Mantispa pulchella (Banks), 1912:179 [Man- 
ti spill a]. 

Holotype female: USA, Utah, Eureka (MCZ). 

TAXONOMY. — Transferred to Mantispa (see 
Redborg, 1976:17). 

DISTRIBUTION. — USA: CO 4 , GA 3 , IL 2 , NC 3 , 



SC J , UT 1 ('Banks, 1912:179; "Redborg, 
1976:18; 3 Hoffman & Brushwein, 1989:8; 
4 Welch & Kondratieff, 1991:74). 
Mantispa sayi Banks, 1897a:23. 

Lectotype male: USA, Texas, Brazos Co. 
(MCZ). 

= Mantispa fuscicornis Banks, 191 1:347. Lec- 
totype male: USA, Florida, Kissimmee (MCZ). 

= Mantispa uhleri Banks, 1943:79. Holotype 
female: USA, Pennsylvania (MCZ). 

TAXONOMY. — M. fuscicornis = M. sayi, M. 
uhleri = M. sayi; lectotypes designated for M. 
sayi and M. fuscicornis (see Hoffman, 1989:638, 
639). 

Distribution. 
CT 9 FL 1 , GA 9 , IL 5 , IN 8 , KS 2 , MD 9 , MI 3 , MN 7 , 
MO 6 , MS 9 , NC 4 , NE 9 , NM 9 , OH 9 , OK 9 , PA 5 , 
SC 9 , TX 1 , UT 9 , WS 5 ('Banks, 1897a:23; 2 Smith, 
1925: 166; 3 Kaston, 1938: 148, as M. fuscicornis; 
4 Brimley, 1938:29; 5 Banks, 1943:79; 6 Froesch- 
ner, 1947:124; 7 Parfin, 1952:423; 8 Lawson & 
McCafferty 1984:130, as M. uhleri; 9 Hoffman, 
1989:637; lD Welch & Kondratieff, 1991:72). 
Mantispa scabrosa (Banks), 1912:179 [Man- 
tispilla]. 

Holotype female: USA, New Mexico, Mesilla 
(MCZ). 

TAXONOMY. — Transferred to Mantispa (see 
Welch & Kondratieff, 1991:75). 



USA: AR 9 , AZ 9 , CO 10 



PENNY, ADAMS AND STANGE: SPECIES CATALOG 



75 



DISTRIBUTION. — USA: AZ", NM 1 , TX" 
('Banks, 1912:179; 2 Welch & Kondratieff, 
1991:75). 
Mantispa viridis Walker, 1853:227. 

Holotype (sex unknown): USA, East Florida 
(BMNH). 

Distribution. — USA: FL 1 , SC\ TX~, VA J , 
WS 4 ('Walker, 1853:227: 2 Banks, 1907c:23; 
3 Parfin, 1958:203; 4 Throne, 1972:121; Hoff- 
man & Brushwein, 1992:161). 

REMARKS. — Mantispa gnlosus Taylor, 
1 862:294 (nomen dubium); Mantispa brevicollis 
Banks, 1905a:88 (nomen nudum). 

Myrmeleontidae 
Myrmeleontidae, or ant-lions and doodle- 
bugs, are among the largest, and in southern parts 
of the country, probably the most familiar neu- 
ropterans to most people. The larvae of one ge- 
nus, Myrmeleon, form pits in sheltered areas of 
dry soil and can only move backward. They wait 
at the bottom of the pits with only their heads 
exposed, waiting for insects to fall in. To make 
sure that the prey cannot escape up the loose sides 
of the funnel, the ant-lion larva throws loose sand 
or soil at it with its head. Then, the larva drags 
the insect under the soil surface where the vic- 
tim's movements are inhibited. The prey is 
drained of body liquids, and the hollow body 
thrown out of the pit by the powerful head and 
mandibles. Other genera of ant-lions move for- 
ward and backward (except Vella) and live just 
under the soil surface, in rock crevices, in caves, 
or in tree-holes. Larvae pupate under the soil 
surface, and a damselfly-like adult later emerges. 
Adults generally rest quietly on twigs and grass 
stems during the day and become active at night. 
However, some species are active during the 
daytime and have rather bright pigment patterns 
on the wings. Ant-lions are most abundant in hot, 
dry regions, but can also be found in moist, 
forested regions. North American species of 
Myrmeleontidae were reviewed by Banks (1927) 
and Stange and Miller (1990). Current views on 
higher categories of Myrmeleontidae are to be 
found in Stange (1994). Keys to genera of 
Glenurini are to be found in Stange (1970b) and 
genera and species of Nemoleontini in Stange 
(1970a). 



ACANTHACLISINI 
Paranthaclisis Banks, 1907a:275 

Paranthaclisis congener (Hagen), 1861:224 
[Acanthaclisis]. 

Holotype female: USA, "Western Texas," Pe- 
cos River (now New Mexico) (MCZ). 

TAXONOMY. — To Paranthaclisis (see Banks, 
1907a:275). 

Distribution. — USA: AZ 2 , CA J b NM 1 , 
OR 3 , UT 3 WA 3 ('Hagen, 1861:224; "Cunie, 
1903:274; 3 Banks, 1927:81). 
Paranthaclisis hageni (Banks). 1899a: 170 
[Acanthaclisis]. 

Lectotype male: USA, Arizona. Phoenix, V 
(MCZ). 

TAXONOMY. — To Paranthaclisis (see Banks. 
1907a:275); lectotype designated by Stange, 
1961a:674. ^ 

DISTRIBUTION. — USA: AZ 1 , NM". TX 2 
( 'Banks, 1899a:170; 2 Banks, 1927:81). 
Paranthaclisis nevadensis Banks. 1939:4. 

Holotype male: USA, Nevada, Esmeralda Co., 
2 mi. S Silver Peak, 4350', 22.VIII.1924. Rehn 
& Hebard (ANSP). 

DISTRIBUTION. — USA: CA 2 , NV 1 ('Banks, 
1939:4; 2 Stange, NEW STATE RECORD). 



Vella Navas. 1913e:46 

Vella americana (Drury), 1 773: unpaginated in- 
dex [Myrmeleon]. 

Type(s) (sex unknown): USA, New York (de- 
pository unknown). 

= Acanthaclisis striata Hagen, 1861:324 (no- 
men nudum). Type(s) (sex unknown): Colombia 
(depository unknown). 

TAXONOMY. — To Myrmecoleon (see Bur- 
meister, 1839:996); to Acanthaclisis (see Ram- 
bur, 1842:380): to Myrmeleon (see Walker, 
1853:3 17); returned Xo Acanthaclisis (see Hagen, 
1861:223); to Heoclisis (see Taschenberg, 
1 879: 1 86); A. striata = A. americana (see Hagen 
1887:136); to Grypoplectron (see Esben-Pe- 
tersen, 1928:75); to Vella (see Stange, 1980:3). 

Distribution. — USA: FL 3 , GA 3 , NC 3 , NJ", 
NY 1 , SC 1 , VA 3 ('Hagen, 1861:223; "Smith, 
1900:56; 3 Banks, 1927:83). 



76 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Vella fallax haitiensis Smith, 1931:815. 

Holotype female: Haiti, Jacmel, V. 1927, G. N. 
Wolcott (MCZ). 

= Vella fallax cubana Hagen, 1860:363 [Acan- 
thaclisis] (nomen nudum). Type(s) (sex un- 
known): unknown locality, unknown depository. 

Taxonomy. — To Vella (see Navas, 
1913e:46); Vella fallax cubana = Vella fallax 
haitiensis (Stange, NEW SYNONYMY). 
DISTRIBUTION. — USA: FL (Stange, NEW NA- 
TIONAL RECORD) 

Vella fallax texana (Hagen), 1887:147 [Acan- 
t hacl is is]. 

Lectotype male: USA, Texas, Dimmit Co., 
Carrizo Springs (MCZ). 

= Acanthaclisis hesperus Banks, 19 14b: 6 18. 
Syntype males: USA, Utah, Eureka and New 
Mexico, Jemez Mts., 28.VII and 4.VIII, Spalding 
and Woodgate (MCZ). Stange, NEW SYNON- 
YMY. 

= Vella texana minor Banks, 1943:75. Holo- 
type (sex unknown): USA, Arizona, Phoenix 
(MCZ). Stange, NEW SYNONYMY. 

Taxonomy. — To Vella (see Navas, 
1913e:46); lectotype designated by Stange, 
1961:674. 

DISTRIBUTION. — USA: AZ 3 , CA\ NM 2 , 
TX 1 , UT 2 ('Hagen, 1887:147; 2 Banks, 
1914b:618; 3 Stange, NEW STATE RE- 
CORDS) 

DENDROLEONTINI 

Dendroleon Brauer, 1866:42 

Dendroleon obsoletus (Say), 1839:44 [Formi- 
ca I eo]. 

Type(s) (sex unknown): USA (specimen lost). 

= Myrmeleon nigrocinctus Rambur, 1 842:398. 
Holotype (sex unknown): no locality data (de- 
pository unknown). 

TAXONOMY. — To Myrmecoleon as M. ocel- 
latus Borkhausen: 179 1:161 (see Burmeister, 
1839:995); to Glenurus (see Hagen, 1866:405); 
D. obsoletus not = D. ocellatus Borkhausen (see 
Hagen, 1888:185) to Dendroleon (see Hagen, 
1888:187); M. nigrocinctus = M. obsoletus (see 
Hagen, 1866:445). 

Distribution. — USA: CT 3 , FL 3 , IA 7 , IL 3 , 
IN 4 , KS 2 , MA 3 , MD 3 MI 3 , MO 5 , NC 3 , NH 3 , 
NJ 1 , NY 3 , VA 3 , WS 6 (' Smith, 1900:56; 2 Smith, 



1925:169; 3 Banks, 1927:7; Montgomery & 
Trippel, 1933:260, 5 Froeschner, 1947:128; 
6 Throne, 1972: 124; 7 Penny, NEW STATE RE- 
CORD). 
Dendroleon speciosus Banks, 1905b:7. 

Holotype female: USA, Colorado, Boulder 
(MCZ). 

DISTRIBUTION. — USA: AZ 2 , CA 2 , CO 1 , 
NM 2 ('Banks, 1905b:7; 2 Banks, 1927:8). 



NEMOLEONTINl 
Eremoleon Banks, 1901a:366 

Eremoleon femoralis (Banks), 1942:146 
[Psammoleon]. 

Holotype (sex unknown): 20 mi N. W. La Paz, 
Lower California, 1 6. VII. 1 938, Michelbacher & 
Ross (CAS). 

TAXONOMY. — To Eremoleon (see Stange, 
1970b:21). 

Distribution. — USA: AZ (Stange. NEW 
NATIONAL RECORD) 
Eremoleon gracile Adams, 1957b:90. 

Holotype female: USA, California, Riverside, 
31. VIII. 1939, P. DeBach (CAS). 

Distribution. — USA: CA (Adams, 
1957b:90). 
Eremoleon insipidus Adams, 1957b:88. 

Holotype male: 5 mi S. San Miguel, Lower 
California, 20.VII.1938, Michelbacher & Ross 
(MCZ). 

Distribution. — USA: CA (Adams, 
1957b:89). 

Eremoleon macer (Hagen), 1861:236 [Myrme- 
leon]. 

Holotype (sex unknown): Mexico (Vienna). 

= Segura vitreus Navas, 19 14g: 1 8. Holotype 
male: Mexico, Morelos, Cuernavaca, 1871, 
Bilimek (MNHN). 

= Novulga mexicana Navas, 1925c: 189. Holo- 
type male: Mexico, Veracruz, Veracruz, 1921 
(MNHN). 

= Hesperoleon atomarius Navas, 1933c: 105. 
Holotype (sex unknown): Mexico, Morelos, 
Cuernavaca, 1929-30 (Hamburg, specimen de- 
stroyed). 

TAXONOMY. — To Eremoleon (see Banks, 
1901:366); S. vitreus = E. macer (see Banks, 
1927:7 1); N. mexicana = E. macer, H. atomarius 
= E. macer (see Stange, 1970:21). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



77 



Distribution. — USA: AZ (Banks, 
1927:71). 
Eremoleon nigribasis Banks, 1920:329. 

Syntypes male and female: USA, Utah, St. 
George, VI 5,6 (MCZ). 

= Eremoleon affine Banks, 1942:144. Holo- 
typemale: Mexico, Baja California Sur. Miraflo- 
res, VII.8. 1938 (CAS). 

TAXONOMY. — E. affine = E. nigribasis (see 
Adams, 1957b:91). 

Distribution. — USA: AZ 2 , NM 2 , UT 1 
('Banks, 1920:329; 2 Adams, 1957b:91). 
Eremoleon pallens Banks, 1941b: 101. 

Syntypes (sex unknown): USA, Arizona, 
Pichaco Peak. VII. 23, Bequaert, Tinkham, Flock 
(MCZ). 

Distribution. — USA: AZ (Banks, 
1941b:101). 

Glenurus Hagen, 1866:372,405 

Glenurus gratus (Say), 1839:45 [Formicaleo]. 

Holotype (sex unknown): USA, Indiana, 
Evansville, Corson (depository unknown). 

— Myrmecoleon roseipennis Burmeister, 
1839:995. Holotype (sex unknown): Nord 
Amerika (Halle). 

Taxonomy. — To Myrmeleon (see Walker, 
1853:392); to Glenurus {see Hagen, 1866:405); 
to Dendroleon (see Banks, 1892:360); returned 
to Glenurus (see Banks, 1907c:30); M. roseipen- 
nis = M. gratus (see Hagen, 1861:225). 

Distribution. — USA: FL^IN ' , KY 3 , MO 4 , 
NJ 2 , OH 3 , TN 3 ( ' Say, 1 839:45; "Smith, 1900:56; 
3 Banks, 1927:68; 4 Froeschner, 1947:128). 

Remarks. — Carpenter (1992:538) has indi- 
cated that Burmeister's Handbuch der Entolo- 
mologie was apparently published in two parts. 
The first 400 pages were published in 1839, 
while the remainder, including all Neuroptera, 
were published in 1838. If this is found to be 
correct, it could lead to serious nomenclatorial 
instability. The synonymy of Glenurus gratus 
and Glenurus roseipennis would be reversed. 
Almost certainly the International Commission 
on Zoological Nomenclature would be petitioned 
to declare, for nomenclatorial puiposes, that the 
entire book was published in 1839. 
Glenurus luniger Gerstaecker, 1894:125. 

Holotype female: Panama, Chiriqui (Greif- 
swald). 



Distribution. — USA: AZ (Stange, 
1970b:22). 
Glenurus snowii Banks, 1907b: 100. 

Holotype female: USA, Arizona, Baboquivari 
Mountains (MCZ). 

Distribution. — USA: AZ (Banks, 
1907b: 100). 

Psammoleon Banks, 1899b:69 

Psammoleon albovaria (Banks), 1942:146 
[Puren]. 

Holotype (sex unknown): Mexico, Baja Cali- 
fornia, Venancio, 17. VII. 1938, Michelbacher & 
Ross (CAS). 

TAXONOMY. — To Psammoleon (see Stange, 
1970b:23). 

Distribution. — USA: CA (Stange, NEW 
NATIONAL RECORD) 
Psammoleon arizonensis Banks, 1935:53. 

Three syntype females: USA, Arizona, Tucson 
(one syntype); Phoenix (two syntypes) (MCZ). 

Distribution. — USA: AZ (Banks, 
1935:53). 

Psammoleon bistictus (Hagen), 1861:235 
[Myrmeleon]. 

Holotype (sex unknown): Cuba, Poey (deposi- 
tory unknown). 

Taxonomy. — To Nelees (see Navas, 
1921:117); to Psammoleon (see Banks, 
1927:61). 

Distribution. — USA: FL (Banks, 
1927:62). 

Psammoleon connexus (Banks), 1920:329 
[Puren]. 

Holotype male: USA, California, San Jacinto 
Mountains, 25.VI (MCZ). 

TAXONOMY. — To Psammoleon (see Banks, 
1927:64). 

Distribution. — USA: AZ 2 , CA 1 , TX 2 
('Banks, 1920:329; 2 Banks, 1927:65). 
Psammoleon decipiens Banks, 1935:53. 

Two syntypes: USA, Georgia, Scriven Co., 
Mi 1 1 in (one male); Louisiana, Shreveport (one of 
unknown sex) (MCZ). 

Distribution. — USA: GA, LA (Banks, 
1935:53). 
Psammoleon guttipes Banks, 1906a:99. 

Holotype male: USA, North Carolina, Tryon 
(MCZ). 

Distribution. — USA: FL 3 , GA 3 , KS 2 , LA 3 , 
MO 4 , NC 1 , NJ 3 , TX 3 ('Banks, 1906a:99; 



78 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



2 Smith, 1925:170; 3 Banks, 1928:64; 4 Froesch- 

ner, 1947:128). 

Psammoleon inscriptus (Hagen), 1861:230 

[Myrmeleon]. 

Holotype male: USA, western Texas (now 
New Mexico), Pecos River, Pope (MCZ). 

TAXONOMY. — To Psammoleon (see Banks, 
1904b: 106); to Puren (see Banks, 1927:66); re- 
turned to Psammoleon (see Stange, 1970b:24). 

Distribution. — USA: CA , CO 2 , NM 1 , 
UT 3 ('Hagen, 1851:230; 2 Hagen, 1875:922; 
3 Banks, 1927:66; 4 Stange, NEW STATE RE- 
CORD). 
Psammoleon minor Banks, 1927:62. 

Lectotype female: USA, Florida, Loggerhead 
Key, Dry Tortugas, VI. 19 17 (MCZ). 

TAXONOMY. — Lectotype designated by 
Stange, 1961:674. 

Distribution. — USA: FL (Banks, 
1927:63). 
Psammoleon normalis Banks, 1942:145. 

Holotype female: Mexico, Baja California, 
Venancio, 17.VII.1938 (CAS). 

Distribution. — USA: AZ, CA (Stange, 
NEW NATIONAL RECORDS) 
Psammoleon sinuatus Currie, 1903:275. 

Holotype male: USA, Arizona, Santa Rita 
Mountains, 31. V(USNM). 

= Psammoleon serpentinus Navas, 1923:185. 
Holotype (sex unknown): USA, New Mexico, 
Jemez Springs, VII. 19 16, Woodgate (MZB). 

TAXONOMY. — P. serpentinus = P. sinuatus 
(see Banks, 1927:5). 

DISTRIBUTION. — USA: AZ 1 , CO 1 , NM 2 , 
UT 2 (' Currie, 1903:275; 2 Banks, 1927:64). 

BRACHYNEMURINi 

Abatoleon Banks, 1924:436 

Abatoleon dorsalis (Banks), 1903c:240 
[Brae In mem urns] . 

Holotype female: USA, Texas, Laredo (MCZ). 

= Brachynemurus curriei McClendon, 
1906a:93. Holotype male: USA, Texas, Browns- 
ville (USNM). 

= Netroneurus pygmaeus Esben-Petersen, 
1933:1 1 1. Holotype male: USA, Texas. Dallas, 
Boll (Hamburg, specimen destroyed) 

TAXONOMY. — To Austroleon (see Banks, 
1928:54); \o Abatoleon (see Stange, 1994:80); B. 



curriei = B. dorsalis (see Banks, 1927:54); N. 
pygmaeus = B. dorsalis (see Stange, 1970a:66). 

Distribution. — USA: TX (Banks, 
1903c:240). 

Abatoleon indiges (Walker), 1860: 189 [Myrme- 
leon]. 

Holotype male: Haiti (BMNH). 

= Nelees cubitalis Navas, 1 92 1 : 1 1 7. Holotype 
female: Cuba, Habana, PlayaChivo. 7.VII.1919, 
Cervera (MCZ). 

TAXONOMY. — To Clathroneuria (see Navas, 
1922b: 168); to Austroleon (see Smith, 
1931:816); to Abatoleon (see Stange, 1994:80); 
N. cubitalis = A. indiges (see Smith, 193 1 :8 16). 

Distribution. — USA: FL (Stange, 
1970a:68). 

Atricholeon Stange, 1994:75, 82 

Atricholeon tuberculatus (Banks), 1899b:70 
{Brachynemurus]. 

Holotype female: USA, New Mexico, Mesilla 
(MCZ). 

TAXONOMY. — To Atricholeon (see Stange, 
1994:82). 

Distribution. — USA: AZ 2 , NM 1 ('Banks, 
1899b:70; 2 Banks, 1928:49). 

Brachynemurus Hagen, 1888:34 

Brachynemurus abdominalis (Say), 1823:163 
[Myrmeleon]. 

Neotype male: USA, Arkansas, Hope, 8. VI, 
Knobel (MCZ). 

= Myrmeleon iniquus Walker, 1853:330. 
Holotype male: (no locality data) (BMNH). 

= Myrmeleon salvus Hagen, 1861:227. Lecto- 
type male: USA, D. C, Washington (MCZ). 

= Myrmeleon juvencus Hagen, 1861:234. 
Holotype female: USA, western Texas (now 
New Mexico), Pecos River, VII, Pope (MCZ). 

= Brachynemurus tenuis Banks, 1898:204. 
Lectotype male: USA, New Mexico, Mesilla, 
28.VI.1897, Pope (MCZ). 

= Hesperolcon placidus Navas, 19 1 5a:5 1 . 
Holotype male: USA, New Jersey, Pt. Pleasant 
(depository unknown). 

Taxonomy. — To Macronemurus (see 
Hagen, 1866:424); to Brachynemurus (see 
Hagen, 1888:57); to Hesperoleon (see Banks, 
1913b:65); returned to Brachynemurus (see 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



79 



Stange, 1970a:76); M. juvencus = M. abdomi- 
nalis (see Hagen, 1888:5759); M. salvus = B. 
abdominalis (see Banks, 1907c:3 1 ); H. placidus 
= B. abdominalis (see 'Banks, 1927:5); M iniquus 
= B. abdominalis, B. tenuis = B. abdominalis; 
neotype for A/, abdominalis and lectotype for A/. 
salvus designated (see Banks, 1938d:413-414); 
lectotype for B. tenuis designated (see Stange, 
1961a:677). 

Distribution. — CAN: BC 6 , MB 4 ON 4 , 
USA: AR 8 , AZ 4 CA 4 CO 4 , CT 4 DC 4 GA 4 , 
IA 8 , ID 8 , IL 8 , IN 5 , KS, LA 8 , MA 4 , MD, MI 8 , 
MN 4 , MO 7 , NC 4 , ND 4 , NE 4 , NJ 2 , NM 1 , NV 8 , 
NY 4 , OK 8 OR 4 , PA 8 , SD 8 , TN 8 , TX 4 , UT 4 , 
VA 4 , WA 4 , WS 9 , WY 8 ('Hagen. 1861:234; 
2 Smith, 1900:56; 3 Smith, 1925:169; 4 Banks, 
1928:39; ^Montgomery & Trippel. 1933:260; 
6 Spencer, 1942:28; Troeschner, 1947:128; 
8 Stange, 1970a:map 7; 9 Throne, 1972:123). 
Brachynemurus blandus (Hagen), 1861:235 
[Myrmeleon]. 

Holotype female: USA, western Texas (now 
New Mexico), Pecos River, Pope (MCZ). 

TAXONOMY. — To Brachynemurus (see 
Hagen, 1888:73); to Hesperoleon (see Banks, 
1913b:65); returned to Brachynemurus (see 
Stange, 1970a:77). 



Distribution. - 
KS 2 , NM 1 , NV 



AZ 4 . CA 4 , 
UT 3 , "'* 4 



CO- 



USA: 

, tx 4 , ut j , wa; wy- 

( T Hagen, 1861:235: 2 Smith, 1925:170: J Banks, 
1927:29; 4 Stange, 1970a:78). 
Brachynemurus californicus Banks, 

1895a:519. 

Lectotype male: Mexico, Baja California Sur, 
San Jose del Cabo (MCZ). 

= Brachynemurus fraternus Banks, 
1895a:520. Lectotype male: Mexico. Baja Cali- 
fornia Sur, San Jose del Cabo (MCZ). 

= Comptesa ambitiosa Navas. 1 9 1 5b:464. 
Holotype male: Mexico, Baja California 
(MNHN). 

= Belluga implexa Navas, 1920: 195. Holotype 
female: Mexico, Baja California Sur, La Paz, 
1914, Diguet (MNHN). 

TAXONOMY. — B. fraternus = B. californicus, 
C. ambitiosa = B. californicus, B. implexa = B. 
californicus (see Banks, 1942:139); lectotypes 
designated for B. californicus and B. fraternus 
(see Stange. 1961:675,676). 

Distribution. — USA: AZ. CA (Stange, 
1970a:93). 



Brachynemurus carolinus Banks, 191 1:349. 

Lectotype female: USA, North Carolina, 
Southern Pines (MCZ). 

TAXONOMY. — To Netroneums (see Banks, 
1927:42); returned to Brachynemurus (see 
Stange, 1961a:675); lectotype designated (see 
Stange, 1961a:675). 

Distribution. — USA: FL, GA, NC (Banks, 
1927:43). 
Brachynemurus elongatus Banks, 1904b: 105. 

Lectotype male: USA, New Mexico, Mesilla, 
30.VI.1897, Morse (MCZ). 

Taxonomy. — Lectotype designated (see 
Stange, 1961:676). 

Distribution. — USA: AZ 1 , CA 2 , CO 1 , 
NE 2 , NM 1 , NV 2 , UT 1 ('Banks, 1928:48; 
2 Stange. 1970a:88). 

Brachynemurus ferox (Walker), 1853:332 
[Myrmeleon]. 

Holotype male: USA, California, Hartweg 
(BMNH). 

= Brachynemurus brunneus Currie, 1898:273. 
Holotype male: USA, Wyoming, Yellowstone 
National Park, Fountain, 10.VIII.1896, Currie 
(USNM). 

= Brachynemurus centralis Banks, 1898:204. 
Lectotype female: USA, Colorado, Estes Park 
(MCZ). 

Taxonomy. — To Macronemurus (see 
Hagen, 1866:424); to Hesperoleon (see Banks. 
1913b:64); to Brachynemurus (see Stange, 
1970a:81); B. centralis = B. brunneus (see 
Banks, 1907c:31); B. brunneus = B. ferox (see 
Stange, 1970a:81). 

Distribution. — CAN: BC 4 , USA: AZ 3 ^ 
CA 1 , CO 3 , NM 3 , NV 3 , OR 5 , UT 3 , WY" 
('Walker, 1853:332; 2 Currie 1898:204; 3 Banks, 
1928:25, as H. brunneus; 4 Spencer, 1942:28; 
5 Stange, 1970a:81). 

Brachynemurus fuscus (Banks). 1905b:6 
[Calinemurus]. 

Lectotype male: USA, Arizona, Nogales, 
6.VII. 1903, Oslar (MCZ). 

TAXONOMY. — To Brachynemurus (see 
Stange, 1970a:93); lectotype designated (see 
Stange,1961a:677). 

Distribution. — USA: AZ (Banks, 
1905b:6). 

Brachynemurus henshawi (Hagen), 1887:216 
[Maracanda]. 

Holotype male: USA, Oregon, Umatilla, 
24.VI.1882, Henshaw (MCZ). 



80 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



TAXONOMY. — To Oyptolcon (see Banks, 
1907c:32); to Brachynemurus (see Stange, 
1970a:82). 

DISTRIBUTION. — USA: CA 2 , NV 2 , OR 1 
('Hagen, 1887:216; 2 Stange, 1970a:82). 
Brachynemurus hubbardi Currie, 1898:241. 

Lectotype male: USA, Arizona, Fort Grant, 
22.VII.1897, Hubbard (MCZ). 

= Brachynemurus cockerelli Banks, 1902:86. 
Holotype female: USA, New Mexico, Lone 
Mountain (MCZ). 

= Brachynemurus hubbardi nubeculipennis 
Currie, 1903:277. Lectotype male: USA, Ari- 
zona, Phoenix (USNM). 

= Brachynemurus hubbardi curtus Banks, 
1927:31. Lectotype male: USA, Kansas, Seward 
Co., 18.VIII.191 1, Williams (MCZ). 

TAXONOMY. — To Hesperoleon (see Banks, 
1913:65); returned to Brachynemurus (see 
Stange, 1970a:88); B. cockerelli = B. hubbardi 
(see Currie, 1903:277); B. h. nubeculipennis = B. 
h. hubbardi (see Banks, 1907c:31); B. h. curtus 
= B. h. hubbardi (see Stange, 1970a:88); lecto- 
type designated for B. h. curtus (see Stange, 
1961a:677); for B. hubbardi and B. h. nubeculi- 
pennis (see Stange, 1970a:88). 

DISTRIBUTION. — USA: AZ 1 , CO 5 , KS 3 , 
NM 2 , NV 5 , OK 5 , TX 4 , UT 4 (' Currie, 1898:241; 
2 Banks, 1902:86; 3 Smith, 1925:169; 4 Banks, 
1927:30, 31; 5 Stange, 1970a:89). 
Brachynemurus irregularis Currie, 1906:186. 

Holotype female: USA. Texas, Columbus. 
VI. 1 879, Schwarz (USNM). 

= Mastega texanus Navas, 1 9 1 4b: 1 3 . Holotype 
female: USA, Texas (Vienna). 

TAXONOMY. — B. irregularis to Hesperoleon 
and M. texanus to Ciyptoleon (see Banks, 
1927:5, 36); returned to Brachynemurus; M. tex- 
anus = B. irregularis) (see Stange, 1970a:78). 

Distribution. — USA: IL 1 , KS 2 , OK 3 , TX 1 
('Currie, 1905:186; 2 Smith, 1925:170; 3 Stange, 
1970a:79). 

Brachynemurus longicaudus (Burmeister), 
1839:994 [Myrmecoleon]. 

Syntypes male and female: "Nord-Amerika," 
Winthem Coll. (MCZ). 

= Myrmecoleon irroratus Burmeister, 
1839:995. Two syntype females: USA, South 
Carolina (Halle). 

Taxonomy. — To Mynneleon (see Hagen, 
1860:365); to Macronemurus (see Hagen, 
1866:424); to Brachynemurus (see Hagen, 



1888:35); M. longicaudus = M. abdominalis, M. 
irroratus = M. conspersus (see Hagen, 
1860:365); M. irroratus = M. talpinus (see 
Hagen, 1 86 1 :226); M. irroratus = M. conspersus 
(see Taschenberg, 1879:2 13); B. longicaudus not 
= B. abdominalis, B. irroratus not = B. consper- 
sus, B. irroratus = B. longicaudus (see Hagen. 
1888:3536). 

Distribution. — USA: FL 3 , GA 2 , IN 5 , MS 4 , 
NC 3 , NJ 3 , SC 1 , VA 4 ('Burmeister, 1839:995; 
2 Hagen, 1861:227; 3 Banks, 1927:46; 4 Stange, 
1970a:83; 5 Lawson & McCafferty, 1984:130). 
Brachynemurus mexicanus Banks, 1895a:520. 

Lectotype female: Mexico, Tepic (MCZ). 

TAXONOMY. — B. mexicanus = B. versutus 
(see Banks, 1901:365); B. mexicanus not = B. 
versutus (see Banks, 1927:32); lectotype desig- 
nated by Stange, 1961:676. 

Distribution. — USA: AZ 1 , NM 2 ('Banks, 
1927:33; 2 Stange, 1970a:91). 
Brachynemurus nebulosus (Olivier), 181 1:127 
[Mynneleon]. 

Type(s) (sex unknown): USA, New York, M. 
Bote (depository unknown). 

= Mynneleon talpinus Hagen, 1866:448 (no- 
men nudum). Type(s) (sex unknown): USA, 
South Carolina, Zimmerman (depository un- 
known). 

= Myrmecoleon contaminants Burmeister, 
1839:995. Syntype females: USA, South Caro- 
lina, Zimmerman (Halle). 

= Mynneleon conspersus Rambur, 1842:387. 
Holotype female: "FAmerique septentrionale" 
(IRSNB). 

TAXONOMY. — To Macronemurus (see 
Hagen, 1866:424); to Maracanda (see Hagen, 
1 887:2 12); to Ciyptoleon (see Banks, 1901:330); 
to Brachynemurus (see Stange, 1970a:84); M. 
irroratus (?) = M. conspersus (see Walker, 
1853:329); M. irroratus = M. abdominalis, M. 
conspersus = M. abdominalis, M. talpinus = M. 
abdominalis (see Hagen, 1861:226); M. talpinus 
= M. nebulosus, M. contaminants = M. nebulo- 
sus, M. irroratus = M. nebulosus, M. conspersus 
= M. nebulosus (see Hagen, 1887:212). 

Distribution. — CAN: ON 4 , USA: FL 4 , 
GA 4 , IL 6 , IN 7 , LA 6 , MI 4 , MN 5 NC 4 , NE 6 , NJ 3 , 
NY ' , OH 4 , OK 6 . PA 6 , SC 2 , TX 6 , WS 6 ( ' Olivier, 
1811:127; 2 Burmeister, 1839:995; 3 Smith, 
1900:56, as M. conspersa; 4 Banks, 1927:11: 
5 Parfin, 1952:425; 6 Stange, 1970a:84; 7 Lawson 
& McCafferty, 1984:130). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



Brachynemurus pulchellus Banks, 191 1:348. 

Holotype female: USA, California, Brown, 
10.X, Grinnell (MCZ). 

= Clathroneuria exigua Navas, 1920:197. 
Holotype male: Mexico, Baja California Sur, La 
Paz, 1914, Diguet (MNHN). 

= Netroneurus pulchellus pallescens Banks, 
1942:140. Holotype male: Mexico, Baja Califor- 
nia, San Domingo (CAS). 

Taxonomy. — To Netroneurus (see Banks, 
1927:42); returned to Brachynemurus (see 
Stange, 1970a: 102); C. exigua = N. pulchellus 
(see Banks, 1942:138); N. p. pallescens = B. 
pulchellus (see Stange, 1970a: 102). 

Distribution.— ^USA: AZ 2 , CA 1 , NV 2 
('Banks, 191 1:348; 2 Stange, 1970a:104). 
Brachynemurus ramburi Banks, 1907c:31 (re- 
placement name). 

Holotype female: "^Ameriqueseptentrionale ,, 
(IRSNB). 

= Myrmeleon nebulosus Rambur, 1 842:387. A 
junior primary homonym of M. nebulosus 
Olivier, 1811:127. 

Taxonomy. — To Macronemurus (see 
Hagen, 1866:444); to Brachynemurus (see 
Hagen. 1888:36). 

Distribution. — USA: FL, GA, NC (Banks, 
1927:47). 
Brachynemurus sackeni Hagen, 1888:94. 

Lectotype female: USA, Texas, Dallas, Boll 
(MCZ). 

= Bollenga dinamitensis Navas, 1932:14. 
Holotype female: Mexico, Dinamita (Turin). 

= Hesperoleon brevipilis Banks, 1938d:414. 
Holotype male: USA, Arizona, Colorado River 
(MCZ). 

TAXONOMY. — To Hesperoleon (see Banks. 
1913b:65); returned to Brachynemurus (see 
Stange, 1970a: 144); B. dinamensis = B. sackeni, 
H. brevipilis = B. sackeni (see Stange. 
1970a: 144); lectotype designated by Stange, 
1961a:676. 

Distribution. — USA: AZ 2 , CA 3 CO 3 
KS 4 , NE 4 , NM 3 . NV 3 , OR 4 , TX 1 , UT 3 , WY 4 
('Hagen, 1888:94; 2 Banks, 1938d:414; 3 Banks, 
1928:34; 4 Stange, 1970a: map 28). 
Brachynemurus seminolae Stange, 1970a:86. 

Holotype male: USA, Florida, Bay Co., St. 
Andrews State Park, 30.VIII.1960, L. A. Stange 
(UCD). 

Distribution. — USA: FL, NC (Stange, 
1970a:86). 



Brachynemurus signatus (Hagen), 1887:215 
[Maracanda]. 

Lectotype female: USA, Michigan, Lake Su- 
perior, Whitefish Point (MCZ). 

Taxonomy. — To Cryptoleon (see Banks, 
1907c:32); to Brachynemurus (see Stange, 
1970a:79); lectotype designated by Banks, 
1928:12. 

Distribution. — USA: CO 5 , DE 5 , IL 3 KS 2 , 
MI ' MN 4 , ND 5 , NE 5 , NJ 5 , NY 3 , OH 3 , PA 5 , RI 3 , 
WS 6 ('Hagen, 1887:215; 2 Smith, 1925:170; 
3 Banks, 1928:12; 4 Parfin, 1952:425; 5 Stange, 
1970a:80: 6 Throne, 1972:123). 
Brachynemurus versutus (Walker), 1853:331 
[Myrmeleon]. 

Holotype male: Mexico (BMNH). 

= Hesperoleon spegazzinius Navas, 1934:72. 
Two syntype females: Mexico, Texcoco, 
20.X.1930; Ixcapalapa, 15.IX.1931. Spegazzini 
(depository unknown). 

TAXONOMY. — To Macronemurus (see 
Hagen, 1866:449); to Hesperoleon (see Banks, 
1928:13); to Brachynemurus (see Stange, 
1970a:91); H. spegazzinius = B. versutus (see 
Stange 1970a:91). 

Distribution. — USA: AZ, CO, NM 
(Stange, 1970a:91). 

Chaetoleon Banks, 1920:328 

Chaetoleon pumilis (Burmeister), 1839:995 
[Myrmecolcon]. 

Type(s) (sex unknown): USA, South Carolina 
(Halle). 

= Myrmeleon angusticollis Rambur, 1 842:399. 
Holotype female: "Bombay" (IRSNB). 

TAXONOMY. — To Macronemurus (see 
Hagen, 1866:424); to Creagris (see McLachlan, 
1873b: 137); to Brachynemurus (see Banks, 
1892:361); to Dendroleon (see Banks, 
1907c:30); to Chaetoleon (see Banks. 
1920:328); returned to Brachynemurus (see 
Stange, 1970a: 100); returned to Chaetoleon (see 
Stange, 1994:84); M. angusticollis = B. pumilis) 
(see Stange, 1970a: 10). 

Distribution. — USA: FL 3 , GA 3 ,^ NC 3 , 
NY 2 . SC 1 ('Burmeister, 1839:995; 2 Smith, 
1900:56; 3 Banks, 1927:57). 
Chaetoleon pusillus (Currie), 1899:363 
[Brachynemurus]. 

Lectotype female: USA, New Mexico. Mesilla 
Park, 13. VI, Cockerell (USNM). 



82 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



TAXONOMY. — To Scotoleon (see Banks. 
1913b:65); to Chaetoleon (see Banks, 
1920:328); returned to Brachynemurus (see 
Stange, 1970a: 100); returned to Chaetoleon (see 
Stange, 1994:84); lectotype designated by 
Stange, 1970a: 101. 

Distribution. — USA: AZ 2 , CA 3 , CO 3 , 
NM 1 , NV 3 , TX 3 , UT 2 ('Currie, 1899:363; 
2 Banks, 1927:58; 3 Stange, 1970a:101). 
Chaetoleon tripunctatus (Banks), 1922:60 
[Hesperoleori]. 

Lectotype male: USA, Arizona, Palmerlee, VI 
(MCZ). 

Taxonomy. — To Austroleon (see Banks, 
1928:55); to Brachynemurus (see Stange, 
1970a:98); to Chaetoleon (see Stange, 1994:84); 
lectotype designated by Stange, 1961:677. 

Distribution. — USA: AZ (Banks, 
1922:60). 

Clathroneuria Banks, 19 13b:65 

Clathroneuria arapahoe Banks, 1938d:418. 

Holotype male: USA, Arizona, Tucson 
(MCZ). 

TAXONOMY. — To Brachynemurus (see 
Stange, 1970a: 105); returned to Clathroneuria 
(see Stange, 1994:85). 

Distribution. — USA: AZ (Banks, 
1938d:418). 

Clathroneuria coquilletti (Currie), 1898:93 
[Brachynemurus]. 

Holotype male: USA, Arizona, San Simon, 
5.VII.1897, Hubbard (USNM). 

= Brachynemurus maculosus Banks, 
1 899a: 1 70. Holotype male: USA, California, Te- 
hama (MCZ). 

Taxonomy. — To Hesperoleon (see Banks, 
1913b:65); to Brachynemurus (see Stange, 
1970a: 106); to Clathroneuria (see Stange, 
1994:85); B. coquilletti = B. blandus (see Banks, 
1899a: 171 ); B. coquilletti not = B. blandus)(see 
Currie, 1903:276); B. maculosus = B. coquilletti) 
(see Stange, 1970a: 106). 

Distribution. — USA: AZ 1 , CA 1 , KS 2 , 
NM 2 , NV 3 ^ TX 2 , UT 2 , WA 2 , WY 3 (' Currie, 
1898:93; "Banks, 1927:32, 37; 3 Stange, 
1970a:Map 17). 
Clathroneuria navajo Banks, 1938d:4 18. 

Holotype female: USA, Arizona, Pinery Can- 
yon (MCZ). 



TAXONOMY. — To Brachynemurus (see 
Stange, 1970a: 107); returned to Clathroneuria 
(see Stange, 1994:85). 

DISTRIBUTION. — USA: AZ 1 , CA 2 ('Banks, 
1938d:418; 2 Stange, 1970a:107). 
Clathroneuria schwarzi (Currie), 1903:280 
[ Br achy mem uri is ] . 

Holotype male: USA, Arizona, Flagstaff 
(USNM). 

TAXONOMY. — To Clathroneuria (see Banks, 
1913b:63); returned to Brachynemurus (see 
Stange, 1970a: 107); returned to Clathroneuria 
(see Stange, 1994:85). 



,3 



DISTRIBUTION. — USA: AZ ' CA", CO, ID J , 
3 OR 3 , TX 3 ,UT",WY 3 (^Currie, 



,3 



OK : 



'Banks, 1927:52; J Stange, 



NM",NV . 

1903:280; 

1970a: 107). 

Clathroneuria westcotti (Stange), 1970a: 109 

[Brachynemurus]. 

Holotype male: USA, Arizona, Cochise Co., 
west side of Willcox Dry Lake, 7. VIII. 1959, 
R. L. Westcott (LACM). 

TAXONOMY. — To Clathroneuria (see 
Stange, 1994:85). 

Distribution. — USA: AZ (Stange, 
1970a: 110). 

Mexoleon Stange, 1994:75, 86 

Mexoleon papago (Currie), 1899:361 
[Brachynemurus]. 

Holotype male: USA, Arizona, Santa Rita 
Mountains, Madera Canyon, 7.VI.1898, 
Schwarz (USNM). 

= Clathroneuria orioles Banks, 1941b: 103. 
Holotype male: USA, California, Riverside, 
25.IX.1940, G. & H. Sperry (MCZ). 

TAXONOMY. — To Hesperoleon (see Banks, 
1913b:65); returned to Brachynemurus (see 
Stange, 1970a:97); to Mexoleon (see Stange, 
1994:86); C. orioles = B. papago (see Stange, 
1970a:97). 

Distribution 
('Currie, 1899:361; "Banks, 1941b: 103; 
3 Stange, 1970a:97). 



USA: AZ',CA 2 ,OR 3 ,TX 3 



Scotoleon Banks, 19 13b:65 

Scotoleon carrizonus (Hagen), 1888:93 
[Brachynemurus]. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



83 



Lectotype male: USA, Texas, Dimmit Co., 
Carrizo Springs (MCZ). 

= Hesperoleon douglasi Banks, 1927:20. Lec- 
totype male: USA, Arizona, Nogales, 
6.VII.1903,Oslar(MCZ). 

Taxonomy. — To Hesperoleon (see Banks, 
19 1 3b:64); returned to Brachynemurus (see 
Stange, 1970a: 11 7); to Scotoleon (see Stange, 
1994:87); H. carrizonus = H. peregrinus (see 
Banks, 19 13b:64); H. carrizonus not = H. pere- 
grinus (see Banks, 1927:16); H. douglasi - B. 
carrizonus) (see Stange, 1970a: 117); lectotype 
designated by Stange. 1961a:675. 

Distribution. — USA: AZ 2 , CA 3 .^ NM 2 , 
NV 3 . OK 3 , TX',UT 3 ('Hagen, 1888:93; 2 Banks, 
1927:20; 3 Stange, 1970a: 120). 
Scotoleon deflexus (Adams), 1957b:98 [Hes- 
peroleon]. 

Holotype male: USA, California, San Ber- 
nardino Co., Baker, 23.VIII.1952 (CAS). 

TAXONOMY. — To Brachynemurus (see 
Stange. 1970a: 120); to Scotoleon (see Stange, 
1994:87). 

DISTRIBUTION. — USA: CA 1 , NV 2 ('Adams, 
1957b: 1 00; 2 Stange, 1970a: 121). 
Scotoleon dissimilis (Banks), 1903d: 175 
[Brachynemurus]. 

Lectotype male: Mexico, Baja California Sur, 
San Jose del Cabo (MCZ). 

= Brachynemurus plectus Navas, 19 13e:49. 
Lectotype male: Mexico, Basse-Califomie, 
Diguet(MNHN). 

= Hesperoleon planus Navas, 1920:200. Holo- 
type male: Mexico, Baja California Sur, La Paz, 
1914, Diguet (MNHN). 

= Hesperoleon curriei Banks, 1938d:416. 
Holotype female: USA, Arizona, Santa Catalina 
Mountains (MCZ). 

Taxonomy. — To Hesperoleon (see Banks, 
1942:136); to Scotoleon (see Stange, 1994:87); 
B. plectus and H. planus probably = B. dissimilis) 
(see Banks, 1942: 135); B. curriei = B. dissimilis 
(see Stange, 1970a: 121); lectotypes designated 
fori?, dissimilis (see Stange, 1961a:675) and B. 
plectus (see Stange, 1 970a: 121. 

DISTRIBUTION. — USA: AZ 1 , TX 2 ('Banks, 
1938d:416, as H curriei; 2 Stange, 1970a: 122). 
Scotoleon eiseni Banks, 1908c:33 [Brachyne- 
murus]. 

Lectotype male: Mexico, Baja California Sur, 
San Jose del Cabo (MCZ). 



Taxonomy. — To Hesperoleon (see Banks, 
1942:137); to Scotoleon (see Stange, 1994:87); 
lectotype designated (see Stange, 1961a:676). 

Distribution. — USA: AZ, CA, NV 
(Stange, 1970a: 123). 

Scotoleon expansus (Navas), 1913e:48 
[Brachynemurus]. 

Holotype female: Mexico, Baja California, 
Diguet (MNHN). 

= Hesperoleon apache Banks, 1938d:417. 
Holotype female: USA, Arizona, Globe (MCZ). 

TAXONOMY. — To Hesperoleon via synon- 
ymy with H. sackeni (see Banks, 1942:138); 
returned to Brachynemurus (see Stange, 
1970a: 125); to Scotoleon (see Stange, 1994:87); 
B. expansus = H. sackeni (see Banks, 1942: 138); 
B. expansus not = B. sackeni, H. apache = B. 
expansus (see Stange, 1970a: 125). 

DISTRIBUTION. — USA: AZ 1 , CA 2 ^ NM 2 . 
TX 2 ('Banks, 1938d:417 as H apache; 'Stange, 
1970a:125). 

Scotoleon fidelitas (Adams), 1957b:94 [Hes- 
peroleon]. 

Holotype male: USA, Arizona, Santa Rita 
Mountains, Box Canyon, 21. VIII. 1949 (MCZ). 

TAXONOMY. — To Brachynemurus (see 
Stange. 1970a: 126); to Scotoleon (see Stange, 
1994:87). 

Distribution. — USA: AZ, CA (Adams, 
1957b:97). 

Scotoleon infuscatus (Adams), 1957b: 104 
[Hesperoleon]. 

Holotype male: USA, California, Contra Costa 
Co., Antioch, 24.V.1949, MacSwain (CAS). 

TAXONOMY. — To Brachynemurus (see 
Stange, 1970a: 127); to Scotoleon (see Stange, 
1994:87). 

Distribution. — USA: CA (Adams, 
1957b:105). 

Scotoleon intermedins (Currie), 1903:283 
[Brae In nemurus] . 

Holotype male: USA, Arizona, Phoenix 
(USNM). 

Taxonomy. — To Hesperoleon (see Banks, 
1927:18); returned to Brachynemurus (see 
Stange, 1970a: 127); to Scotoleon (see Stange, 
1994:87); B. intermedins = B. texanus (see 
Banks, 1907c:31); B. intermedins not = B. tex- 
anus (see Banks, 1927:18). 

Distribution. — USA: AZ 1 , CA 2 , NV 3 , UT 3 
('Currie, 1903:283; 2 Banks, 1927:19; 3 Stange, 
1970a:128). 



84 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Scotoleon longipalpis (Hagen), 1888:95 
[Brachynemurus], 

Lectotype male: Mexico, Baja California Sur, 
Cabo San Lucas, 1860, Xanthus (MCZ). 

= Scotoleon congener Banks, 1942: 138. Holo- 
type male: Mexico, Baja California, Chapala Dry 
Lake, 2 1 .VI, Michelbacher & Ross (CAS). 

Taxonomy. — To Scotoleon (see Banks, 
19 1 3b:64); returned to Brachynemurus (see 
Stange, 1970a: 128); returned to Scotoleon (see 
Stange, 1994:87); B. congener = B. longipalpis) 
(see Stange, 1970a: 129); lectotype designated 
(see Stange, 1961a:676). 

DISTRIBUTION. — USA: AZ 1 , CA 2 . NM 2 , 
NV 2 , TX 2 ('Currie, 1903:276; 2 Stange, 
1970a:Map 19). 

Scotoleon marshi (Stange), 1970a: 130 
[Brae In nemurus] . 

Holotype male: USA, Texas, Brewster Co., 22 
miles south of Marathon, 3.IX.1950, Stange & 
Marsh (UCD). 

TAXONOMY. — To Scotoleon (see Stange, 
1994:87). 

Distribution. — USA: TX (Stange, 
1970a: 130). 

Scotoleon minusculus (Banks). 1898:203 
[Brachynemurus]. 

Lectotype male: USA, California, Lancaster, 
VII, Morge (MCZ). 

= Brachynemurus texanus Banks, 1903d: 175 
Lectotype male: USA, Texas, Laredo, VIII, 
McClendon (MCZ). 

TAXONOMY. — To Hesperoleon (see Banks, 
1927:24); returned to Brachynemurus (see 
Stange, 1961a:676); to Scotoleon (see Stange, 
1994:87); B. texanus = B. minusculus (see 
Stange, 1970a: 13 1); lectotype designated for B. 
minusculus (see Stange, 1961a:676), and for B. 
texanus (see Stange, 1970a: 131). 

Distribution. — USA: AZ 3 , CA 1 , KS 5 , NE 5 , 
NM 4 , NV 5 . OK 5 , TX 2 , UT 5 ('Banks, 1898:203; 
2 Banks, 1903d: 175; 3 Currie, 1903:278; 4 Banks, 
1928:24; 5 Stange, 1970a: 131). 
Scotoleon minutus (Adams), 1957b: 100 [Hes- 
peroleon]. 

Holotype male: USA, California, Cathedral 
City, 24.VII.1950, Isaak (CAS). 

TAXONOMY. — To Brachynemurus (see 
Stange, 1970a: 132); to Scotoleon (see Stange, 
1994:87). 



Scotoleon niger (Currie), 1898:134 [Brachyne- 
murus]. 

Holotype female: USA, Arizona, Fort Grant, 
20.VII.1897, Hubbard (USNM). 

= Hesperoleon huachuca Banks, 1938d:415. 
Holotype male: USA, Arizona, Reef (MCZ). 

TAXONOMY. — To Hesperoleon (see Banks, 
1927:26); returned to Brachynemurus (see 
Stange, 1970a: 133); to Scotoleon (see Stange, 
1994:87); B. huachuca = B. niger (see Stange, 
1970a: 133). 

Distribution. — USA: AZ 1 , CA 2 , NV 2 
('Currie, 1898:134; 2 Stange, 1970a:134). 
Scotoleon nigrescens (Stange), 1970a: 135 
[Brachynemurus]. 

Holotype male: USA, Arizona, Cochise Co., 
13 miles east of Douglas, San Bernardino Ranch, 
12.VI.1959, L. Stange (UCD). 

TAXONOMY. — To Scotoleon (see Stange, 
1994:87). 

Distribution. — USA: AZ, TX (Stange, 
1970a: 136). 

Scotoleon nigrilabris (Hagen), 1888:72 
[Brachynemurus]. 

Lectotype female: USA, Colorado, Manitou 
(MCZ). 

TAXONOMY. — To Hesperoleon (see Banks, 
1913b:65); returned to Brachynemurus (see 
Stange, 1970a: 136); to Scotoleon (see Stange, 
1994:87); lectotype designated (see Banks, 
1927:27). 

Distribution. 

3 »™ T 5 »^6 



- USA: AZ 2 , CA 6 , CO 1 , ID 6 , 
NE 4 , NM 4 , NV 4 , SD 4 , TX 6 , 



Distribution. 



USA: AZ', CA', NV 
('Adams, 1957b: 102, 2 Stange, 1970a:133). 



KS J , MN J , Mr 

UT 4 , WY 6 ('Hagen. 1888:72; -Currie, 
1903:275; 3 Smith, 1925:169; 4 Banks, 1927:28; 
5 Parfin, 1952:425; 6 Stange, 1970a: 137). 
Scotoleon nivatensis (Navas), 1915a:52 [Hes- 
peroleon]. 

Holotype male: USA, Nevada, Lyon Co., Yer- 
ington (despository unknown). 

TAXONOMY. — To Brachynemurus (see 
Stange, 1970a: 138); to Scotoleon (see Stange, 
1994:87); H. nivatensis = H. singularis (see 
Banks, 1927:5): B. nivatensis not = B. singularis 
(see Stange, 1970a: 138). 

Distribution. — USA: AZ 2 , CA 2 , ID 2 , NV ' , 
OR 2 ('Navas, 1915a:52; 2 Stange, 1970a:138). 
Scotoleon pallidus (Banks), 1899a: 171 
[Brachynemurus]. 

Holotype female: USA, Arizona, Phoenix, IX, 
Kunze (MCZ). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



85 



TAXONOMY. — To Hesperolcon (see Banks, 
1913:65); returned to Brachynemurus (see 
Stange, 1970a: 139): to Scotoleon (see Stange, 
1994:87). 

Distribution. — USA: AZ 1 , CA 2 ^ NM 2 , 
NV 2 , TX 2 ('Banks, 1899a: 171; 



Stange. 
861:234 [Myr- 



1970a:139). 

Scotoleon peregrinus (Hagen), 

m el 'eon]. 

Lectotype female: USA, California. Havilah 
(MCZ). 

= Brachynemurus assimilis Banks, 1903d: 174 
Lectotype male: USA, California, Tehama, 
28.VIII. 1897 (MCZ). 

TAXONOMY. — To Brachynemurus (see 
Hagen, 1888:59); to Hesperoleon (see Banks. 
19 13b:64 as synonym of H. ferox); returned to 
Brachynemurus (see Stange, 1 970a: 14 1 ); to Sco- 
toleon (see Stange, 1994:87); lectotype desig- 
nated for H. peregrinus (see Banks, 1927: 16); for 
B. assimilis (see Stange. 1961:675); B. pere- 
grinus = B. ferox (see Banks, 1903d: 175); B. 
peregrinus = B. ferox (see Currie, 1903:276); H. 
carrizonus = H. peregrinus (see Banks, 
1938d:4 14); H. carrizonus not = H. peregrinus 
(see Stange, 1961a:676); B. ferox not = B. pere- 
grinus, B. assimilis = B. peregrinus (see Stange, 
1970a: 141). 

Distribution. — CAN: BC 2 , USA: AZ 2 , 
CA ' , CO 2 , NE 2 , NM 2 , N V 2 , UT 2 , WY 2 ( ' Hagen, 
1861:325; 2 Stange, 1970a:142). 
Scotoleon quadripunctatus (Currie), 1898:136 
[Brachynemurus]. 

Holotype female: USA, California, San Ber- 
nardino Co., Coquillett (USNM). 

TAXONOMY. — To Hesperoleon (see Banks, 
1927:21); returned to Brachynemurus (see 
Stange, 1970a: 143); returned to Scotoleon (see 
Stange, 1994:87). 

Distribution. — USA: AZ 2 , CA 1 , NV 3 
('Currie. 1898:136; 2 Banks, 1927:22; 3 Stange, 
1970a: 144). 

Scotoleon singularis (Currie), 1903:284 
[Brachynemurus]. 

Holotype male: USA, Arizona, Phoenix 
(USNM). 

TAXONOMY. — To Hesperoleon (see Banks, 
1913b:65); returned to Brachynemurus (see 
Stange. 1970a: 145); to Scotoleon (see Stange, 
1994:87); H. nivatensis = H. singularis (see 
Banks, 1927:5); B. nivatensis not = B. singularis 
(see Stange, 1970a: 145). 



Distribution. — USA: AZ 1 , CA, NV J , 
NM 3 ,UT 2 (' Currie, 1903:284; 2 Banks, 1927:18; 
3 Stange. 1970a: 146). 

Scotoleon yavapai (Currie), 1903:281 
[Brachynemurus]. 

Holotype male: USA, Arizona, Yavapai Co., 
Hot Springs (USNM). 

TAXONOMY. — To Hesperoleon (see Banks. 
1913b:65); returned to Brachynemurus (see 
Stange. 1970a: 148); returned to Scotoleon (see 
Stange, 1994:87). 

Distribution. — USA: AZ 1 , CA 3 , NV 3 , UT 2 
('Currie, 1903:281; " 
1970a: 148). 



'Banks, 1927:35; 'Stange 



Gnopholeontini 
Gnopholeon Stange, 1970a: 148 

Gnopholeon barberi (Currie), 1903:282 
[Brachynemurus]. 

Holotype female: USA, Arizona, Hot Springs 
(USNM). 

TAXONOMY. — To Scotoleon (see Banks, 
1913b:65); to Austroleon (see Banks, 1927:55); 
to Gnopholeon (see Stange, 1970a: 150). 

Distribution. — USA: AZ 1 , CA 2 (' Currie. 
1903:282; 2 Stange, 1970a:150). 
Gnopholeon delicatulus (Currie), 1903:279 
[Brachynemurus']. 

Holotype male: USA, Arizona, Phoenix, 
ll.V. 1898, Kunze (USNM). 

TAXONOMY. — To Scotoleon (see Banks, 
1913b:65); to Clathroneuria (see Banks, 
1927:52); to Gnopholeon (see Stange, 
1970a: 151). 

Distribution. — USA: AZ 1 , CA 2 . NV 2 
('Currie, 1903:279; 2 Stange, 1970a:152). 

Menkeleon Stange, 1970a: 154 

Menkeleon bellulus (Banks). 1905b:7 [Mara- 
canduld]. 

Lectotype female: USA, California. Three 
Rivers (MCZ). 

= Maracandula bellula Banks, 1905:90 (no- 
men nudum). 

= Maracandula minima Banks, 1942:140. 
Holotype male: USA. Mexico, Baja California. 
San Miguel (CAS). 



86 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



TAXONOMY. — To Menkeleon (see Stange, 
1970a: 155); M. minima = M. bellula (see Stange, 
1970a: 155); lectotype designated (see Stange, 
1970a:155). 

USA: AZ 2 , CA 1 . NM 3 . 



Distribution 
UT 2 ('Banks, 1905b:7; 
3 Stange, 1970a: 157). 



"Banks, 1927:59; 



Tyttholeon Adams, 1957b:106 

Tyttholeon puerilis Adams, 1957b: 107. 

Holotype male: USA, California, Riverside 
Co., Palm Springs, 27.VI.1939, Isaacs (CAS). 

Distribution. — USA: AZ, CA (Adams, 
1957b:108). 

MYRMELEONTFN1 

Myrmeleon Linnaeus, 1767:913 

Myrmeleon arizonicus Banks, 1943:76. 

Syntypes (sex unknown): USA, Arizona, Tuc- 
son, 6,20.VI; Texas, Brownsville, IV (MCZ). 

Distribution. — USA: AZ, TX (Banks, 
1943:76). 
Myrmeleon californicus Banks, 1943:77. 

Holotype female: USA, California, Gavilan, 
11. VII, G. &J. Sperry (MCZ). 

Distribution. — USA: CA (Banks, 
1943:77). 
Myrmeleon carolinus Banks, 1943:75. 

Syntypes (sex unknown): USA, North Caro- 
lina, Southern Pines, Will, Manee; Georgia, 
Scriben Co., Millin, VII, Morrison (MCZ). 

Distribution. — USA: GA, NC (Banks, 
1943:75). 
Myrmeleon crudelis Walker, 1853:388. 

Holotype (sex unknown): USA, E. Florida, St. 
John's Bluff, E. Doubleday (BMNH). 

= Myrmeleon tectus Walker, 1853:378. Lecto- 
type male: USA, E. Florida, St. John's Bluff, E. 
Doubleday (BMNH). 

TAXONOMY. — M. tectus = M. crudelis 
Stange, NEW SYNONYMY 

Distribution. — USA: FL 1 , GA 2 , MO 3 , 
NC 2 , NJ 2 ,NY 2 , TX 2 , VA 2 (' Walker, 1853:378; 
2 Banks, 1927:78; 3 Froeschner, 1947:128). 
Myrmeleon exitialis Walker, 1853:376. 

Holotype (sex unknown): USA, California, 
Hartweg's coll. (BMNH). 



= Myrmeleon immaculatus occidentalis Cur- 
rie, 1903:274. Holotype male: USA, Arizona, 
Williams (USNM). 

= Myrmeleon immaculatus coloradensis 
Banks, 1905a:90(nomen nudum). Holotype (sex 
unknown): USA, Nevada, Ormsby Co. (MCZ). 

TAXONOMY. — M. immaculatus occidentalis 
= M. exitialis, M. immaculatus coloradensis = M. 
exitialis) Stange, NEW SYNONYMY. 

Distribution. — CAN: BC 4 , USA: AZ 2 , 
CA 1 , NV 3 , ('Walker, 1853:376; 2 Currie, 
1903:274; 3 Banks, 1905a:90; 4 Spencer, 
1942:28, as M. immaculatus occidentalis). 
Myrmeleon immaculatus DeGeer, 1773:564. 

Type(s) (sex unknown): USA, Pennsylvania, 
Acrelius (NRS). 

= Myrmeleon melanocephalum Olivier, 
1811:127. Type(s) (sex unknown): USA, New 
York, environs of New York, M. Bote (deposi- 
tory unknown). 

Taxonomy. — To Neleon (see Navas, 
1915a:53); returned to Myrmeleon (see Banks, 
1928:74); M. melanocephalum = M. immacula- 
tus (see Hagen, 1888:188). 

Distribution. — USA: DC 3 , FL 5 , GA 3 , IN 6 , 
KS 4 , MD 5 , MI 5 , MN 8 , MO 7 , NC 5 , NH 5 . NY 2 , 
OH 5 , OK 5 , PA ' , VA 3 , WS 9 ( ' DeGeer, 1 773:564; 
2 01ivier, 1811; 3 Hagen, 1861:232; 4 Smith, 
1925: 170; 5 Banks, 1927:74 [western records ex- 
cluded as probably pertaining to M. exitialis]; 
Montgomery & Trippel, 1933:260; Froesch- 
ner, 1947:128; 8 Parfin, 1952:425; 9 Throne, 
1972:123). 
Myrmeleon insertus Hagen, 1861:233. 

Syntypes (sex unknown): Cuba, Poey; Haiti, 
Port-au-Prince (MCZ). 

Distribution. — USA: FL (Lucas & Stange, 
1981:213). 
Myrmeleon invisus Walker, 1853:379. 

Holotype (sex unknown): no locality data 
(BMNH). 

= Myrmeleon heriocles Banks, 19 14b:6 19. 
Syntypes (sex unknown): USA, North Carolina, 
Southern Pines, May (MCZ). 

TAXONOMY. — M. heriocles = M. invisus 
Stange, NEW SYNONYMY. 

Distribution. — USA: FL 2 , NC 1 , NJ 2 
('Banks, 1914b:618, as M. heriocles; 2 Banks, 
1927:76, as M. heriocles). 
Myrmeleon mexicanus Banks, 1903c:241. 

Syntypes (sex unknown): Mexico, Guadala- 
jara, McClendon (MCZ). 



PENNY. ADAMS and STANGE: SPECIES CATALOG 



87 



MO 4 , 



Distribution. — USA: AZ (Banks, 
1938d:420). 
Myrmeleon mobilis Hagen, 1888:204. 

Syntypes male and female: USA, Alabama, 
VII. 1883, Lyon (MCZ). 

= Myrmeleon mobilis Hagen, 1860:368 (no- 
men nudum). 

DISTRIBUTION. — USA: AL 1 , FL 5 . GA 2 , 
NC 3 , ('Hagen, 1888:204; "Banks, 
1927:75; 3 Brimley, 1938:30; 4 Froeschner, 
1947: 128; 5 Lucas & Stange, 1981:214). 
Myrmeleon rusticus Hagen, 1861:233. 

Lectotype (sex unknown): USA, western 
Texas (now New Mexico), Pecos River, 4. VIII, 
Pope (MCZ). 

- Myrmeleon diversus Hagen, 1873:729. 
Holotype (sex unknown): USA, Wyoming, Yel- 
lowstone National Park (MCZ). 

= Myrmeleon distans Banks, 1898:206. Holo- 
type (sex unknown): USA, California, Coronado 
Beach, VII, Morse (MCZ). 

= Myrmeleon agriope Banks, 19 14b:6 1 8. Syn- 
types (sex unknown): USA, California, Clare- 
mont; Arizona, Nogales; Arizona, Phoenix 
(MCZ). 

Taxonomy. — M. rusticus = M. crudelis (see 
Banks, 1907c:29); M. rusticus not = M. crudelis, 
M. distans = M. rusticus, M. agriope = M. ?-us- 
ticus (see Banks, 1927:78); M. diversus = M. 
rusticus Stange, NEW SYNONYMY; lectotype 
designated by Banks. 1927:78. 

DISTRIBUTION. — USA: AZ 4 , CA 3 , NM 1 , 



1861:233; "Hagen, 
:206; 4 Banks, 1927:78). 



Ur, WY" ('Hagen 

1873:729; 3 Banks, 1898 

(A New Jersey record by Smith, 1900:56 must 

certainly be a misidentification). 

Myrmeleon texanus Banks, 1900:596. 

Holotype (sex unknown): USA, Texas, 
Galveston, VI. 1900 (MCZ). 

Distribution. — USA: TX (Banks, 
1900:596). 

POLYSTOECHOTIDAE 
Polystoechotids. or giant lacewings, are 
among the largest of North American neuropter- 
ans, and among the least known. Only the first 
instar larva has been described (Welch, 1914), 
and no one has yet been able to get the young 
larvae to feed. Pupal skins of Platystoechotes 
have been found around the roots of incense 
cedar trees in the Sierra Nevada Mountains of 
California. Adult Platystoechotes emerge in 



early summer, while adult Polystoechotes 
emerge in late summer and early fall. Adult 
Polystoechotes are most frequently seen around 
the smoke of campfires, and sometimes are at- 
tracted to lights. The geographical distribution of 
Platystoechotes is entirely in the Sierra Nevada 
of California. The range of Polystoechotes at one 
time included almost all of northern North Amer- 
ica and much of California south to Panama. 
However, this genus cannot now be found over 
much of its former range, and today appears 
confined to remote mountainous areas. 

Platystoechotes Carpenter, 1940:270 

Platystoechotes lineatus Caipenter. 1940:271. 

Holotype male: USA, California, Wolverton, 
Sequoia National Park, 7000-9000', 
14VI.1929, E. C. Van Dyke (CAS). 

Distribution. — USA: CA (Caipenter, 
1940:271). 

Polystoechotes Burmeister, 1839:982 

Polystoechotes punctatus (Fabricius), 1793:73 
[Semblis]. 

Holotype (sex unknown): without locality data 
(BMNH). 

= Hemerobius nebulosus Fabricius, 1798:202. 
Type(s) (sex unknown): (locality unknown), (de- 
pository unknown). 

= Hemerobius irroratus Say, 1824:306. 
Type(s) (sex unknown): USA (depository un- 
known). 

= Hemerobius vittatus Say, 1824:307. Type(s) 
(sex unknown): USA: New Jersey; Pennsylvania 
(ANSP). 

= Polystoechotes sticticus Burmeister, 
1839:982. Type(s) (sex unknown): North Amer- 
ica (Halle). 

= Osmylus validus Walker, 1853:233. 5 syn- 
types (sex unknown): USA (BMNH). 

TAXONOMY. — To Polystoechotes (see 
Hagen, 1861:206); H. nebulosa = P. punctatus, 
H. irroratus - P. punctatus, P. sticticus = P. 
punctatus, O. validus = P. punctatus (see Hagen, 
1861:206); H. vittatus = P. punctatus (see Car- 
penter, 1940:268269). 

Distribution. — CAN: AB 7 , BC 7 ON 7 , 
PQ 7 , USA: AK M , AZ 7 , CA 7 , CO 2 , GA 7 , IA 11 , 
ID, IN 10 , KS 4 , MA 7 , MD 7 , ME 7 , MI 1 ', MN 8 . 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



MT 7 NC 6 NE 11 , NH 7 , NJ 12 , NM 2 , NV 7 , NY 5 , 
OH 11 , OR 7 , PA 3 , SD 1 ', TX 1 , UT 7 , WA 7 , WS 9 , 
WY 7 ('Hagen, 1861:206; 2 Hagen, 1875:920; 
3 Smith, 1900:55; 4 Smith, 1925:167, 5 Leonard, 
1928:40; 6 Brimley, 1938:30; Carpenter, 
1940:269; 8 Parfin, 1952:425; 9 Throne, 
197 lb: 86; 10 Lawson & McCafferty, 1984:130; 
1 'Penny, NEW STATE RECORDS; l2 S Mar 
shall, NEW STATE RECORD) 



SlSYRIDAE 
Sisyrids, or spongilla-flies, are larval predators 
of freshwater sponges, sucking out the liquid 
contents of the cells with their long, straight, 
hollow mandibles. This is the only truly aquatic 
family of Neuroptera, although some Osmylidae 
are semi-aquatic. The larvae bear external, ab- 
dominal gills in second and third instars. When 
mature, larvae leave the water and climb nearby 
vegetation, where they spin a silken cocoon. 
Often, the cocoon is surrounded by a distinctive, 
outer, mesh-like covering. Adults usually 
emerge in late summer, and are often attracted to 
lights, sometimes at a considerable distance from 
water. Eggs are laid above water, and when they 
hatch, the young larvae fall directly into the 
water. The taxonomy of North American 
spongilla-flies was dealt with by Parfin and Gur- 
ney (1956). 

Climacia McLachlan, 1869a:21 

Climacia areolaris (Hagen), 1861:199 [Mi- 
cromus}. 

Neotype male: USA Oklahoma, Grant, 
l.VII. 1937, Standish and Kaiser (MCZ). 

r Climacia dictyona Needham, 1901:558. 
Lectotype male: USA, New York, Saranac Inn, 
28.VI. 1900 (Cornell). 

= Sisyra lampra Navas, 1914c:60. Type(s) 
(sex unknown): USA, New Jersey, Lakehurst 
(depository unknown). 

Taxonomy. — To Climacia (see McLachlan, 
1869:21); S. lampra = C. areolaris (see Navas, 
1935:33); C. dictyona = C. areolaris, S. lampra 
= S. vicaria (see Carpenter, 1940:254, 255); S. 
lampra = C. areolaris (see Parfin & Gurney, 
1956:486). Hagerfs original type was apparently 
lost in shipment from Europe, and Caipenter 
(1940:256) designated a neotype. Parfin and 



Gurney (1956:488) designated a lectotype from 
Needham's syntype series of C. dictyona. 

DISTRIBUTION. — CAN: ON 4 , PQ 4 , USA: 
AL 7 AR 7 , CO 7 , CT 4 , FL 1 , GA 4 , IL 4 , IN 7 , LA 4 , 
MA 4 , MD 4 , ME 4 , MI 4 MN 6 , MO 5 , MS 4 , NC 2 , 
NH 4 , NJ 4 , NM 4 , NY ,OH 4 , OK 4 , PA 4 , TX 4 , 
VA 4 , VT 4 , WS 7 ('Hagen, 1861:199- 2 Brimley, 
1938:29; 3 Leonard, 1928:39; Caipenter, 
1940:256; 5 Froeschner, 1947:132; 6 Parfin, 
1952:424; 7 Parfin & Gurney, 1956:489). 
Climacia californica Chandler, 1953:183. 

Holotype male: USA, California, Lake Co., 
Clear Lake, 19.V.1949, 1318 ft., H. P. Chandler 
(CAS). 

DISTRIBUTION. — USA: CA 1 , ID 2 ('chan- 
dler, 1953: 182; 2 Clark, 1985:391). 
Climaciachapini Parfin and Gurney, 1956:495. 

Holotype male: USA, Texas, Columbus, E. A. 
Schwarz (USNM). 

Distribution. — USA: NM 1 OK 2 , TX 1 
(' Parfin & Gurney, 1956:498; 2 Penny, NEW 
STATE RECORD). 

Sisyra Burmeister, 1839:975 

Sisyra apicalis Banks, 1908a:261. 
Holotype female: Cuba, Havana (MCZ}. 

DISTRIBUTION. — USA: FL',GA', LA", MS 3 
(' Parfin & Gurney, 1956:472; 2 Poirrier, 
1969:574; 3 Lago, 1981:28). 
Sisyra fuscata (Fabricius), 1793:84 [Hemero- 
bius]. 

Holotype female: Denmark (ZMC). 

= Hemerobius con/mis Stephens, 1836:115. 
Type(s) (sex unknown): England (BMNH). 

= Sisyra morio Burmeister, 1839:976. Type(s) 
(sex unknown): Germany (Halle). 

= Sisyra nigripennis Wesmael, 1841:213. 
Type(s) (sex unknown): Belgium (IRSNB). 

= Branchiotoma spongillae Westwood, 
1842:108. Type(s) (sex unknown): larva, Eng- 
land (Oxford). 

= Hemerobius fumatus Motschulsky, 1853:20. 
Type(s) (sex unknown): Russia (Moscow). 

Taxonomy. — To Sisyra (see Burmeister, 
1839:976); H. nitidulus = S. fuscata, H. confinis 
= S. fuscata, S. morio = S. fuscata, S. nigripennis 
= S. fuscata, B. spongillae = S. fuscata, H. fuma- 
tus = S. fuscata (see Parfin & Gurney, 1 956:468). 

DISTRIBUTION. — CAN: BC 1 , ON 1 , PQ , 
USA: AK 1 , IN 2 , MA 1 , ME 1 , Ml', MN f , NY 1 , 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



89 



WS 1 ('Parfin & Gurney, 1956:470; "Lawson & 

McCafferty, 1984:130). 

Sisyra vicaria (Walker), 1853:297 [Hemero- 

bius]. 

Lectotype male: USA, Georgia, John Ab- 
bot(BMNH). 

= Sisvra umbrata Needham, 1901:555. Lecto- 
type male: USA, Illinois, Lake Forest, 2.VI. 1 899 
(Cornell). 

Taxonomy, — To Sisyra (see Hagen, 
1861:197); S. umbrata = S. vicaria, S. lampra = 
S. vicaria (see Carpenter, 1940:254); S. lampra 
not = 5. vicaria (see Parfin & Gurney, 1956:486). 

DISTRIBUTION. — CAN: BC 5 , NS 5 , ON 5 , 
PQ 5 , USA: AZ 9 CT 5 , DC 5 , FL 5 , GA 1 , IL 5 , IN 9 , 
KS 2 , KY 9 , MA 5 MD 5 ME 6 MI 5 , MN 8 , MO 7 , 
MS 10 NC 4 , NY 5 OK 9 OR 5 , PA 5 , RI 9 , TN 9 , 
TX 5 , VA 9 , WA.WS, (' Walker, 1853:297; 
2 Smith, 1925:1 66; 3 Leonard, 1928:39; 4 Brimley, 
1938:29; 5 Carpenter, 1940:254; 6 Procter, 
1946:42; 7 Froeschner, 1947:132; 8 Parfin, 
1952:424; 9 Parfin & Gurney, 1956:464; 10 Poir- 
rier&Holzenthal, 1980:1). 

Raphidioptera 
Snake flies are elongate as larvae, living as 
predators under the surface of tree bark, under 
fallen logs, or in leaf litter. Adults may be asso- 
ciated with a variety of coniferous and broad- 
leafed trees and shrubs. Life histories of most of 
our species are still unknown. In America north 
of Mexico, they do not occur east of the Rocky 
Mountains, except in Texas. The taxonomy of all 
species of Raphidioptera was treated by Aspock 
(1974, 1975), and H. and U. Aspock and Rausch 
(1991). 

INOCELLIDAE 
Inocellid snake-flies can be separated from 
raphidiids by the absence of ocelli, longer thicker 
antennae, and darker thicker pterostigmal spot. 
Only a single genus, Negha Navas 1916b, is 
found in North America. 

Negha Navas, 1916b: 5 10 

Negha inflata (Hagen), 1861:196 [Raphidia]. 

Neotype male: USA, California, Yosemite Na- 
tional Park, Mariposa Grove, 12.VII.1927,F. M. 
Carpenter (MCZ). 



= Inocellia hageni Albarda, 1891:171. Holo- 
type female: USA, California, San Francisco 
(BMNH). 

TAXONOMY. — To Inocellia (see Albarda, 
1891:167); to Negha (see Navas, 1916b:510); 
returned to Inocellia (see Carpenter, 1936:138); 
to Inocellia (Negha) (see Aspock, 1975:540); 
returned to Negha (see Aspock, 1988:107); /. 
hageni = I. inflata (see Carpenter, 1936:139); /. 
longicornis = /. inflata (see Aspock, 1975:540). 

DISTRIBUTION,. — CAN: BC 2 , USA: CA 1 , 
ID 3 , MT 3 , NV 2 , OR 2 , UT 3 , WA 3 ('Hagen, 
1861:196; 2 Carpenter, 1936:140; 3 Aspock, 
1975:540). 

Negha longicornis (Albarda), 1 89 1 : 1 69 [Inocel- 
lia]. 

Holotype male: USA, California (BMNH). 

TAXONOMY. — To Negha (see Navas, 
1919a:75); returned to Inocellia (see Carpenter, 
1936:140); returned to Negha (see Aspock, 
1988:107); /. longicornis = /. inflata (see 
Aspock, 1975:540);/. longicornis not = /. inflata 
(see Aspock, 1988:107). 

Distribution. — CAN: BC 2 , USA: CA 1 , 
NV 2 , OR 2 , WA 2 ( 'Albarda, 1891:169; Carpen- 
ter, 1936:141). 
Negha meridionalis U. Aspock, 1988:107. 

Holotype male: USA, California, Riverside 
Co., 35 miles south of Palm Springs, 26.11.1977, 
S. Johnson (SDNHM). 

Distribution. — USA: CA (Aspock, 
1988:107). 

Raphidiidae 
The biology of raphidiids is similar to that of 
inocellids. For subgeneric characters, see H. and 
U. Aspock and Rausch ( 1 99 1 ). 

Agulla (Agulla) Navas, 1914d:66 

Agulla (Agulla) arnaudi (U. Aspock), 
1973:234 [Raphidia {Agulla)}. 

Holotype male: Mexico, Baja California, San 
Pedro Martir, trail La Joya to La Zanja, 
10.6.1953, P. H. Arnaud, Jr. (CAS). 

Taxonomy. — To Agulla (see Aspock, 
1986:25). 

Distribution. — USA: CA (Aspock, 
1973:234). 

Agulla (Agulla) assimilis (Albarda). 1891:144 
[Rhaphidia]. 



90 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Holotype female: Canada, British Columbia, 
Vancouver Island (BMNH). 

= Raphidia arizonica (Banks), 1911:338. 
Holotype male: USA, Arizona, Phoenix, Oslar 
(MCZ). 

= Agulla bagnalli Navas, 1914d:67. Holotype 
male: Canada, British Columbia, Vancouver Is- 
land, Victoria (Oxford). 

TAXONOMY. — To Agulla (see Navas, 
1914d:66); to Raphidia {Agulla) (see Aspock, 
1975:538); returned to Agulla (see Aspock, 
1986:25); A. bagnalli = R. assimilis (Navas, 
1918c:26); R. arizonica = R. assimilis (Aspock, 
1975:538). 

DISTRIBUTION. — CAN: AB 3 , BC 1 USA: 

AZ 2 CA 3 , CO 4 , ID 4 , MT 4 . OR 3 , UT 4 , WA 4 , 

WYTAlbarda, 1891:144; 2 Banks, 191 1:338, as 

3 4 

R. arizonica; Carpenter, 1936:128; Carpenter, 

1936:1 18, as/?, arizonica). 

Agulla (Agulla) astuta (Banks), 1911:338 

[Raphidia]. 

Three syntype males: USA. California, 
Pasadena, 31.V.1919, F. Grinnell, Jr.; San 
Gabriel Mountains, 17. VI; Mount Wilson, 
5.VI. 1904 (MCZ). 

= Agulla singularis Carpenter, 1936:132. 
Holotype male: USA, California, Banning, 
29.V.1928, E. C. Van Dyke (CAS). 

Taxonomy. — To Gloria (see Navas, 
1919a:30); to Agulla (Agulla) (see Carpenter, 
1936:131); to Raphidia {Agulla) (see Aspock, 
1975:538); returned to Agulla (see Aspock, 
1986:25); A. singularis = A. astuta (see Aspock, 
1975:538). 

Distribution. — USA: AZ 2 , CA 1 ('Banks, 
191 1:338; 2 Aspock, 1975:538). 
Agulla (Agulla) barri (U. Aspock), 1973:238 
[Raph idia (Agulla ) ] . 

Holotype male: USA, California, Inyo Co., 
Westgard Pass, 11.6.1 969, W. F. Barr, on pinyon 
pine (CAS). 

Taxonomy. — To Agulla (see Aspock, 
1986:25). 

Distribution. — USA: CA, NV (Aspock, 
1973:239). 

Agulla (Agulla) bicolor (Albarda), 1891:152 
[Rhaphidia]. 

Ten syntypes (four males, six females): USA, 
Colorado (BMNH). 

= Raphidia occulta Banks, 1905a:88 (nomen 
nudum). 



= Raphidia occulta Banks, 1905b:4. Syntypes 
(sex unknown): USA, California, Claremont, 
Baker; Arizona, Prescott, Oslar; New Mexico, 
Pecos (MCZ). 

TAXONOMY. — To Raphidilla (see Navas, 
1919a:51); to Agulla (Agulla) (see Carpenter, 
1936:126); to Raphidia (Agulla) (see Aspock, 
1975:538); returned to Agulla (see Aspock, 
1986:25);/?. occulta = A. bicolor (see Carpenter, 
1936:126). 

Distribution. — CAN: BC 3 ,^USA: AZ 2 , 
CA 2 , CO 1 , NM 2 , NV 3 , OR 3 , UT 3 , WA 3 ('Al- 
barda, 1891:152; 2 Banks, 1905b:4; 3 Carpenter, 
1936:126). 
Agulla (Agulla) bractea Caipenter, 1936:130. 

Holotype male: USA, California, San Gabriel 
Mountains, 3500', 11.VI.1910, F. Grinnell, Jr. 
(MCZ). 

= Agulla neglecta Carpenter, 1936: 132. Holo- 
type male: USA, California, East Lake, VI. 1883, 
Turner (USNM). 

Taxonomy. — To Raphidia (Agulla) (see 
Aspock, 1975:538); returned to Agulla (see 
Aspock, 1986:25); ^4. neglecta = A. bractea (see 
Aspock, 1975:538). 

Distribution. — USA: CA (Carpenter, 
1936:130). 

Agulla (Agulla) crotchi (Banks), 1924:429 
[Raphidia]. 

Holotype male: USA, California, Crotch 
(MCZ). 

TAXONOMY. — To Agulla (Agulla) (see Car- 
penter, 1936:123). 

Distribution. — USA: CA (Banks, 
1924:429). 
Agulla (Agulla) faulkneri U. Aspock, 1987:1. 

Holotype male: USA, California, Riverside 
Co., Idylwild-Pine Cove, 26.VI.1977, J. W. 
Brown (SDNHM). 

Distribution. — USA: CA (Aspock, 
1987:1). 
Agulla (Agulla) flexa Carpenter, 1936:125. 

Holotype male: USA, New Mexico, Jemez 
Mountains, 18.VI, Woodgate (MCZ). 

TAXONOMY. — To Raphidia (Agulla) (see 
Aspock, 1975:539); returned to Agulla (see 
Aspock, 1986:25). 

Distribution. — USA: AZ, NM (Carpenter, 
1936:125). 

Agulla (Agulla) herbsti (Esben-Petersen), 
1912:273 [Raphidia]. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



Holotype male: "Chile, Concepcion," 
29.XII. 1908, P. Herbst (HMB). Note: This local- 
ity is certainly in error, the species is only found 
in North America. 

= Raphidia bifurcaBanks, 1920:327. Syntypes 
male and female: Canada. British Columbia. 
Wellington, 9.V.1898; 3.V.1897 (MCZ). 

= Agnlla occidentis Carpenter, 1936:129. 
Holotype male: USA, California, Pinecrest, 
Johnson (CAS). 

Taxonomy. — To Raphidia (Agulla) (see 
Aspock, 1975:538); R. bifurca = R. herbsti, A. 
occidentis = R. herbsti (see Aspock, 1975:538). 

Distribution. — CAN: AB 3 , BC 1 , USA: 
CA 2 , ID 3 , OR 2 , MT 3 , NV 3 , WA 3 ('Banks, 
1920:327, as R. bifurca: 2 Carpenter, 1936:130, 
as A occidentis; Aspock. 1975:539). 

Agulla (Californoraphidia) Aspock et al.. 
1991:464 

Agulla (Californoraphidia) nigrinotum Wo- 

glum and McGregor, 1964:201. 

Holotype male: USA, California, Kern Co., 
Mill Potrero, 1. VII. 1963, R. S. Woglum (CAS). 

Taxonomy. — To Raphidia (Agulla) (see 
Aspock, 1975:539); to Agulla (see Aspock. 
1986:25); to Agulla (Californoraphidia) by 
Aspock et al., 1991:464. 

Distribution. — USA: CA (Woglum & 
McGregor. 1964:202). 

Agulla (Franciscoraphidia) Aspock et al., 
1991:462 

Agulla (Franciscoraphidia) directa Carpenter, 
\936:l24[Agulla (Agulla)]. 

Holotype male: USA, California, Oroville, 
17.IV. 1928, H. Kiefer(CAS). 

Taxonomy. — To Raphidia (Agulla) (see 
Aspock, 1975:539); returned to Agulla (see 
Aspock, 1986:25); to Agulla (Francisco- 
raphidia) (see Aspock et al., 1991:462). 

Distribution. — USA: CA (Carpenter, 
1936:125). 

Agulla (Glavia) Navas, 1916b:? 

Agulla (Glavia) adnixa (Hagen), 1861:195 
[Raphidia]. 



Lectotype female: USA, California; Oregon, 
Willcox (HMB). 

= Raphidia oblita Hagen, 1 86 1 : 195. Holotype 
male (destroyed): USA, California . 

Taxonomy. — To Glavia (see Navas, 
1916b:509); to Agulla (Agulla) (see Carpenter, 
1936:116); to Raphidia (Agulla) (see Aspock, 
1975:539); to Agulla (Glavia) (see Aspock et al.. 
1991 :450); R. oblita = A. adnixa (see Carpenter. 
1936:1 17); lectotype designated (see Aspock, et 
al., 1991:452). 

Distribution. — CAN: ^ AB 2 ,^ BC 2 ,^ USA 
AZ 2 , CA 1 . CO 2 ID 2 , MT 2 , NM 2 , NV 2 , OR 1 , 
UT 2 , WA 2 . WY" ('Hagen, 1861:195: Carpen- 
ter, 1936:116). 

Agulla (Glavia) modesta adryte U. Aspock, 
1982:98. 

Holotype male: USA, California, Inyo Co., 7 
miles north of Parcher's Camp, 30. VI. 1 96 1 , J. K. 
Drew (UCB). 

Distribution. — USA: CA (Aspock, 
1982:98). 

Agulla (Glavia) modesta aphynphte U. As- 
pock, 1982:102. 

Holotype male: USA, California, Los Angeles 
Co., Westwood Hills, IV-V.1949 (LACM). 

Distribution. — USA: CA (Aspock, 
1982:102). 

Agulla (Glavia) modesta aphyrte U. Aspock, 
1982:100. 

Holotype male: USA, Arizona, Coconino Co., 
Oak Creek Canyon, 26.VI.1950, L. D. Beamer 
(SEM). 

Distribution. — USA: AZ, NM (Aspock, 
1982:100). 

Agulla (Glavia) modesta banksi Carpenter, 
1936:119 [Agulla (Agulla)]. 

Holotype male: USA, California, Big Bear 
Lake, 26.VII.1932, R. H. Beamer (MCZ). 

TAXONOMY. — To Raphidia (Agulla) (see 
Aspock, 1975:539); to Agulla (Glavia) (see 
Aspock, 1988:98); A. banksi = R. modesta (see 
Aspock, 1975:539); A banksi = subspecies of A. 
modesta (see Aspock, 1988:98). 

Distribution. — USA: CA (Carpenter, 
1936:120). 

Agulla (Glavia) modesta modesta Carpenter, 
1936:1 19[Agulla (Agulla)]. 

Holotype male: USA, Utah, Beaver Canyon 
(USNM). 



92 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Taxonomy. — To Raphidia (Agulla) (see 
Aspock, 1975:539); to Agulla (Glavia) (see 
Aspock, 1982:97). 

DISTRIBUTION. — USA: CO 2 , UT 1 ('Carpen- 
ter, 1936:120; 2 Aspock, 1982:98). 
Agulla (Glavia) paramerica U. Aspock, 
1982:104. 

Holotype male: USA, California, Plumas Co., 
Johnsville, 11.VII-27.VIII. 1964, H.Pini(UCD). 

Distribution. — USA: CA (Aspock, 
1982:104). 

Agulla (Glavia) unicolor Carpenter, 
l936:l2\[Agulla (Agulla)]. 

Holotype male: USA, California, Yosemite 
National Park, Mariposa Grove, 20. VII. 1932, 
F. M. Carpenter (MCZ). 

TAXONOMY. — To Raphidia (Agulla) (see 
Aspock, 1975:539); to Agulla (Glavia) (see 
Aspock, 1982:102). 

DISTRIBUTION. — CAN: BC, USA: CA, ID, 
MT, NV, OR, UT, WA (Carpenter, 1936:121). 

Alena (Alena) Navas, 1916b:510 

Alena (Alena) distincta (Banks), 1911:338 
[Raphidia]. 

Holotype male: USA, California, Kern Co., 
Mt. Pinos, 6.VI. 1904, F. Grinnell, Jr. (MCZ). 

Taxonomy. — To Alena (see Navas, 
19 16b:5 10); to Agulla (Alena) (see Carpenter, 
1936:134); to Raphidia (Alena) (see Aspock, 
1975:539); returned to Alena (see Aspock, 
1986:25). 

Distribution. — USA: CA (Banks, 
1911:338). 

Alena (Aztekoraphidia) U. Aspock and H. 
Aspock, 1970:709 

Alena (Aztekoraphidia) minuta (Banks), 
1903b:239 [Raphidia]. 

Holotype male: USA, Arizona, Williams, 
l.VI, Barber and Schwarz (USNM). 

TAXONOMY. — To Alena (see Navas, 
19 16b:5 10); to Agulla (Alena) (see Carpenter, 
1936: 133); to Raphidia (Sombreroraphidia) (see 
Aspock, 1975:539); to Alena (Aztekoraphidia) 
(see Aspock etal., 1991:471). 

Distribution. — USA: AZ, NM (Banks, 
1903b:239). 



Megaloptera 
corydalidae 
Corydalidae, or dobson-flies and fish-flies, are 
among our largest neuropteroids. Males of the 
dobson-fly Corydalus cornutus can have a wing- 
span of five and one-half inches ( 14 cm) and bear 
elongate, sickle-shaped mandibles more than one 
and one-half inches (4 cm) long. The larvae are 
aquatic predators known as hellgrammites, 
which feed on aquatic insects, tadpoles, and 
small fish. Larvae may go through seven or eight 
instars and take two or three years or longer to 
reach maturity. Larvae of some species of Proto- 
chauliodes live in intermittent streams and form 
a cell in the soil of the stream bed during the 
months when the stream is dry. Despite their 
ferocious appearance, adults either do not feed, 
or feed on small quantities of nectar and fruit 
juices. The taxonomy of this group has been 
treated by Chandler (1956), Contreras-Ramos 
(1996), Evans (1984), Flint (1965), and Merritt 
and Cummins (1978). 

CORYDALINAE 

Corydalus Latreille, 1802:290 

Corydalus cornutus (Linnaeus), 1758:551 [He- 
merobius]. 

Type(s) (sex unknown): USA, Pennsylvania, 
DeGeer (London). 

= Corydalus cognatus Hagen, 1861:193 (as 
Corydalis [sic] cognata). Holotype female: 
USA, western Texas (now New Mexico), Pecos 
River, 4.V1I, Pope (MCZ). 

= Corydalus luteus Hagen, 1861:193 (as Cory- 
dalis [sic] luted). Syntype males: Mexico, Salle; 
Cordova, Saussure, one in Selys Collection 
(IRSNB). 

= Corydalus crassicornis McLachlan, 
1867:233 (as Coryalis [sic] crassicornis). Holo- 
type (sex unknown): USA, Texas (MNHN). 

= Corydalus inamabilis McLachlan, 1867:235 
(as Corydalis [sic] inamabilis). Type(s) (sex un- 
known): USA, Texas (BMNH). 

= Nevromus pallidas Davis, 1903:470 (as 
Neuronitis [sic] pallidas). Holotype male: (local- 
ity and depository unknown). 

= Corydalus texanus Banks, 1903:239 (as Co- 
rydalis [sic] texana). Holotype female: USA, 
Texas (depository unknown). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



93 



TAXONOMY. — To Raphidia (see Linnaeus, 
1 767:9 1 6); returned to Hemerobius (see DeGeer, 
1773:559); to Corydalus (see Latreille, 
1802:290); C. crassicornis = C. cornutus, C. 
inamabilis = C. cornutus, C. cognata = C. cor- 
nutus, C. lutea = C. cornutus, C. pallidus = C. 
cornutus, C. texana = C. cornutus (see Weele, 
1910:13); C. cognatus not = C. cornutus (see 
Evans, 1972:77). 

DISTRIBUTION. — CAN: PQ 4 , ^USA: AZ 5 , 
CA 7 , DC 4 , IA 8 , IL 2 , IN 6 , KS 4 , MD 2 , MN 4 , MO 4 , 
NC 4 , NM 5 , NV 7 ,NY 2 , OH 4 , PA 1 , SC 4 , TX 3 , 
UT 7 , VA 4 ('Linnaeus, 1758:551; 2 Hagen, 
1861:192; 3 McLachIan, 1867:233: 4 Davis, 
1903:478, as C. cornutus; 5 Davis, 1903:479, as 
C. cognata; Montgomery & Trippel. 1933:259; 
Evans, 1972:78, as C. cognata; Penny, NEW 
STATE RECORD) 



Chauliodinae 



Chauliodes Latreille, 1796:102 

Chauliodes pectinicornis (Linnaeus), 1763:412 
[Hemerobius]. 

Type(s) (sex unknown): (locality unknown) 
(Tring). 

= Hemerobius virginiensis Drury, 1773: un- 
paginated index. Type(s) (sex unknown):(local- 
ity unknown) (BMNH). 

TAXONOMY. — To Scmblis (see Fabricius, 
1781:386); to Chauliodes (see Latreille, 
1802:290); H. virginiensis = C. pectinicornis 
(see Davis, 1903:461). 

Distribution. — CAN: BC 7 , USA: DE 2 , 
FL 4 , IA 9 , IL 9 , IN 5 , KS 9 , LA 4 . MA 2 MD 3 , ME 8 , 
MI 4 MO 4 , MS 9 , NC 6 , NE 9 , NH, NJ 3 , NY 4 
OH 4 , PA 3 , SC 4 , RI 9 , TX 9 , VA 1 , VT 9 , WS 4 
('Drury, 1773:2; 2 Walker, 1853:198; 3 Hagen, 
1861:189; 4 Davis, 1903:462; Montgomery & 
Trippel. 1933:259; 6 Brimley, 1938:28; 
7 Spencer, 1942:23; 8 Procter, 1946:42; 9 Tarter, 
Watkins et al., 1976:223) 
Chauliodes rastricornis Rambur, 1842:444. 

Holotype female: no published locality data, 
Selys Collection (IRSNB). 

= Hermes indecisus Walker, 1853:204. 
Type(s) (sex unknown): no locality data 
(BMNH). 

TAXONOMY. — H. indecisus = C. rastricornis 
(see Davis, 1903:460). 



Distribution. — USA: AL 7 , FL 2 , GA 2 , IA 7 , 
IL 1 IN 2 , KS 3 ME 5 , MN 6 , MO 2 , MS 7 NC 4 , 
NE 7 NH 7 , NY 2 , PA 2 , RI 7 , SC 2 , SD 7 , TN, VA 7 , 
WS ('Hagen, 1863:181; 2 Davis, 1903:460; 
3 Smith, 1925:166; 4 Brimley 1938:28; 5 Procter, 
1946:42; 6 Parfin, 1952:422; 7 Tarter, Watkins, et 
al., 1976:224). 



Dysmicohermes Munroe, 1953:191 

Dysmicohermes disjunctus (Walker), 
1866:334 [Chauliodes]. 

Holotype female: Canada, British Columbia. 
Vancouver Island, Chalukweyuh Lake. IX. 1859 
(BMNH). 

Taxonomy. — To Neohermes (see Weele, 
1910:54); to Dysmicohermes (see Munroe, 
1953:190). 

DISTRIBUTION. — CAN: BC 1 , USA: CA 2 , 
OR 2 , WA 2 ('Walker, 1866:334; 2 Evans, 
1972:87). 
Dysmicohermes ingens Chandler, 1954:105. 

Holotype male: USA. California, Mariposa 
Co., Miami Ranger Station, 27. VII. 1946, 
5000 ft., H. P. Chandler (CAS). 

Distribution. — USA: CA (Chandler, 
1954:106). 

Neohermes Banks, 1908d:29 

Neohermes angusticollis (Hagen), 1861:191 
[Chauliodes]. 

Neotype male: USA, Georgia, Atlanta, 
1 1. VI. 1939, P. W. Fattig (USNM). 

TAXONOMY. — To Neohermes (see Banks, 
1908d:29); N. angusticollis = N. californicus(see 
Weele, 1910:53);^. angusticollis is valid species 
(see Flint, 1965:259); neotype designated (see 
Flint, 1965:260). 

Distribution. — USA: GA 1 , SC 2 ('Flint, 
1965:260; 2 Tarter et al., 1976:225). 
Neohermes californicus (Walker), 1853:199 
[Chauliodes]. 

Holotype female: USA, California (BMNH). 

TAXONOMY. — To Neohermes (see Banks, 
1908d:29). 

Distribution. — USA: CA',NV 2 (' Walker, 
1853:199; 2 Flint, 1965:258). 
Neohermes concolor (Davis), 1903:462 
[Chauliodes]. 

Lectotype female: (no locality data) (Cornell). 



94 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Taxonomy. — N concolor = N. califomicus 
(see Weele, 1910:53); N. concolor is valid spe- 
cies (see Flint, 1965:262); lectotype designated 
(see Flint, 1965:262). 

DISTRIBUTION. — USA: AR 2 ^ DC 2 . DE 2 , 
GA 3 , IL 3 IN'^KS 4 , KY 2 , MA", MD 2 , MO 2 , 
MS 3 , NC-, NJ 2 , NY 2 , OH 3 , OK 3 , PA", TN 3 , 
VA 2 , VT 3 ('Montgomery & Trippel, 1933:259; 
2 Flint, 1965:262; 3 Tarter, Watkins et al., 
1976:225; 4 Roble, 1984:69). 
Neohermes filicornis (Banks), 1903b:238 
[Chauliodes]. 

Holotype male: USA, Arizona, Jerome, 
24.VI.1902, Oslar(MCZ). 

TAXONOMY. — To Neohermes (see Banks, 
1908d:29); N. filicornis = N. califomicus (see 
Weele, 1910:53); N. filicornis is valid name (see 
Van Dyke, 1945:110). 

DISTRIBUTION. — USA: AZ 1 , CA 2 , NM 2 
('Banks, 1903b:238; 2 Flint, 1965:257). 
Neohermes matheri Flint, 1965:260. 

Holotype male: USA, Mississippi, Hinds Co., 
Clinton, 1 2. VI. 1960, Bryant Mather (USNM). 

Distribution. — USA: MS (Flint, 
1965:260). 

Nigronia Banks, 1908d:30 



(Walker), 1853:201 



USA, New York 



Nigronia fasciata 

[Chauliodes]. 

Two syntype males: 
(BMNH). 

= Chauliodes lunatus Hagen, 1863:180. Syn- 
types (sex unknown): USA, Illinois; Maryland; 
New York; Pennsylvania (depository unknown). 

TAXONOMY. — To Nigronia (see Banks, 
1908d:30); C lunatus = Cfasciatus (see Davis, 
1903:458). 

Distribution. — USA: AL 6 , AR 3 DC 3 , 
DE 6 , IL 2 , LA 6 , MD 2 , ME 5 , MN 3 , MO 3 , MS 7 , 
NC 4 , NH 6 NY 1 , OH 3 , PA 2 , SC 6 (' Walker, 
1853:201; "Hagen, 1863:181; 3 Davis, 1903:458; 
4 Brimley, 1938:28; 5 Procter, 1946:42; 6 Tarter, 
Watkins et al., 1976:224; 7 Stark & Lago, 
1983:357). 

Nigronia serricornis (Say), 1824:307 
[Chauliodes]. 

Type(s) (sex unknown): USA: Arkansas; Min- 
nesota, Lake of the Woods; Pennsylvania; Mis- 
souri; Canada: Saskatchewan, Red River of Lake 
Winnipeg (depository unknown). 



= Nevromus maculatus Rambur, 1842:442. 
Holotype female: (no locality data) (IRSNB). 

TAXONOMY. — To Nigronia (see Banks, 
1908d:30); C. maculatus = C. serricornis (see 
Hagen, 1863:181). 

Distribution. — CAN: MB 1 , USA: AL 9 , 
ar 1 , ct 8 , dc 2 , fl 11 , ga 1 , in 4 ks 3 la 8 , 
ma 1 ,md , ,me 6 ,mn 7 ,mo 1 ,ms i0 ,no\ny 1 , 

OH 2 , PA 1 , RI 8 , SC 8 , TN 8 ('Hagen, 1861:191; 
2 Davis, 1903:460; 3 Smith, 1925:166; 
Montgomery & Trippel, 1933:259; Brimley, 
1938:28; 6 Procter, 1946:42; 7 Parfin, 1952:422; 
8 Tarter, Watkins et al., 1976:224; 9 Scheiring. 
1979: 176; l0 Stark & Lago, 1983:358; M Stange, 
NEW STATE RECORD) 

Orohermes Evans, 1984:1 

Orohermes crepusculus (Chandler), 1954:107 

[Dysmicohermes] . 

Holotype male: USA, California, El Dorado 
Co., Pyramid Ranger Station, 20.VIII.1952, 
J. W. MacSwain (CAS). 

TAXONOMY. — To Orohermes (see Evans, 
1984:1). 

DISTRIBUTION. — USA: CA, OR (Chandler. 
1954:108). 

Protochauliodes Weele, 1909b:257 

Protochauliodes aridus Maddux. 1954:70. 

Holotype male: USA, California, Butte Co., 
Neal Road, 7 miles southeast of Chico, 
15.V.1951, D.Maddux (CAS). 

Distribution. — USA: CA (Maddux, 
1954:70). 
Protochauliodes cascadius Evans, 1984:1. 

Holotype male: USA, Oregon, Marion Co., ca. 
8 miles west of Mill City, along Little Santiam 
River, 26.VII.1963, S. Jewett (CAS). 

Distribution. — USA: OR (Evans, 1984:2). 
Protochauliodes minimus (Davis), 1903:463 
[Chauliodes]. 

Holotype male: USA, California, San Rafael, 
26V, O. Sacken (MCZ). 

= Neohermes infuscatus Caudell, 1933:125. 
Holotype female: USA, California, Solano Co., 
Vacaville, 1. VI. 1932, A. S. Harrison (USNM). 

= Neohermes nigrinis Van Dyke, 1945:110. 
Holotype male: USA, California, Butte Co., 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



95 



Richardson's Spring, 12.V.1944, E. C. Van 
Dyke (CAS). 

TAXONOMY. — To Protochauliodes (see 
Chandler, 1954: 1 10); C. minimus - N. californi- 
cus (see Weele. 1910:53); P. minimus not = N. 
californicus (see Chandler, 1954:1 10); N. infus- 
catus = P. minimus, N. nigrinis = N. minimus (see 
Evans, 1972:128). 

Distribution. — USA: CA (Davis, 
1903:463). 

Protochauliodes montivagus Chandler, 
1954:111. 

Holotype male: USA, California, Siena Co., 
St. Charles Hill, 7.VII.1921. E. H. Nast (CAS). 

Distribution. — USA: CA (Chandler, 
1954:111). 
Protochauliodes simplus Chandler, 1954: 1 10. 

Holotype male: USA, California. Los Angeles 
Co., San Dimas Experimental Forest, Tanbark 
Flat. 13.VII.1950. J. D. Paschke (CAS). 

Distribution. — USA: CA (Chandler, 
1954:110). 
Protochauliodes spenceri Munroe, 1953:190. 

Holotype male: Canada, British Columbia, 
Duncan, 1 .VIII. 1 9 1 8, W. Downes (CNC). 

Distribution. — CAN: BC 1 , USA: CA 2 , 
OR", WA" 
1972:134). 



(Munroe, 1953:190; "Evans, 



SlALIDAE 
Sialidae. or alder-flies, as adults are rather 
uniform insects about one inch long, with smoky 
black wings. Head markings can vary from or- 
ange to black. They live along the margins of 
rivers, streams, and lakes, never getting more 
than several yards away from water. Larvae live 
in the bottom substrate of these same water 
courses, and often survive in heavily polluted 
waters. The immatures are predators, feeding on 
other aquatic arthropods. The taxonomy of Sialis 
Latreille. 1802, was treated by Ross ( 1937). 

Sialis Latreille, 1802:290 

Sialis aequalis Banks, 1920:326. 

Lectotype male: USA, Virginia, Falls Church, 
25.IV,N. Banks (MCZ). 

TAXONOMY. — Lectotype designated (see 
Ross. 1937:77). 

Distribution. — USA: CT 4 , DE 4 GA 5 , 
KY 5 , MD 2 , ME 5 , MI 5 , MN 4 , NC 2 , NT", NY 4 , 



OH 4 , PA 2 ^ SC 4 , VA 1 , VT 5 , WV 3 ('Banks, 

1920:326; -Ross, 1937:77; 3 Tarter & Woodrum, 

1973b: 147; 4 Tarter, et al.. 1978:231; 5 Whiting, 

1991b:51). 

Sialis americana Rambur, 1842:447 [Semblis]. 

Holotype female: (no locality data), Selys Col- 
lection (IRSNB). 

= Sialis ferrugineus Walker, 1853:195. Holo- 
type (sex unknown): USA, Georgia (BMNH). 

Taxonomy. — To Sialis (see Hagen, 
1861:188); to Protosialis (see Weele, 1909b: 
263); returned to Sialis (see Ross, 1937:70); S. 
ferrugineus = S. americana (see Banks, 
1907c:22). 

Distribution. — USA: CT 9 , DC 6 , FL 9 , GA 1 , 
IN 8 LA 9 ,MD 4 ,MO M MS 10 NC 5 ,NH I1 ,NJ 11 , 
OH , PA 2 , SC 9 , TX 9 , VA 6 , WS 7 (' Walker, 



1853:195 as S ferrugineus; "Hagen, 1861:1 
ks, 1907c:22; ^Ross, 1937:71; 5 Brin 



J Banks 



mley. 



1938:29; 6 Flint, 1964: 12; 7 Tennessen, 1968:185; 
8 Lawson & McCafferty, 1984:129; 9 Tarter, et 
al., 1978:231; l0 Stark & Lago, 1983:117; 
"Whiting, 1991b:51). 
Sialis arvalis Ross, 1937:68. 

Holotype male: USA, California, Mokelumne 
Hill, IV, F. E. Blaisdell (CAS). 

Distribution. — USA: CA 1 , OR 2 ('Ross, 
1937:68; 2 Evans. 1972:23). 
Sialis bilobata Whiting, 1991a:41 1. 

Holotype male: USA, California, Los Angeles 
Co., Brents Mountain Crags, 20.V.1939 
(LACM). 

Distribution. — USA: CA (Whiting, 
1991a:412). 
Sialis californica Banks, 1920:326. 

Lectotype male: USA. California, Kern Co., 
San Emigdio Canon, 3. VI. 1904, F. Grinnell, Jr. 
(MCZ). 

Taxonomy. — Lectotype designated (see 
Ross, 1937:69). 

Distribution. — CAN: AB 2 , BC 2 , USA: 
CA 1 , OR 2 , WA 2 ('Banks, 1920:326; 2 Ross, 
1937:69). 
Sialis concava Banks, 1897a:22. 

Holotype male: USA, New York, Ithaca 
(MCZ). 

Distribution. — CAN: BC 6 , ON 2 , USA: 

4 X 



MD". ME, NC-\ NY 1 



BC 
VA 5 , WV" ('Banks, 
1897a:22; "Ross, 1937:73; 3 Brimley, 1938:29; 
4 Tarter et al., 1977:104; 5 arter et al., 1978:232; 
6 Whiting, 1991b:51). 



96 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Sialis contigua Flint, 1964:10. 

Hololype male: USA, Virginia, Highland Co., 
bridge on route 220 over East Branch Potomac 
River, 19.V.1963, O. S. Flint, Jr. & W. D. Field 
(USNM). 

Distribution. — USA: TN 2 , VA 1 ('Flint, 
1964:12; 2 Tarteretal., 1978:232). 
Sialis cornuta Ross, 1937:69. 

Holotype male: USA, Oregon, Blue Mts., 
Horseshoe Lake, 7500', 26.VII.1929, H. A. 
Scullen (INHS). 

Distribution. — CAN: AB 1 , USA: ID 1 , 
MT 2 , OR 1 , UT 1 , WA 2 , WY 2 ('Ross, 1937:70; 
2 Evans, 1972:32). 
Sialis dreisbachi Flint, 1964:9. 

Holotype male: USA, Michigan, Schoolcraft 
Co., 5.VI.1959, R. & K. Dreisbach (USNM). 

DISTRIBUTION. — USA: MI 1 , MN 2 , WS 3 
('Flint, 1964:10; 2 Tarteretal., 1978:232; 3 Whit- 
ing, 1991b:52). 
Sialis glabella Ross, 1937:71. 

Holotype male: USA, Illinois, Mt. Carmel, 
28.V. 1884 (INHS). 

DISTRIBUTION. — USA: IL 1 , Y 2 , MS 3 (' Ross, 
1937:71; Canterbury & Neff, 1980:415; 3 Stark 
&Lago, 1980:118). 
Sialis haniata Ross, 1937:70. 

Holotype male: USA, Utah, Logan, 
16.IV. 1933, J. A. Meacham (INHS). 

Distribution. — CAN: AB 1 , BC 1 , USA: 
ID 2 , MT 1 , NV 2 , OR 1 , UT 1 , WA 2 , WY 1 ("Ross. 
1937:70); 2 Evans, 1972:34). 
Sialis hasta Ross, 1937:74. 

Holotype male: USA, Michigan, Lovells, 
along Au Sable River, 22.V. 1936, Frison & Ross 
(INHS). 

Distribution. — AR 2 , IN 3 , KY 4 , MI 1 , MO 3 , 
PA 1 ('Ross, 1937:74; 2 Flint, 1964:12; 3 Tarteret 
al., 1978:232; Canterbury & Neff, 1980:413). 
Sialis infumata Newman, 1838:500. 

Holotype female: USA, New Jersey, Trenton 
Falls, Doubleday (BMNH). 

Distribution. — CAN: ON 10 , USA: AR 7 , 
IL 2 , IN 7 , KS 2 , KY 8 , ME 4 , MI 2 , MN 7 , MO 7 , NC 3 , 
NJ 1 NY 2 , OH 2 , OK 9 , PA 2 , SC 7 , VA 5 , WS 6 , 
WV 10 ('Newman, 1838:500; 2 Ross, 1937:73; 
3 Brimley, 1938:29; 4 Procter, 1946:42; 5 Flint, 
1964:13; 6 Tennessen, 1968:185; 7 Tarter et al., 
1978:232; Canterbury & Neff, 1980:4 14; 9 Stark 
&Lago, 1980:121; i0 Whiting, 1991b:52). 



Sialis iola Ross, 1937:68. 

Holotype male: USA, Pennsylvania, Pitts- 
burgh (INHS). 

Distribution. — CAN: Pq',^USA: CT 3 , 
DC 1 , GA 6 , IN 5 ME 3 , MS 4 , NC 3 , NH 1 , NJ 1 , 
NY 1 , OH 3 , PA 1 SC 3 , VA 2 ('Ross, 1937:68; 
2 Flint, 1964:13; 3 Tarter et al., 1978:232; 4 Stark 
& Lago, 1980:118; 5 Lawson & McCafferty, 
1984:129; 6 Whiting, 1991b:53). 
Sialis itasca Ross, 1937:72. 

Holotype male: USA, Illinois, Momence. 
along Kankakee River, 1. VI. 1937, B. D. Burks 
(INHS). 

Distribution. — CAN: ON 1 , PQ 1 ; USA: 
AR 5 , DC ' GA 2 , IL 1 , IN 5 , KS ' , MD 2 , MI ' , MN 2 , 
MO 1 , NC 5 , ND 1 , NY 1 , OH 1 , OK 5 , PA^TN 5 , 
TX 5 , VA 1 , WS 3 , WV 4 , ('Ross, 1937:72; "Flint, 
1964:13; 3 Tennessen, 1968: 185; 4 Tarter. Ashley 
etal., 1976:32; 5 Tarter et al., 1978:232). 
Sialis joppa Ross, 1937:67. 

Holotype male: USA, North Carolina, Great 
Smoky Mountains National Park, Newfound 
Gap, 28.V. 1934, T. H. Frison (INHS). 

Distribution. — CAN: ON 6 , USA: CT 4 , 
DE 4 , FL 6 , IL 1 , KY 5 , LA 4 , MD 2 , ME ' , MI 4 , NC ' 
NH 1 , NY 1 , OH 3 , PA 1 , VA 2 , VT 4 , WS 4 , WV 3 
('Ross, 1937:68; 2 Flint, 1964:13; 3 Tarter & 
Woodrum, 1973a: 165; 4 Tarter et al., 1978:232; 
5 Canterbury & Neff, 1980:413; 6 Stange, 
1990:2). 
Sialis mohri Ross, 1937:74. 

Holotype male: USA, Wisconsin, Boulder 
Junction, on Trout River, 20. VI. 1934, Frison & 
Mohr (INHS). 

Distribution. — CAN: NB 1 , ON 1 PQ 1 , 
USA:AR 2 ,CT I ,IL'.iN',KS 3 ,KY 1 ,MA 1 ,ME 3 , 
Ml' MN 1 , MO 3 , MS 4 , NE 5 , NH 3 , NJ 1 , NY 1 . 
OH, OK 2 ,^PA', RI 3 , TN 3 , VT 5 , WS 1 ('Ross, 
1937:74; 2 Flint, 1964:13; 3 Tarter et al., 
1978:232; 4 Stark & Lago. 1980:1 18; 5 Whiting, 
1991b:53). 
Sialis nevadensis Davis, 1903:450. 

Two syntypes, male and female: USA, Ne- 
vada, Reno (USNM). 

= Sialis morrisoni Davis, 1903:450. Holotype 
female: USA, Nevada, Reno (MCZ). 

TAXONOMY. — S. morrisoni = S. nevadensis 
(see Banks, 1907c:22). 

Distribution. — CA 2 , NV 1 ('Davis, 
1903:450; 2 Ross, 1937:76). 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



97 



Sialis nina Townsend, 1939:224. 

Holotype male: USA, Kentucky. Lexington, 
along North Elkhorn Creek, 1. IV. 1938, P.O. 
Ritcher and L. H. Townsend (USNM). 

Distribution. — KY (Townsend, 

1939:225). 
Sialis occidens Ross, 1937:69. 

Holotype male: USA, California, Sequoia Na- 
tional Park, Wolverton, 7000-9000 ft., 
25.VI.1929, E. C. Van Dyke (CAS). 
^ Distribution. — CA,NV 2 (' Ross. 1937:69; 
2 Evans, 1972:37). 
Sialis rotunda Banks. 1920:327. 

Lectotype male: Canada, British Columbia, 
Bon Accord, 20.V (MCZ). 

Taxonomy. — Lectotype designated (see 
Ross, 1937:75). 

USA: CA 3 

Ross, 

al., 



Distribution. — CAN: BC 

, WS 4 ('Banks, 1920:327; 
3 Evans, 1972:43; 4 Tarter et 



OR", WA 

1937:75; 

1978:233). 

Sialis spangleri Flint, 1964:12. 

Holotype male: USA, Maryland, Garrett Co., 
Swallow Falls State Park near Oakland, 
14.V. 1963. P. J. Spangler (USNM). 

Distribution. — MD (Flint, 1964: 12). 
Sialis vagans Ross, 1937:76. 

Holotype male: USA, Indiana, Columbia City, 
along Eel River, 19. V. 1936, Frison & Ross 
(INHS). 

Distribution. — CAN: NB 1 , NS 1 , ON 1 , 
PQ 1 ; USA: AR 4 CT 3 , FL 6 , GA 3 , IL 1 , IN 1 , KS 7 , 



KY 5 , MA 1 , MD 7 



ME 



Ml',MN 2 , MS 4 , NC 4 , 
NH 1 , NJ 1 , NY 1 , OH 4 , PA 1 , VA 3 , VT 4 WS 1 
('Ross. 1937:77; 2 Parfin, 1952:422; 3 Flint, 
1964: 13; 4 Tarter et al., 1978:233; ^Canterbury & 
Neff, 1980:415; 6 Stange, 1990:2; 7 Whiting, 
1991b:54). 
Sialis velata Ross, 1937:71. 

Holotype male: USA, Michigan. Houghton 
Lake, 15-1 8.VI. 1935, T. H. Frison (INHS). 

Distribution. — CAN: AB 1 , BC 1 , MB 1 , 
ON 1 , PQ 1 , SK 1 , USA: CO 5 CT 5 , ID 4 , IL 1 , IN 2 , 
KS 1 , MA 1 , MDj, ME 1 , MI 1 , MN 1 , MO 1 , MT 5 , 
NC 3 , ND 1 , NE 3 NH 1 , NJ 5 , NY 1 , TN 3 , TX 1 , 
UT 5 , VA 1 ^, VT 3 , WS 1 , WV 1 , WY 5 ('Ross, 
1937:71; 2 Lawson & McCafferty, 1984:129; 
3 Tarter et al., 1978:233; 4 Stark & Lago, 
1980:121; 5 Whiting, 1991b:55). 



Acknowledgments 

The authors wish to thank John D. Oswald for 
pointing out that the names Megulomus specio- 
sus and Ululodes quadrimaculata were junior 
primary homonyms. He also is to be thanked for 
very carefully reviewing an earlier version of this 
manuscript, spending an enormous amount of 
time helping the authors. The Ernst Mayr Fund 
at Harvard University is to be thanked for a grant 
to allow one of us (NDP) to study type material 
in the Museum of Comparative Zoology. 

Literature Cited 

Adams, P. A. 1 957a. A new genus and new species of 
Chrysopidae from the western United States, with 
remarks on the wing venation of the family. Psyche 
(Camb., Mass.) 63:67-74, 5 figs. 

. 1957b. New ant-lions from the southwestern 



United States (Neuroptera: Myrmeleontidae). Psy- 
che (Camb., Mass.) 63:82-108, 46 tigs. 

— . 1958. A synonym in the genus Chrysopa 



(Neuroptera: Chrysopidae). Psyche (Camb., Mass.) 
63(2):45. 



— . 1 962. A stridulatory structure in Chrysopidae 
(Neuroptera). Pan-Pacific Entomologist 

38(3): 1 78-1 80. 

— . 1963. Taxonomy of Hawaiian Chrysopa 



(Neuroptera: Chrysopidae). Proceedings of the Ha- 
waiian Entomological Society 1 8(2):22 1 —223. 

— . 1967. A review of the Mesochrysinae and 



Nothochrysinae (Neuroptera: Chrysopidae). Bulle- 
tin of the Museum of Comparative Zoology, Har- 
vard University 135(4)215-238. 

. 1969. New species and synonymy in the 



genus Meleoma (Neuroptera, Chrysopidae), with a 
discussion of genitalic homologies. Postilla 
136:1-18. 

. 1970. A review of the New World Dilaridae. 



Postilla 148:1-30, 14 figs. 
. 1 975. Status of the genera Ungla and Mallada 



Navas (Neuroptera: Chrysopidae). Psyche (Camb.. 
Mass.) 82:167-173. 



98 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



. 1977. Taxonomy of United States Leuco- 

chrysa (Neuroptcra: Chrysopidae). Psyche (Camb., 
Mass.) 84(1 ):92- 102. 

. 1982a. Ceraeochrysa, a new genus of 

Chrysopinae (Neuroptera) (Studies in New World 
Chrysopidae, Part II). Neuroptera International 
2:69-75. 



— . 1982b. Plesiochrysa, a new subgenus of 
Chrysopa (Neuroptera) (Studies in New World 
Chrysopidae, Part I). Neuroptera International 

2:27-32. 

Adams, P. A. and J. A. Garland. 1 98 1 . A new spe- 
cies of Chrysopiella Banks from western North 
America (Neuroptera: Chrysopidae). Canadian En- 
tomologist 1 13:1-4, 13 figs. 

. 1982. A review of the genus Mallada in the 

United States and Canada, with a new species 
(Neuroptera: Chrysopidae). Psyche (Camb., Mass.) 
89(3-4):239-248, 20 figs. 



Neuropteroidea). 2 vols., 730 and 550 pp. Goecke 
and Evers, Krefeld, Germany. 

Aspock, U. 1 973. Zwei neue Raphidiiden-Spe/ies aus 
Nordamerika (Neuroptera, Raphidioptera). Polskie 
Pismo Entomologic/ne 43:233-240, 4 figs. 

— . 1974. Die Raphidiopteren der Nearktis (In- 



secta, Neuropteroidea). Dissertation from The Uni- 
versity of Wien. 2 vols., 238 pp., 777 figs, 20 maps. 



. 1975. The present state of knowledge on the 

Raphidioptera of America (Insecta, Neurop- 
teroidea). Polskie Pismo Entomologiczne 
45:537-546. 

. 1982. Polymorphisms und polytypic bei 



nearktischen Raphidiiden: Agiilla (Gloria) modesta 
Carpenter, Agulla (Glavio) unicolor Carpenter und 
Agulla (Glavia) paramerica n.sp. (Neuropteroidea: 
Raphidioptera). Zeitschrift der Arbeitsgemein- 
schaft osterreichischer Entomologen 

33(3/4):95-l 12. 



Adams, P. A. and N. D. Penny. [1987] 1985. 
Neuroptera of the Amazon Basin. Part I la. Intro- 
duction and Chrysopini. Acta Amazonica 
15:413-479. 

Agnew, C. W., W. L. Sterling and D. A. Dean. 
198 1 . Notes on the Chrysopidae and Hemerobiidae 
of eastern Texas with keys for their identification. 
The Southwestern Entomologist. Supplement 4. 
20 pp. 

Alayo, P. 1968. Los Neuropteros de Cuba. Poeyana 
Scr. B(2):l-127. 

Albarda, H. 1891. Revision des Rhaphidides. 
Tijdschrift voor Entomologie 34:65-1 84. 

Ashmead, W. H. 1880. Orange insects; a treatise on 
the injurious and beneficial insects found on the 
orange trees of Florida. Ashmead Bros., Jackson- 
ville, FL. 

— . 1895. Notes on the cotton insects found in 



Mississippi. Insect Life 7:25-29. 

Aspock, H., U. Aspock and H. Rausch, 1991. Die 
Raphidiopteren der Erde. Eine monographische 
Darstellung der Systematik, Taxonomie, Biologic, 
Okologie und Chorologie der rezenten Raphidiop- 
teren der Erde, mit einer zusammenfassenden Uber- 
sicht der fossilen Raphidiopteren (Insecta: 



. 1987. Agulla faulknei n.sp. - eine neue 

Raphidiiden-Spezies aus Kalifornien (Neurop- 
teroidea: Raphidioptera: Raphidiidae). Zeitschrift 
der Arbeitsgeminschaft osterreichischer Entomolo- 
gen 39(1/2): 1-4. 

— . 1988. Negha meridional is n.sp. - eine neue 



Inocelliden-Spezies aus Siid-Kalifornien (U. S. A.) 
und Baja California (Mexiko) (Neuropteroidea: 
Raphidioptera: Inocelliidae). Zeitschrift der Ar- 
beitsgemeinschaft osterreichischer Entomologen 
39(3/4): 107-1 12. 

Aspock, U. and H. Aspock. 1970. Untersuchungen 
iiber die Raphidiopteren Mexikos (Insecta, Neurop- 
teroidea). Polskie Pismo Entomologiczne 

40:705-725. 

Bagnall, R. S. 1915. Conwentzia cryptoneuris sp.n., 
a neuropteron (Coniopterygidae) new to the British 
Fauna. Entomologists Monthly Magazine 
51:192-193. 

Banks, N. 1892. A synopsis, catalogue, and bibliog- 
raphy of the neuropteroid insects of temperate North 



spteroid insects of temperate North 
nuiciita. i impactions of the Anr 
logical Society 19:327-373, 6 figs. 



1 tip I IV VI U IV- lll_UIV'pi\-lVMU II IJV.Vl.1 w 

America. Transactions of the American Entomo- 



— . 1895a. Some Mexican Neuroptera. Proceed- 
ings of the California Academy of Sciences 
(2)5:515-522. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



99 



— . 1895b. New neuropteroid insects. Transac- 
tions of the American Entomological Society 
22:313-316. 

— . 1 896. A new species of Meleoma. Entomo- 



logical News 7:95-96. 

— . 1897a. New North American neuropteroid 



insects. Transactions of the American Entomologi- 
cal Society 24:21-31 . 

— . 1 897b. Three new species of Chrysopidae. 



Proceedings of the Entomological Society of Wash- 
ington 4:173-175. 

— . 1 897c. A Leucochrysa from Florida. Entomo- 



logical News 8:183-184. 



. 1898 [1899]. Descriptions of new North 

American neuropteroid Insects. Transactions of the 
American Entomological Society 25:199-218. 

— . 1899a. New Mynneleonidae. Entomological 
News 10:170-172. 



— . 1899b. A Classification of the North Ameri- 
can Myrmeleonidae. Canadian Entomologist 

31:67-71. 



— . 1900. A new species of Myrmeleon from 
Texas. Entomological News 1 1 :596. 

. 1901a. A List of neuropteroid insects from 



Mexico. Transactions of the American Entomologi- 
cal Society 27:361-371, 1 pi. 

— . 1901b. Some insects of the Hudsonian zone 



in New Mexico. VI. Neuropteroid insects. Psyche 
(Camb., Mass.) 9:286-287. 



— . 1902. A new species of Brachynemurus. En- 
tomological News 13:86. 

. 1903a. A revision of the Nearctic Chrysopi- 



dae. Transactions of the American Entomological 
Society 29:137-162, 1 pi. 



— . 1903b. Neuropteroid insects from Arizona. 
Proceedings of the Entomological Society of Wash- 
ington 5:237-245, 1 pi. 

— . 1 903c. Some new neuropteroid Insects. Jour- 



nal of the New York Entomological Society 
11:236-243. 



. 1903d. Notes on Brachynemurini of the B. 

ferox group. Proceedings of the Entomological So- 
ciety of Washington 5:173-1 77, 1 pi. 

. 1904a. New species of Hemerobius. Cana- 



dian Entomologist 36:61-63. 
. 1904b. Neuropteroid insects from New Mex- 



ico. Transactions of the American Entomological 
Society 30:97- 110, 1 pi. 

. 1904c. A list of neuropteroid insects, exclu- 



sive of Odonata, from the vicinity of Washington, 
D. C. Proceedings of the Entomological Society of 
Washington 6:201-217. 

. 1905a. Notes on neuropteroid insects from 



the Pacific Coast of North America, with descrip- 
tions of new species. Invertebrata Pacifica 1 :85-92. 

. 1905b. Descriptions of new Nearctic neurop- 



teroid insects. Transactions of the American Ento- 
mological Society 32:1-20, 2 pis. 

— . 1905c. A revision of the Nearctic Hemerobi- 
idae. Transactions of the American Entomological 
Society 32:21-51. 3 pis. 

. 1906a. Three new species of Neuroptera. 



Psyche (Camb., Mass.) 13:98-100, 1 fig. 
. 1906b. A Revision of the Nearctic Coniop- 



terygidae. Proceedings of the Entomological Soci- 
ety of Washington 8:77-86, 2 pis. 

1 906c. A revision of the Nearctic Conioptery- 



gidae. Proceedings of the Entomological Society of 
Washington 8:77-86, 2 pis. 

. 1906d. A revision of the Nearctic Hemerobi- 



idae. Transactions of the American Entomological 
Society 32:21-51, 3 pis. 

— . 1907a. A new genus and a new species of 



Neuroptera. Entomological News 18:275. 
. 1907b. A new ant lion-fly from Arizona. 



Entomological News 18:100-101. 
. 1907c. Catalogue of the neuropteroid insects 



(except Odonata) of the United States. American 
Entomological Society, Philadelphia. 53 pp 

. 1908a. Neuropteroid insects — notes and de- 



scriptions. Transactions of the American Entomo- 
logical Society 34:255-267, 3 pis. 



100 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



. 1908b. A new genus and species of Neurop- 

tera. Entomological News 19:203-204. 

. 1908c. New tropical American Neuroptera. 



Proceedings of the Entomological Society Wash- 
ington 9:30-34. 

— . 1908d. On the classification of the Corydali- 



nae, with description of a new species. Proceedings 
of the Entomological Society of Washington 
10:27-30. 



. 1924. Descriptions of new neuropteroid in- 
sects. Bulletin of the Museum of Comparative Zo- 
ology, Harvard University 65:421-455, 4 pis. 

. 1 927. Revision of the Nearctic Myrmeleonti- 



dae. Bulletin of the Museum of Comparative Zool- 
ogy, Harvard University 68( 1 ): 1 -84, 4 pis. 

. 1930a. New neuropteroid insects from the 



United States. Psyche (Camb., Mass.) 37:223-233, 
lpl. 



. 1910. New South American neuropteroid 

insects. Proceedings of the Entomological Society 
of Washington 12:146-160. 

. 191 1. Descriptions of new species of North 



American neuropteroid insects. Transactions of the 
American Entomological Society, 37:335-360, 
3 pis. 

. 1912. Notes on Nearctic Mantispidae. Pro- 



ceedings of the Entomological Society of Washing- 
ton 14:178-179. 

— . 1 91 3a. New exotic neuropteroid insects. Pro- 



ceedings of the Entomological Society of Washing- 
ton 1 5 -"l 3 7^ 143. 

— . 1913b. The genus Brachynemurns (Neurop- 



tera). Entomological News 24:63-65. 

— . 1914a. New American Chrysopidae. Cana- 



dian Entomologist 46:24-27. 
. 1914b. New neuropteroid insects, native and 



exotic. Proceedings of the Academy of Natural Sci- 
ences of Philadelphia 66:608-618, 1 pi. 

— . 1915a. Two new names in the Ascalaphidae 



(Neur.). Entomological News 26:350. 

— . 1915b. New neuropteroid insects, native and 



exotic. Proceedings of the Academy of Natural Sci- 
ences of Philadelphia 66:619-632, 1 pi. 

. 1920. New neuropteroid insects. Bulletin of 



the Museum of Comparative Zoology, Harvard 
University 64:299-362, 7 pis. 

— . 1922. South American Glenunis and some 



other Myrmeleonidae. Canadian Entomologist 
54:58-60. 



. 1930b. Some new neotropical neuropteroid 

insects. Psyche (Camb., Mass.) 37: 183-191, 6 figs. 

. 1 93 1 . A new species of Chiysopa. Psyche 

(Camb., Mass.) 38: 174. 



. 1935. A few new North American Neurop- 
tera. Psyche (Camb., Mass.) 42:53-57. 

. 1938a. New native neuropteroid insects. Psy- 



che (Camb., Mass.) 45:72-79, 1 pi. 
. 1 938b. New Chrysopidae and species new to 



the United States. Canadian Entomologist 
70:118-122. 

. 1938c. New West Indian neuropteroid in- 



sects. Revista de Entomologia, Rio de Janeiro 
9:285-304, 29 figs. 

. 1938d. Notes on native Myrmeleonidae. An- 



nals of the Entomological Society of America 
31:413^*20, 1 pi. 

. 1939. On some new and previously-known 



Neuroptera in the collection of the Academy of 
Natural Sciences of Philadelphia. Notulae Naturae 
(Philadelphia), No. 32. 5 pp., 2 figs. 

. 1941a. New neuropteroid insects from the 



Antilles. Memorias de la Sociedad Cubana de His- 
toria Natural — Felipe Poey 15:385-402, 3 pis. 

— . 1 94 1 b. Three new species of Myrmeleontidae 



(Neuroptera). Psyche (Camb., Mass.) 48:101-104. 
— . 1942. Contributions toward a knowledge of 



the insect fauna of Lower California. No. 4. Myr- 
meleontidae. Proceedings of the California Acad- 
emy of Sciences (4) 24: 1 33-1 52, 1 7 figs. 

— . 1 943. New Neuroptera and Trichoptera from 



the United States. Psyche (Camb., Mass.) 50:74-8 1 , 
1 tig. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



101 



. 1944. Neuroptera of northern South America. 

Part III. Boletin de Entomologia Venezolana 
3:1-34. 

— . 1946. A review of the Chrysopidae (Notho- 



chrysidae) of Central America. Psyche (Camb., 
Mass.) 52(3-4): 139-1 74. 



— . 1948. Chrysopidae (Nothochrysidae) col- 
lected in Mexico by Dr. A. Dampf (Neuroptera). 
Psyche (Camb., Mass.) 55(4): 1 5 1-1 77, 3 pis. 

— . 1950. Notes and descriptions of western 



Chrysopidae (Neuroptera). Psyche (Camb., Mass.) 
57:45-67, 2 pis. 

Beatty, G. H. and A. F. J. M. Runner. 1970. The 
occurrence of Ascalaphidae in Pennsylvania 
(Neuroptera). Proceedings of the Pennsylvania 
Academy of Sciences 44:153-155. 

Bei.kin, J. N. 1954. Additional records of Oliarces 
clara in California and Arizona (Neuroptera, Ithoni- 
dae). Bulletin of the Southern California Academy 
of Sciences 53(2):65-68. 

Bickley, W. E. 1941. Records of Tennessee 
Chrysopidae (Neuroptera). Proceedings of the En- 
tomological Society of Washington 43(8): 1 87-1 89. 

Bickley, W. E. and E. G. Macleod. 1 956. A Synop- 
sis of the Nearctic Chrysopidae with a key to the 
genera (Neuroptera). Proceedings of the Entomo- 
logical Society of Washington 58(4): 177-202. 

Bram, R. A. and W. E. Bickley. 1963. The green 
lacewings of the genus Chrysopa in Maryland. 
Maryland agricultural Experiment Station Bulletin 
A-124:l-18. 

Brauer, F. 1864. Entomologische Beitrage. B. 
Beitrage zur Kenntniss der Neuroptera. 1. Gattung 
Isoscelipteron Costa. Verhandlungen der kaiser- 
lich-koniglichen zoologisch-botanischen Gesell- 
schaftin Wien 14:898-899. 



. 1866. Reise der Osterreichischen Fregatte 

Novara um die Erde, in ... 1857-59, unter den 
Befehl des Commodore B. von Wiillerstorf-Urbair, 
H.K. von Scherzer, ed. Zoologischer Theil, Bd. 2, 
No. 4 (Neuroptera). Wien. 104 pp. 

Brimley, C. S. 1938. The insects of North Carolina. 
North Carolina Department of Agriculture, Raleigh, 
N.C. 



Brooks, S. J. and P. C. Barnard. 1990. The green 
lacewings of the world: a generic review (Neurop- 
tera: Chrysopidae). Bulletin of the British Museum 
of Natural History (Entomology) 59:1 17-286. 

Brushwein, J. R. 1987a. Bionomics of Lomamyia 
hamata (Neuroptera: Berothidae). Annals of the 
Entomological Society of America 80(5):671-679. 

. 1987b. Observations on Lomamyia longicol- 

lis (Neuroptera: Berothidae) in South Carolina. 
Journal of the Kansas Entomological Society 
60(1): 150- 152. 

Burmeister, H. C. C. 1839.HandbuchderEntomolo- 
gie. Bd. 2, Abt. 2 (pp. 397-1050). T. C. F. Enslin, 
Berlin. 

Canterbury, L. E. and S. E. Neff. 1980. Eggs of 
Sialis (Sialidae: Megaloptera) in eastern North 
America. Canadian Entomologist 1 12:409^119. 

Carpenter, F. M. 1936. A revision of the Nearctic 
Raphidiodea (Recent and Fossil). Proceedings of 
the American Academy of Arts and Sciences 
71:89-157, 2 pis. 

. 1940. A revision of the nearctic Hemerobii- 



dae, Berothidae, Sisyridae, Polystoechotidae and 
Dilaridae (Neuroptera). Proceedings of the Ameri- 
can Academy of Arts and Sciences 74(7): 193-280. 

. 1992. Treatise on invertebrate paleontology. 



Part R. Arthropoda 4. Vols. 3 and 4: Superclass 
Hexapoda. The Geological Society of America and 
the University of Kansas, xxxi + 1-277 (vol. 3), ii 
+ 279-655 (vol. 4). 265 figs., 2 tables. 

CAUDELL, A. N. 1933. Neohermes infuscatus, a new 
sialid from California. Pan-Pacific Entomologist 
9(3): 125-1 26. 

Chandler, H. P. 1953. A new species of Climacia 
from California (Sisyridae, Neuroptera). Journal of 
the Washington Academy of Sciences 43: 1 82-1 84, 
lfig. 



. 1954. Four new species of dobsonflies from 

California. Pan-Pacific Entomologist 

30(2): 105-1 11, 10 figs. 



. 1956. Megaloptera. Pp. 229-233 in Aquatic 

Insects of California R. L. Usinger, ed., University 
of California Press, Berkeley. 



102 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Clark, W. H. 1985. First record of Climacia califor- 
nica (Neuroptera: Sisyridae) and its host sponge, 
Ephydatia mulleri (Porifera: Spongillidae), from 
Idaho with water quality relationships. Great Basin 
Naturalist 45(3):39 1-394. 

Contrf.ras-Ramos, A. 1 996. A Revision of the genus 
Corydalus. Ph.D. Thesis from the University of 
Minnesota. 

COQUILLETT, D. W. 1 890. "Lace-wiriged fly" and 
"brown He me rob his.'"' Annual Report of the Cali- 
fornia State Board of Horticulture 1 890:288-289, 6 
figs., 1 pi. 

CURRIE, R. P. 1898. New species of North American 
Myrmeleonidae. I— IV. Canadian Entomologist 
30:93-96, 134-140, 241-243, 273-276. 



. [ 1 900] 1 899. New species of North American 

Myrmeleonidae. Canadian Entomologist 
31:361-365. 

— . [1900] 1899. Neuroptera. In Insects of New 



Jersey. A list of species occurring in New Jersey, 
with notes on those of economic importance, J. B. 
Smith, ed. Trenton, New Jersey. 755 pp. [Insects of 
New Jersey was published as a supplement to the 
27th Annual Report of the New Jersey State Board 
of Agriculture.] 

— . 1903. Myrmeleonidae from Arizona. Pro- 



ceedings of the Entomological Society of Washing- 
ton 5:272-284. 

— . 1904a. Notes on some Hemerobiidae from 



Arizona and California. Proceedings of the Entomo- 
logical Society of Washington 6:79-81 . 

— . 1904b. Hemerobiidae from the Kootenay 
District of British Columbia. Proceedings of the 
Entomological Society of Washington 6:81-90. 

— . 1906. A new Myrmeleonid from the United 



States. Proceedings of the Entomological Society of 
Washington 7: 186-1 88. 

Curtis, J. 1 834. British entomology, being illustra- 
tions and descriptions of the genera of insects found 
in Great Britain and Ireland: containing coloured 
figures from Nature of the most rare and beautiful 
species, and in many instances of the plants upon 
which they are found. London. [Issued between 
1824 and 1839 in 16 volumes containing 769 color 
plates with unpaginated descriptive letterpress.] 
Coniopteryx psociformis is in vol. 1 1: pi. 528. 



Davis, K. C. 1 903. Sialidae of North and South Amer- 
ica. Bulletin 68. New York State Museum of Ento- 
mology 18:442-486. 

DeGeer, C. 1773. Memoires pour servir a l'histoire 
des Insectes. Vol. 3. Stockholm. 696 pp. 

DRURY, D. 1773. Illustrations of natural history 
wherein are exhibited upwards of two hundred and 
forty figures of exotic insects, ... interspersed with 
remarks and reflections on the nature and properties 
of many of them. Vol. 2. London. 90 pp. 

Eisner, T., K. Hicks, M. Eisner, and D. S. Robson. 
1 978. "Wolf-in-sheep 1 s clothing" strategy of a pre- 
daceous insect larva. Science, New York 
199:790-794, 4 figs. 

Enderlein, G. 1905a. Conwentzia pineticola nov. 
gen., nov. spec, eine neue Neuroptera aus West- 
preussen. Bericht des Westpreussischen Botanis- 
chen-Zoologischen Vereins 26-27 

(Anlagen):10-12. 



. 1905b. Klassifikation der Neuropteren-Fami- 
Coniopterygidae. Zoologischer Anzeiger 



lie 

29:225-227 



. 1905c. Ein neuer zu den Coniopterygiden 

gehoriger Neuropteren-Typus aus der Umgebung 
von Berlin. Wiener entomologische Zeitung 
24:197-198. 



— . 1906. Monographic der Coniopterygiden. 
Zoologische Jahrbucher (Abt. fur Systematik, 
Geographie und Biologie der Tiere) 23:1 73-242. 

. 1907. Nachtrage zur Monographic der 



Coniopterygiden. Stettiner entomologische Zeitung 
68:10-13. 

. 1908. Neuroptera: Fam. Coniopterygidae. In 



Wytsman, P., Genera Insectorum, fasc. 67. 18 pp. 
Bruxelles. 2 pis., 25 figs. 

. 1910. Klassifikation der Mantispiden nach 



dem Material des Stettiner Zoologischcn Museums. 
Stettiner Entomologische Zeitung 71:341-379. 

. 1930. Die Klassifikation der Conioptery- 



giden auf Grund der recenten und fossilen Gattun- 
gen. Archiv fur klassifikatorische und 
phylogenetische Entomologie 1:98-1 14. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



103 



Esben-Petersen, P. 1912. A few new Neuroptera 
Planipennia. Entomologische Mitteilungen 

1:267-273, 7 tigs. 

— . 1 928. Neue und wenig bekannte Neuropteren 



des Hamburger Museums. Deutsche entomologis- 
che Zeitschrift 1928:73-77. 

— . 1933. New and little-known Neuroptera. Vi- 



denskalbelige Meddelelser fra Dansk Naturhis- 
torisk Forening, Kjobenhaven 94:109-123, 13 tigs. 

. 1 934. The Norwegian zoological expedition 



to the Galapagos Islands, 1925, conducted by Alf 
Wollebaek. XII. Neuropterous insects from Gala- 
pagos Is. Nytt Magasin for Naturvidenskapenne 
74:291-294,3 figs. 

Evans, E. D. 1 972. A study of the Megaloptera of the 
Pacific coastal region of the United States. Unpub- 
lished Ph.D. thesis from Oregon State University, 
June 1 972. 2 1 pp. 

— . 1984. A new genus and a new species of 
dobsonfly from the far western United States 
(Megaloptera: Corydalidae). Pan-Pacific Ento- 
mologist 60: 1-3,3 figs. 

Fabricius. J. C. 1781. Species insectorvm exhibentes 
eorvm differentias specificas, synonymaavctorvm, 
loca natalia, metamorphosin adiectis observa- 
tionibvs, descriptionibvs. Hamburgi et Kilonii. 
Tome 1 . 552 pp. 

— . 1793. Entomologia systematica emendata et 
aucta secundum classes, ordines, genera, species 
adjectis synonimis, locis observationibus, descrip- 
tionibus. Hafniae. Vol. 2. 519 pp. 

Faulkner, D. K. 1990a. Phantom of the Desert. Biol- 
ogy of the little-known moth lacewing. Environ- 
ment West 1(2): 17- 19. 

. 1990b. Current knowledge of the biology of 



the moth-lacewing Oliarces clara Banks (Insecta: 
Neuroptera: Ithonidae). Advances in Neuropterol- 
ogy. Proceedings of the Third International Sympo- 
sium of Neuropterology, pp. 197-203. 

Fitch, A. [ 1 855] 1 854. Report [upon the noxious and 
other insects of the state of New York]. Transactions 
of the New York State Agricultural Society 
14:705-880. 

Fleschner, C. A. and D. W. Ricker. 1953. Food 
habits of coniopterygids on citrus in southern Cali- 



fornia. Journal of Economic Entomology 
46(3):458-461. 

Flint, O. S., Jr. 1964. New species and new state 
records of Sialis (Neuroptera: Sialidae). Entomo- 
logical News 75:9-13, 1 1 figs. 

. 1 965. The genus Neohermes. Psyche (Camb., 



Mass.) 72:255-263, 19 tigs. 



. 1974. A coniopterygid (Aleiiropteryx 

juniperi), — Virginia. Cooperative Economic Insect 



Report 24:703. 

Froeschner, R. C. 1947. Notes and keys to the 
Neuroptera of Missouri. Annals of the Entomologi- 
cal Society of America 40: 1 23-136. 

Garland, J. A. 1985. Identification of Chrysopidae 
in Canada, with bionomic notes (Neuroptera). Ca- 
nadian Entomologist 1 17:737-762. 

Garland, J. A. and B. D. Marshall. 1980. Confir- 
mation of Ululodes quadriinacidota in Canada 
(Neuroptera: Ascalaphidae). Canadian Entomolo- 
gist 112:637-638. 

Gerstaecker, A. 1894. Uber neue und weniger 
gekannte Neuropteren aus der Familie Megaloptera, 
Burm. Mitteilungen des Naturwissenschaftlichen 
Vereins fur Neu-Vorpommern und Rugen im Greif- 
swald 25:93-173. 

Goszy, G. Von. 1 852. Beobachtungen der Verwan- 
dlungsgeschiche und Beschreibung einiger neuen 
Allen der Gattung Hemerobius Leach. Sitzung- 
berichte der Kaiserlichen Akademie der Wissen- 
schaften. Mathematisch-Naturwissenschaftliche 
Class., Wien 8:344-348. 

Guilding, L. 1825. The natural history of Phasma 
cornutwn, and the description of a new species of 
Ascalaphus. Transactions of the Linnean Society of 
London 14:137-141. 

Gurney, A. B. 1948. Notes on Nearctic Hemerobii- 
dae, with descriptions of two new species (Neurop- 
tera). Annals of the Entomological Society of 
America 41:21 3-222, 23 figs. 

Hagen, H. A. 1860. Beitrag zur Kenntniss der Myr- 
meleon-arten. Stettiner Entomologische Zeitung 
21:359-369. 



104 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



. 1861. Synopsis of the Neuroptera of North 

America. With a list of the South American species. 
Smithsonian miscellaneous Collections 4:1-347. 



1863. Ohservations on certain N.A. Neurop- 



tera. Proceedings of the Entomological Society of 
Philadelphia 2: 167-182. 

. 1866. Hemerobidarum synopsis synonymica. 

Stettiner Entomologische Zeitung 27:369^462. 

. 1 873. Odonata from the Yellowstone. Annual 



Report. F. V. Hayden, ed. United States Geological 
Survey of the Territories 6:727-729. 

— . 1 875. Report upon the collections of Neurop- 



tera and Pseudo-Neuroptera made in portions ot 
Colorado, New Mexico and Arizona, during the 
years 1 872, 1 873 and 1 874. Pp. 909-922 in Report 
upon Geographical and Geological Explorations 
and Surveys West of the 100th Meridian. Vol. 5 
(Zoology). Chapter XIV. 



. 1877. Symphrasis, eine neue Mantispiden- 

Gattung. Stettiner entomologische Zeitung 
38:208-211. 



— . 1886. Monograph of the Hemerobiidae. Part 
II. Proceedings of the Boston Society of Natural 
History 23:276-292. 

— . 1 887. Stray notes on Myrmeleonidae. Cana- 



dian Entomologist 19:89-93, 110-112, 133-136, 
147-156,209-216. 

. 1888. Stray notes on Myrmeleonidae. Cana- 



dian Entomologist 20:34-38, 57-60, 72-74, 93-97, 
185-191,204-211. 

Hale Carpenter, G. O. 1938. Notes on insects col- 
lected in West Greenland by the Oxford University 
Greenland Expedition, 1936. With descriptions of a 
new species of Angitia (Hymenoptera, Ichneumoni- 
dae) by A. Roman and of Fannia (Diptera, An- 
thomyiidae) by J. E. Collin. Annals and Magazine 
of Natural History (1 1)1:529-553. 

HANDSCHIN, E. 1960. Zur Revision sud-amerikanis- 
cher Mantispiden. Revue Suisse de Zoologie 
67:523-560. 

Henry, C. S. 1977. The behavior and life histories of 
two North American ascalaphids. Annals of the 
Entomological Society of America, 70: 1 79- 195, 10 
tigs. 



. 1983. Acoustic recognition of sibling species 

within the holarctic lacewing Chrysoperla cornea 
(Neuroptera: Chrysopidae). Systematic Entomol- 
ogy 8:293-301. 



. 1985. Sibling species, call differences, and 

speciation in green lacewings (Neuroptera: 
Chrysopidae: Chrysoperla). Evolution 

39(5):965-984. 

. 1993. Chrysoperla johnsoni (Neuroptera: 



Chrysopidae): Acoustic evidence for full species 
status. Annals of the Entomological Society of 
America 86(1): 14-25. 

Henry, C. S.,M. M.Wells, and R. J. Pupedis. 1993. 
Hidden taxonomic diversity within Chrysoperla 
plorabunda (Neuroptera: Chrysopidae): Two new 
species based on courtship songs. Annals of the 
Entomological Society of America 86(1 ):l-13. 

Henry, T. J. 1974. Beneficial insects. A conioptery- 
gid (Aleuropteryx juniperi Ohm). Cooperative Eco- 
nomic Insect Report (Plant Protection and 
Quarantine Programs) 24:659. 

Hoffman, K. M. 1989. Taxonomic status of Manlispa 
sayi, Manlispa fuscicornis and Mantispa uhleri 
(Neuroptera: Mantispidae). Proceedings of the En- 
tomological Society of Washington 91:637-639. 

. 1 990. Distributional notes on North and Cen- 
tral American Dilaridae (Neuroptera). Entomologi- 
cal News 101(3): 155-157. 

Hoffman, K. M.andJ. R. Brushwein. 1989. Species 
of spiders (Araneae) associated with the immature 
stages of Mantispa pulchella (Neuroptera, Man- 
tispidae). Journal of Arachnology 17:7-14. 

. 1 992. Descriptions of the larvae and pupae of 



some North American Mantispinae (Neuroptera: 
Mantispidae) and development of a system of larval 
chaetotaxy for Neuroptera. Transactions of the 
American Entomological Society 1 1 8(2): 159-1 96. 

[LLIGER, J. K. W. 1798. [Genus Mantispa]. P. 499 in 
Verzeichnis der Kafer Preussens, entworfen von 
Johann Gottlieb Kugelann ... ausgearbeitet von Jo- 
hann Karl Wilhelm Illiger. mit einer Vorrede des 
Professors und Pagenhofmeisters Helwig in 
Braunschweig, und dem angehangten Versuche 
einer natiirlichen Ordnungs- und Gattungs-Folge 
der Insekten. Halle. 510 pp. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



105 



Johnson, J. B. and K. S. Hagen. 1 98 1 . A neuropter- 
ous larva uses an allomone to attack termites. Nature 
289:506-507. 

Johnson, V. 1976. A new genus and species of 
Coniopterygidae (Neuroptera) from New Mexico. 
Psyche (Camb., Mass.), 83(2): 1 92- 1 95, 9 figs. 

— . 1981a. Review of the Coniopterygidae 



(Neuroptera) of North America with a revision of 
the genus Aleuropteryx. Psyche (Camb., Mass.) 
87(3-4):259-298, 15 tigs. 

. 1981b. New species of Coniopterygidae 



(Neuroptera) from North America. Psyche (Camb., 
Mass.) 87(3-4): 18 1-1 92, 7 tigs. 

Kaston, B. J. 1938. Mantispidae parasitic on spider 
egg sacs. Journal of the New York Entomological 
Society 46:147-153. 

Kevan, D. K. McE. 1989. Ms. Taylor's Walking- 
sticks: "Queen Bess" and odd "orthopterians" of 
1862 Mantodea, Phasmatodea, Dermaptera, aredu- 
viid Hemipteran and mantispid Neuroptera). Pro- 
ceedings of the Entomological Society of Ontario 
120:1-9. 

Kevan, D. K. McE. and J. Klimaszewski. 1986. 
Notes on the Hemerobiidae or brown lacewing flies 
(Neuroptera) of Canada and Alaska. Neuroptera 
International 4( 1 ):7— 22. 

— . 1987. The Hemerobiidae of Canada and 
Alaska. Genus Hemerobius L. Giornale Italiano di 
Entomologia 16:305-369. 94 figs., 1 1 maps. 

KlLLlNGTON, F. J. 1937. A monograph of the British 
Neuroptera. Vol. 2. Ray Society, London. 306 pp. 

Kimmins, D. E. 1928. New and little known Neurop- 
tera of Central America. EOS: Revista espanola de 
Entomologia 4:363-370, 7 figs. 



. 1932. Hemerobius simulons Walk., H. oroty- 

pus Wallingr., and H. crispus Steph. Walk. 
(Neuroptera). Entomologist 65:87-88. 

. 1 935. Notes on the genera Megolomus Ram- 



bur and Nesobiella, gen. nov. (Neuroptera), with 
descriptions of new species. Annals and Magazine 
of Natural History (10)16:602-619, 11 figs. 

940. Notes on some types of Chrysopidae 



(Neuroptera) in the British Museum collections 



Annals and Magazine of Natural History 
11(5):442^449. 

K.LAPALEK, F. 1894. Is Aleuropteryx lutea, Low, iden- 
tical with Coniopteryx lutea Wallg.? Entomolo- 
gist's Monthly Magazine 5:121-122. 

K.LIMASZEWSK.I, J. AND D. K. Mce. Kevan. 1 985. The 
brown lacewing tlies of Canada and Alaska 
(Neuroptera: Hemerobiidae). Part 1 The Genus He- 
merobius Linnaeus: Systematics, Bionomics and 
Distribution. Lyman Entomological Museum and 
Research Laboratory Memoir No. 15. 1 19 pp. 

— . 1987a. The brown lacewing flies of Canada 
and Alaska (Neuroptera; Hemerobiidae). Part lib. 
The genus Wesmaelius Kriiger. Neuroptera Interna- 
tional 4:245-274. 



. 1987b. The brown lacewing flies of Canada 

and Alaska (Neuroptera; Hemerobiidae). Part II[a]. 
The genus Wesmaelius Kriiger. Neuroptera Interna- 
tional 4:153-204. 167 figs., 8 maps. 

— . 1988a. The brown lacewing flies of Canada 



and Alaska (Neuroptera: Hemerobiidae). Part III. 
The genus Micromus Rambur. Giornale italiano di 
Entomologia 19:3 1-76, 42 figs., 4 maps. 

— . 1988b. New distribution data for Canadian 



Hemerobiidae (Neuroptera): Genera: Hemerobius 
Linnaeus, Micromus Rambur and Wesmaelius 
Kriiger. Supplement [1st]. Revue d'EntomoIogiedu 
Quebec 32:9-15. 

— . 1989. Distribution data for some Nearctic 



species of the genus Hemerobius Linnaeus and Wes- 
maelius Kriiger (Neuroptera: Hemerobiidae,). Sup- 
plement 2. Neuroptera International 5(4):205-210. 

. 1990a. New distribution data for some 



American species of the genus Hemerobius Lin- 
neaus [sic], with a redescription of Hemerobius 
alpestris Banks (Neuroptera: Hemerobiidae). An- 
nals of the Transvaal Museum 35:151-155, 6 figs. 

. 1990b. The genus Micromus Rambur in 



North America, with special reference to Canada 
and Alaska — a synopsis (Insecta: Neuroptera: He- 
merobiidae). Pp. 63-66 in Advances in Neurop- 
terology. M. W. Mansell and H. Aspock, eds. 
Proceedings of the Third International Symposium 
on Neuropterology, South African Department of 
Agricultural Development: Pretoria. 

Klug, J. C. F. [ 1 838] 1 836. Versuch einer systematis- 
chen Feststellung der lnsekten-Familie:Panorpatae 



106 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



und Auseinandersetzung ihrer Gattungen und 
Arten. Abhandlungen der koniglichen Akademie 
der Wissenschaften zu Berlin. 1836:81-108. 



Leonard, M. D. 1928. A list of the insects of New 
York. Cornell University Agricultural Experiment 
Station. Memoir 101. 



KOLBE, J. 1 888. Die geographische Verbeitung der 
Neuroptera und Pseudoneuroptera der Antillen, 
nebst einer Obersicht iiber die von Henn Consul 
Krug auf Portoriko gesammelten Arten. Archiv fur 
Naturgeschichte 54: 1 53- 1 78. 

KRUGER, L. 1922. Hemerobiidae. Beitrage zu einer 
Monographic der Neuropteren-Familie der He- 
merobiiden. Stettiner entomologische Zeitung 
83:138-172. 



LlCHTENSTElN, A. A. H. 1796. Catalogus rerum natu- 
ralium rarissimarum Hamburgi - auctionis lege dis- 
trahendarum. Gottl. Friedr. Schniebes, Hamburg. 
Vol. 3. xii + 222 pp. 

Linnaeus, C. 1758. Systema nafura per regna tria 
naturae secundum classes, ordines, genera, species, 
cum characteribus, differentiis, synonymis, locis. 
10th Edition. Vol. 1. Salvii, Holmiae, Sweden. 
824 pp. 



Kuwayama, S. 1 956. Further studies on the Neurop- 
tera-Planipennia of the Kurile Islands. Insecta Mat- 
sumurana, 20:77-82, 2 figs. 

Lacroix, J. L. 1926. Quelques insectes Nevropteres 
exotiques. Annates de la Societe Linneenne de Lyon 

72:67-72. 

LAGO, P. K. 1981. Mississippi spongillaflies [Sisyri- 
dae, Neuroptera]: A new state record, additional 
distribution records and A key to the adults. Journal 
of the Mississippi Academy of Sciences 26:27-30. 

Lago, P. K. and S. Testa, III. 1989. Records of 
owlflies (Neuroptera: Ascalaphidae) from Missis- 
sippi, with a key to species. Entomological News 
100(1):11-17. 

Lambkin, K.J. 1986. A revision of the Australian 
Mantispidae (Insecta: Neuroptera) with a contribu- 
tion to the classification of the family. I. General and 
Drepanicinae. Australian Journal of Zoology, Sup- 
plementary Series No. 116:1-142,391 figs. 

Latreille, P. A. 1796. Classe Quatrieme. Nevrop- 
teres. Pp. 96-104 in Precis des caracteres generi- 
ques des insectes disposes dans un ordre naturel. 
Paris. 201 pp. 

— . 1802. Histoire naturelle, generale et particu- 
liere de Crustaces et des Insectes. Vol. 3, Dufart, 
Paris. 467 pp. 

Lawson, H. R. and W. P. McCafferty. 1984. A 
check list of Megaloptera and Neuroptera (Planipen- 
nia) of Indiana. The Great Lakes Entomologist 
17(3): 129-131. 

Leach, W. E. 1815. Entomology. Pp. 57-172 in Ed- 
inburgh Encyclopaedia, D. Brewster, ed. Vol. 9, 
part 1 , Edinburgh. 



— . 1761. Fauna Svecica, sistens animalia 
Sveciae regni: Mammalia, Aves, Amphibia, Pices, 
Insecta, Vermes distributa per classes and ordines, 
genera and species, cum differentiis specierum, 
synonymis auctorum, nominibus natalium, descrip- 
tionibus insectorum. Salvii, Stockholmiae, Sweden. 
578 pp. 

. 1 763. Centuria Insectorum, quam, praeside 



D. D. Car. von Linne, proposuit Boas Johansson, 
Calmariensis. Pp. 384-415 in Amoenitates Aca- 
demicae; seu dissertationes variae physicae, medi- 
cae, botanicae; antehac seorism editae, nunc 
collectae et auctae cum tabulis aeneis. Vol. 6 (No. 
121 ). Salvii, Holmiae, Sweden. 

. 1 767. Systema natura per regna tria naturae 



secundum classes, ordines, genera, species, cum 
characteribus, differentiis, synonymis, locis. 12th 
Ed. Vol. 1, pt. 2 (pp. 533-1327). Salvii, Holmiae, 
Sweden. 

Low, F. 1885. Beitrag zur Kenntnis der Conioptery- 
giden. Sitzungsberichte der Akademie der Wissen- 
schaften, Wien. Mathematisch- Naturwissen- 
schaftliche Klasse (Abt. I) 91 : 73-89. 

Lucas, J. R. and L. A. Stange. 1981. Key and de- 
scriptions to the Myrmeleon larvae of Florida 
(Neuroptera: Myrmeleontidae). Florida Entomolo- 
gist 64(2):207-2 16, figs. 1-6. 

MacGillivray, A. D. 1894. New species of Notho- 
chrysa. Canadian Entomologist 26: 169-1 71 . 

MacLeod, E. G. and L. A. Stance. 1981. The brown 
lacewings of Florida. Florida Department of Agri- 
culture and Consumer Services, Division of Plant 
Industry, Entomology Circular 227: 4 unnumbered 
pages, 1 7 figs. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



107 



MacNeill, C. D. 1962. Observations on the flight 
behavior of an ascalaphid of the genus Ulalodes 
(Neuroptera: Ascalaphidae). Pan-Pacific Ento- 
mologist 38:186-188. 

Maddux, D. E. 1954. A new species of dobsonfly 
from California (Megaloptera: Corydalidae). Pan- 
Pacific Entomologist 30:70-71. 

Makarkin, V. N. 1985. A Review of the Hemerobii- 
dae (Neuroptera) of the USSR. 1. The genera He- 
merobius L., Micromus Ramb., and Paramicromus 
Nakah. [in Russian]. Entomologicheskoe 
Obozreniye 64:158-170. 49 figs. (English transla- 
tion: 1985: Entomological Review 64(2):99-l 13.) 



. 1891. Descriptions of new species of holo- 

phthalmous Ascalaphidae. Transactions of the En- 
tomological Society of London 1 89 1 :509-5 1 5. 

Meinander, M. 1972. A revision of the family 
Coniopterygidae (Planipennia). Acta Zoologica 
Fennica 136:1-357, 223 figs. 

. 1974. Coniopterygidae from western North 



America (Neuroptera). Entomologica Scandinavica 
5(3-4):2 17-232, 13 figs. 

1975. Coniopterygidae from North America 



(Neuroptera). Notulae Entomologicae, 55( 1 ):28-32 
4 figs. 



McClendon, J. F. 1901. A new species of Chrysopa 
from Texas. Psyche (Camb., Mass.) 9:215-216, 1 

— . 1906a. Notes on the true Neuroptera. 1. 



Brachynemurus curriei, a new ant-lion from Texas. 
Entomological News 17:93. 

-. 1906b. Notes on the true Neuroptera. 3. A 



catalogue of Texas Neuroptera. Entomological 
News 17:169-173. 

McLachlan, R. [1867] 1868. New genera and spe- 
cies, &c, of neuropterous insects; and a revision of 
Mr. F. Walker"s British Museum Catalogue of 
Neuroptera, part ii ( 1 853), as far as the end of the 
genus Mynnelcon. Journal of the Linnean Society 
of London, Zoology 9:230-28 1 . 

— . 1868. A monograph of the British Neurop- 
tera — Planipennia. Transactions of the Entomologi- 
cal Society of London 1 868: 145-224. 

. 1 869. New species, &c, of Hemerobiina; 

with synonymic notes (first series). Entomologist's 
Monthly Magazine 6:21-27, 1 fig. 



— . 1 873a [ 1 87 1 ]. An attempt towards a system- 
atic classification of the family Ascalaphidae. Jour- 
nal of the Linnean Society of London, Zoology 
11:219-276. 



. 1873b. Notes sur les Myrmeleonides decrits 

par M. le Dr. Rambur. Annales de la Societe de 
Entomologie de Belgique 16:127-141. 



— . 1990. The Coniopterygidae (Neuroptera, 
Planipennia). A check-list of the species of the 
world, descriptions of new species and other new 
data. Acta Zoologica Fennica 1 89: 1-95, 29 tigs. 

Merritt, R. W. and K. W. Cummins. 1978. An in- 
troduction to the aquatic insects of North America. 
Kendall/Hunt Publishing Company, Dubuque, 
Iowa. 441 pp. 

Montgomery, B. E. and A. W. Trippel. 1933. A 
preliminary list of Indiana Neuroptera. Entomologi- 
cal News 44:258-26 1 . 

Monserrat, V. J. 1 993. New data on some species of 
the genus Micromus Rambur, 1842. Annali del 
Museo Civico di Storia Naturale "G. Dona" 
89:477-516. 

— . 1996. Revision del Genero Hemerobius de 
Latinoamerica (Neuroptera, Hemerobiidae). Frag- 
menta Entomologica, Roma, 27(2):399-523. 

Motschuesky, V. Von. 1853. Neuroptera, Pp. 
211-212 in Etudes Entomologiques. Helsingfors. 

Munroe, E. G. 1953. Chauliodes disjunctus Walker: 
a correction, with the descriptions of a new species 
and a new genus (Megaloptera: Corydalidae). Ca- 
nadian Entomologist 85:190-192, 4 figs. 

Nakahara, W. 1960. Systematic studies on the He- 
merobiidae (Neuroptera). Mushi 34:1-69, 16 pis. 

. 1966. Contributions to the knowledge of the 



— . 1881. A North American species of Dilar. 
Entomologist's Monthly Magazine 18:55. 



Hemerobiidae of western North America (Neurop- 
tera). Proceedings of the United States National 
Museum 1 16:205-222, I pi., 4 figs. 



108 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



Navas, L. 1905. Notas zoologicas. VIII. Mis excur- 
siones durante el verano de 1904. Boletin de la 
Sociedad Aragonesa de Ciencias Naturales, 
Zaragoza 4: 107-131. 

— . 1908. Neuropteros nuevos. Memorias de la 



real Academia de Ciencias y Artes de Barcelona 
(3)6:401-423, 29 figs. 

— . 1909a. MonografiadelafamiliadelosDilari- 



dos (Ins. Neur. ). Memorias de la real Academia de 
Ciencias y Artes de Barcelona (3)7:619-671 . 



— . 1909b. Sur deux Hemerobides (insectes 
nevropteres) nouveaux. Annales de la Societe 
Scientifique de Bruxelles 33:2 1 5-220. 

— . 1911. Chrysopides nouveaux. Annales de la 



Societe Scientifique de Bruxelles 35:266-282. 
. 1912a. Neuropteros nuevos de America. 



[1A]. Broteria (Serie zoologica) 10:194-202. 

. 1912b. Crisopidos y Hemerobidos (Ins. 

Neur.) nuevos 6 criticos. Broteria (Serie zoologica) 
10:98-113. 



. 1913a. Les Chrysopides (Ins., Nevr.) du 

Musee de Londres. [1A]. Annales de la Societe 
scientifique de Bruxelles 37:292-330. [For the con- 
clusion of this article, see Navas, 1914e.l 



. 1913b. Nevropteres nouveaux de rAmerique 

du Nord. [1 ]. EntomologischeZeitschrift, Frankfurt 
am Main 27:19-20. 



— . 1913c. Crisopidos Sudamericanos. Broteria 
(Serie zoologica) 11:73-104, 149-168. 

— . 1913d. [Notas entomologicas, 2. a serie]. 5. 



Cuatro pequenas colecciones de Neuropteros de la 
peninsula Iberica. Boletin de la Sociedad Aragonesa 
de Ciencias Naturales, Zaragoza 12:77-89, 2 figs. 

— .191 3e. Neuropteros nuevos de America, [lb]. 



Broteria (Serie zoologica) 1 1 :45-53, 4 figs. 



— . 1914a. Neuropteros sudamericanos. Primera 
serie. Broteria (Serie zoologica) 12:45-56, 
215-234. 



— . 1914b. New Neuroptera from the United 
States. Bulletin of the Brooklyn Entomological So- 
ciety 9: 1 3-20, 5 figs. 



— . 1914c. Some Neuroptera from the United 
States. Bulletin of the Brooklyn Entomological So- 
ciety 9:60-62. 



— . I9l4d. Algunos Neuropteros del Museo de 
Oxford. I Serie. Boletin de la Sociedad Aragonesa 
de Ciencias Naturales, Zaragoza 13:61-68. 

— . 1914e. Les Chrysopides (Ins., Nevr.) du 



Musee de Londres. [IB]. Annales de la Societe 
scientifique de Bruxelles 38:73-1 14. [For the be- 
ginning of this article, see Navas, 1913a.] 

. 19I4L Mantispidos nuevos. (Segunda [II] 



serie). Memorias de la real Academia de Ciencias y 
Artes de Barcelona (3)1 1:83-103, 16 figs. 

. 1914g. Nevropteres nouveaux de rAmerique 



du Nord. Heme serie. Entomologische Zeitschrift, 
Frankfurt am Main 28: 1 8-20, 25-26. 

. 1915a. Some Neuroptera from the United 



States. Bulletin of the Brooklyn Entomological So- 
ciety 10:50-54, 2 figs. 

. 1915b. Neuropteros nuevos 6 poco conoci- 



dos. (Quinta [V] serie). Memorias de la real Acade- 
mia de Ciencias y Artes de Barcelona 
(3)11:455^180. 

. 1915c. Neuropteros nuevos 6 poco conoci- 



dos. (Sexta [VI] serie). Memorias de la real Acade- 
mia de Ciencias y Artes de Barcelona 
(3)12:119-136. [Separate: (3)1 2(7): 1-20.] 

. 1916a. Neuropteros sudamericanos. Tercera 



[III] serie. Neuropteros del Brasil recogidos por el 
R.P. Joaquin da Silva Tavares S.J. Broteria (Serie 
zoologica) 14: 14-35. 

— . 1916b. Notas sobre el orden de los Rafidiop- 



teros (Ins.). Memorias de la real Academia de Cien- 
cias y Artes de Barcelona (3)1 2:507-5 13,3 figs. 

. 1917a. Insectanova. 1 and 2 series. Memorie 



dell' Accademia Pontifica dei Nuovi Lincei, Roma 
3:1-22. 



. 1917b. Neue Neuropteren. Dritte Serie. En- 
tomologische Mitteilungen (Berlin) 6(7/9): 
274-282, 5 figs. 

. 1918a. Neuropteros nuevos 6 poco conoci- 



dos. (Decima serie). Memorias de la real Academia 
de Ciencias y Artes de Barcelona (3)14:339-366, 
20 figs. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



109 



. 1918b. Insectanova. IV Serie. Memorie dell' 

Accademia Pontificia dei Nuovi Lincei, Roma 
(2)4:13-23, 5 tigs. 



. 1925a. Neuropteren aus Brasilien. Mitteilun- 

gen der Miinchner entomologischen Gesellschaft 
15:64-68, 2 figs. 



. 1919a. Monografia de l'ordre dels Rafidiop- 

ters (Ins). Arxius [Arxivs] de l'lnstitute [d'Estudis 
Catalans, Seccio] de Ciencias, Barcelona. 90 pp., 42 
figs. 

. 1919b. Algunos insectos Neuropteros de la 

Republica Argentina. Serie Tercera. Revista de la 
real Academia de Ciencias Exactas, Fisicas y Naru- 
rales de Madrid 1 7:287-305, 6 figs. 

— . 1920. Sur des Nevropteres nouveaux ou cri- 



tiques II. Annales de la Societe Scientifique de 
Bruxelles39:l89-203, 5 figs. 

. 1 92 1 . Insectos americanos nuevos 6 criticos. 



Broteria (Serie zoologica) 19:1 13-124, 5 figs. 
. 1922a. Insectos exoticos. Broteria (Serie 



. [1925b] 1924. Communicaciones Ento- 

mologicas. 7. Neuropteros del Museo de Berlin. 
Revista de la real Academia de Ciencias Exactas, 
Fisico-Quimicas y Naturales de Zaragoza 
( 1 )9:20-34, 8 figs. 

. 1925c. Insectos exoticos nuevos 6 pococono- 

cidos. Segunda [II] serie. Memorias de la real Aca- 
demia de Ciencias y Artes de Barcelona 
(3)19:181-200, 11 figs. 



. 1925d. Crisopidos (Ins. Neur.) neotropicos. 

Segunda [II] serie. Revista chilenade Historia natu- 
ral 29:8-13, 2 figs. 

. 1925e. Entomologische Ergebnisse der 



zoologica) 20:49-63, 7 figs. 



schwedischen Kamtchatka-Expedition 1920-1922. 
6. Neuroptera et Mecoptera. Arkiv tor Zoologi 
18B(2):1-L 2 figs. 



. 1922b. Sur des Nevropteres nouveaux ou 

critiques. Sixieme [VI] serie. Annales de la Societe 
Scientifique de Bruxelles 42: 1 66-1 73. 

. 1 922c. Insectos nuevos 6 poco conocidos. [I]. 

Memorias de la real Academia de Ciencias y Artes 
de Barcelona 17:383-400. 



— . 1927a. Veinticinco formas nuevos de insec- 
tos. Boletin de la Sociedad iberica de Ciencias natu- 
rales 26:48-75, 9 figs. 

. 1927b. Insecta Nova. Series XII. Memorie 



dell' Accademia Pontificadei Nuovi Lincei, Roma 
(2)10:1-10, 4 figs. 



— . 1923. Estudis sobre Neuropters (Insectes). 
Arxius [Arxivs] de l'lnstitute [d'Estudis Catalans, 
Seccio] de Ciencias, Barcelona 7: 1 79-203, 2 figs. 

. 1924a. Algunos insectos de Cuba, recogidos 

por don Fermin Z. Cervera. [I]. Revista de la real 
Academia de Ciencias Exactas Fisicas y Naturales 
de Madrid (2)21:323-332. 



. 1924b. Algunos insectos de Cuba. Segunda 

[11] Serie. Revista de la real Academia de Ciencias 
Exactas, Fisicas y Naturales de Madrid 
(2)21:333-340. 



. 1924c. Crisopidos (Ins. Neur.) neotropicos. 

[I]. Revista Chilena de Historia Natural 
27:110-116. 



Estudios: Revista Mensual (Academia literaria del 
Plata, Buenos Aires) 22:358-368, 4 figs. 



. 1928a. Insectos del Museo de Estocolmo. 

Revista de la real Academia de Ciencias Exactas, 
Fisicas y Naturales de Madrid (2)24:28-39, 6 figs. 

. 1 928b. Insectos neotropicos. 3a serie. Revista 



chilena de Historia natural 31:31 6-328, 7 figs. 
. 1929a. Monografia de la Familia de los 



Berotidos (Insectos Neuropteros). Memorias de la 
Academia de Ciencias Exactas, Fisico-Quimicas y 
Naturales de Zaragoza 2: 1-107, 44 figs. 

1929b. Insectos Neuropteros del Museo de 



I924d. Insectos de la Argentina y Chile. (2)12:24-32, 3 figs 



Hamburgo. Memorias de la Real Sociedad Espanola 
de Historia Natural 15:315-322, 3 figs. 

. 1929c. Insecta Nova. Serie XIV. Memorie 

dell' Accademia pontifica dei Nuovi Lincei, Roma 



. 1931. Decadas de Insectos Nuevos. Decada 

3. Revista de la real Academia de Ciencias Exactas, 
Fisicas y naturales de Madrid 26:79-86. 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



— . 1932. Alcuni insetti del museo di zoologia 
dellareal Universitadi Torino. Bollettinodei Musei 
di Zoologia e di Anatomia Comparata della R. Uni- 
versitadi Torino (3)42(26): 1-38, 32 figs. 

— . 1933a. Insectos Suramericanos. Septima 



[VII] serie. Revistade la real AcademiadeCiencias 
Exactas, Fisicas y Naturales de Madrid 30:303-3 14, 
7 figs. 



. 1933b. Neurotteri e Tricotteri del "Deutches 

Entomologisches Instituf di Berlino-Dahlem. 
Bollettino della Societa Entomologica Italiana 
65:105-1 13, 8 figs. 

. 1933c. Decadas de insectos nuevos. Decada 



24. Broteria(Ciencias naturales) 2:101-1 10, 7 figs. 
934. Insectos de Mejico. Boletin de la So- 



ciedad Entomologica de Espana 16:68-75, 2 figs 

— . 1935. Monografia de la familia de los sisiri- 
dos (Insectos neuropteros). Memorias de la Acade- 
mia de Ciencias Exactas, Fisico-Quimicas y 
Naturales de Zaragoza 4: 1-87, 40 figs. 

Needham, J. G. 1 901 . Order Neuroptera, Pp. 540-560 
In Needham and Betten, Aquatic Insects in the 
Adirondacks. New York State Museum Bulletin 47 
(Entomology 13). 6 text figs., 1 pi. 

— . 1905. Systematic notes on Hemerobiidae. 



Bulletin of the New York Museum 86 (Entomology 
23): 1 5-1 7, 2 pis. 

Newman, E. 1838. Entomological Notes. Entomo- 
logical Magazine 5: 1 68-1 81 , 372-402, 483-500. 

Ohm, P. 1968. Vorlaufige Beschreibung einer neuen 
europaischen Aleuroptetyx-AvX (Neuroptera, 
Coniopterygidae). Entomologisches Nachrichten- 
blatt, Wien 15:12-15, 4 figs. 



. 1973. Zwei neue Coniopterygiden-Arten aus 

Siidwesteuropa (Planipennia, Coniopterygidae). 
Reichenbachia 14(30):237-243, 5 tigs. 

Olivier, G. A. 1811. 'Myrmeleon, Myrmeleon.' 
Pp.1 15-128 and 'Nemoptere, Nemoptera.' Pp. 
175-179 in Encyclopedic Methodique dTTistoire 
Naturelle, Insectes. Vol. 8. Paris. 

Oswald, J. D. 1 987. A new species of Micromus from 
the western United States (Neuroptera: Hemerobii- 
dae). Journal of the New York Entomological Soci- 
ety 95(1): 1^, 11 figs. 



. 1988. A revision of the genus Sympherobius 

Banks (Neuroptera: Hemerobiidae) of America 
north of Mexico with a synonymical list of the world 
species. Journal of the New York Entomological 
Society 96(4):390-45 I, 1 14 figs. 

. 1993. Revision and cladistic analysis of the 



world genera of the family Hemerobiidae (Insecta: 
Neuroptera). Journal of the New York Entomologi- 
cal Society 101:143-299, 222 tigs, 10 tables, 5 
appendices. 

Oswald, J. D. and Penny, N. D. 1991. Genus-group 
names of the Neuroptera, Megaloptera and 
Raphidioptera of the World. Occasional Papers of 
the California Academy of Sciences 1 47: 1 -94. 

Parfin, S. I. 1952. The Megaloptera and Neuroptera 
of Minnesota. American Midland Naturalist 
47:421^134. 

. 1956. Taxonomic notes on Kimminsia 



(Neuroptera: Hemerobiidae). Proceedings of the 
Entomological Society of Washington 58:203-209, 
1 5 figs. 

. 1958. Notes on the Bionomics of the Man- 



tispidae. Entomological News 69:203-207. 

Parfin, S. I. and A. B.Gurney. 1956. The spongilla- 
flies, with special reference to those of the Western 
Hemisphere (Sisyridae, Neuroptera). Proceedings 
of the United States National Museum 
105:421-529, 24 text figs., 3 pis., 24 figs. 

Penny, N. D. [1982a] 1981. Review of the generic 
level classification of the New World Ascalaphidae 
(Neuroptera). Acta Amazonica 11:391-406, II 
figs. 

. 1982b. Review of the generic level classifi- 



cation of New World Mantispidae (Neuroptera). 
Acta Amazonica 1 2( 1 ):209-223, 22 figs., 2 tables. 

. 1996. A remarkable new genus and species 



of Ithonidae from Honduras (Neuroptera). Journal 
of the Kansas Entomological Society 69( 1 ):8 1—86. 

Poirrier, M. A. 1969. Some fresh-water sponge hosts 
of Louisiana and Texas spongilla-flies with new 
locality records. American Midland Naturalist 

81:573-575. 

Poirrier, M. A. and R. W. Holzenthal. 1980. Re- 
cords of spongilla-flies (Neuroptera: Sisyridae) 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



111 



from Mississippi. Journal of the Mississippi Acad- 
emy of Sciences 25:1-2. 

Poole, R. W. and P. Gentili. 1996. Nomina Insecta 
Nearctica, A Check List of the Insects of North 
America. Volume 2: Hymenoptera, Mecoptera, 
Megaloptera, Neuroptera, Raphidioptera, Trichop- 
tera. Entomological Information Services. 

PROCTER, W. 1946. Biological survey of the Mount 
Desert Region incorporated. Part VII. The insect 
fauna. The Wistar Institute of Anatomy and Biol- 
ogy, Philadelphia. 566 pp. 

PROVANCHER, L. 1 875. Description de plusieurs insec- 
tes nouveaux. Le Naturaliste Canadien 7:247-251. 

. 1 877. Faune Canadienne (Petite Faune Ento- 

mologique du Canada). Les Nevropteres. Le Natu- 
raliste Canadien 9:38-43, 84-90, 118-123, 
173-176,201-205,209-217,241-244,257-269. 



Roble, S. M. 1984. First records of Limnephilus sub- 
monilifer (Trichoptera: Limnephilidae) and Neo- 
hermes concolor (Megaloptera: Corydalidae) for 
Kansas. Transactions of the Kansas Academy of 
Science 87(1 -2):69-70. 

ROSS, A. 1949. Les Mantispides de la Province de 
Quebec. Le Naturaliste Canadien 61:5-14. 

Ross, H. H. 1 937. Studies of Nearctic aquatic insects. 
I. Nearctic alderflies of the genus Sialis (Megalop- 
tera, Sialidae). Bulletin of the Illinois Natural His- 
tory Survey 21:57-78. 

Say, T. 1 823. Descriptions of insects belonging to the 
order Neuroptera Linn. Latr., collected by the expe- 
dition authorized by J. C. Calhoun, etc. under the 
command of Major S. H. Long. Godman's Western 
Quarterly Reporter of Medical Surgical, and Natu- 
ral Science 2(2): 160-1 65. 



Rambur, J. P. 1842. Histoire naturelle des insectes, 
Nevropteres. Librairie encyclopedique de Roret. 
Fain et Thunot, Paris. 534 pp. 

Redborg, K. E. 1976. Species of IVlantispidae 
(Neuroptera) at Dixon Springs Agricultural Center. 
Illinois Agricultural Experiment Station, Dixon 
Springs Agricultural Center Bulletin 4: 17-18. 

Redborg, K. E. and MacLeod, E. G. 1983. Clima- 
ciella brunnea (Neuroptera: Mantispidae): a man- 
tispid that obligately boards spiders. Journal of 
Natural History 17:63-73, 1 fig., 6 tables. 



— . 1985. The developmental ecology of Man- 
tispa uhleri Banks (Neuroptera: Mantispidae). Illi- 
nois Biological Monographs. 53:1-130. 1 8 figs, 30 
tables. 

Rehn,J. W. H. 1939. Studies in North American Man- 
tispidae (Neuroptera). Transactions of the Ameri- 
can Entomological Society 65:237-264, 22 figs. 

REUTER, O. M. 1894. Neuroptera Fennica. Forteckn- 
ing och Beskrifning ofver Finlands Neuropterer. 
Acta Societatis pro Fauna et Flora fennica 
9(8): 1-36. 

Rice, M. E. 1987. Morphological variation in Plega 
dactylota (Neuroptera: Mantispidae) and status of 
its subspecies. Journal of the Kansas Entomological 
Society 60:341-344. 



— . 1 824. Order Neuroptera. Pp. 303-3 1 in Nar- 
rative of an expedition to the source of the St. Peter's 
River, Lake Winnepeek, Lake of the Woods, &c. 
performed in the year 1 823, by order of the Hon. 
J.C. Calhoun, Secretary of War, under the command 
of Stephen H. Long, Major U.S.T.E., W.H. Keating, 
ed. Vol. 2. Carey and Lea, Philadelphia. 



. 1825. Mantispa. Plate 25 and unpaginated 

letterpress [7 pp.] in American Entomology, or de- 
scriptions of the insects of North America. Vol. 2. 
Philadelphia. 1 pi., 7 figs. 

— . 1839. Descriptions of new North American 



neuropterous insects, and observations on some al- 
ready described. Journal of the Academy of Natural 
Sciences of Philadelphia 8:9-46. 

Scheiring, J. F. 1979. New state record of fish fly 
Nigronia serricornis (Say) in Alabama (Megalop- 
tera: Corydalidae). Entomological News 90:176. 

Schimer, H. 1 865. Description of the imago and larva 
of a new species of Chrysopa. Proceedings of the 
Entomological Society of Philadelphia 4:208-2 12. 

Schneider, W. G. 1 85 1 . Symbolae ad monographiam 
generis Chrysopae, Leach. Hirt, Vratislaviae. 178 
pp., 60 pis. 

Shetlar, D. 1 977. The biosystematics of the Nearctic 
Ascalaphidae (Insecta: Neuropteroidea, Planipen- 



112 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



nia), with notes on biology and morphology. Un- 
published Ph.D. Thesis, Pennsylvania State Univer- 
sity, x + 233 pp., 47 figs. (University Microfilms 
Internationa] No. 78-3369.) 

Smith J. B. [1900] 1899. Order Platyptera [= Mega- 
loptera], Pp. 51-52, 2 figs, and Order Neuroptera, 
Pp. 53-57, 5 figs, in Insects of New Jersey. A list of 
species occurring in New Jersey, with notes on those 
of economic importance. Trenton, New Jersey. 
755 pp. 

Smith, R. C. 1922. The biology of the Chrysopidae. 
Memoirs of the Cornell University Agricultural Ex- 
periment Station 58:1287-1372, 10 text figs., 14 
pis., 122 figs. 

. 1 925. The Neuroptera and Mecoptera of Kan- 
sas. Bulletin of the Brooklyn Entomological Society 
20:165-171. 



. 1931. The Neuroptera of Haiti, West Indies. 

Annals of the Entomological Society of America 
24:798-823, 2 pis., 21 figs. 

. 1 932. The Chrysopidae (Neuroptera) of Can- 



ada. Annals of the Entomological Society of Amer- 
ica 25:579-601, 1 pi., 6 figs. 

Spencer, G. J. 1 942. A preliminary list of the Neurop- 
tera of British Columbia. Proceedings of the Ento- 
mological Society of British Columbia 38:23-28. 

Stance, L. A. 1961a. Lectotype designations in the 
New World Myrmeleontidae. Canadian Entomolo- 
gist 93:674-677. 

. 1 96 1 b. New records for the genus Nallachins 



Navas (Neuroptera: Dilaridae). Pan-Pacific Ento- 
mologist 37:144. 

. 1970a. Revision of the ant-lion tribe 



Brachynemurini of North America (Neuroptera: 
Myrmeleontidae). University of California Publica- 
tions in Entomology 25:1-192, 231 figs., 31 maps, 
2 tables. 

-. 1970b. A generic revision and catalog of the 



Western Hemisphere Glenurini with the description 
of a new genus and species from Brazil (Neuroptera: 
Myrmeleontidae). Los Angeles County Museum, 
Contributions in Science 186:1-28, 8 figs. 

— . 1980. The ant-lions of Florida. I. Genera. 



. 1 990. The alderflies of Florida (Megaloptera: 

Sialidae). Florida Department of Agriculture and 
Consumer Services, Division of Plant Industry, En- 
tomology Circular 329:2 unnumbered pages, 6 figs. 



. 1994. Reclassification of the New World 

antlion genera formerly included in the tribe 
Brachynemurini (Myrmeleontidae). Insecta Mundi 
8(1 ):67-l 19. 



. 1995. Entomology Section: Insect Detection. 

Tri-ology 34(2):7. 

Stange, L. A. and R. B. Miller. 1 990. Classification 
of the Myrmeleontidae based on larvae (Insecta: 
Neuroptera). Pp. 151-169 in Advances in Neurop- 
terology. M. Mansell and H. Aspock, eds. Proceed- 
ings of the Third International Symposium on 
Neuropterology. Department of Agricultural De- 
velopment, Republic of South Africa. 

Stark, B. P. and P. K. Lago. 1980. New records of 
Nearctic Sialis (Megaloptera: Sialidae), with em- 
phasis on Mississippi fauna. Entomological News 
91(4):1 17-121, 12 figs. 

. 1983. Studies of Mississippi fishflies (Mega- 



loptera: Corydalidae: Chauliodinae). Journal of the 
Kansas Entomological Society 56(3):356-364, 17 
figs. 

Steinmann, H. 1 964. The Chrysopa species (Neurop- 
tera) of Hungary. Annales Historico-Naturales 
Musei Nationalis Hungarici 56:257-266. 

Stephens, J. F. 1 829. A systematic catalogue of Brit- 
ish insects, being an attempt to arrange all the hith- 
erto discovered indigenous insects in accordance 
with their natural affinities; containing also refer- 
ences to every English writer on entomology, and 
to principal foreign authors; with all the published 
British genera to the present time. 2 vols. 416, 388 
pp. London. 

. 1836. Illustrations of British entomology; or, 



Florida Department of Agriculture and Consumer 
Services, Division of Plant Industry, Entomology 
Circular 2 1 5:4 unnumbered pages, 2 1 figs. 



a synopsis of indigenous insects: containing their 
generic and specific distinctions; with an account of 
their metamorphoses, times of appearance, locali- 
ties, food, economy, Mandibulata. Baldwin and 
Cradock, London. Vol. 6. 240 pp. 

Stitz, H. 1913. Mantispiden der Sammlung des Ber- 
liner Museums. Mitteilungen aus dem Zoologis- 
chen Museum in Berlin 7:1—49,41 figs. 



PENNY, ADAMS and STANGE: SPECIES CATALOG 



113 



Strom, H. 1788. Nogle Insect Larver med deres For- 
vandlinger. Ny Samling af det Kongelige Norske 
VidenskabersSelskabsSkrifter2:375-400cl0p1s., 
48 tigs. 

Tarter, D. C, D. L. Ashley and C. K. Lilly. 1976. 
New record of the alderfly Sialis itasca Ross, for 
West Virginia (Megaloptera: Sialidae). Entomo- 
logical News 87(1 /2):32. 

Tarter, D. C, W. D. Watkins, D. L. Ashley and 
J. T. Goodwin. 1978. New state records and sea- 
sonal emergence patterns of alderflies east of the 
Rocky Mountains (Megaloptera: Sialidae). Ento- 
mological News 89(9-1 0):23 1-234. 

Tarter, D. C, W. D. Watkins, M. L. Little and 
D. L. Ashley. 1977. New state record of the alder- 
fly Sialis concava Banks from Cranberry Glades, 
West Virginia. Entomological News 88(3/4): 104. 

Tarter, D. C, W. D. Watkins, M. L. Little and 
J. T. Goodwin. 1 976. New state records of fish flies 
(Megaloptera: Corydalidae). Entomological News 
87(7/8):223-228, 5 figs. 

Tarter, D. C. and J. E. Woodrum. 1973a. First re- 
cord of the alderfly, Sialis joppa Ross (Megalop- 
tera: Sialidae), in West Virginia. Proceedings of the 
West Virginia Academy of Sciences 
45(2): 165-167. 

. 1973b. Distribution and new record of the 



alderfly Sialis (Megaloptera: Sialidae) in West Vir- 
ginia. Entomological News 84:147-148. 

Taschenberg, E. L. 1879. Die Arten der Gattung 
Myrmecoleon Br. und Ascalaphus des Zoologisches 
Museums der Universitat Halle. Zeitschrift fiir die 
gesamte Naturwissenschaft 52: 1 74-23 1 . 

Tauber, C. A. 1969. Taxonomy and biology of the 
lacewing genus Meleoma (Neuroptera: Chrysopi- 
dae). University of California Publications in Ento- 
mology 58:1-94, 67 figs., 7 pis. 

. 1975. Larval characteristics and taxonomic 

position of the lacewing genus Suarius. Annals of 
the Entomological Society of America 
68(4):695-700, 4 figs. 

Tauber, C. A. andM. J. Tauber. 1973. Diversifica- 
tion and secondary intergradation of two Chrysopa 
carnea strains (Neuroptera: Chrysopidae). Cana- 
dian Entomologist 105(9): 1 153-1 167, 4 figs, 5 ta- 
bles. 



Taylor, C. 1862. An orthopterian defense. Harper's 
New Monthly Magazine 24:490^496. 

Tennessen, K. J. 1968. Four new species records of 
Sialis (Megaloptera: Sialidae) for Wisconsin. Wis- 
consin Academy of Sciences 53: 1 85-1 96. 

Throne, A. L. 1971a. The Neuroptera - Suborder 
Planipennia of Wisconsin. Part I - Introduction and 
Chrysopidae. Michigan Entomologist 4:65-78, 15 
figs., 3 tables. 



-. 1971b. The Neuroptera - Suborder Planipen- 
of Wisconsin. Part II - Hemerobiidae, 



Polystoechotidae and Sisyridae. Michigan Ento- 
mologist 4:79-87, 18 figs. 

. 1 972. The Neuroptera - Suborder Planipennia 



of Wisconsin. Part III - Mantispidae, Ascalaphidae, 
Myrmeleontidae and Coniopterygidae. The Great 
Lakes Entomologist 5(4): 1 19-128, 17 figs, 1 table. 

TlLLYARD, R. J. 1923. Descriptions of new species and 
varieties of lacewings (Order Neuroptera Planipen- 
nia) from New Zealand, belonging to the families 
Berothidae and Hemerobiidae. Transactions of the 
New Zealand Institute, Wellington 54:217-225, 5 
figs. 

Tjeder, B. 1960. Neuroptera from Newfoundland, 
Miquelon, and Labrador. Opuscula Entomologica 
25:146-149. 

. 1967. Kullabergs natvingar, nabbslandor, 



nattslandor och harkrankar. Kullabergs Natur 
12:1-21, 1 fig. 

Townsend, L. H. 1939. A new species of Sialis 
(Megaloptera, Sialidae) from Kentucky. Proceed- 
ings of the Entomological Society of Washington 
41:224-226, 6 figs. 

TSUKAGUCHI, S. 1 995. Chrysopidae of Japan ( Insecta, 
Neuroptera). Osaka, Japan. 223 pp. (Privately pub- 
lished.) 

Tullgren, A. 1906. Zur Kenntnis schwedischer 
Coniopterygiden. Arkiv for Zoologi 3(5): 1-1 5, 9 
figs. 

Van Dyke, E. C. 1 945. A new dobsonfly (egaloptera) 
from California. Pan-Pacific Entomologist 20:1 10. 

Vogtsberger, R. C. 1990. First United States record 
of Ascalobyas (Neuroptera: Ascalaphidae), a range 



14 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 3 



extension from northeastern Mexico to Texas. En- 
tomological News 101 (3): 1 58-1 60. 

Walker, F. 1853. Catalogue of the specimens of 
neuropterous insects in the collection of the British 
Museum, Part II (pp. 193-476). (Sialides -Nemop- 
terides). British Museum (Natural History), Lon- 
don. 

— . 1860. Characters of undescribed Neuroptera 



in the collection of W. W. Saunders. Transactions 
of the Entomological Society of London (N. S.) 
5:176-199. 

— . 1 866. Order Neuroptera. P. 334 in The Natu- 



ralist in Vancouver Island and British Columbia, J. 
K. Lord, ed. Vol. 2. Richard Bently, London. 

Wallengren, H.D.J. 1870. Anteckningar i Ento- 
mologi. Ofversigt af Kongl. Vetenskaps- 
Akademiens Forhandlinger, Stockholm 

27:145-182. 



Welch. P. S. 191 4. The early stages of the life history 
of Polystoechotes punctatus Fabr. Bulletin of the 
Brooklyn Entomological Society 9:1-6. 

Wesmael, C. 1841. Notice sur les Hemerobides de 
Belgique. Bulletin de FAcademie Royaledes Scien- 
ces et Belles-Lettres de Bruxelles 8( 1 ):203-22 1 . 

WESTWOOD, J. O. 1842. Description of some insects 
which inhabit the tissue of Spongilla fluviatilis. 
Transactions of the Royal Entomological Society of 
London (1)3: 105-108. 1 pi, 12 Figs. 

Wheeler, A. C, Jr. 1980. First United States record 
of Aleuropteryx simillima, a predator of scale in- 
sects on ornamental juniper (Neuroptera: Coniop- 
terygidae). The Southwestern Entomologist 
5(l):51-52. 

Whiting, M. F. 1991a. New species of Sialis from 
southern California (Megaloptera: Sialidae). Great 
Basin Naturalist 51 (4):41 1-413, 3 figs. 



. 1871. Skandinaviens Neuroptera. Kongl. 

svenska Vetenskaps-Akademiens Handlingar (N. 
F.) 9(8): 1-76. 



. 1991b. A distributional study of 5W/s( Mega- 
loptera: Sialidae) in North America. Entomological 
News 102(l):50-56. 



Weele, H. W. Van Der. [ 1 909a] 1 908. Ascalaphiden. 
Monograph isch Bearbeitet. Collections zoolo- 
giques du Baron Edmond de Selys Longchamps. 
Catalogue Systematique et Descriptif. Fasc. VIII. 
326 pp., 254 figs., 2 pis. 

— . 1 909b. New genera and species of Megalop- 



tera Latr. Notes from the Leyden Museum 
30:249-264. 

Welch, J. L. and B.C. Kondratieff. 1991. The 
Mantispidae (Neuroptera) of Colorado. Journal of 
the Kansas Entomological Society 64( 1 ):69-76, 16 
figs. 



Wise, K. A. J. 1973. New records in the New Zealand 
Neuroptera: Hemerobiidae. New Zealand Ento- 
mologist 5:181-1 85. 

Woglum, R. S. andE. A. MacGregor. 1964. Anew 
snakefly from southern California (Neuroptera - 
Raphidiidae). Bulletin of the Southern California 
Academy of Sciences 63:201-203, 1 fig. 

Zetterstedt, J. W. 1840. Ordo VII. Neuroptera. Col- 
umns 1037-1074 in Insecta Lapponica. Voss, Lip- 
siae. 1140 columns. This work printed in 1140 
individually numbered, double columns. 



CALIFORNIA ACADEMY OF SCIENCES, 1997 
Golden Gate Park 
San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 4, pp. 1 15-138, 9 figs., 9 tables. December 9, 1997 



A NEW LATE MIOCENE BALAENOPTERID WHALE 

(CETACEA: MYSTICETI), PARABALAENOPTERA BAULINENSIS, 

(NEW GENUS AND SPECIES) FRC^M THE SANTA CRUZ 

MUDSTONE, POINT REYES PENINSULA, CALIFORNIA 



By 

Carl V. Zeigler 



DEC 1 8 1997 

Woods Hole, MA 02543 



Cetacean fossil technician, College of Marin, Kentfield, California 94904; deceased, 



Gordon L. Chan 

Professor of Biology; College of Marin, Kentfield, California 94904; deceased, 1996. 

and 

Lawrence G. Barnes 

Section of Vertebrate Paleontology, Natural Histoiy Museum of Los Angeles County, 
900 Exposition Boulevard, Los Angeles, California 90007 



A fossil balaenopterid whale, Parabalaenoptera baulinensis, new genus and species, is named on 
the basis of a partial skeleton that was collected from the Late Miocene Santa Cruz Mudstone at 
Bolinas Point, Point Reyes Peninsula, California. Diatoms from the Santa Cruz Mudstone indicate 
an age of 6.8 to 6.0 Ma. The skeleton includes the cranium, dentaries, cervical and dorsal 
vertebrae, pectoral limb bones, and ribs, and the animal is estimated to have been 10 m in length 
when alive. The skull of Parabalaenoptera baulinensis has pronounced interdigitation of the 
rostral and cranial elements and an abrupt slope from the intertemporal region to the 
supraorbital process, indicating that it is a balaenopterid. However, the elongate and very narrow 
nasals, narrow intertemporal region, long and posteriorly tapering ascending processes of the 
maxillae, conspicuously elevated or swollen anterior portions of the parietals, short postglenoid 
processes, and long and sloping coronoid crest of the dentary distinguish P. baulinensis from fossil 
and Recent members of the subfamilies Megapterinae and Balaenopterinae. Its unique characters 
indicate its separation at the subfamily level, and we assign it to the new subfamily 
Parabalaenopterinae. 

Received September 11, 1995. Accepted April 15, 1996. 



Marine vertebrate remains occur within the sirenian, and Domning (1978) reported a du- 

Santa Cruz Mudstone exposed at the southern tip gongid sea cow from these exposures. From far- 

of the Point Reyes Peninsula, California (Fig. 1), ther north on the Point Reyes Peninsula, in the 

where rapid erosional retreat of the coastal cliffs same formation, Barnes (1976) reported a sperm 

frequently uncovers fossils. Galloway (1977) whale (cf. Scaldicetus du Bus, 1867); a small 

reported fragments of whale bone and an uniden- rorqual (cf. Burtinopsis Van Beneden, 1872); 

tified skull, possibly that of a cetacean or Wilson (1937) reported a dolphin; and Repen- 

[115] 



16 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



ning and Tedford (1977) reported an imago- 
tariine pinniped (pseudo-sea lion). 

The Bolinas Point area has yielded two major 
discoveries, relatively near each other, only 
about 375 m apart. The dugongid sea cow re- 
ported by Domning (1978) was found in 1972. 
The second specimen, discovered 12 August 
1976 by the senior author, began with the seren- 
dipitous discovery of a vertebra that had eroded 
from the cliff about 10 m above the beach. Fur- 
ther excavation along the cliff face yielded the 
partial skeleton of the balaenopterid which is the 
subject of this paper. We describe this fossil 
balaenopterid herein and name it as a new genus 
and species in a new subfamily of the family 
Balaenopteridae. 

Materials and Methods 

The location of the holotype skeleton in the sea 
cliff made its excavation difficult. Collection was 
dangerous because of the steepness of the cliff 
face (10 m above the beach), frequent landslides 
due to the crumbly nature of the mudstone, and 
strong surf action against the cliff face during 
high tides. To solve the problem of recovery, we 
decided to remove the fossil bone as pieces were 
sequentially (and naturally) exposed in the rap- 
idly retreating cliff face. During preliminary 
excavation we found that the vertebral column 
and skull were in no immediate danger of ero- 
sion. The entire collecting process required 
weekly, and sometimes daily, monitoring of the 
site over a period often years. 

Excavated fossil pieces, sometimes encased 
within the mudstone matrix, were brought to the 
College of Marin Biology Laboratory in Kent- 
field, California, for preparation. Removal of the 
bone from the surrounding matrix was time-con- 
suming due to the highly fractured condition of 
the fossil, its variable state of preservation, and 
the hardness (cementation) of the rock. Small 
fossil fragments were bonded with white glue or 
Glyptal cement. An epoxy compound proved to 
be most effective for the larger pieces, especially 
the dentaries. Final reconstruction of the entire 
specimen was completed in 1986. 

The skull and dentaries were not assembled as 
single units, but as closely fitting blocks or sec- 
tions that can be easily disassembled for study. 



The assembled skull rests on a fiberglass cradle 
with the dentaries alongside (Fig. 2). 

The descriptive anatomical terminology and 
measurements (e.g., Tables 1-8) follow Kellogg 
(1922, 1928, 1931, 1934) and Miller (1923). The 
classification follows that of Barnes, Domning 
and Ray (1985) and Fordyce and Barnes (1994). 
The anatomical drawings were made by a grid 
and measurement method. The authors and dates 
of publications are provided for all cetacean taxa, 
and those of all fossil taxa are listed in the LIT- 
ERATURE CITED. The estimate of body length 
(10 m) was made by taking the mean of How- 
ell's (1930:146) formula for rorquals, based 
on the length of the cervical series of verte- 
brae. 

The abbreviation CASG is for the Department 
of Geology, California Academy of Sciences, 
Golden Gate Park, San Francisco, California, 
U.S.A. 

Abbreviations used in the figures are as fol- 
lows: 

apm - ascending process of maxilla 

be - basioccipital crest 

cc - coronoid crest 

Bo - basioccipital bone 

cp - coronoid process of dentary 

earn - channel for external auditory meatus 

Eoc - exoccipital bone 

fm - foramen magnum 

fn - nutrient foramina 

Fr - frontal bone 

ft - temporal fossa 

gf- glenoid fossa 

Mx - maxillary bone 

n - naris 

Na - nasal bone 

Oc - occipital bone 

occ- occipital condyle 

Pa - parietal bone 

Pal - palatine bone 

pgp - postglenoid process 

Pmx - premaxillary bone 

pp - posterior process of periotic 

sop - supraorbital process of frontal 

Sq - squamosal bone 

Tb- tympanic bulla 

Vo - vomer bone 

zyg - zygomatic process of squamosal 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENSIS 



117 




Figure 1 . Type locality of Parabalaenoptera baulinensis, new genus and species, indicated by arrow, at Bolinas Point, Point 
Reyes Peninsula, Marin County, California. 



SYSTEMATICS 

Class Mammalia Linnaeus, 1758 

Order Cetacea Brisson, 1762 

Suborder Mysticeti Flower, 1864 

Family Balaenopteridae Gray, 1864 

Included Subfamilies. — Parabalaenopter- 
inae, new subfamily; Megapterinae (Gray, 1866) 
Gray, 1868; and Balaenopterinae (Gray, 1864) 
Brandt, 1872. 

Parabalaenopterinae, new subfamily 

Diagnosis of Subfamily. — A subfamily of 
the family Balaenopteridae differing from 



Megapterinae and Balaenopterinae by having 
cranium with a transversely narrower intertem- 
poral region; nasal bones elongate (approaching 
a length to width ratio of 4: 1), narrow, and nearly 
parallel-sided, terminating anterior to the level of 
the antorbital notch, rather than short, terminat- 
ing posterior to the antorbital notch, and distal ly 
wide; ascending process of maxilla tapered pos- 
teriorly rather than lobate or expanded trans- 
versely; exposure of frontals on dorsal surface of 
intertemporal region between apex of occipital 
shield and posterior ends of nasals, premaxillae, 
and maxillae nearly twice as wide anteroposteri- 
orly; parietals on either side of cranial vertex 
swollen and elevated above level of adjacent 
frontals; postglenoid process of squamosal short, 



1 1 8 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 

TABLE 1. Measurements (in mm) of cranium of holotype of P. baulinensis, new genus and species. 

1. Greatest length of skull, anterior end of right premaxilla to level of posteroexternal angles of 

exoccipitals 2240± 

2. Distance between anterior end of right premaxilla and apex of supraoccipital shield .... 1700± 

3. Distance between posterior angles of exoccipitals to apex of supraoccipital shield 540 

4. Distance from apex of supraoccipital shield to posterior end of left nasal bone 50 

5. Transverse distance between outside margins of premaxillae at level of anterior ends of nasal 

bones 94 

6. Maximum width of right premaxilla at level of anterior end of right maxilla 80 

7. Transverse diameter or rostrum at level of widest point of nasal fossa 620+ 

8. Greatest anteroposterior diameter of left supraorbital process 440 

9. External anteroposterior diameter of left supraorbital process 210 

10. Transverse diameter of cranium between outer surfaces of zygomatic processes 980 

11. Transverse diameter of cranium between margins of exoccipitals 690 

12. Width of occipital condyles 198 

13. Distance from upper margin of formen magnum to apex of supraoccipital shield 455 

14. Vertical distance from basioccipital to level of apex of suproccipital shield 250± 

15. Distance from anterior margin right zygomatic process to posterior margin exoccipital . . . 360 

16. Distance from anterior margin left zygomatic process to posterior margin exoccipital 475 

17. Greatest breadth of basioccipital between inner surfaces of notches for jugular leash 248 

18. Greatest length of right palatine bone 490+ 

19. Greatest width of combined palatine bones, anteriorly 275+ 

20. Greatest height of left occipital condyle 131 

21. Greatest width of left occipital condyle 76 

22. Vertical diameter of foramen magnum 93 

23. Transverse diameter of foramen magnum 64 

24. Greatest width of basioccipital between tympano-periotic processes 245 



extending only slightly ventral to the ventral whale, and Greek pteron, for fin; in reference to 

surface of the basioccipital crest; and dentary the apparent separation between this taxon and 

with long, sloping coronoid crest approaching the lineage that includes the living balaenopter- 

coronoid process rather than an abruptly ascend- ine whales, 
ing coronoid process; coronoid process with a 

spindle-shaped, posteriorly-directed apex; and Parabalaenoptera baulinensis, new species 

anterior margin of mandibular foramen smooth, (Figs. 2-7, 8b) 
lacking notch. 

Diagnosis of Species. — Because the genus 

Parabalaenoptera, new genus Parabalaenoptera is at present monotypic, the 

diagnosis of the type species, Parabalaenoptera 

DIAGNOSIS OF GENUS. — Because the sub- baulinensis, shall remain identical to that of the 

family Parabalaenopterinae is at present mono- genus Parabalaenoptera until further species are 

typic, the diagnosis of the genus shall remain diagnosed in the genus. 

identical to that of the subfamily Parabalaenop- HOLOTYPE. — CASG 66660, a partial associ- 

terinae until further genera are diagnosed in the ated skeleton consisting of most of the cranium 

subfamily. with tympanic bullae and periotics, both dentar- 

Type AND ONLY INCLUDED SPECIES. — Para- ies, fragments of the left scapula, both humeri, 23 
balaenoptera baulinensis, new species, Late vertebrae in varying states of preservation, in- 
Miocene of California. eluding 7 cervicals, 15 thoracic vertebrae, a lum- 

ETYMOLOGY. — From the Greek para, for be- bar vertebra, and numerous sections of 

side; plus balaenoptera, from Latin balaena, for incomplete ribs. 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAUL1NENS1S 



19 




Figure 2. Parabalaenoptera baulinensis, new genus and species, assembled holotype cranium and dentaries, C ASG 66660, 
oblique anterodorsolateral view. 




Figure 3. Parabalaenoptera baulinensis, new genus and species, oblique anterodorsal view of part of holotype skull, CASG 
66660, showing elongate nasal bones. 



120 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



Type LOCALITY. — The sea cliff, above the 
high tide mark near Bolinas Point (37° 54' 6" N, 
122° 43' 15" W), Point Reyes Peninsula, Marin 
County, California, U. S. A (Fig. 1). 

FORMATION AND AGE. — Santa Cruz Mud- 
stone, of latest Miocene age, between approxi- 
mately 6.8 and 6.0 Ma. 

The holotype skeleton of Parabalaenoptera 
baulinensis was parallel to the bedding planes in 
shale of the Santa Cruz Mudstone, dipping 45° 
to the southwest. The interbedded siltstone con- 
tains faint parallel laminations in beds ranging 
from 5 to 25 cm thick. Concretions and carbon- 
ate-cemented horizons are common and occur 
parallel to the bedding planes. Formerly consid- 
ered part of the Monterey Formation, these rocks 
are now assigned to the Santa Cruz Mudstone 
(Clark etal. 1984). 

Although not abundant, fossils such as gastro- 
pods, pelecypods, echinoderms, isopod frag- 
ments, crab claws, decapod shrimp parts, fish 
teeth and bones, and isolated mammal bones 
have been collected from this formation. Many 
of the fossils, including the fossil whale, are 
better preserved in the carbonate-cemented con- 
cretions, although fossils occur throughout the 
Santa Cruz Mudstone. Fossil mollusks in the 
same strata as the holotype of Parabalaenoptera 
baulinensis were tentatively identified as limids, 
possibly Acesta sp. These bivalves generally re- 
quire water depths of 500 m and water tempera- 
tures of 8°C or less (Ellen J. Moore, Branch of 
Paleontology and Stratigraphy, U. S. Geological 
Survey. Menlo Park, California, 5 May 1984. 
pers. comm.). 

An assemblage of fossil diatoms was recov- 
ered from carbonate concretions in the same bed- 
ding plane adjacent to the fossil whale. The 
diatom assemblage is assignable to "Subzone A" 
of the Late Miocene Nitzschia reinholdi Zone, 
and implies an estimated age of 6.8 to 6.0 Ma for 
the fossil whale (John A. Barron, Branch of 
Paleontology and Stratigraphy, U. S. Geological 
Survey, Menlo Park, California, 10 January 
1985, pers. comm.). 

ETYMOLOGY. — The species name, baulinen- 
sis, is in reference to the type locality at Bolinas 
Point, Point Reyes Peninsula, Marin County, 
California; Bolinas is named after and was part 
of the 1860 historical land title of Rancho las 
Baulines. 



DESCRIPTION. — Cranium. — Characters 
that distinguish the cranium of Parabalaenop- 
tera baulinensis from those of Balaenoptera 
Lacepede, 1 804, and Megaptera Gray, 1 846, are: 
the more narrow intertemporal region (Figs. 4, 
6), the long and narrow nasals (Figs. 3, 4, 6), and 
the laterally swollen and elevated portions of the 
parietals on either side of the intertemporal re- 
gion (Fig. 7b). 

The entire rostrum is not preserved and some 
large sections are missing. In order to estimate 
the original total length of the skull, we calcu- 
lated the ratios of rostral length to total skull 
length in large extant species of rorquals. The 
average of these gives a rostral length that is 
approximately 74% of the total skull length. We 
applied this ratio to the fossil to obtain its esti- 
mated original skull length and body length. 

The rostrum is very flat and gradually in- 
creases in depth posteriorly. The lateral margin 
of the maxilla is thin and blade-like. The maxilla 
thickens medially as it curves medially and ven- 
trally to enclose the vomer. 

The distal ends of the premaxillae are missing 
anterior to the distal ends of the maxillae (Fig. 6). 
The portion of the left premaxilla anterior to the 
narial opening has been pressed ventral to and 
against the right premaxilla as a result of geologi- 
cally induced right-lateral compression of the 
cranium. In life, these bones were definitely 
separated, and probably somewhat parallel, up to 
their point of divergence at the anterior margin 
of the vertical orientation as they delineate the 
long, broad narial fossa. Lateral to the nasals, the 
premaxillae become progressively narrower pos- 
teriorly, only being 10 mm wide at the posterior 
ends of the nasals, and terminate 45 mm from the 
apex of the supraoccipital. 

The maxillae are fractured and are missing 
some portions on their lateral margins (Fig. 6). 
What is preserved of the rostrum indicates that it 
was very broad, not arched anteroposteriorly, 
and that its lateral margins were thin and curved 
ventrally. The posterior parts of the rostral mar- 
gins are fragmentary and their exact relationship 
to the antorbital processes is difficult to deter- 
mine. The long, ascending (posterior) processes 
of the maxillae, overlapping and penetrating the 
frontals, are cuneate in shape, tapering gradually 
posteriorly to within 30 mm of the anterior mar- 
gin of the supraoccipital, and are each 15 mm 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENSIS 



121 



fracture 




Figure 4. Parabalaenoptera baulinensis, new genus and species, dorsal view of holotype cranium and dentaries, CASG 
66660; fractured area of left dentary, discussed in text, is indicated. 




FIGURE 5. Parabalaenoptera baulinensis, new genus and species, lateral view of holotype cranium and right dentary, CASG 
66660; see Materials and Methods for abbreviations. 



wide at the level of the posterior margin of the 
nasals. 

The nasals (see Figs. 3, 6) are distinctive by 
being extremely long (maximum length = 250 
mm) and narrow for a balaenopterid, and this is 
a primitive character. Their combined anterior 
width, as preserved now is 57 mm, but we esti- 
mate that this width was 64 mm prior to geologic 
distortion. The combined posterior transverse 
width of the nasals is 29 mm. The nasal bones are 
tightly wedged between the ascending processes 
of the premaxillae, and terminate 50 mm from 
the apex of the supraoccipital. 



The length-to-anterior width ratio of the com- 
bined nasals is approximately 4:1 (Fig. 3). By 
contrast, among the living balaenopterids, this 
ratio ranges from approximately 2:1 in 
Balaenoptera borealis Lesson, 1828, (Sei whale; 
see Fig. 8c), to approximately 1:1 in Balaenop- 
tera physalus (Linnaeus, 1758) (fin whale; see 
Fig. 8d), which has exceptionally short nasals (a 
derived character). The anterior ends of the 
nasals in Parabalaenoptera baulinensis extend 
anteriorly farther than the anterior margin of the 
supraorbital processes. Using the supraorbital 
process as a point of reference for comparison 
with living balaenopterids, only in the blue whale 



122 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 




FIGURE 6. Parabalaenoptera baulinensis, new genus and species, dorsal view of holotype cranium, CASG 66660; see 
Materials and Methods for abbreviations. 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENSIS 



123 




Eoc 





occ 



Figure 7. Parabalaenoptera baulinensis, new genus and species, braincase of holotype, CASG 66660; a. posterior view; 
b. lateral view; c. ventral view; see Materials and Methods for abbreviations. 



124 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



Balaenoptera musculus (Linnaeus, 1758); see 
Fig. 8g) do the nasal bones extend more pos- 
teriad. 

Concomitant with the elongate nasals is the 
conspicuously anterior position of the narial 
opening, which has its maximum width of 
140 mm at a point 180 mm anterior to the ante- 
rior ends of the nasals. The dorsoventral depth of 
the narial opening at this point, from the dorsal 
surfaces of premaxillae to the vomer, is 135 mm. 
The anteroposterior diameter of the narial open- 
ing, taken from the point of divergence of the 
premaxillae to the anterior ends of the nasals, is 
520 mm. The choanae, ventral to the nasals, are 
filled with matrix and not visible for inspection. 

Both supraorbital processes of the frontals are 
fragmented, especially the right process, which 
crumbled during excavation; however, the left 
supraorbital process is nearly complete (Fig. 6). 
Unfortunately, the relationship of the antorbital 
process to the lacrimal bone cannot be deter- 
mined as the lacrimal bones are both missing. As 
is typical of balaenopterids, the supraorbital 
process is broad, somewhat flat, approximately 
rectangular in dorsal view, and its dorsal surface 
slopes ventrally away from the higher cranial 
vertex. The supraorbital process is broad antero- 
posteriorly at its medial part. The lateral orbital 
margin is moderately arched dorsally. The flat 
plate of the supraorbital process is very long 
anteroposteriorly at the base and is abruptly de- 
pressed well below the dorsal level of the interor- 
bital region. The lateral margin of the 
supraorbital process is slightly arched over the 
orbit, and the postorbital process is small as in 
Recent balaenopterids. The optic nerve canal is 
enclosed proximally, but as it proceeds laterally, 
the canal opens and flares out to form the slightly 
arched roof of the orbit. 

In the intertemporal region the parietals are 
swollen on either side of the frontals and con- 
spicuously raised laterally above the nearly ver- 
tical lateral margins of the interorbital region. An 
anterior projection of the anteroexternal portion 
of the parietals extends over the frontals on either 
side of the intertemporal region and the cranial 
vertex, and they extend on either side of the 
intertemporal region anteriorly beyond the pos- 
terior ends of the nasals, the premaxillae, and 
ascending processes of the maxillae. 

The cranial vertex has the typical configuration 
of a balaenopterid, with the rostral bones deeply 



interdigitating with the frontals. There is a small, 
somewhat rectangular exposure of the frontals on 
the vertex, but it is still larger than in any Recent 
balaenopterid (Fig. 8). The narrow (for a 
balaenopterid) interorbital part of the frontals is 
exposed between the posterior ends of the nasals, 
the maxillae, and the premaxillae. 

The triangular occipital shield tapers to a trun- 
cated apex, which extends anteriorly to a point 
beyond the level of the anterior extremities of the 
zygomatic processes of the squamosals, ap- 
proaching a line drawn transversely through the 
centers of the orbits. The anterior extent of the 
supraoccipital shield and the parietals in P. 
baulinensis exceeds that of B. musculus, and 
closely approaches that of the other Recent spe- 
cies of Balaenopteridae. The apex of the occipital 
shield is not clearly defined, due to apparent 
abrasion prior to deposition. The essentially 
straight lateral margins of the occipital shield 
project laterally over the temporal fossae. There 
is no concavity to the lateral margin of the occipi- 
tal shield such as is present in all living species 
of Balaenoptera (Fig. 8). The mid-part of the 
occipital shield is depressed, attaining a maxi- 
mum depression of 85 mm below the lateral mar- 
gins. A lambdoidal crest, formed by the 
conjunction of the lateral margins of the supraoc- 
cipital with the posterior margins of the 
squamosals and the parietals, arcs dorsomedially 
toward the truncated apex of the supraoccipital 
shield. The braincase of P. baulinensis is shallow 
dorsoventral ly (Fig. 5), notwithstanding the geo- 
logic compression of the skull. The wide exoc- 
cipitals flare ventrolaterally (Fig. 7a). The 
exoccipitals protrude only slightly beyond the 
posterior margin of the squamosal, and comprise 
the posterolateral corners of the cranium. 

The parietal suture with the squamosal is ob- 
scure, however, where the parietal overrides the 
supraorbital process of the frontal, the bounda- 
ries of this bone can be traced. The anterior-most 
extension of the parietal lies 100 mm distal to the 
posterior level of the ascending process of the 
maxilla. 

The zygomatic process of the squamosal is 
large, excavated ventrally, and arches upward at 
a steep angle from the shallow postglenoid proc- 
ess. It terminates in a stout, anteromedially- 
directed anterior tip. 

The occipital condyles are widely separated 
dorsally, but are closely approximated ventrally. 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENSIS 



125 










f 



g 



h 



Figure 8. Dorsal views of skulls of fossil and Recent Balaenopteridae; a. Plesiocetus cortesii (after Van Beneden, 1 875:fig. 
1); b. Parabalaenoptera baulinensis, new genus and species; c. Balaenoptera borealis (after Miller, 1924b:pl. 1); d. 
Balaenoptera physalus (after True, 1904:pl. 1, fig. 3); e. Balaenoptera «/e«/ (after Omura, 1959:pl. l,fig. l);f. Balaenoptera 
acutorostrata (after True, 1 904:pl. 22, fig. 1 ); g. Balaenoptera musculus (after Miller, 1 924a:pl. 1 ); h. Megaptera novaeangliae 
(after True, 1904:pl. 29); not to scale. 



Their articular surfaces are narrowest dorsally 
and broadest near the ventral margin of the fora- 
men magnum. There is a slight constriction or 
neck around the bases of the occipital condyles, 
which have slightly rugose articular surfaces. 
The foramen magnum is posterodorsally ori- 
ented. 

The flat ventral surfaces of the basioccipital 
crests are on the same plane as the ventral mar- 



gins of the exoccipitals. The postglenoid proc- 
esses (these are wide but very shallow dorsoven- 
trally), extend only about 30 mm ventrally. 
Although there is some evidence of minor abra- 
sion of the ventral surface of the postglenoid 
processes, it is not enough to affect their dor- 
soventral dimension. 

A vertically oriented slot-like foramen 
pseudovale (37 mm * 13 mm) is open anterolat- 



126 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



erally as in Recent rorquals. It is located in the 
squamosal immediately above the ventral junc- 
ture with the pterygoid at about the same level as 
the anterior margin of the scaphoid fossa. 

The ventral surface of the rostrum is not as 
complete nor as well preserved as the dorsal 
surface. The infraorbital processes of the maxil- 
lae, characteristic of all mysticetes, have been 
broken off and are missing. The boundaries of the 
palatines can only be barely discerned, but their 
anterior and lateral margins can be faintly traced. 
Their posterior margins are obscure, except for 
the divergent medial margins that reveal a shal- 
low vomer (20 mm deep at this point), which 
forms the vertical septum between the narial 
passages (Fig. 7c). The relatively small cross- 
sectional area of the nasal choanae is a primitive 
condition, and may indicate a less efficient div- 
ing ability compared to the living rorquals. The 
exposed vomer septum, extending posteriorly on 
the basicranium, decreases in height posteriorly 
until it disappears approximately 220 mm from 
the posteromedial margins of the palatines. 

The distributions of some of the basicranial 
bones are unclear because of fusion of the sutures 
between them and the incomplete preservation. 
The basioccipital is transversely concave be- 
tween the descending basioccipital crests, which 
have relatively flat ventral surfaces and have a 
dorsoventral dimension of 57 mm. The greatest 
transverse distance between the medial margins 
of the ventral surfaces of the basioccipital crests 
is 116 mm. 

Both tympanic bullae were recovered and the 
left bulla remains in place in the basicranium. 
The right bulla measures 80 mm anteroposteri- 



orly, 52 mm transversely, and 41 mm in depth. 
The tympanic bullae extend approximately 10 
mm below the basioccipital crests (Fig. 7a). The 
relatively long posterior processes of the peri- 
otics are wedged between the squamosals and 
exoccipitals (Fig. 7c). 

The postglenoid process is broad transversely 
and very shallow dorsoventrally. The slightly 
posteriorly directed postglenoid process ob- 
scures (in ventral view) the central part of the 
deep external acoustic meatus. This meatus is 
straight and is almost at 90° to the sagittal plane. 
The zygomatic process of the squamosal projects 
laterally and anteriorly about 45° to the sagittal 
plane of the cranium. It arches dorsally, terminat- 
ing in an enlarged, medially directed anterior 
extremity. 

The occipital condyles do not extend so far 
posteriorly as do the posterolateral corners of the 
exoccipitals. The exoccipitals are very com- 
pressed anteroposteriorly, being not much 
thicker in this dimension than the lateral end of 
the posterior process of the periotic. 

Dentary. — Both dentaries (Figs. 4, 5) are 
nearly complete, except for a ± 100 mm section 
of the right horizontal ramus about 120 mm from 
the anterior tip, and a few small fragments miss- 
ing from the coronoid processes and the dorsal 
margins along the dental foramina (see Table 2). 

There is evidence that the left dentary was 
broken in life. An apparent fracture (Fig. 4) is 
located about 550 mm from the anterior end. It 
appears that the break was at least partially 
healed, leaving a ± 25 mm offset of the anterior 
portion. Subsequent transverse fracturing of the 
fossilized bone in this area is unclear. The injury 



TABLE 2. Measurements (in mm) of right dentary of holotype of Parabalaenoptera baulinensis, 
new genus and species. 

1 . Greatest length of dentary along outside curvature 2340± 

2. Greatest length of dentary in a straight line 2240+ 

3. Greatest vertical diameter of dentary at level of symphysis 74 

4. Greatest transverse diameter of dentary at level of symphysis 47 

5. Greatest vertical diameter of dentary at a point 850 mm anterior to hinder face of condyle. 150 

6. Greatest vertical diameter of dentary at coronoid process 124 

7. Least vertical diameter of dentary behind coronoid process 245 

8. Greatest vertical diameter of condyle 1 80 

9. Greatest transverse diameter of condyle 127 

10. Greatest dorsoventral diameter of portal to mandibular canal 66 

11. Greatest transverse diameter of portal to mandibular canal 43 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENSIS 



127 



to the left dentary might account for its being The fossa that is confluent with the mandibular 

more slender than the right dentary. foramen begins near the mandibular condyle, and 

Seven nutrient foramina (Figs. 4, 5) are located is directed slightly ventrally, and deepens as it 

along the dorsolateral side of each dentary. The progresses anteriorly. At a point just posterior to 

spacing between these foramina varies between the coronoid process, the mandibular foramen 

the two dentaries. enters the dentary with an arch-like margin curv- 

The dentary is conspicuously bowed laterally ing smoothly ventrally and slightly posteriorly; 

(Figs. 4, 5), with no evidence of torsion or lateral consequently, there is no notch in the anterior 

flattening of the anterior portion. It increases in margin of the foramen as in Balaenoptera and 

transverse thickness very gradually to about the Megaptera. 

level of the posterior margin of the coronoid Vertebrae. — The preserved anterior part of 

process, where it then narrows to become con- the vertebral column includes the seven cervi- 

stricted between the coronoid process and the cals, fifteen thoracic vertebrae in series, and an 

mandibular condyle. isolated lumbar vertebra. The centra of these 

In addition to the difference in thickness be- vertebrae are mostly well preserved and show 
tween the two dentaries, the coronoid regions of little evidence of abrasion, 
the two dentaries show some differences. The The centra of vertebrae in the thoracic and 
right coronoid process is somewhat longer an- lumbar regions are uniformly separated by ma- 
teroposteriorly, and has a greater lateral curva- trix that filled the inter-vertebral spaces. The 
ture than does the left. This difference in neural arches are only fragmentary and the trans- 
curvature is probably due to distortion pressure verse processes have been broken away from the 
from the left that straightened the left coronoid, centra of most of the vertebrae. Cervicals 6 and 
while exaggerating the curvature of the right 7, and the first and second thoracic vertebrae 
coronoid (Fig. 5). If this is the case, the distortion remain in a block of concretionary matrix. The 
was very uniformly distributed, because the out- vertebral epiphyses are completely fused to all of 
ward arc of the right coronoid process is smooth the centra, indicating that the holotype was an 
throughout its entire length. adult individual. 

At a point on the dentary about two-thirds of Atlas. — The neural arch of the atlas (see Ta- 

its length from the anterior tip, the coronoid crest ble 3) is missing, as is the right parapophysis. The 

begins its long, sloping elevation from the dorsal posterior articular surfaces are convex. Along the 

margin of the ramus. As the dorsal margin of the ventral margin of the atlas is a lip-like projection, 

coronoid crest sweeps gradually upward, it also most prominent at the midline, which extends 

begins arcing laterally. At its apex, the coronoid posteriorly a short distance under the ventral 

process forms a pointed and posteriorly directed surface of the axis, and conforms closely to the 

tip creating a boss (Fig. 5). contours of the latter. The imperforate left trans- 

TABLE 3. Measurements (in mm) of atlas of holotype of Parabalaenoptera baulinensis, new genus 
and species. 

Greatest breadth across transverse processes 354+ 

Distance ventral margin midline to dorsal margin adontoid fossa 50 

Greatest height of right anterior articular surface 150 

Greatest breadth of right anterior articular surface 85 

Greatest breadth of left anterior articular surface 75 

Greatest transverse diameter of neural canal 80 

Maximum breadth across posterior articular surfaces 218 

Greatest height of right posterior articular surface Ill 

Greatest height of left posterior articular surface Ill 

Greatest breadth of right posterior articular surface 80 

Greatest breadth of left posterior articular surface 80 

Greatest anteroposterior diameter 68 

Maximum anteroposterior diameter at ventral midline 57 



128 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



TABLE 4. Measurements (in mm) of axis of holotype of Parabalaenoptera baulinensis, new genus 
and species. 

1 . Greatest breadth across transverse processes 500± 

2. Height of right transverse process at exterior margin of transverse foramen 143 

3. Height of left transverse process at exterior margin of transverse foramen 130 

4. Transverse diameter of right transverse foramen 67 

5. Transverse diameter of left transverse foramen 62 

6. Dorso-ventral diameter of right transverse foramen 38 

7. Dorso-ventral diameter of left transverse foramen 32 

8. Distance of external margin right articular surface to internal margin right transverse foramen 12 

9. Distance of external margin left articular surface to internal margin right transverse foramen 15 

10. Greatest breadth of anterior articular surfaces 204 

1 1. Height of right anterior articular surface 121 

12. Height of left anterior articular surface 120 

13. Maximum extension of odontoid process above the posteriormost level or articular surface . 18 

14. Distance of anteriormost margin of odontoid process above midline ventral margin 50 

15. Greatest posterior transverse diameter of centrum 160 

16. Greatest posterior dorsoventral diameter of centrum Ill 

17. Greatest anteroposterior diameter of centrum at dorsal level 50 

18. Greatest anteroposterior diameter of centrum at ventral level 37+ 



verse process is short and stout, tapering to a 
rounded end. This process extends 55 mm be- 
yond the lateral margin of the left posterior ar- 
ticular surface at a level slightly below the dorsal 
margin of this facet. 

Axis. — The neural arch of the axis (Table 4) 
and a small portion of the right transverse process 
are missing. There is a rudimentary odontoid 
process projecting anteriorly only 18 mm from 
the concave anterior articulating surfaces. The 
long and deep, wing-like, transverse processes 
are posteriorly deflected approximately 30° from 
the transverse plane of the centrum. 

Third to Seventh Cervical Vertebrae. — 
The third cervical (see Table 5) is a very fragile 
vertebra compared to the robust axis. The supe- 
rior and inferior transverse processes meet to 
delineate the relatively large transverse foramen. 
The left foramen has an estimated transverse 
diameter of 100 mm and a dorsoventral diameter 
of 75 mm. The neural canal is 95 mm in trans- 
verse diameter. The greatest breadth across the 
transverse processes would appear to approach 
that of the axis. 

The fourth cervical is the most anteroposteri- 
orly compressed cervical vertebra. Its neural ca- 
nal has an estimated dorsoventral diameter of 
50 mm and an estimated transverse diameter of 
95 mm. 



The fifth cervical vertebra reverses the trend of 
anteroposterior compression from the atlas to- 
ward cervical 4, with the sixth and seventh cerv- 
icals becoming progressively thicker antero- 
posteriorly. The neural arches are crushed upon 
the centra of all three of these vertebrae. 

Thoracic and Lumbar Vertebrae. — Due to 
the fragmentary condition or absence of trans- 
verse processes on the series of 15 vertebrae 
posterior to the cervicals, it is not possible to 
determine the transition point from the thoracic 
to lumbar vertebrae. It is possible that the series 
are all thoracic vertebrae, inasmuch as the num- 
bers of thoracic vertebrae in balaenopterid 
whales range from 1 1 to 15 (Kellogg 1925). With 
this variability in mind, we refer to the vertebrae 
as the thoracic-lumbar series (Table 6); the more 
complete twelfth vertebra in that series is treated 
more extensively (Table 7). The first thoracic has 
a centrum that is larger dorsoventrally than any 
others in the thoracic-lumbar series, 3 through 
15, and the width of the anterior face of its 
centrum is exceeded only slightly by the second 
thoracic. There is a progressive increase in length 
of the centra posteriorly in the column. The iso- 
lated lumbar vertebra is smaller in all dimensions 
compared with the last vertebra in the preceding 
series. 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENSIS 



129 



TABLE 5. Measurements (in mm) of cervical vertebrae 3-7 of holotype of Parabalaenoptera 
baulinensis, new genus and species. 



Cervical vertebrae 



1 . Length of centrum at dorsal level 

2 . Height of anterior face of centrum 

3 . Width of anterior face of centrum 

4. Length of centrum at ventral level 

Ribs. — The ribs are very fragmentary, but 
parts of eight right ribs and one left rib have been 
assembled. The right rib that was found adjacent 
to the right humerus measures 820 mm in length 
along its exterior margin. The narrowest part of 
this rib is at the neck, where it measures 26 mm 
anteroposterior^, and the thickest part of this rib 
in this dimension is 43 mm, midway between the 
tuberosity and the exterior margin of the angle. 
In anterior view the rib is widest (80 mm) at the 
angle, and the transverse diameter at the broken 
distal portion is 68 mm. 

Scapula. — Only fragments of the left scapula 
were collected. The largest of these is the distal 
portion of the scapula bearing the glenoid area. 
The glenoid fossa has a maximum anteroposte- 
rior diameter of 1 50 mm. There is a protuberance 
with a broken surface about 20 mm above the 
anterior margin of the glenoid fossa that may be 
the base of the coronoid process; it is directed 
anteriorly and slightly medially. 

Humerus. — Both humeri are relatively intact 
and are fairly complete (Table 8). The margins 
of the tuberosities and distal portions are eroded 
and a small central section of the left shaft is 
missing. The large, globular head is smooth and 
connected by a short but prominent neck to the 
shaft, which is quite flat mediolaterally. The dis- 
tal end has two trochlear facets, which are 
slightly concave, for articulation with the radius 
and ulna. On the right humerus, the radial facet 
is at an angle of approximately 48° to the plane 
of the ulnar facet. Erosion has made it difficult to 
estimate the dimensions of the trochlea. 



C-3 



C-4 



C-5 



C-6 



C-7 



41 


35 


39 


40 


- 


105 


123 


155 


150 


150± 


157 


150 


- 


163 


160± 


38 


34 


Discussion 


41 


47 



Balaenopterid Taxonomy and the Problem of 
Disparate Parts 

Not all named fossil balaenopterids are based 
on directly comparable elements. Regarding 
mysticete taxonomy, Kellogg (1931) has stated 
that the classification of fossil taxa must be based 
upon cranial characters, an opinion that was 
shared by Barnes (1976). For our comparisons, 
we examined 14 fossil species of mysticetes in 
14 genera. Nine of these species are represented 
by parts of the cranium, and three of these also 
include parts of dentaries. The remaining five 
species are represented by only a single dentary 
or by parts of dentaries. In contrast, the holotype 
of P. baulinensis is represented by about 90% of 
the cranium and dentaries, along with both 
humeri and 23 vertebrae, making it one of the 
most complete fossil mysticete specimens yet 
described. Because of its completeness, it was 
possible to compare it to and distinguish it from 
all other relevant mysticete taxa. 

Other Described Fossil Balaenopterids 

Fossil balaenopterids are relatively abundant 
in late Cenozoic marine deposits around the 
world (Barnes 1976). In publications, some of 
these fossils have been assigned to living genera, 
but have not been more precisely identified. Most 
of these less precisely identified fossils are 
clearly referable to the subfamilies Balaenopter- 
inae and Megapterinae and are not directly rele- 
vant to the present study. It is, however, 
necessary to consider previously named 
balaenopterids in arguing that Parabalaenoptera 
baulinensis is a previously unrecognized taxon 
in anew subfamily. 

The extinct balaenopterids that were described 
by Van Beneden (e.g., 1872, 1880, 1882, 1885) 



130 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



-3 
C 



Q 

2 






It 



E 



o 



E 



° t: 

H > 



O 

&B E 
Up — 

- 3 J3 



-H 

o 



<4- <U 

I! 

op 5 
'5 - c 



■±3 i- P 

T3 O S 

^ ' C £ 



-J c 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENSIS 



131 



TABLE 7. Measurements (in mm) of 12th vertebrae of dorsal-lumbar of holotype of Parabalaenop- 
tera baulinenis, new genus and species. 

Distance from front of anterior zygapophysis to posteriormost margin of neural spine -190 

Distance from level of ventral margin anterior face of centrum to dorsal level of anterior 

zygapophysis 205 

Greatest dorsoventral diameter of neural canal at anterior dorsal margin of centrum 60 

Greatest transverse diameter of neural canal 55 

Greatest length of neural canal 100 

Greatest width of neural spine at dorsal base of anterior zygopophysis 27 

Length of base of right transverse process 115 

Height of base of right transverse process 34 

TABLE 8. Measurements (in mm) of humeri of holotype of Parabalaenoptera baulinensis, new 
genus and species. 

Right Left 

1. Greatest length 305 — 

2. Greatest width posterior margin of shaft to anterior 

margin of greater tuberosity — 135 

3. Anteroposterior diameter of distal end — -130 

4. Exterointernal diameter of distal end — 67 

5. Anteroposterior diameter of head 132 130 

6. Proximal-distal diameter of head 101 115 



from the Antwerp Basin in Belgium belong to the 
closely related balaenopterine genera Balaenop- 
tera and Burtinopsis, to the megapterine genus 
Megaptera, and to the primitive balaenopterid 
genus Plesiocetus Van Beneden, 1859. Most of 
the species that Van Beneden described are based 
on rather complete associated skeletons, but in 
most cases the specimens include only parts of 
the crania and mandibles. Included among the 
Belgian fossils are the extinct balaenopterines 
Balaenoptera musculoides Van Beneden, 1880; 
Balaenoptera borealina Van Beneden, 1880; 
and Balaenoptera rostratella Van Beneden, 
1 880; and the closely related species Burtinopsis 
similis Van Beneden, 1872, and Burtinopsis mi- 
nutus Van Beneden, 1880. Balaenoptera muscu- 
loides and B. borealina both have parallel dorsal 
and ventral margins of the horizontal ramus of 
the dentary, without the long ascending coronoid 
crest that characterizes Parabalaenoptera 
ban linens is, and in this regard they are like the 
living species of Balaenoptera. Likewise, B. ros- 
tratella has characters of the genus Balaenop- 
tera, and is approximately one-half the size of 
Parabalaenoptera baulinensis. Species of Burti- 



nopsis are similar to Balaenoptera, and in fact, 
Demere (1986) has considered Burtinopsis to be 
a junior synonym of Balaenoptera. 

Because the known morphologies of these fos- 
sil balaenopterids are consistent with those of the 
living species of Balaenoptera and Megaptera, 
we accept Van Beneden's original generic 
allocations for them and assign them to the sub- 
families Balaenopterinae and Megapterinae, re- 
spectively. Each of them can be distinguished 
from Parabalaenoptera by the characters that we 
use to diagnose the subfamily Parabalaenopteri- 
nae (Table 9). 

A species in the supposed balaenopterid genus 
Mizuhoptera Hatai, Hayasaka, and Masuda, 
1963, was described on the basis of isolated 
tympanic bullae of Pliocene age from Japan. 
Oishi and Hasegawa (1995) concluded that the 
type material actually represents the cetotheriid 
genus Herpetocetus Van Beneden, 1872. There- 
fore, the genus Mizuhoptera is invalid and is not 
a balaenopterid. 

A Late Miocene mysticete, Balaenoptera ry- 
ani Hanna and McLellan, 1924, from the Mon- 
terey Formation in central California, was named 



132 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



TABLE 9. Differences between Parabalaenopterinae, new subfamily, and the subfamilies of 
Megapterinae and Balaenopterinae. 



Parabalaenopterinae 



Megapterinae and 
Balaenopterinae 



CRANIUM 

1. Cranial vertex 

2. Nasals 

3. Shape of ascending process of maxilla 

4. Anterior portion of parietal 

5. Postglenoid process 

DENTARY 

6. Length of coronoid ridge anterio- 
posteriorly relative to mandible length 

7. Direction of terminus of coronoid process 

8. Terminus of coronoid process 

9. Mandibular foramen 



narrow 
long, narrow 
cuneate 
raised, swollen 
shallow 



long 

posterior 

bullet-shaped boss 
lacks notch 



broad 

short, broad 

flared 

flat, feathered 

deep and moderately deep 



short 
dorsal 

smoothly tapered 
notched 



on the basis of a braincase. It is not a member of 
the genus Balaenoptera, and is not even a 
balaenopterid. Instead, it is a typical cetotheriid 
(see Barnes, 1976:330) because its parietals are 
exposed in a wide band across the top of the 
intertemporal region, and it does not have the 
type of interfingering of the rostral and cranial 
bones that is diagnostic of the family Balaenop- 
teridae. 

Only two fossil species of balaenopterids have 
been previously named from the Pacific coast of 
North America. Megaptera miocaena Kellogg, 
1922, from the Late Miocene Monterey Forma- 
tion in Santa Barbara County, California, is a 
large species of balaenopterid (Barnes, 
1976:329). It has short and wide nasal bones and 
a wide intertemporal region like both Megaptera 
and Balaenoptera, but the concave lateral mar- 
gins of its occipital shield are like members of the 
subfamily Balaenopterinae. Its zygomatic arches 
are relatively narrow, a primitive character, and 
it appears to be a primitive species of the subfam- 
ily Balaenopterinae. 

Balaenoptera davidsonii Cope, 1872, is a 
small mysticete from the Late Pliocene age San 
Diego Formation at San Diego, California. It was 
first named as a species of Eschrichtius (gray 
whales), but Barnes (1976:333, table 6) removed 
it from the Eschrichtiidae, pointing out that the 
holotype dentary did not have characters of the 



family, and Demere (1986) placed it in the genus 
Balaenoptera. Thus, both previously named fos- 
sil balaenopterids from the eastern North Pacific 
realm are apparently members of the subfamily 
Balaenopterinae. 

The Relationships of Parabalaenoptera 
baulinensis 

Parabalaenoptera baulinensis is demonstra- 
bly a member of the family Balaenopteridae, 
which includes the living humpback {Megaptera 
novaeangliae (Borowski, 1781)), various species 
of rorquals of the genus Balaenoptera, and vari- 
ous extinct taxa as mentioned above. Para- 
balaenoptera baulinensis possesses six derived 
cranial characters and one derived mandibular 
character that are diagnostic of the family 
Balaenopteridae (see also Miller 1923; Kellogg 
1931). These derived characters, which define 
the family Balaenopteridae, are listed in the leg- 
end (Node 1) of the cladogram (Fig. 9), and are 
discussed as follows: 

1) The parietals are excluded from medial ex- 
posure on the dorsal surface of the intertemporal 
region, but they extend anteriorly along either 
side of the frontals at least to a point beyond the 
posterior ends of the rostral bones (the nasals, 
maxillae, and premaxillae). 

2) The apex of the occipital shield extends 
anteriorly at least as far as the level of the poste- 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENS1S 



133 



rior border of the supraorbital process and the 
anterior extremity of the zygomatic process of 
the squamosal. 

3) The frontal has only a small exposure on the 
middle of the dorsal surface of the interorbital 
region. 

4) The lateral part of the squamosal is rela- 
tively large and expanded, and the postglenoid 
process is large. 

5) The dorsal surface of the supraorbital proc- 
ess of the frontal is very depressed medially, 
descending abruptly from the more elevated dor- 
sal surface of the interorbital region. 

6) The maxillary part of the rostrum is trans- 
versely expanded and relatively flat, except in its 
median part, which has a keel that is created by 
the vomer, and is not conspicuously arched an- 
teroposteriorly. 

7) The dentary is conspicuously bowed later- 
ally, beyond the curvature of the rostral margin. 

Parabalaenoptera baulinensis has five primi- 
tive characters that exclude it from the derived 
subfamilies Balaenopterinae and Megapterinae, 
and these contribute to the definition of the new 
subfamily Parabalaenopterinae. These five char- 
acters are discussed as follows: 

1) The narrow intertemporal region of Para- 
balaenoptera baulinensis is a retained primitive 
character that is shared with members of the 
family Cetotheriidae and with the primitive 
balaenopterid Plesiocetus cortesii (Capellini, 
1865) (see discussion of Plesiocetus below). 
Members of the Balaenopterinae and Megapter- 
inae (see Kellogg 1928; Barnes and McLeod 
1984) have relatively wider intertemporal re- 
gions than P. baulinensis, in some cases wider by 
a factor of more than two. 

2) The dorsal exposure of the frontals on the 
middle of the dorsal surface of the intertemporal 
region between the posterior ends of the rostral 
bones and the apex of the occipital shield in 
Parabalaenoptera baulinensis is relatively large 
for a balaenopterid, forming an approximately 
rectangular patch. This larger exposure is a 
primitive character that is shared with members 
of the family Cetotheriidae, and contrasts with 
the much reduced and nearly absent dorsal expo- 
sure of the frontals in skulls of animals in the 
subfamilies Balaenopterinae and Megapterinae. 

3) The nasals of Parabalaenoptera baulinensis 
are longer (4: 1 length to width ratio) than those 



of any extant balaenopterid, extending anteriorly 
to a point anterior to the level of the antorbital 
notches. In members of the subfamilies 
Balaenopterinae and Megapterinae, however, the 
nasals are short and terminate posterior to the 
level of the antorbital notches. The nasals of P. 
baulinensis are nearly as long as those of such 
Late Oligocene and Early Miocene cetotheriids 
(sensu Kellogg 1931) as Mauicetus parki (Ben- 
ham, 1937) (nasal length to width ratio of 5.5:1) 
and Aglaocetus moreni (Lydekker, 1894) (nasal 
length to width ratio of 5: 1 ). Therefore, the long 
and narrow nasals of Parabalaenoptera 
baulinensis are a retained primitive character. 

4) The posteriorly tapering, narrow, ascending 
processes of the maxillae of P. baulinensis differ 
from the lobate posterior ends of the maxillae in 
the Balaenopterinae and Megapterinae. In P. 
baulinensis these processes become progres- 
sively more narrow posteriorly and terminate in 
rounded points. This condition we refer to as 
"cuneate," in order to distinguish it from the 
transversely flared or lobate shape of the ascend- 
ing processes, which was considered by Barnes 
and McLeod (1984) to be a balaenopterid char- 
acter. Henceforth, the posteriorly expanded as- 
cending processes of the maxillae shall be 
considered a derived character that is shared by 
the subfamilies Balaenopterinae and Megapteri- 
nae. The cuneate processes are a primitive char- 
acter of Parabalaenoptera baulinensis, which is 
shared with cetotheriids, and distinguishes the 
subfamily Parabalaenopterinae from the two 
other balaenopterid subfamilies. 

5) The short postglenoid process of Para- 
balaenoptera baulinensis is a primitive character 
that is shared with species of Cetotheriidae and 
primitive balaenopterids of the genus Plesio- 
cetus. It differs from the bulbous postglenoid 
process of the derived balaenopterids, and is a 
defining character of the subfamily Para- 
balaenopterinae. 

The derived character states of the above five 
structures (the opposite polarity of the character 
states in Parabalaenoptera baulinensis) are 
shared by the subfamilies Balaenopterinae and 
Megapterinae, and indicate that the latter two are 
sister taxa. They appear at Node 5 on the 
cladogram (Fig. 9). 

Parabalaenoptera baulinensis has three auta- 
pomorphies, which appear at Node 4 in the 



1 34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 

Cetotheriidae Plesiocetus Parabalaenoptera Balaenopterinae Megapterinae 

cortesii baulinensis 

.7 




Figure 9. Cladogram showing postulated relationships of Parabalaenoptera baulinensis, new genus and species, among 
fossil and Recent Balaenopteridae. The subfamily Cetotheriinae is the outgroup. Characters marking the numbered dichotomies 
and end points are as follows: 

1 . Anterior border of supraorbital process of frontal depressed relative to dorsal surface of maxilla and forming a coved recess; 
parietals excluded from dorsal surface of intertemporal region and extending anteriorly on lateral sides of the frontals in the 
intertemporal region to a point beyond the posterior ends of rostral bones; apex of occipital shield extended anteriorly at least 
as far as the level of the posterior border of the supraorbital process and the anterior extremity of the zygomatic process of the 
squamosal; lateral part of the squamosal relatively large and expanded; supraorbital process of the frontal abruptly depressed 
at its medial part to a level well below that of the dorsal surface of the interorbital region; dentary conspicuously bowed laterally 
[family Balaenopteridae]. 

2. Nasals short, terminating anteriorly at a point posterior to the level of the antorbital notch [Plesiocetus cortesii]. 

3. Contact between the posterior border of the maxilla and the anterior margin of the supraorbital process of the frontal curved 
anteriorly; postorbital process of frontal reduced; zygomatic process of squamosal inflated and curved medially (concomitant 
with shorter postorbital process); mandibular condyle spherical and bulbous. 

4. Parietal on either side of the intertemporal region swollen and elevated dorsal to the level of the adjacent frontal; coronoid 
crest of dentary elevated and inclined toward the coronoid process; apex of coronoid process posteriorly directed and 
spindle-shaped [Parabalaenoptera baulinensis]. 

5. Nasal bones short and terminating posterior to the antorbital notch; posterior end of the ascending process of the maxilla 
lobate or transversely expanded; intertemporal region expanded transversely; dorsal exposure of the frontals between posterior 
ends of the rostra! bones and apex of the occipital shield reduced; parietals on either side of intertemporal region thin and 
interfingering with frontals anteriorly; postglenoid process of the squamosal large, bulbous, and extending ventrally from the 
cranium. 

6. Concave lateral margin of the occipital shield where it overhangs the temporal fossa [subfamily Balaenopterinae]. 

7. Braincase transversely expanded; coronoid process of dentary reduced in size; olecranon process of ulna lost [subfamily 
Megapterinae]. 



cladogram (Fig. 9). These contribute to the defi- 
nition of the taxon, and are discussed as follows: 
1 ) In Parabalaenoptera baulinensis, as in all 
Balaenopteridae, the parietals are excluded from 
exposure in the dorsal surface of the intertempo- 
ral region, and they extend anteriorly along both 
sides of the frontals to a point as far as the 
posterior ends of the nasals. However, a unique 
derived character of P. baulinensis is the infla- 
tion and elevation of this lateral part of the parie- 
tals above the level of the frontals. Both the right 



and left parietals exhibit this feature to about the 
same degree, indicating that it is normal and not 
pathological. This contrasts with the flat, inter- 
fingering condition of the parietals in the sub- 
families Balaenopterinae and Megapterinae. 

2) The long, gradually ascending coronoid 
crest on the dentary that approaches the coronoid 
process (see Fig. 5) is a unique derived character 
of P. baulinensis. The structure is absent in all 
other Balaenopteridae, in which the coronoid 
process rises abruptly from the straight dorsal 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAUUNENSIS 



135 



other Balaenopteridae, in which the coronoid 
process rises abruptly from the straight dorsal 
margin of the horizontal ramus. 

3) The apex of the coronoid process is posteri- 
orly directed and spindle shaped. This is unlike 
the condition in Cetotheriidae and in the 
Balaenopterinae and Megapterinae. 

To summarize the points above, Para- 
balaenoptera baulinensis is a member of the 
family Balaenopteridae, because it has the suite 
of diagnostic osteological characters that are 
presently used to recognize the family. It also has 
some plesiomorphic characters that are not pre- 
sent in members of the two highly evolved 
balaenopterid subfamilies. Balaenopterinae and 
Megapterinae: narrow intertemporal region, 
larger exposure of frontal in intertemporal re- 
gion, long and narrow nasal bones, tapered as- 
cending processes of the maxillae, and short 
postglenoid process of the squamosal. These 
primitive characters are shared with members of 
the family Cetotheriidae, and they add to evi- 
dence for derivation of the Balaenopteridae from 
the Cetotheriidae. Parabalaenoptera baulinensis 
also has three unique derived characters that 
separate it from other Balaenopteridae: the ele- 
vated and swollen parietals on either side of the 
intertemporal region, the long, ascending 
coronoid crest anterior to the coronoid process on 
the dentary, and the spindle-shaped posteriorly- 
directed apex of the coronoid process. The com- 
bination of the five primitive characters and three 
unique derived characters of Parabalaenoptera 
baulinensis serve to define the taxon and the new 
subfamily Parabalaenopterinae that contains it. 

The Genus Plesiocetus Van Beneden, 1859 

The family assignment of the extinct mysticete 
genus Plesiocetus Van Beneden, 1859, has var- 
ied, and it has been assigned to both the Cetoth- 
eriidae and the Balaenopteridae. At least 13 
species of Late Miocene and Pliocene age from 
throughout the world have been assigned to the 
genus. While it is beyond the scope of the present 
study to resolve the issues affecting the compli- 
cated typology and systematics of Plesiocetus 
and the species that have been assigned to it, it is 
useful to mention some of the problems that 
surround it (see also Cabrera 1926; Kellogg 
1968:103-104). 

Both Winge ( 1 9 1 0) and True (1912), who pre- 
sented a generic diagnosis, considered Plesio- 



cetus to be a valid balaenopterid genus, but Kel- 
logg (1968:104) wrote that "Sufficient diagnos- 
tic criteria for the recognition of Plesiocetus as a 
valid genus are not presently known." Brandt 
(1872:407) assigned the genus Plesiocetus to 
what he regarded as the balaenopterid subfamily 
Cetotheriinae, the group now recognized as the 
family Cetotheriidae. Van Beneden (1872:242) 
fixed the type species as Plesiocetus garopii Van 
Beneden, 1859, and considered the genus to be 
related to balaenopterines. Kellogg (1925:51), 
however, noting that in 1 885 Van Beneden omit- 
ted P. garopii from a listing of species in the 
genus Plesiocetus, fixed the type species as Ple- 
siocetus hupschii Van Beneden, 1859, and dis- 
cussed it in the context of the family 
Cetotheriidae. Later (1934:79) Kellogg, re- 
stricted the genus name Plesiocetus to Plesio- 
cetus garopii (Van Beneden's type species), and 
considered it to be a balaenopterine. 

Comparisons with Plesiocetus cortesii 

Regardless of the status and systematics of the 
genus Plesiocetus, one species that has been re- 
ferred to the genus is of direct relevance to the 
present study because it helps to establish the 
polarity of the characters of Parabalaenoptera 
baulinensis and its taxonomic position. This is 
the species Plesiocetus cortesii, from Pliocene 
deposits in Italy, the skull and dentary of which 
were illustrated by Van Beneden (1875). 

The skull of Plesiocetus cortesii (see Fig. 8a) 
resembles that of Parabalaenoptera baulinensis 
(see Fig. 8b) in its general proportions, but it 
differs in some details. 

Compared to Parabalaenoptera baulinensis, 
P. cortesii (Fig. 8a) has the following more 
primitive character states: 

1 ) The contact between the posterior border of 
the maxilla and the anterior margin of the su- 
praorbital process of the frontal is transversely 
straight rather than curved anteriorly. 

2) The postorbital processes of the frontals are 
larger and project posterolaterally. 

3) The occipital shield is smaller and has more 
nearly the shape of an equilateral triangle. 

4) The zygomatic processes are narrower and 
straighter. 

5) The mandibular condyle is smaller and not 
so spherical. 

The derived character states of the above five 
structures (the opposite polarity of the character 



136 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



states in Plesiocetus cortesii) are shared by Para- 
balaenoptera baulinensis and the subfamilies 
Balaenopterinae and Megapterinae, and indicate 
that the latter three are sister taxa (Node 3 on the 
cladogram, Fig. 9). 

Compared to Parabalaenoptera baulinensis, 
P. cortesii has nasal bones that are much shorter, 
terminating at a point posterior to the level of the 
antorbital notches, and this is a derived character 
that defines P. cortesii and is listed at Node 2 on 
the cladogram (Fig. 9). The nasal bones are simi- 
larly short in Recent Balaenopterinae and 
Megapterinae, but the nasals of P. cortesii differ 
by being narrower and not expanded anteriorly. 
Because of the numerous other primitive charac- 
ters of P. cortesii, we conclude that short nasal 
bones evolved convergently in P. cortesii and in 
both of the derived subfamilies Balaenopterinae 
and Megapterinae. 

Plesiomorphies that Plesiocetus cortesii shares 
with Parabalaenoptera baulinensis include the 
narrow posterior terminations of the maxillae, 
the narrow anterior ends of the nasals, and the 
narrow intertemporal region. Plesiocetus cortesii 
is a primitive balaenopterid, and in most of its 
characters is more primitive than Parabalaenop- 
tera baulinensis. 

Conclusions 

1 . Parabalaenoptera baulinensis, a new genus 
and species of extinct whale in the family 
Balaenopteridae, is based on a partial fossil 
skeleton collected from the sea cliff at Bolinas 
Point, Point Reyes Peninsula, in Marin County, 
California. 

2. The holotype specimen is from the Late 
Miocene age Santa Cruz Mudstone, and analysis 
of diatoms from the rock unit indicates an age of 
between approximately 6.8 and 6.0 Ma. 

3. The holotype skeleton includes the skull, 
both dentaries, cervical, thoracic, and one lumbar 
vertebrae, pectoral limb bones, and ribs, making 
this one of the most complete fossil balaenopterid 
specimens recorded in the published literature. 
The animal is estimated to have had a total body 
length of approximately 10 m in life. 

4. Parabalaenoptera baulinensis is a relatively 
primitive balaenopterid, and it lacks synapomor- 
phies of the derived balaenopterids of the sub- 
families Balaenopterinae and Megapterinae. It 
has a suite of autapomorphies (conspicuously 



elevated or swollen anterior portions of the pa- 
rietals on either side of the intertemporal region, 
long and ascending coronoid crest on the dentary 
approaching the coronoid process, and posteri- 
orly-directed apex of the coronoid process), and 
we classify it in the new subfamily Parabalaenop- 
terinae. 

5. Parabalaenoptera baulinensis shares sym- 
plesiomorphies with the primitive family Cetoth- 
eriidae (narrow intertemporal region, larger 
exposure of frontals on the dorsal surface of 
intertemporal region, long nasals, long and pos- 
teriorly tapering ascending processes of the max- 
illae, and short postglenoid processes of the 
squamosals), and these shared primitive charac- 
ters lend support to the theory that the Balaenop- 
teridae evolved from the Cetotheriidae. 

6. Species of the genus Plesiocetus Van 
Beneden, 1859, pose many problems of classifi- 
cation and relationships. One species in the ge- 
nus, Plesiocetus cortesii (Capellini, 1865), is 
possibly a sister taxon of a less inclusive clade 
containing Parabalaenoptera baulinensis, the 
subfamily Balaenopterinae, and the subfamily 
Megapterinae. 

7. Because other, more highly derived taxa of 
Balaenopteridae are contemporaneous with P. 
baulinensis, it may be considered to be a relict 
descendant of a lineage that was ancestral to both 
the subfamilies Megapterinae and Balaenopteri- 
nae, and thus to the extant Megaptera and 
Balaenoptera. The origin of Parabalaenoptera 
baulinensis must lie among pre-Late Miocene 
mysticetes that are closer in morphology to 
primitive members of the family Cetotheriidae. 

ACKNOWLEDGMENTS 

The third author regrets to report the deaths of 
the first two authors, and dedicates this work to 
the memories of Carl V. Zeigler, who discovered 
the holotype of Parabalaenoptera baulinensis 
and was the foundation of the College of Marin 
whale project, and of Gordon L. Chan, who had 
a love of the paleontology, geology, and natural 
history of Marin County and supported the whale 
project. 

We thank the many College of Marin students 
and laboratory assistants who helped excavate 
and assemble the holotype of Parabalaenoptera 
baulinensis. We especially acknowledge Ron 
Melander, who spent many hours working on the 



ZEIGLER, CHAN, AND BARNES: PARABALAENOPTERA BAULINENSIS 



137 



holotype and preparing the illustrations, and 
David Ferguson, the whale project coordinator at 
College of Marin. 

We acknowledge consultants William N. Orr 
of the Geology Department at the University of 
Oregon; John A. Barron and Ellen J. Moore of 
the U. S. Geological Survey in Menlo Park, Cali- 
fornia; and Terrence Gosliner and Peter U. 
Rodda of the California Academy of Sciences. 
Marjorie A. Chan, Professor of Geology at the 
University of Utah, shepherded this manuscript 
to completion to fulfill her father's wishes. Ac- 
cordingly, Dr. Chan took care of correspondence 
related to this publication, helped edit the text, 
and prepared the illustrations. 

Finally, we thank the agencies that have 
funded our work: Marin Community College 
District, Chancellor's Office of the State of Cali- 
fornia Community Colleges, The Marin Founda- 
tion, and a host of private individual contributors. 
Zeigler and Chan initiated the manuscript and 
Barnes completed and modified it for publica- 
tion. 

Literature Cited 

Barnes, L. G. 1976. Outline of eastern North Pacific 
fossil cetacean assemblages. Syst. Zool. 
25:321-343. 

Barnes, L. G., D. P. Domning and C. E. Ray. 1985. 
Status of studies on fossil marine mammals. Mar. 
Mamm. Sci. 1(1): 15-53. 

Barnes, L. G. and S. A. McLeod. 1984. The fossil 
record and phyletic relationships of gray whales. 
Chapter 1 . Pp. 3-32 in The gray whale: Eschrichtius 
robustus. M. L. Jones, S. L. Swartz, and S. Leather- 
wood, eds. Academic Press, Orlando, Florida. 

Benham, W. B. 1 937. Fossil Cetacea of New Zealand. 
II — On Lophocephalus, a new genus of zeuglo- 
dont Cetacea. Trans. Roy. Soc. New Zealand 
67:1-7, pis. 1-7. 

Brandt, J. F. 1872. Bericht iiber den bereits vollen- 
deten druckfertigen Theil seiner Untersuchungen 
iiber die fossilen und subfossilen Cetaceen 
Europa's. Bull. Acad. Sci. St. Petersburg, Ser. 3, 
17:401-408. 

Cabrera, A. 1926. Cetaceos fosiles del Museo de la 
Plata. Rev. Mus. La Plata 29:363^11 1 . 



Capellini, G. 1 865. Balenottere fossili del Bolognese. 
Mem. Accad. Sci. Bologna, Ser. 2, 4:3 1 5-336, 3 pis. 

Clark, J. C, E. E. Brabb, H. G. Greene and D. C 
ROSS. 1984. Geology of the Point Reyes Peninsula 
and implications for San Gregorio fault history. 
Tectonics and sedimentation along the California 
margin. Soc. Econ. Paleontol. Mineral., Pacific 
Sect. 38:67-86. 

Cope, E. D. 1872. On an extinct whale from Califor- 
nia. Proc. Acad. Natur. Sci. Philadelphia 1 872 (part 
l):29-30. 

Demere, T. A. 1986. The fossil whale, Balaenoptera 
davidsonii (Cope, 1 872), with a review of other 
Neogene species of Balaenoptera (Cetacea: Mys- 
ticeti). Marine Mammal Science 2(4):277-298. 

Domning, D. P. 1978. Sirenian evolution in the North 
Pacific Ocean. Univ. Calif. Publ. Geol. Sci. 1 8:i-xi, 
1-176, pis. 1-18. 

DU Bus, B. A. L. 1867. Sur quelques mammiferes du 
Cragd'Anvers. Bull. Acad. Roy. Sci. Belgique, Ser. 

2(24):562-577. 

FORDYCE, R. E. AND L. G. Barnes. 1994. The evolu- 
tionary history of whales and dolphins. Pp. 4 1 9-455 
in 1994 Annual Review of Earth and Planetary 
Sciences, Volume 22, G. W. Wetherill, ed. Annual 
Reviews, Inc., Palo Alto, California. 

Galloway, A. J. 1977. Geology of the Point Reyes 
Peninsula, Marin County, California. Calif. Div. 
Mines Geol. Bull. 202:i-v, 1-72, pi. 1 (map). 

Hanna, G D. and M. E. McLellan. 1924. A new 
species of whale from the type locality of the Mon- 
terey Group. Proc. Calif. Acad. Sci., 4th Series, 
13(14):23 7-241, pis. 5-9. 

Hatai, K., S. Hayasaka and K. Masuda. 1963. 
Some fossil tympanies from the Mizuho period of 
northern Japan. Saito Ho-on Kai Mus. Natur. Hist. 
Res. Bull. 32:5-17. 

Howell, A. B. 1930. Aquatic mammals. Their adap- 
tations to life in the water. Charles C Thomas, 
Springfield, Illinois, and Baltimore, Maryland. 

Kellogg, A. R. 1922. Description of the skull of 
Megaptera miocaena, a fossil humpback whale 
from the Miocene diatomaceous earth of Lompoc, 
California. Proc. U. S. Natl. Mus. 61 (14): 1-1 8, pis. 
1-4. 



138 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 4 



. 1925. Additions to the Tertiary history of 

pelagic mammals on the Pacific coast of North 
America. II. Fossil cetotheres from California. 
Carnegie Inst. Washington Publ. 348(2):35-56. 



. 1928. The history of whales — their adapta- 
tion to life in the water. Quart. Rev. Biol. 
3(l,2):29-76, 174-208. 

. 1931. Pelagic mammals from the Temblor 



Formation of the Kern River region, California. 
Proc. Calif. Acad. Sci., 4th Series, 1 9( 1 2):2 1 7-397. 

. 1934. The Patagonian fossil whalebone 



whale, Cetothehum moreni (Lydekker). Carnegie 
Inst. Washington Publ. 447:63-81, pis. 1-4. 

. 1968. Fossil marine mammals from the Mio- 



cene Calvert formation of Maryland and Virginia. 
Part 5. Miocene Calvert mysticetes described by 
Cope. U. S. Nat. Mus. Bull. 247:103-132. 



Omura, H. 1959. Bryde's whale from the coast of 
Japan. Sci. Rep. Whales Res. Inst. 14:1-33. 

Repenning, C. A. and R. H. Tedford. 1 977. Otarioid 
seals of the Neogene. U. S. Geol. Surv. Prof Paper 
992:i-vi, 1-93, pis. 1-24. 

True, F. W. 1 904. The whalebone whales of the west- 
ern North Atlantic. Smithsonian Contrib. Know!. 
33:1-332. 

. 1912. The genera of fossil whalebone whales 



allied to Balaenoptera. Smithsonian Misc. Coll. 
59(6): 1-8. 

Van Beneden, P. J. 1859. Ossements fossiles decou- 
verts a Saint Nicolas en 1 859. Bull. Acad. Roy. Sci. 
Belgique, Sen 2, 8(1 1): 123- 146. 

. 1872. Les baleines fossiles d'Anvers. Bull. 



Acad. Roy. Sci. Belgique, Ser. 2, 34(7):6-20. 



Lydekker, R. 1 894. Contributions to a knowledge of 
the fossil vertebrates of Argentina. 2. Cetacean 
skulls from Patagonia. An. Mus. La Plata, Sec. 
Paleontol. (2):1-13, 6 plates. 

Miller, G. S. 1923. The telescoping of the cetacean 
skull. Smithsonian Misc. Coll. 75(5): 1-71, pis. 1-8. 



. 1875. Le squelette de la baleine fossile du 

Musee de Milan. Bull. Acad. Roy. Sci. Belgique, 
Ser. 2, 40(12):736-758, figs. 1-2. 

. 1 880. Les mysticetes a courts fanons des 



sables des environs d' Anvers. Bull. Acad. Roy. Sci. 
Belgique, Ser. 2, 50(7): 11 -27. 



. 1924a. Some hitherto unpublished photo- 
graphs and measurements of the blue whale. Proc. 
U.S. Natl. Mus. 66(7): 1-4. 

. 1924b. A pollack whale from Florida pre- 



sented to the National Museum by the Miami 
Aquarium Association. Proc. U. S. Natl. Mus. 
66(9): 1-1 5. 

Oishi. M. and Y. Hasegawa. 1995. Diversity of 
Pliocene mysticetes from eastern Japan. The Island 
Arc [Thematic Issue: Evolution and Biogeography 
of Fossil Marine Vertebrates in the Pacific Realm, 
Proceedings of the 29th International Geological 
Congress, Kyoto, Japan, edited by L. G. Barnes, Y. 
Hasegawa, and N. Inuzuka] 3(4):436-452 [for 
1994]. 



. 1 882. Description des ossements fossiles des 

environs d'Anvers, Troisieme Partie, Cetaces. Gen- 
res: Megaptera, Balaenoptera, Burtinopsis & Erpe- 
tocetus. Ann. Mus. Roy. d'Hist. Nat. Belgique, Ser. 
Paleontologique 7:1-90, pis. 40-109. 

. 1 885. Description des ossements fossiles des 

environs d'Anvers, Quatrieme Partie, Cetaces. 
Genre: Plesiocetus. Ann. Mus. Roy. d'Hist. Nat. 
Belgique, Ser. Paleontologique 9:1^0, pis. 1-30. 

Wilson, L. E. 1937. New species of dolphin from 
Point Reyes, California. Abstract, Proc. Geol. Soc. 
Amer. 1936:387. 

WlNGE, H. 1910. Om Plesiocetus og Sqvalodon [sic] 
fra Danmark. Vidensk. Meddel. Naturhist. Foren., 
Copenhagen, 7(1): 1-38, pis. 1-2. 



© CALIFORNIA ACADEMY OF SCIENCES, 1997 

Golden Gate Park 

San Francisco, California 941 18 



PROCEEDINGS 

OF THE '1998 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 5, pp. 139-165, 15 figs. February 11, 1998 



FIVE NEW SPECIES OF CHROMODORIS (MOLLUSCA: 

NUDIBRANCHIA: CHROMODORIDIDAE) FROM THE 

TROPICAL INDO-PACIFIC OCEAN 

By 

Terrence M. Gosliner 

and 

David W. Behrens 

Department of Invertebrate Zoology; California Academy of Sciences 
Golden Gate Park, San Francisco, California 941 18 



Five new species of Chromodoris are described from the tropical Indo-Pacific. Three of these, 
Chromodoris josfti, C. dianae and C. michaeli, are members of the Chromodoris quadricolor 
complex. Chromodoris joshi is known from the Philippines, Indonesia and the Andaman Sea, 
Thailand. Chromodoris dianae is commonly found in the Philippines, Indonesia and Malaysia. 
Chromodoris michaeli is known from the Philippine Islands of Luzon, Cebu and Mindanao. 
Chromodoris hintuanensis is known from the Ryukyu Islands, the Philippines, Indonesia, Papua 
New Guinea and Thailand. It is compared to two similar species, C geometrica and C. conchyliata. 
Chromodoris robot is known from the Ryukyu Islands and Lord Howe Island and Western 
Australia and is similar in color pattern to C. vibrata and C. aureopurpurea. Consistent patterns 
of radular morphology, mantle gland arrangement and reproductive anatomy suggest that 
members of the Chromodoris quadricolor complex may be closely related phylogenetically in 
addition to having a similar color pattern. 

Received September 16, 1997. Accepted December 5, 1997. 

Numerous species of the highly diverse nudi- his 1977 and 1982 works, he has described sev- 
branch genus Chromodoris have been recently eral species with black lines which he included 
described from many localities in the tropical in the Chromodoris quadricolor complex. Three 
Indo-Pacific (Rudman 1977, 1982, 1983, 1984, of the new species described here are members 
1986, 1987; Yonow 1994; Gosliner 1994). Our of this complex. The other two are similar to 
recent field collections in the Philippines as well members of two other color groups of chromo- 
as collections and observations by others from dorids. 
Indonesia, Papua New Guinea, Okinawa, Malay- 
sia and Thailand have yielded specimens of sev- SPECIES DESCRIPTIONS 
eral new species of Chromodoris. This paper 

describes five of these new species and provides Chromodoris joshi sp. nov. 

comparison with other species that are similar in (Figs. 1 A, 2A-D, 3A-D, 4A) 
appearance. 

In several of Rudman's papers he has dealt Chromodoris sp. Debelius, 1996:213, top photo- 

with different color groups of chromodorids. In graph. 

[139] 



140 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 



Chromodohs quadricolor Dakin, 1992:346, bottom 
photograph, not (Ruppell and Leuckart, 1828), 
misidentification. 

Type Material. — Holotype: CASIZ 083806, one 
specimen, Kirby's Rock, Caban Island, Maricaban 
Strait, Philippine Islands, 24 February 1992, Jerry 
Allen. Paratypes: CASIZ 083674, two specimens, Se- 
pok Point, SW side Maricaban Island, Philippine Is- 
lands, 15 m depth, 19 February 1992, T. M. Gosliner. 
CASIZ 096328, two specimens, Sepok Point, SW side 
Maricaban Island, Philippine Islands, 20 m depth, 17 
March 1994, T. M. Gosliner and M. Miller. 
CASIZ 105711, one specimen, Anilao, Batangas 
Province, Luzon, Philippine Islands, M. Miller. 
CASIZ 103750, two specimens, one dissected, Pinna- 
cle Rock, Hamilo Bay, Luzon, Philippine Islands, 10 
m depth, 4 March 1995, Michael T. Ghiselin. CASIZ 
1 0564 1 , two specimens, Escarcia Point, Sabang, Min- 
doro, Philippine Islands, 14 m depth, 1 March 1995, 
T. M. Gosliner. CASIZ 085974, one specimen, Cathe- 
dral Rocks, Balayan Bay, Luzon, Philippine Islands, 
20 m depth, 25 March 1993, T. M. Gosliner. CASIZ 
1 10468, one specimen, buccal mass removed, Philip- 
pine Islands. 

ETYMOLOGY. — Chromodorisjoshi is named 
for the first author's son, Joshua Todd Gosliner, 
a bright enthusiastic student, who has had to put 
up with several missed birthdays while his father 
was conducting field work in the Philippines. 

Distribution. — Thus far, this species is 
known only from the Philippine Islands (present 
study), from a photograph by Jim Black from 
Sumatra, Indonesia (Debelius 1996), and from 
photographs by Mark Strickland of a specimen 
found in the Andaman Sea, Thailand. 

Natural History. — Chwmodoris joshi is 
found on the outer edges of rock walls and reef 
fronts in 5-30 meters of water. 

External Morphology. — Preserved ani- 
mals are 22-60 mm in length. The living animals 
(Fig. 1 A) are buttery yellow with a darker, golden 
yellow margin. White flecks are scattered 
densely within the yellow pigmented areas. A 
wide, black, submarginal band encircles the dor- 
sal surface of the animal, from the anterior end 
of the notum to a short distance behind the gill 



plume. The band is somewhat variable in width 
between specimens and in one specimen the band 
is much wider at the middle of the animal creat- 
ing an hourglass effect. The band may also ap- 
pear as two narrower bands along the sides of the 
body in some animals where the black is broken 
with a narrow gray area. A black stripe of about 
the same width begins anteriorly at or just behind 
the submarginal band and runs longitudinally 
between the rhinophores down the midline of the 
animal (Figs. 1 A and 2 A). The stripe may extend 
to near the base of the gill pocket but in one 
animal the stripe terminates before the midline of 
the body. The stripe may be discontinuous in 
places. The black color fades to gray along the 
edges of the bands. In all specimens seen, gills 
and rhinophores are uniformly pumpkin yellow- 
orange. The gills of the specimen figured by 
Debelius (1966:213) are orange basally, while 
the upper two-thirds are white. The triangular 
foot is the same color as the body and extends 
posteriorly beyond the notum. Two black stripes 
run along either side of the dorsal surface of the 
foot. The mantle glands (Fig. 2B, 4A) are con- 
tinuous around the margin of the notum except 
for the anterior portion of the animal from just 
behind the rhinophores where they are absent. 
The glands consist of spherical lobes that are 
clustered together closely resembling bunches of 
grapes (Fig. 2C). On either side of the head is a 
short, digitiform tentacle. There are 11-15 unip- 
innate gills forming the branchial plume. The 
perfoliate rhinophores bear about 30 lamellae. 

Buccal Armature. — The muscular portion 
of the buccal mass is approximately equal in 
length to the oral tube. At the anterior end of the 
muscular portion of the buccal mass is the chiti- 
nous labial cuticle, which bears numerous jaw 
rodlets. The rodlets (Fig. 3A) are elongate, 
curved and mostly undivided at the apex while a 
few are bifid in one specimen (CASIZ 1 10468). 
The radular formula is 8 1 x 96.0.96 in one speci- 
men examined (CASIZ 103750) and 103 * 
90.0.90 in a second specimen (CASIZ 1 10468). 
There is no trace of a rachidian tooth. The inner- 



-> 

Figure 1. Living animals. A. Chromodohs joshisy. nov., specimen from Sepok Point, NW side Maricaban Island, Philippine 
Islands, photograph by T. M. Gosliner. B. Chromodohs dianae sp. nov., specimen from Batangas, Philippine Islands, photograph 
by Mike Miller. C. Chromodohs dianae sp. nov., specimen from Kirby's Rock, Caban Island, Maricaban Strait, Philippine 
Islands, photograph by T. M. Gosliner. 



GOSLINER AND BEHRENS: CHROMODORIS 



14] 




142 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 




Figure 2. Chromodoris joshi sp. nov. A. Living animal. B. Subcutaneous glandular network. C. Mantle glands, enlarged, 
scale = 1 .0 mm. D. Reproductive system, am = ampulla, be = bursa copulatrix, ej = ejaculatory duct, fgm = female gland mass, 
p = penis, pr = prostate, rs = receptaculum seminis, vg = vestibular gland, scale = 1 .0 mm. 



most lateral teeth (Fig. 3B) are arched with a 
short cusp. There are 0-^4 poorly-developed den- 
ticles on the inner side of the tooth. The outer side 
lacks denticles. The second through fourth lateral 
teeth entirely lack denticles. The lateral teeth 
from the middle of the radular row (Fig. 3C) bear 
1-5 minute, triangular denticles on the outer side 
of the teeth. The outermost lateral teeth (Fig. 3D) 
are elongate with one to four denticles. 

Reproductive System.— (Fig. 2D) The 
ampulla is thick and tubular, narrowing some- 
what before bifurcating into an oviduct and vas 



deferens. The short oviduct enters the female 
gland mass near the albumen gland. The proxi- 
mal prostatic portion of the vas deferens is folded 
over itself once before it narrows slightly into the 
relatively short, muscular, ejaculatory portion. 
The ejaculatory portion narrows abruptly to a 
curved segment and then enters the short penial 
bulb, which is adjacent to the slender vaginal 
duct at the common gonopore. The distal end of 
the vas deferens is devoid of any penial hooks. 
The female gland mass consists of the large mu- 
cous gland and smaller membrane and albumen 



GOSLINER AND BEHRENS: CHROMODORIS 



143 




Figure 3. Chromodoris joshi sp. nov. Scanning electron micrographs. A. Jaw rodlets, scale = 1 5 urn. B. Inner lateral teeth, 
scale = 15 urn. C. Lateral teeth from central portion of half-row, scale = 25 urn. D. Outer lateral teeth, scale = 43 urn. 



144 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 













Figure 4. Mantle gland distribution. A. Chromodoris joshi sp. nov. B. C. qfricana Eliot, 1904. C. C. magnified (Quoy and 
Gaimard, 1832). D. C. strigata Rudman, 1982. E. C. dianae sp. nov. F. C. lochi Rudman, 1982. G. C. wV/an/ Rudman, 1982. 
H. C. elisabethina Bergh, 1877. I. C. michaeli sp. nov. J. C. a««a<? Bergh, 1877. K. C hamiltoni Rudman, 1977. 



GOSLINER AND BEHRENS: CHROMODOR1S 



145 



glands. Near the exit of the mucous gland is a 
small, ovoid vestibular gland. The vagina is rela- 
tively short. At the proximal end of the vagina is 
a short duct to the club shaped receptaculum 
seminis and a long, wide duct to the thin walled, 
spherical bursa copulatrix. The uterine duct 
emerges at a position one third of the length along 
the duct leading to the bursa and enters the female 
gland mass near the albumen gland. 

DISCUSSION. — Chromodoris joshi is most 
similar in external appearance to other members 
of the Chromodoris quadricolor complex that 
have prominent black mantle coloration, namely 
C. africana Eliot, 1904, C. magnified (Quoy and 
Gaimard, 1832), and C. kuiteri Rudman, 1982. 
However, C. joshi lacks an orange marginal man- 
tle band and has yellow rather than white pigment 
in addition to the black. Among these four spe- 
cies, the mantle glands are similar to those ob- 
served in C. africana and C. magnifica (Figs. 4B, 
C, respectively). The glands of C. joshi and C. 
africana are closely crowded in a continuous 
array, while in C. magnifica groups of glands are 
somewhat isolated, especially in the posterior 
region of the mantle. The mantle glands are not 
described for C. kuiteri. 

There are significant differences in the buccal 
armature among the four darkly-pigmented spe- 
cies. In C. joshi, a majority of the jaw rodlets are 
unifid, while in C. africana, C. kuiteri and C. 
magnifica the rodlets are bifid (Rudman 1977, 
fig. 18C; Rudman 1982).Thedenticulationofthe 
radular teeth varies between species. In C. kuiteri 
and C. magnifica all of the radular teeth have 
denticles on their outer side and a vestigial, tri- 
angular rachidian tooth is present. In C. joshi and 
C. africana there is no vestige of a rachidian and 
the inner lateral teeth lack denticles on the outer 
side. In C. joshi the next three teeth lack denti- 
cles. From tooth five to the outer edge of the 
radula minute denticles are present on the outer 
side of the tooth. In C. africana, the first 30 teeth 
lack denticles on the outer side while the remain- 
ing teeth have a few small denticles. 

The reproductive systems of C. magnifica and 
C. africana were described and illustrated by 
Rudman (1977, figs.l7A, C). In both of these 
species the vagina and the ejaculatory portion of 
the vas deferens are much shorter than in C. joshi. 
In C. joshi the ejaculatory segment consists of a 
wide proximal portion and a much thinner distal 
one, while in the above two species it appears to 



be of a uniform diameter throughout its length. 
The arrangement of reproductive organs of C. 
joshi is very similar to that described for C. 
kuiteri (Rudman 1984:137, fig. 13B). 

Chromodoris dianae sp. nov. 
(Figs. IB, C; 4E; 5A-D; 6; 7A-D) 

Casella sp. Wong, 1991:103. 

Chromodoris sp. Allen and Steene, 1994:198, photo- 
graph in second row, first column. 

Chromodoris sp. Colin and Arneson, 1995:183, figs. 
852 and 856. 

Type Material. — Holotype: CASIZ 106464, one 
specimen, Arthur's Rock, Calumpan Peninsula, 
Luzon, Philippine Islands, 1 7 April 1 996, Jerry Allen. 
Paratypes: CASIZ 083686, three specimens, one dis- 
sected, Kirby's Rock, Maricaban Island, Philippine 
Islands, 23 m depth, 18 February 1992, T. M. 
Gosliner. CASIZ 103789, two specimens, Cathedral 
Rock, Batangas Province, Luzon, Philippine Islands, 
22 m depth, 25 February 1 995, T. M. Gosliner. CASIZ 
086002, three specimens, N end of Ligpo Island, 
Balayan Bay, Luzon, Philippine Islands, 26 m depth, 
24 March 1993, T. M. Gosliner. CASIZ 084276, eight 
specimens, N end of Ligpo Island, Balayan Bay, 
Luzon, Philippine Islands, 21 m depth, 25 February 
1992, T. M. Gosliner. CASIZ 076326, one specimen, 
SW corner of north reef, Tubbataha Reefs, Sulu Sea, 
Philippine Islands, 9°09'N, 1 19°55'E, 8-16 m depth, 
13 March 1984, Antonio J. Ferreira. CASIZ 096278, 
three specimens, Kirby's Rock, Caban Island, Mari- 
caban Strait, Philippine Islands, 1 8 m depth, 1 7 March 
1994, T. M. Gosliner. CASIZ 084292, one specimen, 
Bus Stop Reef, Balayan Bay, Luzon, Philippine Is- 
lands, 23 m depth, 21 February 1992, T. M. Gosliner. 
CASIZ 087270, one specimen, Manado, Celebes Sea, 
Sulawesi, Indonesia, April 1988, Pauline Fiene- 
Severns. CASIZ 096266, three specimens, Kirby's 
Rock, Caban Island, Maricaban Strait, Philippine Is- 
lands, 25 m depth, 13 March 1994, T. M. Gosliner. 
CASIZ 076308, three specimens, Jessie Beazley Reef, 
Tubbataha Reefs, Sulu Sea, Philippine Islands, 2-77 
m depth, 9° 02 'N, 119°49'E, 15 March 1984, Antonio 
J. Ferreira. CASIZ 076288, three specimens, Filo- 
mona Shoals, Coyos Island, Sulu Sea, Philippine Is- 
lands, 10°40'N, 120°46'E, 25-33 m depth, 10 March 
1984, Antonio J. Ferreira. CASIZ 076374, one speci- 
men, Ilin Point, Ilin Island, off Mindoro, Philippine 
Islands, 16-23 m depth, 17 March 1984, Antonio J. 
Ferreira. CASIZ 110465, one specimen, Kirby's 
Rock, Caban Island, Maricaban Strait, Philippine Is- 
lands, 25 m depth, 19 April 1997, D. W. Behrens. 
CASIZ 1 10466, one specimen, buccal mass removed, 
Santa Rosa Island, Cebu, Philippine Islands, 20 m 
depth, 27 April 1997, Nora Ross. CASIZ 1 10467, 



146 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 




Figure 5. Chromodoris dianae sp. nov. A. Living animals. B. Subcutaneous glandular network. C. Mantle glands, enlarged, 
scale = 1 .0 mm. D. Reproductive system, am = ampulla, be = bursa copulatrix, ej = ejaculatory duct, fgm = female gland mass, 
p = penis, pr = prostate, rs = receptaculum seminis, vg = vestibular gland, scale = 1 .0 mm. 



GOSLINER AND BEHRENS: CHROMODORIS 



147 














Figure 6. Chromodoris dianae sp. nov. Variation in color pattern. 



148 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 



one specimen, Santa Rosa Island, Cebu, Philippine 
Islands, 30 m depth, 26 April 1997, D. W. Behrens. 

ETYMOLOGY. — Chromodoris dianae is 
named for the second author's wife, Diana Lynn 
Behrens, for her unselfish support and patience 
with this author during his studies of this group 
of animals. 

DISTRIBUTION. — Thus far, this species is 
known only from the Philippines and Indonesia, 
and from photographs by Marc Chamberlain 
from Sipadan Island, Borneo, Malaysia. 

NATURAL HISTORY. — Chromodoris dianae 
has commonly been found on relatively shallow 
walls and reef edges near drop-offs in 10-30 
meters depth. 

External Morphology. — The living ani- 
mals (Figs. IB, C) are up to 60 mm in length. The 
body appears powdery blue due to a fine encrust- 
ing of white over blue ground color. Around the 
margin of the notum is a line of fine, opaque 
white spots, which are continuous in most speci- 
mens examined. A few specimens have a series 
of orange spots along the margin of the notum, 
within the boundaries of the white line. Adjacent 
to the white margin is an area of speckled pow- 
dery blue. A discontinuous black submarginal 
band encircles the medial area of the notum from 
in front of the rhinophores to behind the gill 
circle. Black oval spots or patches may occur 
down the midline of the notum. Patterns of the 
submarginal band and the spots are highly vari- 
able (Figs. 5A, 6). The submarginal band may be 
nearly entire except for the areas adjacent and 
lateral to the rhinophores or may be a series of 
spots and dashes. The width and shape of the 
submarginal band are variable, as well. When 
thickening of the band occurs it is usually at the 
middle of the animal. Towards the middle of the 
animal, the band may occur closer to the longi- 
tudinal midline. All specimens seen have a spot 
or line between the rhinophores. Posteriorly, the 
two black submarginal bands may meet behind 
the gills or may be interupted with a medial black 
spot posterior to the gills. Some specimens have 
two black oval spots down the midline of the 
notum which either occur singly or directly adja- 
cent to each other forming an hourglass shape. 
The rhinophores are usually uniformly orange 
throughout. In one specimen, the upper 2/3 of the 
rhinophores is orange while the lower 1/3 is 
transparent. Gills are white, tinged with orange. 



Orange coloration may be limited to the longitu- 
dinal edges of the gills or may color the upper 1/3 
to 1/2 of the gills. A single specimen has black 
pigmentation along the inner edges of the gills 
and a smudge of color basally. The hyponotum 
is similar in color to the notum, with a white 
marginal line and black submarginal band encir- 
cling the posterior end of the foot. This band may 
be continuous or broken at the junction on the 
tail. 

The subcutaneous mantle glands (Figs. 4E, 5B) 
form a discontinuous submarginal band around 
the notum of the animal. Generally, they are 
absent from the anterior portion of the animal to 
just behind the rhinophores, but are occasionally 
situated alongside the rhinophores. These glands 
are elongate, wide and highly ramified (Fig. 5C). 
The glands situated near the middle of the body 
are wider and longer than those located at the 
anterior and posterior end of the body. 

On either side of the head is a short, digitiform 
oral tentacle. The branchial plume consists of 
9-14 unipinnate gills in four specimens exam- 
ined. The rhinophores are perfoliate with 16-22 
simple lamellae. 

Buccal Armature. — The muscular portion 
of the buccal mass is approximately equal in 
length to the oral tube. At the anterior end of the 
muscular portion of the buccal mass is the chiti- 
nous labial cuticle, which bears numerous jaw 
rodlets. The rodlets (Fig. 7 A) are elongate and 
sharply curved with a deeply bifid apex. The 
radular formula is 76 x 47. 1 .47 and 7 1 x 52. 1 .52 
in two specimens examined (CASIZ 083684 and 
110466, respectively). The vestigial rachidian 
teeth (Fig. 7B) are long and thin. The innermost 
lateral teeth (Fig. 7B) are arched with an elongate 
cusp. There are 2-3 strong denticles on the inner 
side of the tooth and three denticles on the outer 
side. The inner denticles are two to three times 
wider and more rounded than the outer denticles 
which taper to a sharper point. The second lateral 
has 3^4 denticles on its outer side. The lateral 
teeth from the middle of the radular row (Fig. 7C) 
are elongate and sharply curved with four to five 
elongate, triangular denticles on the outer side of 
the teeth. The outermost lateral teeth (Fig. 7D) 
are elongate with 3-5 finger-like denticles. 

Reproductive System. — (Fig. 5D) The 
ampulla is short, thick and tubular, narrowing 
somewhat before bifurcating into an oviduct and 
vas deferens. The short oviduct enters the female 



GOSLINER AND BEHRENS: CHROMODORIS 



149 




Figure 7. Chromodoris dianae sp. nov. Scanning electron micrographs. A. Jaw rodlets, scale = 7.5 urn. B. Inner lateral 
teeth, scale = 20 urn. C. Lateral teeth from central portion of half-row, scale = 20 urn. D. Outer lateral teeth, scale = 30 um. 



150 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 



gland mass near the albumen gland. The proxi- 
mal prostatic portion of the vas deferens is folded 
over itself once before it narrows slightly into the 
short, muscular, ejaculatory portion. The ejacu- 
latory portion narrows gradually to a curved seg- 
ment and then enters the elongate penial bulb. 
The penis is adjacent to the slender vaginal duct 
at the common gonopore. The distal end of the 
vas deferens is devoid of any penial hooks. The 
female gland mass consists of the large mucous 
gland and smaller membrane and albumen 
glands. Near the exit of the mucous gland is a 
small, ovoid vestibular gland. The vagina is rela- 
tively short. At the proximal end of the vagina is 
a short duct to the pyriform receptaculum 
seminis. The thin walled, spherical bursa copula- 
trix connects directly to the junction of the proxi- 
mal end of the vagina and the receptaculum 
seminis. The uterine duct emerges at a position 
immediately proximal of the junction of the re- 
ceptaculum with the bursa and vagina. The uter- 
ine duct is relatively short and enters the female 
gland mass near the albumen gland. 

DISCUSSION. — Chromodoris dianae is simi- 
lar in color to other pale blue members of the 
Chromodoris quadricolor complex that lack an 
orange marginal band. Chromodoris lochi Rud- 
man, 1982 and C. boucheti Rudman, 1982 have 
a narrow, continuous submarginal black band 
and a divided or continuous black longitudinal 
line along the mid-dorsum. In C. dianae, the 
submarginal band is wider and has large areas 
where it is interrupted laterally, anteriorly and 
posteriorly. The mid-dorsal pigment is restricted 
to a few isolated spots or bands. Chromodoris 
willani Rudman, 1 982 also has a continuous sub- 
marginal band and a more or less continuous 
mid-dorsal longitudinal line. It also has gray 
rhinophores and gills with opaque white spots 
while those of C. dianae are orange and white. 
Chromodoris dianae may have orange spots 
along the margin of the mantle; the spots are 
never present in the other three light blue species. 
The mantle glands of Chromodoris dianae are 
well separated from each other and are highly 
ramified with digitate branches. Similar mantle 
glands are found in C. lochi and C. willani (Fig. 
4F, G), but in these two species the glands are 
less branched, especially in C. lochi, where they 
form compact spherical bodies. Mantle glands 
are not known for C boucheti. 



The radula of C dianae contains 71-76 rows 
of radular teeth with 47-52 lateral teeth per side. 
This is similar to the formulae for C. lochi and C. 
willani (Rudman 1982). In contrast, C. boucheti 
has 53-56 rows of teeth with only 40-41 lateral 
teeth per side. The most significant difference in 
the radula of these four species is the prominent, 
well-developed rachidian found in C willani 
(Rudman 1982, fig. 7A, B), while in the remain- 
ing species the rachidian is a low, vestigial plate. 
It appears that the lateral teeth from the middle 
of the radular row of C. dianae are less highly 
arched and have a more elongate cusp than is 
depicted for C. lochi, C. boucheti or C. willani 
by Rudman (1982). In C. dianae and C. willani 
there are a maximum of 5 denticles on the middle 
lateral teeth, while in C. boucheti and C. lochi 
there may be as many as 8 denticles. 

The reproductive system of C. dianae has an 
extremely short ejaculatory segment of the vas 
deferens. The reproductive systems of C. willani 
and C. lochi were not described in detail, but 
specimens of these two species were examined 
here (CASIZ 083679 and 076101, respectively). 
In both of these species the ejaculatory segment 
and vaginal duct are proportionately longer than 
that observed for C dianae and C. boucheti 
(Rudman 1984:136, fig. 13A). The penial bulb 
of C. boucheti is much narrower than the bulbous 
bulb of C dianae. 

Chromodoris michaeli sp. nov. 
(Figs. 41, 8 A, 9A-B, 10, 1 1A-D) 

Chromodoris sp. Colin and Arneson, 1995:183, 
fig. 855. 

Type Material. — Holotype: CASIZ 076692, one 
specimen, Sepok, Maricaban Island, Philippine Is- 
lands, 22 m depth, 26 February 1995, Michael T. 
Ghiselin. Paratypes: CASIZ 079447, one specimen, 
buccal mass removed, Philippine Islands. CASIZ 
085983, one specimen, dissected, unnamed island 20 
mi N of Dakak Resort (Mindanao), Philippine Islands, 
17-22 m depth, 30 March 1993, T. M. Gosliner. 
CASIZ 1 10426, one specimen, Ligpo Island, Balayan 
Bay, Luzon, Philippine Islands, 20 m depth, 23 April 
1997, Bruce Baker. CASIZ 110777, one specimen, 
Kirby's Rock, Caban Island, Maricaban Strait, Philip- 
pine Islands, 20 m depth, 22 April 1 997, Bruce Baker. 

ETYMOLOGY. — Chromodoris michaeli is 
named for the second author's son, Michael 



GOSLINER AND BEHRENS: CHROMODORIS 



LSI 




Figure 8. Living animals. A. Chromodoris michaeli sp. nov , specimen from Liuay Rock, Dakak, Mindanao, Philippine 
Islands, photograph by T. M. Gosliner. B. Chromodoris hintuanensis sp. nov., specimen from Bus Stop Reef, Balayan Bay, 
Luzon, Philippine Islands, photograph by T. M. Gosliner. C. Chromodoris roboi sp. nov., specimen from Onna Village, 
Okinawa, Ryukyu Islands, Japan, photograph by Robert Bolland. 



152 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 







am 



Figure 9. Chromodoris michaeli sp. nov. A. Living animal. B. Subcutaneous glandular network. C. Mantle glands, enlarged, 
scale = 1 .0 mm. D. Reproductive system, am = ampulla, be = bursa copulatrix, ej = ejaculatory duct, fgm = female gland mass, 
p = penis, pr = prostate, rs = receptaculum seminis, v = vagina, vg = vestibular gland, scale = 1 .0 mm. 



David Behrens, an aspiring biologist with keen 
interests in malacology. 

Distribution. — Thus far, this species is 
known only from three localities in the Philip- 
pine Islands: northern Mindanao; Mactan Island, 
Cebu; and Batangas Province, Luzon (Colin and 
Arneson 1995; present study). 



NATURAL HISTORY. — Chromodoris mi- 
chaeli is found on reef surfaces to a depth of 22 
m. We have no information on this species' prey 
or other habits. 

External Morphology. — Preserved ani- 
mals are 24-^46 mm in length. The body of the 
living animal (Figs. 8A, 9A) appears powdery 



GOSLINER AND BEHRENS: CHROMODORIS 



!53 







Figure 10. Chromodoris michaeli sp. nov. Variation of color pattern. 



blue due to a fine speckling of white over its 
blue-brown ground color. A white line edges the 
margin. Inside the white line is a wide submargi- 
nal bright orange band which is uninterrupted. 
Adjacent to the orange band is a narrow band of 
pale body color. Inside this blue area is a black 
band that encircles the animal well posterior to 
the gills and anterior to the rhinophores. This 
black band is discontinuous in two spots, subme- 
dially at the anterior end and again posteriorly in 
all specimens seen. Often the band is wider to- 
wards the middle. Within the boundaries of this 
black band are black spots which vary in size, 
number and position on the notum. One paratype 
specimen has elongate spots along the midline of 
the dorsal surface of the notum appearing as a 
broken line. Additionally, this specimen has ran- 
dom spots. Another specimen has two short lon- 
gitudinal stripes on either side of the midline at 
the middle of the animal, spots circling the gills 
and two elongate spots antero-medially which 
form a broken line. The specimen depicted by 
Colin and Arneson (1995, fig. 855) has two 
spots: one at the base of the gill circle and another 
between the rhinophores. In some specimens, the 
black patterns have thin white margins. The 
range of variation in color pattern is shown in 
Figure 10. The edge of the foot is orange and the 
hyponotum is powdery blue with three lateral 
black stripes in all specimens observed. The gills 
and rhinophores are burnt orange throughout. 
The triangular foot extends posteriorly beyond 
the notum when the animal is crawling. 

The subcutaneous mantle glands form a dis- 
continuous submarginal band around the notum 
of the animal (Figs. 41, 9B). Glands are densely 
lobed and convoluted (Fig. 9C) and are generally 



absent anteriorly from just behind the rhino- 
phores but occasionally may occur lateral to one 
or both rhinophores. 

On either side of the head is a short, digitiform 
oral tentacle. In the three specimens examined 
there are 11-18 unipinnate gills forming the 
branchial plume. The perfoliate rhinophores bear 
21-23 lamellae for the same three specimens. 

Buccal Armature. — At the anterior end of 
the muscular portion of the buccal mass is the 
chitinous labial cuticle, which bears numerous 
jaw rodlets. The rodlets (Fig. 1 1 A) are elongate, 
sharply curved and deeply bifid at the apex. The 
radular formula is 75 x 61. 1.61 in one specimen 
examined (CASIZ 079447) and 75 * 49- 
51.1.49-51 in another (CASIZ 085983). The 
vestigial rachidian teeth (Fig. 1 IB) are long and 
thin. The innermost lateral teeth (Fig. 1 IB) are 
arched with an elongate cusp. There are two to 
three prominent denticles on the inner side of the 
tooth and three to four on the outer side which 
are bluntly pointed. The inner denticles are two 
to three times wider than the outer denticles and 
are more rounded. The lateral teeth from the 
middle of the radular row (Fig. 1 1C) are elongate 
and sharply curved with two to four small, 
sharply pointed triangular denticles on the outer 
side of the cusp. Near the base of the denticles 
the denticle widens abruptly, creating an uneven 
taper. This is consistent in the two specimens 
examined. There are no denticles on the inner 
side. The cusp of the outermost lateral teeth (Fig. 
1 ID) is reduced and bears 4-6 finger-like denti- 
cles on the outer side of the teeth. 

Reproductive System. — (Fig. 9D) The 
ampulla is short, thick and tubular, narrowing 
somewhat before bifurcating into an oviduct and 



154 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 




Figure 1 1 . Chromodoris michaeli sp. nov. Scanning electron micrographs. A. Jaw rodlets, scale = 10 urn. B. Inner lateral 
teeth, scale = 30 urn. C. Lateral teeth from central portion of half-row, scale = 25 urn. D. Outer lateral teeth, scale = 30 um. 



GOSLINER AND BEHRENS: CHROMODORIS 



155 



vas deferens. The short oviduct enters the female 
gland mass near the albumen gland. The proxi- 
mal prostatic portion of the vas deferens is folded 
over itself once before it narrows markedly into 
the short, muscular, ejaculatory portion. The 
ejaculatory portion narrows abruptly to a thin 
curved segment and then enters the elongate 
penial bulb. The distal end of the vas deferens is 
devoid of any penial hooks. The penial bulb is 
adjacent to the slender vaginal duct at the com- 
mon gonopore. The female gland mass consists 
of the large mucous gland and smaller membrane 
and albumen glands. Near the exit of the mucous 
gland is a small, ovoid vestibular gland. A rela- 
tively short vagina leads to the thin-walled, 
spherical bursa copulatrix. A short duct emerges 
from the bursa copulatrix a short distance from 
its junction with the vagina and connects to the 
pyriform receptaculum seminis. The uterine duct 
emerges a quarter of the length along the duct to 
the receptaculum seminis. The uterine duct is 
relatively short and enters the female gland mass 
near the albumen gland. 

Discussion. — The color pattern of C. 
michaeli is similar to C. elisabethina Bergh, 
1877; C. hamiltoni Rudman, 1977; C. quadri- 
color (Ruppell and Leuckart, 1828); C. annae 
Bergh, 1877; and C. westralensis (O'Donoghue 
1924). All of these species have a light blue body 
color with an orange marginal or submarginal 
band and black pigment on the body. The first 
three of these similar species have a continuous 
black medial line and may have additional black 
markings on the notum. Chromodoris westralen- 
sis can be readily distinguished from C. michaeli 
and C. annae by the presence of a broad, black 
medial blotch that connects with the two lateral 
margins of the black ring. Chromodoris annae 
has dark black pigment spots that show through 
gaps in the blue-colored area, while in C. 
michaeli these spots are whitish. The blue pig- 
ment of C. michaeli is lighter than that of C. 
annae. The black submarginal band of C. annae 
is almost always continuous posteriorly, al- 
though Rudman (1982, fig. 13E) illustrated a 
specimen where the lines did not meet posteri- 
orly. In C. annae the black bands do not meet 
anteriorly, but a single black spot is present an- 
teriorly and may extend posteriorly between the 
rhinophores as an uninterrupted medial line. In 
C. michaeli the anterior portion of the bands is 
interrupted and a medial anterior black spot is 



always present. This spot never extends posteri- 
orly as a line. A second spot is present between 
or immediately posterior to the rhinophores. Ad- 
ditional black spots or interrupted black bars are 
usually present on the medial line of the notum. 
An additional black spot is always present ante- 
rior to the anterior margin of the branchial 
pocket. 

The mantle glands of C. michaeli, C. elisabe- 
thina, C. hamiltoni C. quadricolor, and C. annae 
are all similar in degree of branching and the fact 
that the glands are widely separated with ap- 
proximately 5 glands on either side of the mantle 
margin. 

The radular teeth of C. michaeli, C. elisabe- 
thina, C. hamiltoni C. quadricolor, C. annae and 
C. westralensis are similar in many respects. All 
species have a vestigial rachidian tooth. The re- 
maining teeth are all denticulate. In C. westralen- 
sis there are more radular rows and more teeth 
per half row than has been reported for the other 
species (Rudman 1982). The middle lateral teeth 
ofC. michaeli have a maximum of four denticles, 
while C. hamiltoni has 4-5 denticles and the 
remaining species have 6-8. 

The reproductive system is similar in its ar- 
rangement in C. michaeli, C. elisabethina, C. 
hamiltoni and C. annae (Rudman 1977; present 
study). It has not been described for C. quadri- 
color or C. westralensis. In all cases, the ejacu- 
latory portion of the vas deferens is short relative 
to most other described chromodorids. In C. 
elisabethina and C. hamiltoni (Rudman 1977, 
fig. 17B, D), the ejaculatory portion is longer 
than that found in C. michaeli and C. annae 
(present study). The vagina of C. annae is wider 
and more elongate than that found in C. michaeli. 
While only minor anatomical differences 
separate C. michaeli from other blue chromo- 
dorids with an orange submarginal band, the 
differences in coloration, radular tooth morphol- 
ogy and arrangement of reproductive organs are 
consistent and warrant its consideration as a dis- 
tinct species. 

Chromodoris hintuanensis sp. nov. 
(Figs. 8B, 12A-D, 13A-D) 

Chromodoris geometrica Debelius, 1 996:2 1 0, bottom 
left photograph, not Risbec, 1928, misidentifica- 
tion. 



156 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 








Figure 12. A-D, Chromodoris hintuanensis sp. nov. A. Living animal. B. Subcutaneous glandular network. C. Mantle 
glands, enlarged, scale = 1 .0 mm. D. Reproductive system, scale = 1 .0 mm. E. Chromodoris geometrica Risbec, 1 928, 
reproductive system, scale = 1 .0 mm. F. Chromodoris conchyliata Yonow, 1984, reproductive system, scale = 1 .0 mm, am = 
ampulla, be = bursa copulatrix, ej = ejaculatory duct, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum 
seminis, v = vagina, vg = vestibular gland. 



GOSLINER AND BEHRENS: CHROMODORIS 



57 




Figure 13. Chromodoris hintuanensis sp. nov. Scanning electron micrographs. A. Jaw rodlets, scale = 10 urn. B. Rachidian 
and inner lateral teeth, scale = 25 urn. C. Lateral teeth from central portion of half-row, scale = 30 urn. D. Outer lateral teeth, 
scale = 30 urn. 



158 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 



Type Material. — Holotype: CASIZ 1 0649 1 , one 
specimen, Bus Stop Reef, Balayan Bay, Luzon, Phil- 
ippine Islands, 2 m depth, 18 April 1996, M. Miller. 
Paratypes: CASIZ 079286, one specimen, buccal mass 
removed, Seragaki Beach, 1 .3 km ENEof Maeki-zaki, 
Okinawa, Ryukyu Islands, Japan, 8 m depth, 7 April 
1990, Robert F. Bolland. CASIZ 085931, one speci- 
men, Cathedral, Balayan Bay, Luzon, Philippine Is- 
lands, 8 m depth, 25 March 1993, T. M. Gosliner. 
CASIZ 083663, four specimens, one specimen dis- 
sected, Bus Stop Reef, Balayan Bay, Luzon, Philip- 
pine Islands, 2 m depth, 21 February 1992, T. M. 
Gosliner. CASIZ 105728, one specimen, Seafari 
Beach, Anilao, Batangas Province, Luzon, Philippine 
Islands, 10 m depth, 23 February 1995, M. Miller. 
CASIZ 103717, one specimen, La Laguna Munbi 
(point W of Sabang Point), Sabang, Mindoro, Philip- 
pine Islands, 1 March 1995, Mel Segara. 

ETYMOLOGY. — The name hintuanensis 
comes from the Tagalog word hintuan meaning 
"stopping place," referring to the type locality, 
dubbed Bus Stop by the authors, situated off- 
shore from the waiting shed where the bus stops 
to collect passengers going to Manila. 

DISTRIBUTION. — Thus far this species is 
known only from Bali, Indonesia (Debelius 
1996); a photograph by Jim Black from Papua 
New Guinea; Batangas Province, Luzon, Philip- 
pine Islands (present study); Okinawa, Ryukyu 
Islands, Japan (present study); and from photo- 
graphs by Mark Strickland from the Andaman 
Sea, Thailand. 

Natural History. — This species is com- 
monly encountered in shallow subtidal sandy or 
silty habitats, where it is observed crawling in the 
open. When actively crawling, the animal alter- 
nately raises and lowers the anterior portion of 
the body. 

External Morphology. — Preserved ani- 
mals are 11-16 mm in length. The body color of 
the living animals (Figs. 8B, 12A) is white and 
the dorsal surface is covered with irregularly- 
shaped nodules of varying sizes. The mantle 
margin is edged with deep violet which, in areas 
of mantle folding, extends inward from the edge 
to form small patches of color. Small circles of 
deep magenta outline 5-8 of the small, opaque 
white, rounded nodules on the medial area of the 
dorsal surface of the animal and are also found at 
the base of the rhinophore sheath and gill plume. 
The outer edges of the circles fade to a plum color 
which also forms an irregular network on the 
notum in the spaces between nodules. The rhino- 



phores and gills are a dull plum. Crimson lines 
are present along the longitudinal edges of the 
gill plumes and on the rachis and lamellae of the 
rhinophores. In all of the specimens examined, at 
least one small, rounded nodule is found on the 
base of the rhinophoral sheath. The foot is white 
with a deep violet margin and may extend only 
slightly beyond the posterior edge of the notum 
when the animal is crawling. There is a high 
degree of mantle folding. 

The subcutaneous mantle glands (Fig. 12B) 
form a discontinuous submarginal band around 
the notum except for the anterior portion of the 
animal from just behind the rhinophores where 
they are absent. The glands are relatively small 
yet dense, forming highly lobate masses (Fig. 
12D). 

On either side of the head is a short, digitiform 
oral tentacle. There are 10 unipinnate gills form- 
ing the branchial plume. The perfoliate rhino- 
phores bear 28-34 lamellae. 

BUCCAL ARMATURE. — At the anterior end of 
the muscular portion of the buccal mass is the 
chitinous labial cuticle, which bears numerous 
jaw rodlets. The rodlets (Fig. 13 A) are elongate 
and curved. There is a distinct narrowing of the 
rodlets towards the apex which is shallowly bifid. 
The radular formulae of two specimens exam- 
ined is 48 x 43.1.43 (CASIZ 083663) and 47 * 
42.1.42 (CASIZ 079286). The rachidian teeth 
(Fig. 13B) are reasonably well developed. Their 
width is 1/3 to 1/2 that of the first laterals. The 
innermost lateral teeth (Fig. 13B) have a short, 
pointed cusp and bear 1-2 rounded denticles on 
the inner side and 4-5 pointed denticles on the 
outer side which are elongate and curved. The 
arched lateral teeth from the middle of the row 
(Fig. 13C) bear 4-8 elongate pointed denticles on 
the outer side. The cusp on the outermost lateral 
teeth (Fig. 1 3D) is reduced and the teeth bear 2-3 
rounded, elongate denticles on the outer side. 

Reproductive System. — (Fig. 12D) The 
ampulla is thick, long and tubular and narrows 
into a relatively long postampullary duct prior to 
bifurcating into the oviduct and vas deferens. The 
short oviduct enters the female gland mass near 
the albumen gland. The proximal prostatic por- 
tion of the vas deferens is tightly coiled, consist- 
ing of several distinct loops. The duct narrows 
slightly and again expands into the relatively 
long muscular ejaculatory portion, which termi- 
nates at a short, tubular penial bulb. The distal 



GOSLINER AND BEHRENS: CHROMODORIS 



159 



end of the vas deferens is devoid of any penial 
hooks. The penial sheath is adjacent to the wide, 
moderately elongate vaginal duct at the common 
gonopore. The female gland mass consists of a 
large mucous gland and smaller membrane and 
albumen glands. Near the exit of the mucous 
gland is a small, pyriform, elongate vestibular 
gland. The vagina is long and curved, being 
markedly narrower proximally. The thin-walled, 
spherical bursa copulatrix and the pyriform re- 
ceptaculum seminis have a common junction at 
the proximal end of the vagina. The bursa is 
large, nearly twice the size of the receptaculum 
and one-third the size of the female gland mass. 
The uterine duct emerges from the common junc- 
tion of the vagina, the bursa copulatrix and the 
receptaculum seminis, and enters the female 
gland mass near the albumen gland. 

DISCUSSION. — Chromodoris hintuanensis is 
a member of a group of species in which the 
anterior end of the body is rhythmically raised 
and lowered as the animal crawls. Included in 
this group of species are C. geometrica Risbec, 
1 928 and C. conchyliata Yonow, 1 984, which are 
similar in appearance to C. hintuanensis. These 
three species also have a series of tubercular 
warts on the notum and a network of darker lines. 
This similarity has led to confusion of these 
species. Specimens of C conchyliata have been 
erroneously attributed to C. geometrica (Rud- 
man 1973; Gosliner 1987) and specimens of C. 
hintuanensis have been misidentified as C. 
geometrica (Debelius 1996:210, lower left pho- 
tograph). Chromodoris roboi, described here, 
also exhibits the characteristic behavior of rais- 
ing and lowering the head (R. Bolland, pers. 
comm.). However, it lacks the characteristic net- 
work of lines found in the other three species 
described above. 

Owing to this confusion, specimens of all three 
species were examined in this study. Specimens 
of C conchyliata from Natal, South Africa 
(SAM A35449) were examined as was a speci- 
men of C. geometrica from the Philippines 
(CASIZ 103766). The three species differ in 
several aspects of their external morphology and 
color pattern. The notum of C. hintuanensis has 
a network of pale plum areas while in C geomet- 
rica the network is dark brown to black (Yonow 
1994; present study) and in C conchyliata it is 
deep purple (Rudman 1973; Yonow 1984; 
Gosliner 1987). In C hintuanensis the rhino- 



phores and gills have deep red lines along their 
rachis as do the rhinophoral lamellae. In C. con- 
chyliata the gills have red lines along the rachis, 
but the rhinophores are uniformly red. In C. 
geometrica the gills and rhinophores are yel- 
lowish green with opaque white spots. The man- 
tle margin of C. hintuanensis has a distinct purple 
band with a series of indentations around the 
edge of the mantle. A purple margin without 
indentations is found only on the anterior end of 
the mantle margin of C geometrica and is en- 
tirely absent in C conchyliata. Chromodoris hin- 
tuanensis has opaque white spots, which are 
absent in the other two species. In the specimens 
of C. hintuanensis examined here there are al- 
ways 10 gills while in C. conchyliata there are 
7-10 and in C. geometrica there are 5-8. Chro- 
modoris hintuanensis has more rhinophoral la- 
mellae (28-34) than has been observed in present 
or previous records of C geometrica (18-22) or 
C. conchyliata (28-29). The mantle glands of C 
hintuanensis are highly digitate and well scat- 
tered, forming a single row of glands. Chromo- 
doris conchyliata has as a single row of glands 
that are positioned more closely together and are 
less ramified than in C hintuanensis. In C. 
geometrica the glands are arranged in a double 
row and only a few are highly ramified. 

Internally, there are also differences in the 
reproductive systems, especially with respect to 
the length of the vagina and the elaboration of the 
ejaculatory portion of the vas deferens. In C. 
hintuanensis the vaginal duct is of moderate 
length, whereas it is elongate and convoluted in 
C. geometrica (Figure 12E) and short and wide 
in C. conchyliata (Figure 12F). In C. hintuanen- 
sis and C geometrica the receptaculum seminis 
joins the base of the bursa copulatrix while in C. 
conchyliata the receptaculum joins the vagina 
slightly more distally from the bursa. 

Chromodoris roboi sp. nov. 
(Figs. 8C, 14A-D, 15A-D) 

Chromodoris sp. Coleman, 1989:35, top photograph. 

Hypselodoris vibrata Debelius, 1996:239, bottom 

photograph, not (Pease, 1 860), misidentification. 

Type Material. — Holotype: CASIZ 079346, one 
specimen, Horseshoe Cliffs, 1 km NW of Onna Vil- 
lage, Okinawa, Ryukyu Islands, Japan, 26°30'N, 127° 
50.9'E, 47 m depth, 31 August 1991, Robert F. Bol- 
land. Paratypes: CASIZ 070184, one specimen, 



160 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 




Figure 14. Chromodoris roboi sp. nov. A. Living animal. B. Reproductive system, am = ampulla, be = bursa copulatrix, 
ej = ejaculatory duct, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, vg = vestibular gland, 
scale = 1 .0 mm. 



Horseshoe Cliffs, 1 km NW of Onna Village, Oki- 
nawa, Ryukyu Islands, Japan, 26°30' N, 127° 50.9' E, 
40 m depth, 20 May 1989, Robert F. Bolland. CASIZ 
070048, one specimen, Horseshoe Cliffs, 1 kmNWof 
Onna Village, Okinawa, Ryukyu Islands, Japan, 
26°30'N, 127°50.9'E, 37 m depth, 1 April 1989, 
Robert F. Bolland. CASIZ 070120, one specimen, 



dissected, Horseshoe Cliffs, 1 km NW of Onna Vil- 
lage, Okinawa, Ryukyu Islands, Japan, 26°30'N, 
127°50.9'E, 50 m depth, 11 April 1987, Robert F. 
Bolland. CASIZ 070230, one specimen, Horseshoe 
Cliffs, 1 km NW of Onna Village, Okinawa, Ryukyu 
Islands, Japan, 26°30'N, 1 27°50.9'E, 50 m feet depth, 
9 May 1987, Robert F. Bolland. 



GOSLINER AND BEHRENS: CHROMODORIS 



161 




Figure 15. Chromodoris robot sp. nov. Scanning electron micrographs. A. Jaw rodlets, scale = 10 urn. B. Inner lateral teeth, 
scale = 30 |im. C. Lateral teeth from central portion of half-row, scale = 30 |im. D. Outer lateral teeth, scale = 25 urn. 



ETYMOLOGY. — Chromodoris robot is named "robo," synonymous with Robo Cop, a fictitious 

for Robert Bolland, friend and enthusiastic nudi- personality equally as daring and adventurous as 

branch biologist. We have taken the first two our friend, Bob Bolland. 
letters of his first and last names to produce 



162 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 



Distribution. — Thus far this species is 
known only from Lord Howe Island, New South 
Wales, Australia (Coleman 1989), Murion Is- 
land, Western Australia (Debelius 1996, as Hyp- 
selodoris vibrato) and Okinawa, Ryukyu Islands, 
Japan (present study). 

Natural History. — As in C. hintuanensis, 
C. robot raises and lowers its anterior end when 
actively crawling (R. Bolland, pers. comm.). 

External Morphology. — The living ani- 
mals (Figs. 8C, 14A) are 17-50 mm in length. 
This species is easily distinguished by the mag- 
nificent, bright blue blocks of color which appear 
in series separated by black areas of pigment. The 
blue blocks are squarish or rectangular in shape 
with edges that are rounded to varying degree. 
The black color sometimes extends into the outer 
edges of the blue areas, bisecting them to varying 
degree. The blue coloration often extends inward 
from the black marginal band interrupting the 
yellow-orange pigment which covers most of the 
notum of the animal. The yellowish orange dor- 
sal surface of the notum is absent in many areas 
where circles or irregular patches of blue to lav- 
ender are visible. These patches are different 
shapes and sizes. Usually, a large patch is present 
behind each rhinophore. In some specimens, 
dark brown blotches cover the dorsal surface to 
varying degrees. These areas appear wine-col- 
ored or black depending on the amount of pig- 
ment present. The areas bordering the rhinophore 
and gill sheaths are orange and may or may not 
have spots of bright blue, lavender, black or wine 
color. The lamellae and rachis of the rhinophores 
and the longitudinal edges of the gill plumes are 
dark blue to black. In one specimen, the gill 
plumes are entirely blue-black. Coloration of the 
foot is similar to that of the mantle margin. It is 
blue with black stripes radiating from near the 
median line. There are 5-6 elongate, unipinnate 
gills and the rhinophores bear 19-22 lamellae. 

The mantle margins are very thin and not well 
preserved in the present material; thus arrange- 
ment of mantle glands could not be ascertained. 
However, it is evident from a photograph of this 
animal (Debelius 1996:239, lower photograph) 
that small round mantle glands are arranged in 
several distinct rows and are found continuously 
from the posterior end of the animals along the 
lateral margins to about the level of the rhino- 
phores. 



Buccal Armature. — The jaw rodlets (Fig. 
1 5 A) of the labial cuticle are elongate, curved and 
shal lowly bifid. The radular formulae of two 
specimens are 50 * 49.1.49 and 45 x 48.1.48 
(CASIZ 070120 and 070184, respectively). The 
narrow, thin rachidian tooth (Fig. 15B) is elon- 
gate at its base. The innermost lateral teeth (Fig. 
15B) are curved with a short, pointed cusp. One 
to two thick, rounded denticles are on the inner 
side of the tooth and four to five thinner, elon- 
gate, pointed teeth are on the outer side. The 
lateral teeth from the middle of the radular row 
(Fig. 15C) are arched with an elongate cusp and 
bear eight to eleven small elongate, pointed den- 
ticles on the inner side of the tooth. The cusp of 
the outermost lateral teeth (Fig. 15D) is reduced 
and the teeth bear 8-10 finger-like denticles. 

Reproductive System. — (Fig. 14B) The 
ampulla is thick and tubular, narrowing some- 
what before bifurcating into an oviduct and vas 
deferens. The short oviduct enters the female 
gland mass near the albumen gland. The proxi- 
mal prostatic portion of the vas deferens is slen- 
der and forms a few tight loops. The relatively 
long, muscular ejaculatory portion is similar in 
width to the prostatic portion and terminates in a 
relatively short penial bulb. The distal end of the 
vas deferens is devoid of any penial hooks. The 
penial bulb is adjacent to the slender vaginal duct 
at the common gonopore. The female gland mass 
consists of a large mucous gland, and smaller but 
indistinct membrane and albumen glands. Near 
the exit of the mucous gland is a small, ovoid 
vestibular gland. The vagina is relatively long 
and curved. The thin-walled, spherical bursa 
copulatrix and the pyriform receptaculum 
seminis share a common junction at the proximal 
end of the vagina. The uterine duct emerges from 
this junction and enters the female gland mass 
near the albumen gland. 

DISCUSSION. — Chromodoris robot is readily 
distinguishable from all other described species 
of Chromodoris by its unique pattern of colora- 
tion. It is the only species with a yellow-orange 
body color with blue marginal rectangles sepa- 
rated by black spaces. Chromodoris vibrata 
(Pease, 1860) is the most similar species in terms 
of color pattern and has been confused with C. 
robot (Debelius 1996:239, lower photograph). It 
is a yellow-orange animal with white pustules on 
the notum and a white marginal region with 
triangular dark blue interrupting the white. It also 



GOSLINER AND BEHRENS: CHROMODORIS 



163 



has dark blue pigment on the rachis of the gills 
and on the white rhinophores. Chromodoris 
roboi has 5-6 gills which do not appear to vibrate 
(R. Bolland, pers. comm.) while C vibrata has 
8-10 vibratile gills (Kay and Young 1969, as 
Hypselodoris vibrata). 

The radula of C. roboi has more rows of teeth 
than has been described for C. vibrata. A speci- 
men of C. vibrata with 56 rows of teeth had only 
38 teeth per half row (Kay and Young 1969) 
while a specimen of C. roboi with 50 row of teeth 
had 49 teeth per half row. A vestigial rachidian 
tooth is present in C roboi but is apparently 
absent in C. vibrata. The lateral teeth from the 
middle of the radular row of C. roboi have 8-1 1 
denticles, while those of C. vibrata have a maxi- 
mum of four denticles. 

The reproductive systems of C. roboi and C. 
vibrata appear to have some differences, as well. 
Kay and Young (1969) describe a large curved 
receptaculum seminis while the specimen of C. 
roboi described here has a small pyriform recep- 
taculum. The ejaculatory portion of the vas def- 
erens of C. roboi is much larger than that 
described for C. vibrata. No vestibular gland was 
described for C. vibrata. However, additional 
material should be examined to verify this obser- 
vation. 

Chromodoris aureopurpurea Collingwood, 
1881, also bears some resemblance to C. roboi in 
its coloration. This species has a white body with 
dark blue intrusions along the mantle margin and 
orange spots on the notum (Rudman 1987:342, 
fig. 23B). It also has opaque white lines on the 
gills and rhinophores, which are absent in C. 
roboi. Its radular formula is more similar to that 
of C. roboi than that of C vibrata. The radular 
teeth of C. aureopurpurea have a smaller 
rachidian plate and have far coarser, more well 
separated denticles (Rudman 1987:349, fig. 23) 
than those present in C roboi. The ejaculatory 
duct of C. roboi is much wider than that depicted 
for C aureopurpurea (Rudman 1987:351, fig. 
29B). 

Discussion 

Rudman (1982) described several species in 
the Chromodoris quadricolor complex. In this 
paper he stated that members of this color com- 
plex represent an artificial group in a phyloge- 
netic sense, since species of Hypselodoris are 



also included. Since then, Rudman (1984) has 
determined many useful characters, such as 
elaboration of mantle glands, variation in radular 
morphology and elaboration of the reproductive 
system, to distinguish between chromodorid 
genera. However, he questioned the utility of 
employing several of these characters in the sepa- 
ration of species groups within genera. In exam- 
ining the species described in the present paper 
and in comparing them with other taxa, we have 
observed several consistent anatomical features 
within Chromodoris. The mantle glands within 
species of the Chromodoris quadricolor com- 
plex are all relatively large and highly digitate. 
The species with black and white or yellow pig- 
ment (C. africana, C magnifica and C joshi) 
have a more or less continuous band of glands 
around the margins, although they are more sepa- 
rated in C magnifica than in the other two spe- 
cies examined. Members of this group, including 
C. kuiteri Rudman, 1982, have smaller, reduced 
denticles on the middle lateral teeth. These facts 
suggest that the similarities between these spe- 
cies with similar color patterns are not just lim- 
ited to color, but that they may have closer 
phylogenetic proximity to each other than to 
other taxa. Similarly, members of the C. quadri- 
color complex with a blue body color (C. lochi, 
C. willani, C elisabethina, C annae, C hamil- 
toni, C dianae and C michaeli) examined in this 
study tend to have more highly ramified mantle 
glands that are well separated from each other. In 
addition, the reproductive system of all of the 
species examined here in the Chromodoris 
quadricolor complex (C. joshi, C. dianae, C 
michaeli, C lochi, C. willani and C annae) plus 
species previously studied by Rudman (1977) 
(C. magnifica, C elisabethina, C. qfricana and 
C. hamiltoni) and Rudman (1984) (C boucheti 
and C kuiteri) all have an extremely short ejacu- 
latory segment of the vas deferens. The morphol- 
ogy of the reproductive system of these species 
differs from other species of Chromodoris de- 
scribed here (C. hintuanensis and C. roboi) 
which are not part of the C. quadricolor complex. 
These other species have much longer prostatic 
and ejaculatory portions of the vas deferens. 
Elongate prostatic and ejaculatory segments are 
present in most other species of Indo-Pacific 
Chromodoris not included in the C. quadricolor 
complex (Kay and Young 1969; Rudman 1987; 
Gosliner 1994). Other species from southern 



164 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 5 



Australia (Rudman 1987) also have a short ejacu- 
latory segment. 

Based on the examination of the distribution 
and arrangement of mantle glands and reproduc- 
tive characters examined here, there is evidence 
that the species of Chromodoris placed within 
the C. quadricolor color group may represent a 
distinct subclade within this large genus consist- 
ing of several hundred species. Identification of 
monophyletic subunits is essential in building an 
understanding of the phylogenetic relationships 
in order that comparative biological studies of 
biogeography and evolution of color patterns 
may be undertaken. Preliminary data with re- 
spect to variation in distribution of mantle glands 
and reproductive characters present promising 
data to untangle the seemingly overwhelming 
diversity of Chromodoris species and certainly 
warrant further study of anatomical features to 
develop a more comprehensive data set for phy- 
logenetic analysis. 

Acknowledgments 

This work would not have been possible with- 
out the generous support of many individuals. 
We offer sincere thanks to Elizabeth Kools of the 
Department of Invertebrate Zoology and Geol- 
ogy of the California Academy of Sciences for 
her assistance in many aspects of the research and 
manuscript preparation of this work. Field work 
was made possible by financial support from 
Katherine Stewart, United Airlines and the Lind- 
say Field Research Fund of the California Acad- 
emy of Sciences. Our collections were made 
possible through the kind support of our col- 
leagues at the Bureau of Fisheries and Aquatic 
Resources of the Republic of the Philippines. Jim 
Black, Marc Chamberlain, Roger Steene and 
Mark Strickland kindly provided photographic 
records of the species described here. Bob Bol- 
land and Mike Miller kindly provided specimens 
and permitted us to use their photographs for 
preparation of color plates. Dong Lin of the Pho- 
tography Department of the California Academy 
of Sciences provided the scans and final produc- 
tion of scanning electron micrographs. 

Literature Cited 

Allen, G. R. and R. Steene. 1994. Indo-Pacific coral 
reef field guide. Tropical Reef Research, Singapore. 
378 pp. 



Coleman, N. 1 989. Nudibranchs of the South Pacific. 
Neville Coleman's Sea Australia Resource Centre, 
Springwood. 64 pp. 

Colin, P. and C. Arneson. 1995. Tropical Pacific 
invertebrates. Coral Reef Press, Beverly Hills. 
296 pp. 

Dakin, N. 1992. The Macmillan Book of the Marine 
Aquarium. A definitive reference to more than 300 
marine fish and invertebrate species and how to 
establish and maintain a reef aquarium. Macmillan 
Publishing Co., New York. 400 pp. 

Debelius, H. 1996. Nudibranchs and sea snails. Indo- 
Pacific field guide. IKAN - Unterwasserarchiv, 
Frankfurt, Germany. 321 pp. 

GOSLINER, T. 1987. Nudibranchs of southern Africa. 
Sea Challengers, Monterey. 1 36 pp. 

-. 1994. New species of Chromodoris and 



Noumea (Nudibranchia: Chromodorididae) from 
the western Indian Ocean and southern Africa. Pro- 
ceedings of the California Academy of Sciences 
48(12):239-252. 

Kay, E. A. and D. Young. 1969. The Doridacea 
(Opisthobranchia, Mollusca) of the Hawaiian Is- 
lands. Pacific Science 23(2): 1 72-23 1 . 

Rudman, W. B. 1973. Chromodorid opisthobranch 
Mollusca from the Indo-West Pacific. Zoological 
Journal of the Linnaean Society 52(3): 175-1 99. 

. 1977. Chromodorid opisthobranch Mollusca 



from East Africa and the tropical West Pacific. 
Zoological Journal of the Linnaean Society 
61:351-397. 

. 1982. The Chromodorididae (Opisthobran- 



chia: Mollusca) of the Indo-West Pacific: Chromo- 
doris quadricolor, C lineolata and Hypselodoris 
nigrolineata color groups. Zoological Journal of the 
Linnaean Society 76:183-241. 

. 1983. The Chromodorididae (Opisthobran- 



chia: Mollusca) of the Indo-West Pacific: Chromo- 
doris splendida, C aspersa and Hypselodoris 
placida colour groups. Zoological Journal of the 
Linnean Society 78:105-173. 

. 1984. The Chromodorididae (Opisthobran- 



chia: Mollusca) of the Indo-West Pacific: a review 
of the genera. Zoological Journal of the Linnean 
Society 8 1(2,3): 115-273. 



GOSLINER AND BEHRENS: CHROMODORIS 



165 



. 1986. The Chromodorididae (Opisthobran- 

chia: Mollusca) of the Indo-West Pacific: Noumea 
purpurea and Chromodoris decora colour groups. 
Zoological Journal of the Linnean Society 
86:309-353. 

. 1987. The Chromodorididae (Opisthobran- 



chia: Mollusca) of the Indo-West Pacific: Chromo- 
doris epicuria, C. aureopurpurea, C. annulata, C. 
coi and Risbecia tryoni colour groups. Zoological 
Journal of the Linnean Society 90: 305-407. 



Wong, M. P. 1991. Sipadan, Borneo's underwater 
paradise. Odyssey Publishing, Singapore. 216 pp. 

YONOW, N. 1984. Doridacean nudibranchs from Sri 
Lanka, with descriptions of four new species. The 
Veliger26(3):2 14-228. 

. 1994. Opisthobranchs from the Maldive Is- 



lands including descriptions of seven new species 
(Mollusca: Gastropoda). Revue Francaise 
d'Aquariologie 20(4):97-129. 



CALIFORNIA ACADEMY OF SCIENCES, 1998 
Golden Gate Park 
San Francisco, California 941 18 






PROCEEDINGS 

of the FEB 17 1998 

CALIFORNIA ACADEMY OF SCIENCES 






Vol. 50, No. 6, pp. 167-184, 44 pi. figs. 



February 11, 1998 



A REVIEW OF THE COSTA RICAN SPECIES OF 

LEPTOGLOSSUS GUERIN, WITH DESCRIPTIONS OF TWO NEW 

SPECIES (HEMIPTERA: HETEROPTERA: COREIDAE: 

COREINAE: ANISOSCELINI) 



By 

Harry Brailovsky 1 and Ernesto Barrera 

Departamento de Zoologia, Instituto de Biologia, UN AM 
Apdo Postal 70153, Mexico D.F. 



Two new species of Leptoglossus, L. cartagoensis and L. talamancanus, collected in Costa Rica are 
described and illustrated; L. brevirostris Barber, L. humeralis Allen, L. lineosus (St&l), L. 
nigropearlei Yonke, L. subauratus Distant, and L. tetranotatus Brailovsky and Barrera are 
recorded for the first time from Costa Rica; new records for L. ductus (Herrich-Schaeffer), L. 
concolor (Walker), L. phyllopus (Linnaeus), and L. zonatus (Dallas) arc given; and a revised key 
to known species of Leptoglossus from Costa Rica is presented. 

Received May 14, 1997. Accepted September 5, 1997. 



The genus Leptoglossus Guerin is a large com- 
plex group that ranges from southern Canada, 
through the United States, Mexico, Antilles, 
Central America, and South America including 
Chile and Argentina. Leptoglossus gonagra 
(Fabricius) is the only species of Leptoglossus 
that occurs outside of the Western Hemisphere, 
and has been recorded from Africa, Southeast 
Asia, the Pacific Islands, and Australia (Allen 
1969). 

Allen (1969) revised the genus and provided a 
key to the 37 species and one subspecies known 
at that time. Baranowski and Slater (1986) 
synonymized L. australis (Fabricius) with L. go- 
nagra (Fabricius). Close to that period, Alayo 
and Grillo (1977), Brailovsky (1976, 1990), 
Yonke (1981), and Brailovsky and Barrera 
(1994) added nine additional new species, thus, 
increasing to 45 the total number of species. 



Osuna (1984) revised the genera of An- 
isoscelini and separated the genus Leptoglossus 
into five genera: Fabrictilis Osuna, Leptoglossus 
Guerin (sensu strictu), Stalifera Osuna, Theognis 
Stal, and Veneza Osuna, but in that monograph 
he did not treat species or discuss relationships 
within each genus. Because Osuna's (1984) new 
genera have more characters in common than 
they have differences, and there are no good and 
consistent external characters that validate their 
creation, the authors do not follow his classifica- 
tion proposal. 

Previous to this paper only four species of the 
genus Leptoglossus were known from Costa 
Rica. This contribution adds six noteworthy re- 
cords, two new species, and a key to the Costa 
Rican species. 



Research Associate, Department of Entomology, California Academy of Sciences, San Francisco, California 941 1 8. 

[167] 



1 68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 6 

Key to the Species of Leptoglossus from Costa Rica 

1 . Thoracic pleura dark with at least two strongly contrasting yellow maculae or yellow 

maculated fascia; thoracic and abdominal venter without numerous small black spots .2 
Thoracic pleura orange to dark red brown without strong yellow markings; thoracic and 

abdominal venter with numerous small black spots 6 

2.(1) Clavus and corium dark brown with strongly contrasting pale yellow veins 3 

Clavus and corium dark brown with veins concolorous or at most bright red, but never 

pale yellow 5 

3.(2) Pronotum without yellow or orange transverse fascia; humeral angles conspicuously 
acute, raised and expanded laterally; pronotal disc with median longitudinal carina 

strongly dentate L. talamancanus, new species 

Pronotum with narrow or wide yellow or orange transverse fascia, running across humeral 
angles; humeral angles acute but not raised and expanded; pronotal disc with median 

longitudinal carina obsolete 4 

4.(3) Pronotum with wide yellow or orange transverse fascia, occupying most of the anterior 

third of the disc; antennal segments II to IV pale ochraceous; posterolateral margins of 

the pronotum serrate L. subauratus Distant 

Pronotum with yellow or orange transverse fascia very narrow; antennal segments II to IV 

bicolored; posterolateral margins of the pronotum entire L. lineosus (Stal) 

5.(2) Pronotal disc with four, small, yellow to orange spots, two on the anterior lobe, and two 
on the posterior margin; thoracic pleura with six strongly contrasting orange spots 

L. tetranotatus Brailovsky and Barrera 

Pronotal disc without small yellow to orange spots; thoracic pleura with a single yellow 

maculae occupying almost the entire acetabulae of each leg 

L. cinctus (Herrich-Schaeffer) 

6.(1) Humeral areas of the pronotum broadly expanded as long and tapering projections; 

humeral angles subacuminate (Fig. 2) L. humeralis Allen 

Humeral areas of the pronotum not broadly expanded; humeral angles rounded or 

subacuminate but never expanded 7 

7.(6) Pronotal disc, scutellum, clavus and corium with numerous black discoidal spots; 
pronotum orange with humeral angles pale yellow; pronotum with anterolateral 

margins dentate L. nigropearlei Yonke 

Pronotal disc, scutellum, clavus and corium without black discoidal spots; pronotum never 
orange with humeral angles pale yellow; pronotum with anterolateral margins entire . . 8 

8.(7) Transverse yellow fascia on corium always present and straight 9 

Transverse yellow fascia on corium irregular, not straight, or fascia absent 10 

9.(8) Antennal segments II and III orange with apical third black; hind tibiae with inner dilation 
considerably shorter than outer (Fig. 11); connexival segments black and with or with- 
out inner margins dark red L. cartagoensis, new species 

Antennal segments II and III entirely yellow to orange; hind tibiae with inner dilation only 
slightly shorter than outer (Fig. 15); connexival segments black or brown with posterior 

margin yellow L. phyllopus (Linnaeus) 

10.(8) Rostrum short, not extending past abdominal sternite II; rostral segment IV shorter than 

antennal segment I L. brevirostris Barber 

Rostrum longer, usually extending well onto abdomen; rostral segment IV longer than an- 
tennal segment 1 11 

1 1.(10) Pronotum dark with two large and strongly rounded yellow spots, dotted with black, and 
occupying most of the anterior disc; male genital capsule with prominently dorsal 

prongs (Fig. 34) L. zonatus (Dallas) 

Pronotum dark without strongly contrasting yellow spots; male genital capsule without 
dorsal prongs (Fig. 39) L. concolor (Walker) 



BRAILOVSKY AND BARRERA: COSTA RICAN LEPTOGLOSSUS 



169 



Leptoglossus brevirostris Barber 
(Figs. 18,40) 

Leptoglossus brevirostris Barber, 1918:35-36. 

Distribution. — Known only from the 
United States and Mexico. 

Material Examined. — COSTA RICA, 1 
female, Prov. Puntarenas, Peninsula Osa, Bosque 
Esquinas, 200 m, March, 1994 (J. Quesada). De- 
posited in INBIO. 

This small-sized species possesses orange tho- 
racic pleura, with numerous small black spots, 
corium with yellow transverse fascia in a "zig- 
zag" pattern, pronotum with anterolateral margin 
entire, and posterolateral dentate, pronotal disc 
concolorous without strong yellow spots, hind 
tibiae phylliform with at least two deep emargi- 
nations, with outer dilation occupying less than 
70% of the length of hind tibiae (Fig. 1 8), rostrum 
reaching the metasternum or the anterior margin 
of abdominal sternite II, with rostral segment IV 
shorter than antennal segment I, and the male 
genital capsule with a deep median notch, lateral 
angles obliquely straight, and long dorsal prongs 
(Fig. 40). 

Leptoglossus cartagoensis, new species 
(Figs. 1, 11,22-23,33,41) 

TYPES. — Holotype male (deposited in IN- 
BIO, courtesy of CAS) from COSTA RICA, 
Puntarenas, 2.5 km., SW Las Alturas, 08°55'N, 
8 1°51'W, February 17, 1991 (N.D.Penny). 

PARATYPES. — COSTA RICA, 2 males, 3 fe- 
males, Prov. Cartago, Grano de Oro (Chirripo), 
Turrialba, 1 120 m, October, 1992 (J. C. Saborio, 
P. Campos); COSTA RICA, 1 male, 10 km, S 
San Jose, Escazu, November, 1992 (J. M. Maes); 
COSTA RICA, 3 males, 8 females, Prov. Guana- 
caste, Estacion Cacao, SW side of Volcan Cacao, 
1000-1400 m, March, June, September, Novem- 
ber, December, 1990 (R. Blanco, C. Chavez); 
COSTA RICA, 1 female, Prov. Puntarenas, San 
Luis, 1040 m, July, 1992 (Z. Fuentes); COSTA 
RICA, 4 females, Prov. Puntarenas, Estacion Las 
Mellizas, Parque Nacional La Amistad (Finca 
Cafrosa), 1300 m, October, 1989 (M. Ramirez, 
G. Mora). Deposited in BMNH, INBIO, MELN, 
UNAM. 



Description. — Male: Body large and ro- 
bust. Head. Tylus unarmed, rounded apically, 
barely surpassing juga and slightly raised in lat- 
eral view; rostrum reaching anterior third of ab- 
dominal sternite IV. Pronotum. Collar wide; 
each pronotal margin entire; humeral angles 
broadly expanded, subacuminate, flattened, and 
obliquely ascending; calli barely elevated, al- 
most impunctate, without two medial tubercles; 
surface densely punctate; disc posteriorly with 
median longitudinal carina obsolete (Fig. 1). 
Legs. Hind tibiae: Outer dilations wide, phylli- 
form, occupying 72% of the length of hind tibiae, 
width about 1.5 times width of inner dilations; 
inner dilations lanceolate, shorter than outer di- 
lations, furnished with numerous small spine- 
like teeth along margins, occupying 52% of the 
length of hind tibiae; inner margins of undilated 
portions with numerous small spine-like teeth 
(Fig. 11). Scutellum. Triangular, wider than 
long, flat, without median longitudinal carina; 
apex rounded to subtruncate. Genitalia. Genital 
capsule: Posteroventral edge with median notch 
deep and rounded, with lateral angles sinuate; no 
dorsal prongs (Fig. 33). Parameres: Figs. 22-23. 

Dorsal coloration. Head black with three nar- 
row stripes dirty orange, one on the middle line, 
the other two close to eyes; antennal segment I 
black with internal face dirty orange; antennal 
segments II and III dirty orange with apical third 
black; segment IV yellow with basal joint black, 
Pronotal disc bright red brown with each prono- 
tal margin black; calli almost black with the area 
between them bright red brown; anterolateral 
margin with anterior third dirty orange, and pos- 
terior third black. Scutellum bright red brown 
with each margin and the apex black. Hemelytra 
with clavus and corium bright red to bright red 
brown, with costal margin, commissure claval, 
anal suture, and basal endocorium black; corium 
with a light yellow, straight, transverse fascia; 
hemelytral membrane uniformly dark. Abdomi- 
nal segments I to VI dirty orange and VII black; 
connexival segments black (some specimens 
with inner face bright red). 

Ventral coloration. Head black with two nar- 
row, dirty orange, stripes close to eyes; bucculae 
bright orange; rostral segments I and II orange 
with inner and lateral face mostly black; rostral 
segments III and IV dirty brownish yellow, with 
inner face and apex of the segment IV black; 
thorax, abdominal sterna, and genital capsule 



170 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 6 



dirty orange-red, with numerous small black 
spots; anterior lobe of metathoracic peritreme 
dirty orange with black central spot, posterior 
lobe dirty orange; coxae and trochanters shiny 
black, with brownish yellow marks; femora or- 
ange, with posterior half mostly black; fore and 
middle tibiae orange or brownish yellow with or 
without a narrow black stripe near the apical 
third; hind tibiae with outer and inner dilation 
black in dorsal view and orange-red in ventral 
view, each with small irregular yellow spot, with 
the rib and margins bright red, and the undilated 
portion orange-yellow; tarsi orange or brownish 
yellow with or without diffuse narrow black 
stripe running laterally; abdominal spiracles 
black or dark brown. 

Description. — Female: Similar to male. 
Connexival segments VIII and IX, and dorsal 
segments VIII and IX black; genital plates bright 
orange red with inner margins of gonocoxae I, 
paratergite VIII and paratergite IX black. 

Measurements. — Male first, then female: 
Head length 2.73 mm, 3.11 mm; width across 
eyes 2.20 mm, 2.43 mm; interocular space 1.21 
mm, 1.31 mm; interocellar space 0.53 mm, 0.61 
mm; preocular distance 1.59 mm, 1.74 mm; 
length antennal segments: I, 2.50 mm, 2.58 mm; 
II, 4.18 mm, 4.48 mm; III, 2.96 mm, 3.26 mm; 
IV, 4.48 mm, 4.86 mm. Pronotal length 3.26 
mm, 3.72 mm; width across frontal angles 1.74 
mm, 1.97 mm; width across humeral angles 6.23 
mm, 7.52 mm. Hind tibiae: Total length 8.32 
mm, 9.50 mm; length outer dilation 6.15 mm, 
7.22 mm; length inner dilation 4.33 mm, 4.94 
mm; width outer dilation 1.74 mm, 2.35 mm; 
width inner dilation 0.76 mm, 0.82 mm. Scutel- 
lar length 1.97 mm, 2.35 mm; width 2.05 mm, 
2.66. Total body length 17.42 mm, 21.00 mm. 

REMARKS. — This species is most similar to L. 
balteatus (Linnaeus) and L. phyllopus (Lin- 
naeus) with the thoracic pleura orange to dark 
red, and usually with numerous small black 
spots, corium with a yellow transverse fascia, 
always present and straight, and the hind tibiae 
with an inner dilation shorter than the outer one. 

Leptoglossus cartagoensis is a larger and more 
robust species than L. balteatus and L. phyllopus, 
with antennal segments II and III orange with 
apical third black, hind tibiae with inner dilation 
considerably shorter than outer, connexival seg- 
ments black with or without inner margins dark 



red, median notch of male genital capsule 
U-shaped and deep (Fig. 33), humeral angles 
broadly expanded, subacuminate and flattened, 
and pronotum bright red brown, with each mar- 
gin mostly black. The closely related species are 
medium sized and elongate, with antennal seg- 
ments II and III entirely yellow or orange, hind 
tibiae with inner dilation slightly shorter than 
outer (Figs. 11, 15), connexival segments black 
or brown, with posterior margin yellow, medial 
notch of male genital capsule V-shaped and not 
deep (Fig. 38), humeral angles subacute and not 
broadly expanded, and pronotum red brown with 
or without anterolateral margin and posterior 
margin yellow, and with the disc anteriorly con- 
colorous or with two yellow spots. 

ETYMOLOGY. — The species is named for the 
Province of Cartago in Costa Rica. 

Leptoglossus cinctus (Herrich-Schaeffer) 
(Figs. 17, 37) 

Anisoscelis cincta Herrich-Schaeffer, 1836:91. 

Distribution. — This is a widely distributed 
species occurring in Mexico, Central America, 
Greater Antilles, and South America. It has been 
recorded from Pacayas, Costa Rica (Allen 1969). 

Material Examined. — COSTA RICA, 1 
male, Prov. Guanacaste, Estacion Maritza, W 
side of Volcan Orosi, 600 m, July, 1989; COSTA 
RICA, 2 females, Prov. Puntarenas, Sabanas 
Curacucha (Potrero Grande), July, 1989 (A. So- 
ils); COSTA RICA, 3 males, 4 females, Prov. 
Guanacaste, Finca Jenny, 30 km, N Liberia, 
Guanacaste National Park, September, October, 
1988. Deposited in INBIO. 

This is one of the most showy and distinctive 
species of Leptoglossus. It is readily recogniz- 
able by the completely yellow pronotal disc, 
contrasting with the remainder of pronotum and 
hemelytra, by the shining orange thoracic pleura, 
with a single yellow spot occupying almost the 
entire acetabulae of each pleuron, the hind tibiae 
with one or two shallow emarginations, with 
outer dilation occupying 65% of the length of 
hind tibiae (Fig. 17), with clavus and corium dark 
orange with veins concolorous, and for the an- 
terolateral and posterolateral margins of the 
pronotum serrate. 



BRAILOVSKY AND BARRERA: COSTA RICAN LEPTOGLOSSUS 



171 



Leptoglossus concolor (Walker) 
(Figs. 13,39) 

Anisoscelis concolor Walker, 1871:128. 

Distribution. — Widely distributed through 
Mexico, Central America, and Greater Antilles. 
It is recorded from Hamburg Farm and Piedras 
Negras, Costa Rica (Allen 1969). 

Material Examined. — COSTA RICA, 1 
male, Prov. Guanacaste, Rio Gongora, 6 km, NE 
Quebrada Grande de Liberia, 700 m, February, 
1992; COSTA RICA, 2 males, Prov. Puntarenas, 
Peninsula Osa, Rancho Quemado, 200 m, April 
and July, 1992 (A. Marin, D. Brenes); COSTA 
RICA, 1 male, Prov. Guanacaste, Aqua Buena, 
Guanacaste National Park, 220 m, June 1992. 
Deposited in INBIO. 

This species may be recognized by the combi- 
nation of thoracic pleura orange, with numerous 
small black spots, corium with yellow transverse 
fascia in a "zig-zag" pattern, pronotum with the 
anterolateral margin entire, and the posterolater- 
ally dentate, pronotal disc concolorous without 
strong yellow areas, calli with a rough surface 
with numerous thick hairs, hind tibiae phylliform 
with at least two deep emarginations, with outer 
dilation occupying less than 70% of the length of 
hind tibiae (Fig. 13), rostrum reaching abdominal 
sternite IV, antennal segment I bicolorous, ros- 
tral segment I longer than antennal segment I, 
and the male genital capsule with straight median 
notch, without dorsal prongs (Fig. 39). Lepto- 
glossus stigma (Herbst), the most closely related 
species, has the pronotal calli smooth, and the 
median notch of the male genital capsule is 
rounded, with dorsal prongs. 

Leptoglossus humeralis Allen 
(Figs. 2, 10, 24-25, 35) 

Leptoglossus humeralis Allen, 1969:126-127. 

Distribution. — This species previously 
was known only from British Guiana (Kartabo) 
and French Guiana (Cayenne) (Allen 1969). 

Material Examined. — COSTA RICA, 1 
male, Prov. Guanacaste, 14 km, S Canas, July 29, 
1990 (F.D. Parker); COSTA RICA, 1 female, 
Prov. Guanacaste, Estacion Pitilla, 9 km, S Santa 
Cecilia, 700 m, February 27-March 2, 1989 (R. 
Miranda); COSTA RICA, 1 male, Prov. Limon, 



Manzanillo (R. N. F. S Gandoca and Man- 
zanillo), 0-100 m, November 20-30, 1992 (F. A. 
Quesada); COSTA RICA, 1 male, 1 female, 
Prov. Limon, Cuatro Esquinas, Parque Nacional 
Tortuguero, August 26-September 6, 1989 (J. 
Solano) and July, 1990 (U. Chavarria); COSTA 
RICA, 3 males, Prov. Limon, Amubri, A. C. 
Amistad, 700 m, July 5-28, 1993, December 
4-21, 1993 and February 1-19, 1994 (G. Gal- 
lardo). Deposited in INBIO, UNAM, USU. 

This species, originally described from two 
females, may be easily recognized by the unique 
humeral expansions, which are produced as 
broad, long, and tapering lateral projections, 
obliquely ascending, and with the humeral angles 
subacuminate (Fig. 2). Other characters inserted 
to recognize this species are the shape of the hind 
tibiae with the outer dilation phylliform with one 
shallow and two deep emarginations occupying 
84% of the length of hind tibiae, with inner 
dilation lanceolate and considerably shorter than 
outer dilation (Fig. 10); the rostrum reaching the 
posterior margin of the fourth abdominal ster- 
nite; and the pronotal disc, as well as the corium, 
without pale areas. Male genital capsule: Pos- 
tero ventral edge with median notch deep and 
rounded; no dorsal prongs (Fig. 35). Parameres: 
Figs. 24-25. 

Leptoglossus lineosus (Stal) 
(Figs. 3, 19, 30) 

Theognis lineosus Stal, 1862:295. 

DISTRIBUTION. — Only known from Mexico. 

Material Examined. — COSTA RICA, 1 
female, Prov. Cartago, July, 1981 (R. Hernan- 
dez). Deposited in INBIO. 

Closely related to L. subauratus Distant with 
thoracic pleura dark, with two well-defined lon- 
gitudinal yellow stripes, and clavus and corium 
dark brown with strongly contrasting pale yellow 
or ochraceous veins. Leptoglossus lineosus is 
easily identified by the following characters: the 
pronotum has a very narrow, orange-yellow, 
transverse band, antennal segments II to TV are 
bicolored with strongly contrasting pale and dark 
areas, and the posterolateral margins of the 
pronotum are entire, whereas in L. subauratus 
antennal segments II to IV are pale ochraceous, 
the pronotal disc has the orange-yellow trans- 
verse band wider and occupying most of the 



172 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 6 



anterior third, and the posterolateral margins of 
the pronotum are serrate (Figs. 3, 6). 

Leptoglossus nigropearlei Yonke 
(Figs. 8,21,36) 

Leptoglossus nigropearlei Yonke, 198 1:21 3-2 1 7. 

Distribution. — This species was known 
only from Panama (Yonke 1981). 

Material Examined. — COSTA RICA, 2 
males, 2 females, Prov. Guanacaste, Estacion 
Cacao, SW side of Volcan Cacao, 1000-1400 m, 
September, 1989 (R. Blanco, C. Chavez). Depos- 
ited in INBIO, UNAM. 

This peculiar species is recognized by the nu- 
merous, small, black, discoidal spots scattered on 
the pronotal disc (Fig. 8), scutellum, clavus and 
corium; by the orange pronotum with the hu- 
meral angles pale yellow; the thoracic pleura 
orange with numerous black spots; the anterolat- 
eral and posterolateral margins of the pronotum 
dentate; and the male genital capsule with a deep 
median notch, without dorsal prongs (Fig. 36). 

Leptoglossus phyllopus (Linnaeus) 
(Figs. 4, 15,38) 

Cimex phyllopus Linnaeus, 1767:731. 

Distribution. — A wide-ranging species oc- 
curring throughout the United States, Mexico, 
Central America and South America. It is re- 
corded from San Jose and Zarzero, Costa Rica 
(Allen 1969). 

Material Examined. — COSTA RICA, 1 
male, 1 female, Prov. Guanacaste, Finca Jenny, 
30 km, N Liberia, Guanacaste National Park, 
September, October, 1988. Deposited in INBIO. 

This species is distinguishable by the humeral 
angles subacuminate, not broadly expanded; 
pronotum concolorous with posterior margin 
never yellow, disc rarely with yellow areas, co- 
rium with yellow transverse fascia always 
straight; thoracic pleura orange to dark red 
brown, with numerous small black spots; hind 
tibiae with inner dilation slightly shorter than 
outer (Fig. 15); and the male genital capsule with 
a shallow V-shaped median notch (Fig. 38). 

It is very similar to L. balteatus, in both color 
and structure, but in this species the posterior 
margin of the pronotum is always yellow, the 



pronotal disc always has contrasting yellow ar- 
eas, and the distribution is restricted to the 
Greater Antilles. 

Leptoglossus subauratus Distant 
(Figs. 6, 14, 43) 

Leptoglossus subauratus Distant, 1881:1 26. 

DISTRIBUTION. — Described from El Salva- 
dor, and later recorded from Guatemala and 
Nicaragua (Allen 1969). 

Material Examined. — COSTA RICA, 1 
male, Prov. Guanacaste, Estacion Santa Rosa, 
300 m, May, 1989; COSTA RICA, 1 female, 
Prov. Cartago, July, 1981 (R. Hernandez). De- 
posited in INBIO. 

This is a medium-sized species, readly recog- 
nizable by the following characters: antennal 
segments I to IV yellow; pronotal disc with wide, 
yellow, transverse fascia between humeral an- 
gles (Fig. 6); thoracic pleura dark with at least 
two strongly contrasting longitudinal yellow fas- 
cia; clavus and corium dark brown with strongly 
contrasting pale yellow veins; hind tibiae with 
outer dilation lanceolate, occupying 60% of the 
length of hind tibiae, and inner dilation lanceo- 
late and about equal in length to outer dilation 
(Fig. 14); and the male genital capsule has a 
shallow median concavity, with lateral angles 
straight, without dorsal prongs. 

Leptoglossus talamancanus, new species 
(Figs. 7, 20, 28, 29, 32, 42) 

TYPES. — Holotype male (INBIO), COSTA 
RICA, Prov. Alajuela, Cano Negro, 20 m, March 
10-29, 1993 (K. Martinez). 

PARATYPES. — COSTA RICA, 1 male, 3 fe- 
males, same data as holotype; COSTA RICA, 1 
female, Prov. Limon, Amubri-Talamanca, 70 m, 
July 1-22, 1992 (G. Galindo); COSTA RICA, 1 
male, Prov. Cartago, Turrialba (Catie), June 
26-29, 1986 (W. Hanson, G. Bohart). Deposited 
in INBIO, UNAM, USU. 

DESCRIPTION. — MALE: Body medium sized. 
Head. Tylus unarmed, rounded apically, extend- 
ing anteriorly to the juga and slightly raised in 
lateral view; rostrum reaching anterior third of 
abdominal sternite III. Pronotum. Collar wide; 
anterolateral and posterolateral margins dentate; 



BRAILOVSKY AND BARRERA: COSTA RICAN LEPTOGLOSSUS 



173 



anterior and posterior margins entire; humeral 
angles extremely acute, raised, and conspicu- 
ously extended laterally; calli slightly elevated; 
area between calli with two large tubercles; sur- 
face densely punctate; pronotal disc with low 
median longitudinal carina, strongly dentate 
(Fig. 7). Legs. Hind tibiae: Outer dilation short, 
lanceolate, entire, occupying 36-41% of the 
length of hind tibiae, width of outer dilation 
wider to width of inner dilation; inner dilation 
lanceolate, slightly shorter than outer, with few 
strong teeth distally, occupying 34% of the 
length of hind tibiae; undilated portion of hind 
tibiae with double row of small spine-like teeth 
along inner margins (Fig. 20). Scutellum. Trian- 
gular, longer than wide, barely elevated, with low 
median longitudinal carina, slightly dentate; 
apex truncated. Genitalia. Genital capsule: Pos- 
teroventral edge with shallow median notch, with 
lateral angles rounded; no dorsal prongs (Fig. 
32). Parameres: Figs. 28-29. 

Dorsal coloration. Head yellow with four nar- 
row stripes light red brown, two on the postocular 
region, and two above; antennal segments I to III 
dirty yellow, and rv creamy yellow; pronotum, 
scutellum, clavus, and corium light red brown, 
with following areas yellow: each pronotal mar- 
gin, the two tubercles on calli, the median longi- 
tudinal carina of pronotal and scutellar disc, apex 
of scutellum, clavus and corial veins, costal and 
apical margin, and claval commissure; corium 
with pale yellow, straight, transverse fascia; he- 
melytral membrane uniformly dark; calli region 
shiny black; connexival segments III to VI black 
to pale brown, with anterior third yellow, and 
segment VII yellow with black spot close to 
posterior third; upper margin of connexival seg- 
ments III to VII yellow; dorsal abdominal seg- 
ments black. 

Ventral coloration. Head, including the buc- 
culae, yellow, with three longitudinal narrow 
stripes pale red brown; rostral segments yellow 
with apex of IV black; thoracic pleura brownish 
yellow with red brown spots irregularly distrib- 
uted, and at least two usually more strongly yel- 
lowish longitudinal fascia; prosterna and 
mesosterna pale brown, with middle furrow of 
mesosterna yellow; metasterna shiny black; ab- 
dominal sterna III to VI shiny black with poste- 
rior margin and longitudinal lateral fasciae shiny 
yellow; abdominal stemite VII bright yellow 
with pale brown irregular spots; pleural margin 



of abdominal sterna III to VII yellow with pale 
brown spots on posterior third of sterna V and 
VI; genital capsule brownish yellow with pale 
brown irregular spots; fore and middle coxae 
chestnut brown with diffuse yellow spots; fore 
and middle trochanters yellow with one chestnut 
brown spot; fore and middle femora, tibiae, and 
tarsi yellow with small, pale brown, irregular 
spots; hind coxae shiny red brown; hind trochan- 
ter chestnut orange with diffuse yellow spots; 
hind femora with internal face mostly black to 
red brown, and external face dirty yellow an on 
each face the spines are shiny black; hind tibiae 
yellow with inner and outer dilations shiny black, 
and with small irregular spots yellow on each 
dilation; hind tarsi yellow; anterior lobe of 
metathoracic peritreme black, with the margins 
yellow, posterior lobe yellow. 

Description. — Female: Similar to male. 
Connexival segments VIII and IX yellow; ab- 
dominal segments VIII and IX black with poste- 
rior margin yellow; genital plates yellow with 
inner third of paratergite IX and gonocoxae I pale 
brown; abdominal sterna III to VII yellow, and 
densely covered with shiny black spots; fore and 
middle femora yellow with two or three narrow 
stripes pale red brown; hind femora yellow, with 
the spines shiny black and some irregular spots 
and short stripes shiny red brown. 

Variation in Species. — 1. Antennal seg- 
ments II and III yellow with inner face light 
brown. 2. Antennal segment IV creamy yellow 
with apical third pale brown. 3. Hind femora 
mostly shiny black with few external narrow 
stripes yellow. 

MEASUREMENTS. — First male, then female: 
Head length 1.88 mm, 2.12 mm; width across 
eyes 1.68 mm, 1.93 mm; interocular space 0.88 
mm, 1.04 mm; interocellar space 0.34 mm, 0.48 
mm; preocular distance 1.10 mm, 1.26 mm; 
length antennal segments: I, 2.08 mm, 2.12 mm; 
II, 2.96 mm, 3.20 mm; III, 2.24 mm,2.28 mm; 
IV, 3.52 mm, 3.44 mm. Pronotal length 2.12 
mm, 2.60 mm; width across frontal anles 1.40 
mm, 1.72 mm; width across humeral angles 4. 16 
mm, 5.64 mm. Hind tibiae: Total length 5.76 
mm, 6.32 mm; length of outer dilation 2.10 mm, 
2.60 mm; length of inner dilation 1 .98 mm, 2.23 
mm; width of outer dilation 0.62 mm, 0.88 mm; 
width of inner dilation 0.43 mm, 0.68 mm. 
Scutellar length 1.32 mm, 1.60 mm; width 1.28 



174 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 6 



mm, 1.40 mm. Total body length 11.00 mm, 
13.82 mm. 

REMARKS. — Like L. subauratus, this species 
has the thoracic pleura dark, with at least two 
strongly contrasting, longitudinal yellow fascia, 
the clavus and corium dark brown or pale red 
brown with pale yellow veins, hind tibiae with 
outer and inner dilations lanceolate, and antennal 
segments I to III yellow to dirty yellow. The two 
species can be separated on the basis of the 
following combination of characters. In L. 
subauratus the pronotal disc has a wide yellow 
transverse fascia running across humeral angles, 
antennal segment IV orange to orange-yellow, 
humeral angles acute, but not raised and not 
greatly expanded (Fig. 14), median longitudinal 
carina obsolete and rostrum reaching the poste- 
rior margin of abdominal sternite V. In L. tala- 
mancanus, the antennal segment IV is creamy 
yellow, the rostrum reaching anterior third of 
abdominal sternite III, the pronotal disc without 
yellow transverse fascia, with the median longi- 
tudinal carina strongly dentate, and the humeral 
angles extremely acute, raised, and conspicu- 
ously extended laterally (Fig. 20). 

Leptoglossus lineosus has antennal segment I 
mostly black, and segment IV red-brown with 
apex pale yellow, the pronotal disc has a narrow 
yellow transverse fascia across humeral angles, 
and rostral segments I to IV are mostly black. In 
L. talamancanus the rostral segments are yellow 
with apex of segment IV black. 

ETYMOLOGY. — The species is named for the 
City of Talamanca in Costa Rica. 

Leptoglossus tetranotatus Brailovsky and 
Barrera 

(Figs. 5, 16,26-27,31,44) 

Leptoglossus tetranotata Brailovsky and Barrera, 
1994:60-62. 

Distribution. — The only known record 
came from the original description in which the 
holotype and only known specimen was col- 
lected in French Guiana (Brailovsky and Barrera 
1994). 

Material Examined. — COSTA RICA, 1 
male, Prov. Limon, Estacion Cuatro Esquinas, 
Tortuguero National Park, March 27, 1992 and 
April 29, 1992 (D. Garcia); COSTA RICA, 3 
males, Prov. Puntarenas, Peninsula Osa, Bosque 



Esquinas, 200 m, April and May, 1994 (J. Que- 
sada); COSTA RICA, 1 male, 1 female, Prov. 
Puntarenas, Rancho Quemado, April, 1991 (F. 
Quesada); COSTA RICA, 2 males, 2 females, 
Prov. Puntarenas, Estacion Sirena, Corcovado 
National Park, March 21, 1992, and April 21, 
1992 (Z. Fuentes); COSTA RICA, 1 female, 
Prov. Limon, Rio Sardinas, R. N. F. S. Barra de 
Colorado, 10 m. August 14-22, 1993 (F. Araya); 
COSTA RICA, 1 female, Prov. Limon, Man- 
zanillo, R. N. F. S., Gandoca and Manzanillo, 
October 22 and November 1 1, 1992 (K. Taylor); 
COSTA RICA, 1 female, Prov. Limon, Cerro 
Tortuguero, Tortuguero National Park, 100 m, 
November, 1989 (J. Solano). Deposited in CAS, 
INBIO, UNAM. 

This species may be distinguished by the fol- 
lowing characters: pronotal disc with four, yel- 
low to orange, small spots, two on the anterior 
lobe and the other pair on the posterior margin 
(Fig. 5); thoracic pleura with six strongly con- 
trasting orange spots (three on propleura, two on 
mesopleura, and one on metapleura); antennal 
segments I to IV orange-yellow; hind tibiae with 
outer dilation phylliform, with two or three emar- 
ginations, occupying 52% of the length of hind 
tibiae, and inner dilation lanceolate, occupying 
48% of the length of hind tibiae (Fig. 1 6); and the 
male genital capsule with a deep median notch, 
lateral angles almost straight, and dorsal prongs 
barely exposed (Fig. 31). Parameres: Figs. 
26-27. 

Leptoglossus zonatus (Dallas) 
(Figs. 9, 12, 34) 

Anisoscelis zonata Dallas, 1 852:452. 

Distribution. — This is a widely distributed 
species occurring in the United States, Mexico, 
Central America and South America. From Costa 
Rica, it is recorded from Puntarenas, Turrialba 
and San Jose (Allen 1969). 

Material Examined. — COSTA RICA, 1 
female, Prov. Guanacaste, Rio San Josecito, 
Guanacaste National Park, 960 m, April 3-4, 
1987 (Holsenthal, Hamilton, Heyn); COSTA 
RICA, 3 males, 2 females, Prov. Guanacaste, 
Finca Jenny, 30 km, N Liberia, December, 1 988; 
COSTA RICA, 1 female, Prov. Guanacaste, Es- 
tacion Cacao, SW side of Volcan Cacao, 
1000-1400 m, March, 1988; COSTA RICA, 1 



BRAILOVSKY AND BARRERA: COSTA RICAN LEPTOGLOSSUS 



175 



male, Prov. Guanacaste, vicinity Estacion Mur- 
cielago, 8 km, SW Cuajiniquil, 100 m, February, 
1989; COSTA RICA, 1 female, Prov. Guana- 
caste, La Pacifica, February 20, 1983 (G. C. 
Eickwort). Deposited in INBIO, UMSP, USU. 

Recognized by the relatively large size, prono- 
tal disc with two distinct pale yellow rounded 
spots (Fig. 9), corium with a yellow transverse 
fascia in a zig-zag pattern, hind tibiae with outer 
dilation phylliform (Fig. 12), antennal segment 
II bicolorous, and male genital capsule with me- 
dian notch deeply rounded, with dorsal prongs 
prominently acute and projecting medially 
(Fig. 34). 

Acknowledgments 

We gratefully acknowledge the loan of mate- 
rial from the following individuals and institu- 
tions: Janet Margerison-Knight, The Natural 
History Museum, London (BMNH); Norman D. 
Penny, and Keve Ribardo, California Academy 
of Sciences, San Francisco (CAS); Angel Solis 
and Jesus Ugalde, Instituto Nacional de Biodi- 
versidad, Costa Rica (INBIO); Philip J. Clausen, 
University of Minnesota, Insect Collection 
(UMSP); Thomas J. Henry, Systematic Entomol- 
ogy Lab., USDA, c/o Smithsonian Institution, 
Washington D. C. (USNM); Wilford J. Hanson, 
Utah State University (USU). Special thanks to 
Cristina Urbina for the preparation of some of the 
illustrations. 

Resumen 

Se analizo el genero Leptoglossus para Costa 
Rica, describiendose e ilustrandose dos nuevas 
especies L. cartagoensis y L. talamancanus; L. 
brevirostris Barber, L. humeralis Allen, L. lineo- 
sus (Stal), L. nigropearlei Yonke, L. subauratus 
Distant y L. tetranotatus Brailovsky y Barrera 
son citados por primera vez; nuevos registros 
para L. cinctus (Herrich-Schaeffer), L. concolor 
(Walker), L. phyllopus (Linnaeus) y L. zonatus 
(Dallas) son incluidos; se ofrece una clave para 
separar las especies de Costa Rica conocidas. 



Literature Cited 

Alayo P. D. and H. Grillo Ravelo. 1977. Los 
hemipteros de Cuba- XVI. El genero Leptoglossus 
Guerin (Hemiptera:Coreidae) en Cuba. Centro 
Agricola, mayo-agosto 1977:91-1 1 1. 

Allen, R. C 1 969. A revision of the genus Leptoglos- 
sus Guerin (Hemiptera: Coreidae). Entomol. Am. 
45:35-140. 

Baranowski, R. M. and J. A. Slater. 1986. Arthro- 
pods of Florida and Neighboring land areas Volume 
1 2. Coreidae of Florida (Hemiptera: Heteroptera). 
Florida Department of Agriculture and Consumer 
Services. Contribution 630:1-82. 

Brailovsky, H. 1976. Contribution al estudio de los 
Hemiptera-Heteropterade Mexico: VIII. Una nueva 
especie de Leptoglossus Guerin (Coreidae-Corei- 
nae) y datos sobre distribution de las especies mexi- 
canas del genero. An. Inst. Biol. Univ. Nac. Auton. 
Mexico, Ser. Zool. 47:35-42. 

. 1 990. Generos nuevos y Especies nuevas de 

Coreidos Neotropicales (Hemiptera-Heteroptera- 
Coreidae:Acanthocerini, Leptoscelidini y An- 
isoscelidini). An. Inst. Biol. Univ. Nac. Auton. 
Mexico, Ser. Zool. 61:107-123. 

Brailovsky, H. and E. Barrera. 1994. Description 
de cuatro especies y una subespecie nuevas de la 
Tribu Anisoscelidini (Hemiptera-Heteroptera- 
Coreidae). An. Inst. Biol. Univ. Nac. Auton. 
Mexico, Ser. Zool. 65:45-62. 

OSUNA, E. 1984. Monografiade la Tribu Anisoscelid- 
ini (Hemiptera, Heteroptera, Coreidae) I. Revision 
Generica. Bol. Entomol. Venez. N. S. 3 
(5-8):77-148. 

YONKE, T. R. 1 98 1 . Description of two new species of 
Neotropical Leptoglossus Guerin (Hemiptera: 
Coreidae). Proc. Entomol. Soc. Wash. 83:213-221. 



1 76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 6 

PLATE 1 

Figs. 1-9. Pronotum of Leptoglossus spp. L. cartagoensis, new species (1); L. humeralis Allen (2); 
L. lineosus (Stal) (3); L. phyllopus (Linnaeus) (4); L. tetranotatus Brailovsky and Barrera (5); L. 
subauratus Distant (6); L. talamancanus, new species (7); L. nigropearlei Yonke (8); L. zonatus 
(Dallas) (9). 

PLATE 2 

Figs. 10-21. Hind tibiae of Leptoglossus spp. L. humeralis Allen (10); L. cartagoensis, new species 
(11); L. zonatus (Dallas) (12); L. concolor (Walker) (13); L. subauratus Distant (14); L. phyllopus 
(Linnaeus) (15); L. tetranotatus Brailovsky and Barrera (16); L. cinctus (Herrich-Schaeffer) (17); L. 
brevirostris Barber (18); L. lineosus (Stal) (19); L. talamancanus, new species (20); L. nigropearlei 
Yonke (21). 

PLATE 3 

Figs. 22-29. Parameres of Leptoglossus spp. L. cartagoensis, new species (22-23); L. humeralis 
Allen (24—25); L. tetranotatus Brailovsky and Barrera (26-27); L. talamancanus, new species 
(28-29). Figs. 30-40. Caudal view of the male genital capsule of Leptoglossus spp. L. lineosus (Stal) 
(30); L. tetranotatus Brailovsky and Barrera (31); L. talamancanus, new species (32); L. cartagoensis, 
new species (33); L. zonatus (Dallas) (34); L. humeralis Allen (35); L. nigropearlei Yonke (36); L. 
cinctus (Herrich-Schaeffer) (37); L. phyllopus (Linnaeus) (38); L. concolor (Walker) (39); L. brevi- 
rostris Barber (40). 

PLATE 4 

Fig. 4 1 . Leptoglossus cartagoensis, new species. 

PLATE 5 

Fig. 42. Leptoglossus talamancanus, new species. 

PLATE 6 

Fig. 43. Leptoglossus subauratus Distant. 

PLATE 7 

Fig. 44. Leptoglossus tetranotatus Brailovsky and Barrera. 



BRAILOVSKY AND BARRERA: COSTA RICAN LEPTOGLOSSUS 

PLATE 1 



177 





Sj-i ...,.s- &>; ;!: 





P^6 








178 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 6 



PLATE 2 




16 V 




BRAILOVSKY AND BARRERA: COSTA RICAN LEPTOGLOSSUS 

PLATE 3 



179 




180 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 6 



PLATE 4 




BRAILOVSKY AND BARRERA: COSTA RICAN LEPTOGLOSSUS 

PLATE 5 




182 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 6 



PLATE 6 










BRAILOVSKY AND BARRERA: COSTA RICAN LEPTOGLOSSUS 



183 



PLATE 7 




CALIFORNIA ACADEMY OF SCIENCES, 1998 
Golden Gate Park 
San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 7, pp. 185-215, 14 figs., 1 pi., 4 tables. April 27, 1998 



A REVISION OF THE SNAKE-EEL GENUS CALLECHELYS 

(ANGUILLIFORMES: OPHICHTHIDAE) WITH THE 

DESCRIPTION OF TWO NEW INDO-PACIFIC SPECIES AND A 

NEW CALLECHELYIN GENUS 



By 

John E. McCosker 

Department of Aquatic Biology, California Academy of Sciences f\D& /£ 9 13 30 
Golden Gate Park, San Francisco, California 941 18 



Species of the tropical snake-eel genus Callechelys, family Ophichthidae, subfamily Ophichthinae, 
are reviewed. An identification key for the species of the tribe Callechelyini and synonymies, 
illustrations, and diagnoses are provided for each of the valid species of Callechelys: C. bilinearis, 
C. bitaeniata, C. colostoma, C. cliffi, C. eristigma, C. galapagensis, C. guineensis, C. leucoptera, C. 
lutea, C. marmorata, C. muraena, and C. spiingeri. Callechelys papulosa n. sp. is described from 
Papua, New Guinea, and differs in its morphometry, coloration, and vertebral number. 
Callechelys randalli n. sp. is described from Fatu Hiva, Marquesas, and differs from the 
closely-related C. catostoma in its body elongation and higher vertebral numbers. A neotype for 
C. catostoma is designated. Callechelys striatus Smith is synonymized with C. catostoma. 
Callechelys sibogae Weber is referred to Bascanichthys and C. maculata Chu, Wu and Jin is 
provisionally referred to Yirrkala. The status of all nominal species of Callechelys is confirmed. 
Xestochilus gen. nov. is erected for Callechelys nebulosus Smith; it differs from other callechelyins 
in the unique condition of its snout, lips, dentition, and branchiostegal rays. Its range is extended 
to include Polynesia, Micronesia, Indonesia and the Red Sea. The distribution of C catostoma is 
extended to Hawaii, the western Indian Ocean, and the Red Sea. 

Received November 19, 1997. Accepted December 19, 1997. 

Although the snake-eels and worm-eels of the blatt 1972. 1995); the western Indian Ocean 
family Ophichthidae are known to have achieved (Smith 1962); South Africa (McCosker and Cas- 
perhaps the greatest diversity of body form and tie 1986); and Australia (McCosker in prep.), 
adaptive radiation within the anguilliform fishes. The osteology and intrafamilial relationships of 
few can be accused of being beautiful. The spe- several species were described by McCosker 
cies of the genus Callechelys are exceptional. ( 1977) and the leptocephali of the western Atlan- 
however, in their possession of an elongate mane tic species by Leiby (1989). A considerable 
and generally colorful livery, such that Kaup number of specimens of Callechelys have accu- 
(1856a, b) was correct in creating the generic mulated in recent years, allowing for the first 
name to include the Greek KaAAoq, meaning time a revision of the genus. In doing so, I dis- 
beautiful. Since Storey's (1939) review, the spe- covered that C nebulosus, described by Smith 
cies of Callechelys have only received regional (1958) from the western Indian Ocean, is wide- 
treatments: the western Atlantic (McCosker et al. spread in the lndo-Pacific and has affinities else- 
1989); the eastern Atlantic (Blache and Cadenat where within the tribe Callechelyini. 
1971); the eastern Pacific (McCosker and Rosen- 

[185] 



186 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



The purpose of this paper is to provide an 
update of recent taxonomic actions concerning 
Callechelys, a key to their identification, a listing 
of all nominal species and their status, the de- 
scription of two new Indo-Pacific species, and 
the creation of a new generic name for C. nebu- 
losus. 

Materials and Methods 

Measurements are straight-line, made either 
with a 300 mm ruler with 0.5 mm gradations (for 
total length, trunk length, and tail length) and 
recorded to the nearest 0.5 mm, or with dial 
calipers (all other measurements) and recorded to 
the nearest 0. 1 mm. Body length comprises head 
and trunk lengths. Head length is measured from 
the snout tip to the posterodorsal margin of the 
gill opening; trunk length is taken from the end 
of the head to mid-anus; maximum body depth 
does not include the median fins. Head pore 
terminology follows that of McCosker et al. 
(1989:257), such that the supraorbital pores are 
expressed as the ethmoid pore + pores in supraor- 
bital canal, e.g., 1 + 3, and the infraorbital pores 
are expressed as pores along the upper jaw + 
those in vertical part of canal behind eye (the 
"postorbital pores"), e.g., 4 + 2, in that frequently 
the last pore included along the upper jaw is part 
of the postorbital series. Gill arch examination 
was accomplished after removal and clearing and 
counterstaining with alcian blue and alizarin red 
dyes (Dingerkus and Uhler 1977). Vertebral 
counts (which include the hypural) are taken 
from radiographs. The mean vertebral formula 
(MVF) expresses the average of predorsal 
(which begins above the head in all Callechelys, 
indicated in the formula by "H"), preanal, and 
total vertebrae (Bohlke 1982). The number of 
specimens examined for vertebral analysis are 
listed after "total vertebrae" as "n = ". Institu- 
tional abbreviations follow the Standard Sym- 
bolic Codes for Institutional Research 
Collections in Herpetology and Ichthyology 
(Leviton et al. 1985). 



Key to the Species of the Tribe 
Callechelyini 

la. Anal fin absent 2 

lb. Anal fin present 6 

2a. Anterior nostril a hole, without a raised 
rim; four supraorbital pores; head brown 
or black, with white markings (not spot- 
ted), dorsal fin white . Letharchus ... 3 

2b. Anterior nostril tubular; three supraorbital 
pores; head and dorsal fin spotted or mot- 
tled Paraletharchus ... 5 

3a.Trunk and tail cream-colored, overlain 
with a broad brown mid-lateral stripe run- 
ning the entire length of the body; verte- 
brae 157-164 

Letharchus aliculatus McCosker 1974 
(Brazil) 

3b.Trunk and tail uniform dark brown to 
black; vertebral range 135-151 4 

4a. Supraorbital pores linear when viewed 

from above; vertebrae 135-142 

Letharchus velifer Goode and Bean 1882 
(western Atlantic) 

4b. Second supraorbital pore displaced me- 
dially when viewed from above; vertebrae 

144-151 

Letharchus rosenblatti McCosker 1974 
(eastern Pacific) 

5a. Background body color dark tan to dark 
brown, spotted anteriorly, the spots coa- 
lescing along the posterior trunk and tail 
to become uniformly dark; vertebrae 

156-167 

Paraletharchus pacificus (Osburn and 
Nichols 1916) (eastern Pacific) 

5b. Background body color cream to tan, 
overlain with numerous brown spots; ver- 
tebrae 170-180 

Paraletharchus opercularis (Myers and 
Wade 1941) (Galapagos Islands) 

6a. Incision on underside of snout, if present, 
not extending beyond anterior nostril base 
(Figs, lb— d); vomerine and/or anterior in- 
termaxillary teeth absent 7 

6b. Median groove on underside of snout ex- 
tends to and beyond anterior nostril bases 
(Fig. la); vomerine and intermaxillary 
teeth present Callechelys ... 9 

7a. Underside of snout with a raised platform 
before, between, and ending behind ante- 
rior nostril tubes (Figs, lb, c), appearing 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



187 



laterally as a horizontal slit above lip; an- 
terior intermaxillary teeth present; a short 
crease in underside of snout, not extend- 
ing beyond anterior nostril bases; verte- 
brae 135-165 8 

7b. Underside of snout without a raised plat- 
form between tubular anterior nostrils 
(Fig. Id); underside of snout without a me- 
dian crease; anterior intermaxillary and 
vomerine teeth absent; vertebrae 148-152 

Aprognathodon platyventris 

Bohlke 1967 (western Atlantic) 
8a. An elongate Y-shaped platform on under- 
side of snout (Fig. lb); anterior intermaxil- 
lary teeth partially hidden by skin folds; 
snout and nape smooth; vertebrae 

157-165 

Xestochilus nebulosus (Smith 1962) (Red 
Sea and Indian Ocean to central Pacific) 
8b. A V-shaped platform on underside of 
snout (Fig. lc); anterior intermaxillary 
teeth exposed; snout and nape with numer- 
ous papillae; vertebrae 135 

Leuropharus lasiops Rosenblatt and 
McCosker 1970 (eastern Pacific) 
9a. Body evenly colored, marbled or striped, 

not spotted 10 

9b. Body covered with numerous brown to 
black spots 16 

10a. Body uniform brown to black, not striped 
or marbled 11 

10b. Body not uniformly colored, with longitu- 
dinal stripes, marbling, or a combination 
of both 12 

1 la. Body chocolate brown to dark brown, fins 
colorless; face smooth; tail 2.1-2.3 in TL 
.... Callechelys leucoptera (west Africa) 

1 lb. Body dark black, fins black like body (a 
white spot beneath eye); face covered 

with pits; tail 2.7 in TL 

Callechelys papulosa (New Guinea) 

12a. A prominent fleshy ridge from the snout 
tip to dorsal fin origin; vomerine teeth 
neatly biserial; coloration yellow to tan, 
speckled and spotted with darker, coalesc- 
ing spots Callechelys 

marmorata (east Africa to central Pacific) 

12b. Fleshy ridge along mid-snout region ab- 
sent; vomerine teeth few, not in two rows; 
bold stripes along body 13 

13a. Body with two dark lateral stripes along 
body and one along the dorsal fin; body 



moderately elongate, its depth 28-36 in 

TL Callechelys bilinearis 

(western Atlantic eastward to St. Helena 
Island) 

13b. Body with two stripes, the dorsal fin mar- 
gin black, the fin pale, and a single mid- 
body stripe along the length of the trunk 
and tail, the lateral line within the band; 
body variously elongate, its depth 48-76 
in TL 14 

14a. Tail very short, more than 3.5 in TL; a 

pair (rarely two pairs) of large recurved in- 
termaxillary fangs; ethmoidal pore (at 
snout tip) absent 15 

14b. Tail 2.5-2.7 times in TL; four intermaxil- 
lary teeth, none enlarged as recurved 

fangs; ethmoidal pore present 

Callechelys bitaeniata (western Indian 
Ocean) 

15a. A small but conspicuous barbel on lip be- 
tween anterior and posterior nostrils; body 

depth 48-73; vertebrae 192-205 

Callechelys catostoma (east Africa to cen- 
tral Pacific) 

15b. Lip continuous between anterior and pos- 
terior nostrils, no conspicuous barbel; 
body depth 59-73; vertebrae 220-224 

Callechelys randalli 

(Marquesas Islands, central Pacific) 

16a. Color pattern of numerous fine dark spots, 
not much larger than eye diameter, over a 
pale tan or brown body; fins with a dis- 
tinct white or pale edging 17 

16b. Color pattern pale with dark round or ob- 
long spots and blotches, mostly larger 
than snout; fins without a distinct white or 
pale edging 18 

17a. Tail 2.2-2.4 in TL; depth 19-31 in TL; 
color tan with numerous fine brown spots 

on body and fins 

Callechelys cliffi (eastern Pacific) 

17b. Tail 2.5-2.7 in TL; depth 27-38 in TL; 
color pale, overlain with numerous eye- 
sized brown spots and mottling 

Callechelys muraena (western Atlantic) 

18a. Tail very short, 3.2-3.7 in TL; depth 

38-52 in TL; color mostly cream, overlain 
with numerous dark spots about as long as 

snout 

Callechelys eristigma (eastern Pacific) 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



18b. Tail longer, 2.1-3.2 in TL; depth 17-39 in 
TL; color various, but with dark spotting 
over a paler background 19 

19a. Coloration pale to yellow, overlain with 
spots mostly smaller than snout; anal fin 
pale; tail 2.4-2.8 in TL; depth 38-53 in 

TL Callechelys lutea 

(Hawaii to Midway Island) 

19b. Coloration pale, spotting mostly larger 
than snout, coalescing to form large 
blotches in some species; anal fin spotted; 
tail 2.2-3.1 in TL; depth 26-59 in TL 
20 

20a. Tail 2.8-3.1 in TL; depth 45-59 in TL 
21 

20b. Tail 2.2-2.3 in TL; depth 26-38 in TL 
Callechelys galapagensis (Galapagos Is- 
lands) 

21a. Tail 2.8-2.9 in TL; total vertebrae 

169-171; dorsal spots of large individuals 

coalesce into dark saddles 

Callechelys springeri (North Carolina and 
Florida, western Atlantic) 

21b. Tail 2.9-3.1 in TL; total vertebrae 

171-186; dorsal spots of large individuals 

not coalesced into saddles 

Callechelys guineensis (eastern and west- 
ern Atlantic) 

Genus Callechelys Kaup, 1856 

Callechelys Kaup, 1 856a:51 (type species Callechelys 
Guichenoti Kaup, 1 856 = Dalophis marmorata 
Bleeker 1853, by monotypy). 

Cryptopterygium Ginsburg 1951:482 (type species 
Cryptopterygium holochroma Ginsburg 1951 = 
Gordiichthys springeri Ginsburg 1951, by original 
designation and by monotypy). 

Diagnosis. — Ophichthid eels, subfamily 
Ophichthinae, tribe Callechelyini (sensu 
McCosker 1977) with body moderately to very 
elongate, laterally compressed throughout body 
and trunk, and longer than tail. Dorsal fin origin 
on nape, above supraoccipital, well before gill 
openings. Pectoral fins absent. Snout acute, 
rounded at tip, overhanging lower jaw, grooved 
on underside. Gill openings low lateral to entirely 
ventral, converging forward, their length much 
greater than isthmus. Head pores reduced; three 
supraorbital pores, two preopercular pores, and 
three pores in supratemporal canal. Teeth coni- 



cal, slender, small and uniserial on jaws, those of 
intermaxillary larger and separated from those of 
vomer by a gap. Body coloration variable: 
striped, barred, spotted, mottled or uniform. 

ETYMOLOGY. — From the Greek KctAAog 
(beauty) and ey^s^ot; (eel). Treated as feminine 
according to Opinion 9 1 5 of the Bulletin of Zoo- 
logical Nomenclature, 1970. 

REMARKS. — A more comprehensive descrip- 
tion of Callechelys osteology is available in 
McCosker (1977) and McCosker et al. (1989). 

Callechelys bilinearis Kanazawa 
(Fig. 2; Tables 1,2,4) 

Callechelys bilinearis Kanazawa 1952:72 (Bermuda, 
holotype FMNH 48973). 

DIAGNOSIS. — Moderately elongate, body 
depth 28-36 in TL; head 12-14 in TL; tail 
2.6-3.6 in TL; maxillary teeth 8-12, dentary 
teeth 12-19; boldly colored with two dark lateral 
stripes along body and one stripe along dorsal fin; 
MVF H-97-160; total vertebrae 155-167 (n = 
25). 

SIZE. — Reported to 1724 mm by Cervigon 
(1973); the largest observed by McCosker et al. 
(1989) was 871 mm. 

ETYMOLOGY. — From the Latin bilinearis 
(two-lined), an adjective. 

DISTRIBUTION. — An insular species, wide- 
spread throughout the tropical western Atlantic, 
and the most common species of Atlantic 
Callechelys (McCosker et al. 1989:307). Re- 
cords include Bermuda, throughout the West In- 
dies, from islands of Central and South America, 
and from St. Helena and Ascension islands in the 
mid-Atlantic. Not known from North American 
coastal waters or from the eastern Atlantic. Cap- 
tured using rotenone between depths of 1-22 m. 

REMARKS. — This species is unmistakable in 
its appearance and would not be mistaken for any 
of its congeners. 

MATERIAL EXAMINED. — A listing of the ma- 
terial examined is in McCosker et al. 
(1989:307-308); not included was SIO 70-376, 
285 mm, from San Bias, Panama, which was 
captured in 1 m of water, expanding the range of 
depth of capture reported therein. 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



189 




B 










■■■■■-; 




111 
111 

HP I law 




Figure 1. Underside of the snouts of: A) Callechelys catostoma, CAS 91786, 621 mm; B) Xestochihis nebulosus, CAS 
92202, 392 mm; C) Leuropharus lasiops, SU 57313, 174 mm (from Rosenblatt and McCosker 1970, fig. 8); D) Aprognatho- 
don platyventris, ANSP 138770, 244 mm (from McCosker et al. 1989, fig. 289). 



Callechelys bitaeniata (Peters) 
(Fig. 3; Tables 1,2,4) 

Ophichthys bitaeniatus Peters 1878:556, unnumbered 
plate, figs. 2-2a (Mombasa, east Africa, holotype 
ZMB 10052). 

Callechelys bitaeniatus, Storey 1939:68. 



Callechelys canaliculus Smith 1958:836, PI. 27 
(Pinda, Mozambique, holotype RUSI 103). 

DIAGNOSIS. — Elongate, body depth 48-60 in 
TL; head 14.0-16.7 in TL; tail 2.5-2.64 in TL; 2 
pairs of fangs on intermaxillary, 3-5 on vomer, 
maxillary teeth 5-8, dentary 9-10; body colora- 



190 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



Table 1. Tail proportion (in thousandths) and vertebral number of the valid species of Callechelys. 



Species 


TailATL 


Vertebral Range 


Vertebral Mean 


N 


bilinearis 


278-385 


155-167 


160.0 


25 


bitaeniata 


379-400 


185-194 


189.0 


5 


catostoma 


278-312 


192-205 


198.6 


27 


cliffi 


417^55 


149-158 


155.0 


14 


eristigma 


270-312 


154-163 


158.0 


37 


galapagensis 


435^55 


170-174 


172.0 


4 


guineensis 


323-345 


172-182 


178.0 


12 


leucoptera 


435-476 


162-165 


164.0 


4 


lutea 


357-417 


210-219 


214.3 


8 


mannorata 


348-356 


174-183 


180.0 


21 


muraena 


370-400 


139-144 


142.0 


7 


papulosa 


490 


215 


— 


1 


randalli 


278-288 


220-224 


222.0 


2 


springeh 


345-357 


168-171 


169.0 


3 



tion pale to milky yellow, overlain on dorsal 2/5 
of flank by a wide, dark brown stripe from mid- 
head to tail, the stripe stopping at the dorsal fin 
base below a narrow, pale stripe above the base; 
the remainder of the fin dark brown, darkest at 
the margin; anal fin pale, the margin of males 
irregularly dusky in the anterior 2/3 of its length; 
MVFH-109-189.0; total vertebrae 185-194 
(n = 5). 

SIZE. — The largest reported specimen is 820 
mm (Smith 1958, 1962), a male from Pinda, 
Mozambique. 

ETYMOLOGY. — From the Latin bitaeniata 
(two-striped), an adjective. 

DISTRIBUTION. — From Mozambique, Ken- 
ya, and Aldabra Atoll. 

REMARKS. — Smith (1958) 1 described Cal- 
lechelys canaliculars on the basis of five adult 
specimens from east Africa, and later (1962), 
after having compared them to Peters' type, 
placed it in the synonymy of C. bitaeniata. I, too, 
have examined the type of O. bitaeniatus and 
some of Smith's specimens of C. canaliculars 
(including a radiograph of the type), and after 
examining the vertebral counts of those speci- 
mens, agree with his action. The striped appear- 
ance of C. bitaeniata might cause it to be 
mistaken for C. catostoma or C. randalli; how- 



ever, both are very different in their dentition and 
body proportions. 

Material Examined. — ZMB 10052, 311 
mm, Mombasa, the holotype of Ophichthys bi- 
taeniatus. RUSI 103, 780 mm, the holotype of 
Callechelys canaliculars (examined only from 
a radiograph), Mozambique, Pinda. RUSI 5217, 
2(760-764 mm), Mozambique, Pinda. RUSI 
31804, 565 mm (tail broken anJ regrown, head 
and trunk length 360 mm), Mozambique, Baz- 
aruto. 

Callechelys catostoma (Forster) 
(Figs, la, 4; Tables \-4) 

Sphagebranchus catostomus Forster in Bloch and 

Schneider 1801:536, based on Forster manuscript 

(Tahiti, no types known; neotype CAS 91793). 
Callechelys melanotaenia Bleeker 1864:66 (Am- 

boina, holotype RMNH 7180). 
Callechelys melanotaenia Bleeker 1865:pl. 193, 

fig. 2. " 
Ophichthys melanotaenia, Gunther 1870:87. 
Leptenchelys pinnaceps Schultz in Schultz et al. 

1953:79^ fig. 16 (Bikini Atoll, holotype USNM 

141691). 
Callechelys striatus Smith 1958:838, pi. 27, fig. c 

(Baixo Pinda, Mozambique, holotype RUSI 96). 
Callechelys catostomus, Randall and Wheeler 

1991:761. 



Smith's 1958 publication, cited as 1957 by all subsequent authors, was published on 23 April 1958. 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



191 




1 



192 



PROCEEDINGS OF THE CALIFORNIA ACADEMV OF SCIENCES 

Volume 50, No. 7 







J! 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



193 




Figure 4 Callechelys catostoma, BPBM 10190, 520 mm. Photograph by J. E. Randall. 



DIAGNOSIS. — Very elongate, body depth 
48-73 in TL; head 19 in TL; tail 3.2-3.6 in TL; 
teeth recurved, 2 (rarely 3 or 4) large anterior 
intermaxillary fangs within groove between an- 
terior nostrils, maxillary 4-6, dentary 8-10; eth- 
moidal pore absent; skin of throat and vent area 
with prominent longitudinal grooves; a small but 
conspicuous barbel between anterior and poste- 
rior nostrils; coloration cream-colored, with a 
prominent dark brown or black band along back 
from head to near tail tip, the band beginning at 
lateral line and extending to dorsal fin base, the 
ventral surface and median fins pale, except for 
thin dark median edge of dorsal; cephalic pores 
within brown spots, those over branchial basket 



often within pale spots; MVF H- 130- 199; total 
vertebrae 192-205 (n = 27). 

Size. — To 770 mm. 

ETYMOLOGY. — From the Greek Koixco (infe- 
rior) and axou-a (mouth). 

Distribution. — Widespread in the tropics 
from Hawaii, Polynesia and the Phoenix islands 
throughout Oceania, south to Lord Howe and 
north to the Ryukyus in the western Pacific, 
across the Indian Ocean to east Africa and the 
Red Sea. It is found from shallow sand habitats 
to a depth of 32 m. 

My taxonomic decisions, as well as the discov- 
ery of additional specimens, has considerably 
expanded the range of this species. The Hawaiian 



194 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



Table 2. Vertebral counts of holotypes of 
valid species of Callechelys and their synonyms. 



Species 


Preanal 


Total 


Callechelys bilineahs 


98 


163 


Ophichthys bitaeniatus 


107 


185 


Callechelys canaliculars 


106 


194 


Callechelys catostoma 


127 


197 


Callechelys melanotaenia 


134 


207 


Leptenchelys pinnaceps 


- 


204 


Callechelys striatus 


128 


196 


Callechelys cliffi 


80 


155 


Callechelys eristigma 


105 


159 


Callechelys galapagensis 


92 


172 


Ophichthys guineensis 


119 2 


186 


Caecula pantherina 


- 


179 


Callechelys perryae 


114 


177 


Callechelys luteus 


123 


216 


Dalophis marmorata 


107 


174 


Callechelys guichenoti 


110 


183 


Callechelys muraena 


80 


141 


Callechelys papulosa 


126 


215 


Callechelys randalli 


147 


220 


Gordiich tin 's springe ri 


105 


171 


Ciyptoperygium 






holochroma 


108 


168 



Neotype, designated herein 

Preanal count based on lateral line pores 



record is the first for that archipelago and is based 
on BPBM 29292, a 412 mm male with vivid 
coloration, normal dentition, andH-123-193 ver- 
tebrae. It was caught by J. E. Randall and party 
using rotenone over sand in 32 m. 

REMARKS. — A color photograph of C. ca- 
tostoma appears in Randall et al. (1990:44). 

The 9 1 mm-long specimen (CAS 9 1 79 1 ) from 
Kapingamirangi is ajust-transformed juvenile. It 
lacks dentition. Having been in isopropyl alcohol 
for 43 years, it is now pale yellow throughout 
except for numerous patches of dark pigmenta- 
tion, about equal in size to its orbit and which it 
has retained from its larval condition. There are 
eight equally-spaced, stellate patches along the 
ventral midline from behind the gill openings to 
before the anus. Beginning approximately 3/4 
head length behind the anus, there are four pairs 
of equally-spaced patches located about mid- 
way between the lateral line and the anal fin. 



The taxonomy of this colorful Indo-Pacific 
species is fraught with complications, exempli- 
fied by the four nominal species that I have united 
herein. I have examined a large series of this 
form, ranging from Hawaii to the Red Sea, in- 
cluding all of the type specimens of the nominal 
species. Whereas a modest range in vertebral 
number exists from east to west (Table 3), that 
difference is proportionally small. I can find no 
other characters that would indicate differences 
among populations. 

Wheeler (1981) and Randall and Wheeler 
( 1 99 1 ), based on Whitehead's ( 1 978) description 
of Forster's watercolor of a striped eel from 
Tahiti, recognized Schneider's (in Bloch and 
Schneider 1801) description of Sphagebranchus 
catostomus as the senior synonym of Callechelys 
melanotaenia Bleeker (1864). Bleeker described 
melanotaenia in an 1864 fascicie of his serially 
published Atlas Ichthyologique and redescribed 
it in 1865(a). The illustrations appeared in an 
1865(b) fascicle of the Atlas (cf. Boeseman 
1983:4). Confusion exists about a possible syn- 
type of C. melanotaenia ascribed by Giinther 
( 1 870:7 1 ) to be the type. Bleeker did not mention 
the specimen that is now in the British Museum 
collection (BMNH 1867.11.28.291, 481mm), 
and I agree with Storey (1939) who suspected 
that the unique holotype is within the Leiden 
Museum collection (RMNH 7180, 516 mm). 
Bleeker' s holotype of melanotaenia, as illus- 
trated in the Atlas, differed from the norm in its 
dentition. It atypically has flanking pairs, rather 
than the single pair, of recurved intermaxillary 
fangs, which are unique to this species and C. 
randalli (see Figs, la, 12a). (The British Museum 
specimen, also from Ambon, has the normal 
dental condition.) 

Smith (1958) described Callechelys striatus 
from the western Indian Ocean, noting that it had 
but a single pair of intermaxillary fangs; he pre- 
sumed that Klunzinger's (1871:612) mention of 
Ophichthys melanotaenia from the Red Sea was 
also striatus in that it had only two large fangs. 
Smith assumed that melanotaenia was restricted 
to the eastern Indian and Pacific oceans based on 
Bleeker's (1865b, pi. 193, fig. 2, erroneously 
cited as fig. 1 in the text) illustration of the 
dentition of the four-toothed Ambon specimen 
and Schultz's (1943, fig. 2f) illustration of a 
specimen from Canton Island, central Pacific, as 
having three intermaxillary teeth (reproduced by 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



195 



Table 3. Total vertebral counts of specimens of Callechelys catostoma. 



Location 


Range 


Mean 


N 


Hawaii 


193 




1 


Palmyra Id. (Line Is.) 


199-201 


200.0 


2 


Tahiti (Society Is.) 


195-201 


198.0 


4 


Enewetak (Marshall Is.) 


204 


— 


1 


Lord Howe Id. 


193 


— 


1 


Escape Reef (Coral Sea) 


197 


— 


1 


Palau Is. 


196-205 


200.5 


6 


Ishigaki Id. (Ryukyu Is.) 


203 


— 


1 


Ambon Id. (Indonesia) 


200-205 


202.5 


2 


Diego Garcia (Chagos) 


198 


— 


1 


Aldabra/Seychelles 


195-203 


197.3 


6 


Eilat (Red Sea) 


192 


— 


1 


All localities 


192-205 


198.6 


27 



Schultz in 1953, as fig. 1 If, to indicate the den- 
tition of Enewetak specimens). David Smith has 
examined Schultz's Canton Island specimen 
(USNM 1 15939) and advised me that it (and all 
other C. "melanotaenia" in the USNM collec- 
tion) has but a single pair of intermaxillary fangs. 
I have examined 67 specimens of Red Sea, Indian 
and Pacific Ocean specimens of C. catostoma 
and have found seven individuals (some from 
within large collections of 2-fanged conspeci- 
fics) with more than a single pair, indicating that 
such occasional variation does occur. For the 
reasons stated above, Callechelys striatus Smith 
(1958) is also considered to be a junior synonym 
of C. catostoma. 

McCosker (1970) placed Leptenchelys pin- 
naceps Schultz (1953), based on juvenile speci- 
mens from the Marshall Islands, in the synonymy 
of C. melanotaenia; it is herein included in C. 
catostoma. 

In that the type specimen of Sphagebranchus 
catostomus does not exist (Wheeler 1981), I 
herein designate CAS 9 1793 to be the neotype of 
Callechelys catostoma. It is an adult female with 
developing ova (ca. 0.7 mm diameter), in excel- 
lent condition, and from the locality of the holo- 
type. It has a single pair of recurved 
intermaxillary fangs, vertebral formula H-127- 
197, the normal composition of cephalic pores 
for this species, typical coloration, and the fol- 
lowing measurements (in mm): TL 471; head 
length 26.9; trunk length 304.1; tail length 140; 



dorsal fin origin 8.8; length of snout 3.5; upper 
jaw length 8.2; eye diameter 1.5; and body depth 
behind gill openings 8. 

MATERIAL EXAMINED. — NEOTYPE: CAS 
91793, 471 mm, SOCIETY ISLANDS, Tahiti, 
Moorea, lagoon between Papetaoi and Paopao 
bays (field number GVF 1170), collected over 
sand in 0-1.3 m using rotenone by J. E. Randall 
on 28 August 1956. RMNH 7 1 80, 5 1 6 mm, holo- 
type of Callechelys melanotaenia, Amboina. 
BMNH 1867.11.28.291, 481mm, incorrectly 
called the type or a syntype of Callechelys 
melanotaenia, Amboina. RUSI 96, 473 mm, 
holotype of Callechelys striatus, Mozambique. 
Marshall Islands — USNM 141691, 97 
mm, the holotype of Leptenchelys pinnaceps, 
Bikini Atoll. HAWAII — BPBM 29292, 4 1 2 mm, 
Hawaii, off Kailua. POLYNESIA — CAS 91797, 
560 mm; CAS 91794, 4(332^196 mm), col- 
lected with the neotype, Tahiti. BPBM 12017, 
164 mm, Moorea. PHOENIX ISLANDS — BPBM 
3542, 595 mm; BPBM 12502, 550 mm, Palmyra 
Island. Gilbert Islands — AMS IA. 18055, 
9(300-470 mm), Betio Island. PALAU — CAS 
91796, 6(335-503 mm), Agulpelu Reef. CAS 
28686, 28(70-540 mm), Choi Island. CAS 
91785, 2(383-538 mm), Babelthaup Island. 
CAS 91789, 2 (200-154 mm), Taprakl Reef. 
CAS 91795, 420 mm, Ngaruangl Reef. CAS 
91792, 295 mm, Kayangel. BPBM 10190, 
3(520-548 mm), Bairakaseru. CAROLINE IS- 
LANDS— CAS 91786, 621mm, Ifaluk Atoll. 



196 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



Table 4. Status of the nominal species of Callechelys. Sources are: 1. This study, 2. McCosker et al. 1989, 
3. Smith 1962, 4. Randall and Wheeler 1991, 5. McCosker and Rosenblatt 1972, 6. Storey 1939, 7. McCosker 
1977, 8. Blache et al. 1979, 9. Blache and Cadenat 1971, 10. McCosker 1979, 11. McCosker 1970. 



Nominal Species 



Current Status 



Source 



Callechelys bilinearis Kanazxawa 1 952 

Ophichthys bitaeniatus Peters 1878 

Callechelys canaliculars Smith 1958 

Sphagebranchus catostomiis Forster 1 80 1 

Callechelys cliffi Bohlke and Briggs 1954 

Callechelys eristigmus McCosker and Rosenblatt 1972 

Callechelys galapagensis McCosker and Rosenblatt 1972 

Callechelys guichenoti Kaup 1 856 

Callechelys fijiensis Seale 1935 

Ophichthys (Sphagebranchus) guineensis Osorio 1894 

Cryptoptoygium holochroma Ginsburg 1951 

Caecula leucoptera Cadenat 1 954 

Callechelys longissimus Cadenat and Marchal 1963 

Callechelys luteus Snyder 1 904 

Callechelys maculata Chu, Wu and Jin 1981 

Dalophis marmorata Bleeker 1 853 

Callechelys melanotaenia Bleeker 1 864 

Callechelys muraena Jordan and Evermann 1 887 

Callechelys myersi Herre 1932 

Callechelys nebulosus Smith 1962 

Caecula pantherina Cadenat 1 956 

Callechelys papulosa new species 

Callechelys perryae Storey 1939 

Callechelys peninsulae Gilbert 1892 

Leptenchelys pinnaceps Schultz 1953 

Callechelys randalli new species 

Callechelys sibogae Weber 1913 

Gordiichthys springeri Ginsburg 1 95 1 

Callechelys striatus Smith 1958 



Callechelys bilinearis 


2 


C bitaeniata 


1 


C. bitaeniata 


1,3 


C. catostoma 


1,4 


C. cliffi 


1 


C. eristigma 


1 


C. galapagensis 


1 


C. marmorata 


5,7 


Bascan ich thys fijiensis 


6 


C. guineensis 


2,8 


C. springeri 


2 


C. leucoptera 


9 


Phaenomonas longissima 


2 


C. lutea 


1,10 


Yirrkala? maculata 


1 


C marmorata 


1,5 


C. catostoma 


1,4 


C. muraena 


2 


Bascanichthys myersi 


6 


Xestochilus nebulosus 


1 


C guineensis 


2,8 


C papulosa 


1 


C. guineensis 


2,8 


Myrichthys xysturus 


6,7 


C. catostoma 


4,11 


C. randalli 


1 


Bascanichthys sibogae 


1 


C. springeri 


2 


C. catostoma 


1 



CAS 80993, 495 mm; CAS 91790, 4(280^10 
mm); CAS 91788, 2(325-385 mm); CAS 91787, 
2(367-540 mm); CAS 91791, 4(91^97 mm), 
Kapingamirangi Atoll. TIMOR SEA — CAS 
29138, 2 tails only, from the stomachs of sea 
snakes, Ashmore Reef. AUSTRALIA — AMS 
IA.2263 1-036, 4(98-590 mm), Queensland, Es- 
cape Reef. 1.15564, adult (head and trunk), from 
the stomach of a lethrinid {Lethrinus chrysos- 
tomus), Queensland, Fairfax Island. AMS 
IA.3252, 770 mm; 1.5193, 475 mm, Lord Howe 
Island. CHAGOS ARCHIPELAGO — USNM 
3 1385 1 , 270 mm, Diego Garcia Atoll. ALDABRA 
ATOLL — USNM 313852, 4(307-393 mm). 
Red Sea — SIO 71-165, 425 mm, Eilat. 



Callechelys cliffi Bohlke and Briggs 
(Fig. 5; Tables 1,2,4) 

Callechelys cliffi Bohlke and Briggs 1954:275 (Gulf 
of California, Fraile Bay; holotype CAS-SU 

47521). 

DIAGNOSIS. — Moderately elongate, body 
depth 19-3 1 in TL; head 9-13 in TL; tail 2.2-2.4 
in TL; maxillary teeth 7-8, dentary teeth 14-16; 
coloration tan with numerous fine brown spots 
on head, body and fins; median fins with white 
margins; MVF H-85-155; total vertebrae 
149-158 (n= 14). 

SIZE. — The largest specimen reported by 
McCosker and Rosenblatt (1972) was 455 mm. 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



197 




Figure 5. Callechelys cliffi, SIO 62-42, 455 mm (from McCosker and Rosenblatt 1972, fig. 4). 




Figure 6. Callechelys eristigma, SIO 65-263, 503.5 mm (from McCosker and Rosenblatt 1972, Figs. la). 



ETYMOLOGY. — Named for Frank S. Cliff, 
who captured the type specimen; treated as a 
noun in the genitive case. 

Distribution. — Known from the Gulf of 
California, Mexico, to the Gulf of Panama. All 
specimens were either dip-netted at the surface 
under night lights or collected with ichthyocides 
over sand bottoms to 30 m depth. 

Remarks. — A color photograph of C. cliffi 
appears in Allen and Robertson (1994:52). 

Material Examined. — A listing of the ma- 
terial examined is in McCosker and Rosenblatt 
(1972:22), as well as additional specimens from 
the Perlas Archipelago of the Gulf of Panama: 
USNM 3 18277, 105 mm, Isla San Jose; USNM 
3 18293, 349 mm, Isla del Rey. 



Callechelys eristigma McCosker and Rosen- 
blatt 
(Fig. 6; Tables 1,2,4) 

Callechelys eristigmus McCosker and Rosenblatt 
1972:16, fig. 1 (Gulf of California, Isla San Jose, 
holotype SIO 65-263). 

Diagnosis. — Very elongate, body depth 
38-52 in TL; head 12-15 in TL; tail 3.2-3.7 in 
TL; maxillary teeth 5-6, dentary teeth 6-7; col- 
oration mostly cream, overlain with numerous 
dark spots that extend on to dorsal fin, anal fin 
pale; MVF H- 105- 15 8; total vertebrae 154-163 
(n = 37). 

Size. — The largest specimen recorded by 
McCosker and Rosenblatt (1972) was 1126 mm. 

Etymology. — From the Greek spi (very), 
and GTiyu.a (spot), regarded as an adjective. 



198 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



Distribution. — Known from the Gulf of 
California, Mexico, to the Gulf of Chiriqui, Pan- 
ama, and the Cocos Islands, generally over a sand 
and rock bottom at 5-25 m. 

Material Examined. — A listing of the ma- 
terial examined is in McCosker and Rosenblatt 
(1972:18). 

Callechelys galapagensis McCosker and Ros- 
enblatt 
(Fig. 7; Tables 1,2,4) 

Callechelys galapagensis McCosker and Rosenblatt 
1 972: 1 8, fig. 3 (Galapagos Islands, Isla Santa Cruz, 
holotype SIO 72-1, formerly UCLA 64-39). 

Callechelys marmoratus, nee Bleeker, Fowler 1932:3. 
Fowler 1938:251. 

Callechelys luteiis, nee Snyder, Storey 1939:69. 

DIAGNOSIS. — Moderately elongate, body 
depth 26-38 in TL; head 12-14.5 in TL; tail 
2.2-2.3 in TL; maxillary teeth 7-8, dentary teeth 
15-17; mostly cream-colored, overlain with nu- 
merous dark oblong markings that vary in length 
from size of eye to upper jaw, spots extending 
onto fin, spots smaller and more densely aggre- 
gated in anterior head region, the ventral and 
dorsal surface more spotted than the flanks; MVF 
H-91-172; total vertebrae 170-174 (n = 4). 

SIZE. — The largest specimen reported by 
McCosker and Rosenblatt (1972) was 818 mm, 
the holotype. 

ETYMOLOGY. — Named for the type locality. 

DISTRIBUTION. — Known only from the 
Galapagos Islands, from tidepools to 10 m depth. 

REMARKS. — The smallest paratype (248 
mm) of C. galapagensis differed in coloration 
from that of the larger specimens in having a 
chocolate brown background coloration, al- 
though the general spotting was similar. 

Material Examined. — A complete listing 
of the material examined is in McCosker and 
Rosenblatt (1972:20). 

Callechelys guineensis (Osorio) 
(Fig. 8; Tables 1,2,4) 

Ophichthys (Sphagebranchus) guineensis Osorio 
1894:179 (He de Sao Tome, Cape Verde Islands, 
neotypeMNHNB.1635). 

Callechelys perryae Storey 1 939:7 1 , figs. 2, 4 (Sanibel 
Island, Florida, holotype CAS-SU 33027). 



Caecula pantherina Cadenat 1956:1267, fig. 1 
(Senegal, holotype MNHN B.1635) 

Callechelys pantherina, Cadenat 1961:235. 

Callechelys guineensis, Blache, Bauchot and 
Saldanha 1979:97 (C. perryae and C. pantherina 
synonymized with C. guineensis, holotype of 
Caecula pantherina designated as the neotype of 
Ophichthys guineensis). 

DIAGNOSIS. — Elongate, body depth 45-59 in 
TL; head 15-18 in TL; tail 2.9-3.1 in TL; max- 
illary teeth 4-5, 9-11 on dentary; coloration 
cream to tan overlain with numerous dark brown 
spots and blotches; MVF H-l 15-178; total verte- 
brae 172-186 (n= 12). 

SIZE. — The largest reported specimen is 1080 
mm (Blache et al. 1979). 

ETYMOLOGY. — Named for the type locality, 
a former Portuguese colony. 

DISTRIBUTION. — Apparently widely distrib- 
uted throughout the tropical Atlantic from shal- 
low water to a depth of 15 m, but infrequently 
taken (McCosker et al. 1989). In the western 
Atlantic, it is known from eastern and western 
Florida, Puerto Rico, the Bahamas, St. 
Barthelemy, and possibly Venezuela. In the east- 
ern Atlantic it is known from Senegal and the 
Cape Verde Islands. 

Remarks. — McCosker et al. (1989:309) 
cautiously recognized Callechelys springeri as 
distinct from C. guineensis on the basis of its 
lower total and preanal vertebral counts. The 
coloration of C. guineensis is somewhat variable, 
particularly in spot size, as illustrated by Bohlke 
and Chaplin ( 1968: 108). The leptocephalus of C. 
guineensis was described by Leiby (1989). 

Material Examined. — A complete listing 
of the material examined is in McCosker et al. 
(1989:309-310). 

Callechelys leucoptera (Cadenat) 
(Fig. 9; Tables 1,2,4) 

Caecula leucoptera Cadenat 1954:241, fig. 2 
(Senegal, Isle de Goree, lectotype MNHN 1964- 
527). 

Callechelys leucoptera, Cadenat 1961:235. 

Callechelys leucoptera, Blache and Cadenat 1 97 1 : 1 92 
(lectotype established). 

DIAGNOSIS. — Very elongate, body depth 
53-62 in TL; head 13-14 in TL; tail 2.1-2.3 in 
TL; maxillary teeth 6, dentary 10; body uniform 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



199 




Figure 7. Callechelys galapagensis, UCLA 64-40, 767 mm (from McCosker and Rosenblatt 1972, fig. 3a). 



brown to black, snout and chin paler, fins color- 
less; total vertebrae 162-165, mean =164 
(n = 4). 

SIZE. — The largest specimen reported 
(Blache and Cadenat 1 97 1 ) is 730 mm TL. 

Etymology. — From the Greek X&vK.6q 
(white), and Tkepov (fin). 

Distribution. — Eastern Atlantic, from 
Senegal to the Ivory Coast. Blache and Cadenat 
(1971:199) suggest that it lives in shallow water 
to depths of 45 m, in that several specimens were 
taken from the stomachs of a grouper 
{Epinephelus aeneus) and a ray {Rhynchobatus 
lubberti). 

Material Examined. — MNHN 1964-527, 
545 mm, the lectotype, from Goree, Senegal. 



Callechelys lutea Snyder 
(PI. 1; Tables 1,2,4) 

Callechelys luteus Snyder 1904:517, pi. 3, fig. 5 (Ha- 
waiian Islands, south coast of Molokai Island, holo- 
type USNM 50864). 

Ophichthys marmorata, nee Bleeker, Gunther 
(1910:404). 

Callechelys marmoratus, nee Bleeker, Fowler 
(1928:43). 

DIAGNOSIS. — Elongate, depth 38-53 in TL; 
head 13-18 in TL; tail 2.4-2.8 in TL; maxillary 
teeth 5-6, dentary teeth 8-10; coloration yellow 
to cream, overlain with numerous yellow, and 
brown to dark brown spots between size of eye 



200 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 









Figure 8. Callechefys guineensis, ANSP 98333, 722 mm (from McCoskeret al. 1989, fig. 295). 




Figure 9. Calleehelys leucoptera, IFAN 61-19, 730 mm (from Blache and Cadenat 1971, fig. 26). 



and upper jaw. spotting extends onto dorsal fin, 
but ventral surface and anal fin pale; spotting of 
large adults becomes very dense as freckles on 
head (cf Randall 1996:34); MVF H- 124-2 14; 
total vertebrae 210-219 (n = 8). 

SlZF. — The largest specimen examined was 
1038 mm. Randall (1996:34 and pers. comm.) 
photographed the head of a large adult at 24 m 
off Molokini. the proportions of which indicate 
that the specimen was approximately 1.5 m in 
length. 

ETYMOLOGY. — From the Latin luteus (yel- 
low). 

DISTRIBUTION. — Known only from the Ha- 
waiian Islands. Midway Island (McCosker 
1979:63), and French Frigate Shoals. 



REMARKS. — It appears that after larval settle- 
ment, adults of most Calleehelys spend most of 
their lives within the sediment. They do not form 
permanent burrows, as do heterocongrids and 
some ophichthins, but rather move through the 
interstitial habitat. Calleehelys lutea is excep- 
tional and has on rare occasions been dip-netted 
beneath lights at the sea surface at night and has 
been seen with its head protruding from the sand 
by divers at various Hawaiian locations (J. E. 
Randall, pers. comm.). 

During my only diving experience (16 May 
1993) at Molokini Island (20°37'N, 156°30'W), 
a small collapsed caldera 5 km west of Maui 
Island, I observed two individuals at 15 m over 
the sand bottom of the crater. One was much 
darker than the other and was extended approxi- 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



201 







O G. 

— ca 

if & 

F ° 

= O 



0Q n-) 

PQ 3 






a e 

U to 



■c S 

_ ^ 



E .S 



W) 



E.2 



rf ° 

00 c 

si 

CQ in 

0- . 
00 o 



^3 ft. 



C E5, 

o a^ 
f- ft. 



P -B 

< & 1 
►J J" 
ft, ^ 

3 



202 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



mately 15 cm from the sand. Its dorsal fin was 
elevated and it was nervously flapping its lower 
jaw at the lighter specimen, which was extended 
on to the level of its gill openings, and was, in 
like manner, flapping its jaw. The interaction 
went on for several minutes, but I was unable to 
determine its significance. Ed Robinson (in litt. 
to the author, 14 April 1997), a professional dive 
tour operator who has made many dives at 
Molokini during the last 20 years, relates that 

Most of the time I find them very shy, and quick 
to withdraw into the sand. They do seem to be 
social, since I have often observed small indi- 
viduals) in close proximity (within 12") of a 
larger individual, and large individuals within a 
few feet of each other. They don't seem to 
establish permanent burrows since I have rarely 
found them in the same place two days in a row 
. . . they move backwards under the sand, rather 
quickly, I might add. 

Material Examined. — USNM 50864, 830 
mm, the holotype, Hawaiian Islands, south coast 
of Molokai Island. Hawaii — BPBM 7965, 585 
mm, Hawaii, Kona coast. BPBM 8 1 66, 440 mm, 
Oahu, Kaena Point. BPBM 12381, 680 mm, 
Oahu, Coconut Island. BPBM 36873, 2( 1 32-353 
mm), Oahu, Makua. CAS 13978, 2(480^191 
mm); CAS 33488, 190 mm, Oahu, Waikiki. UH 
959 (now at BPBM), 565 mm, Oahu, Kaneohe 
Bay. FRENCH FRIGATE SHOALS — LACM 
47321-1, 3(515-645), S of East Id. Midway 
Island — SIO 68-497, 1038 mm. 

Callechelys marmorata (Bleeker) 
(PI. 1; Tables 1,2,4) 

Dalophis marmorata Bleeker 1 853a:247 (Siboga, Su- 
matra, holotype BMNH 1867.1 1.28.260). 

Sphagebranchusl marmoratus, Kaup 1856b:26. 

Callechelys Guichenoti Kaup 1856b:28, fig. 13 (Ta- 
hiti, holotype MNHN 2126). 

Ophichthys marmoratus, Giinther 1870:88. 

Callechelys marmoratus, Bleeker 1 864:66. 

Diagnosis. — Elongate, depth 37-50 in TL; 
head 13.4-16.3 in TL; tail 2.66-2.87 in TL; a 
prominent fleshy ridge from middle of snout to 
base of dorsal fin; a pair of stout teeth at inter- 
maxillary, surrounded by papillae; jaw teeth mi- 
nute, 8-10 in maxillary, 12-16 on dentary; 
vomerine teeth neatly biserial, 3-4 pairs fol- 
lowed by 1 or 2 single smaller teeth; body col- 



oration yellowish white to cream in life, overlain 
by chocolate to black spots on body and fins; in 
preservative, cream to tan, overlain with large 
dark blotches, coalesced ventrally and in tail 
region (particularly in larger specimens); head 
with smaller, eye-sized spots and freckles on 
chin; fins with black spots onto bases, their mar- 
gins black nearly throughout; MVF H- 109- 180; 
total vertebrae 174-183 (n = 21). 

SIZE. — The largest known specimen is the 
type, approximately 860 mm in length. Its bro- 
ken jaw suggests that it was taken by a fisherman 
with hook and line. Winterbottom et al. ( 1989: 12, 
fig. 51) report on a 767 mm specimen taken with 
a spear in the lagoon at the Chagos Archipelago. 
The largest I examined was 655 mm, a specimen 
taken from the stomach of a sea snake captured 
in the Timor Sea. These facts suggest that large 
individuals of this species, like many species of 
Callechelys, are not easily captured with ro- 
tenone ichthyocides. 

ETYMOLOGY. — From the Latin marmoratus 
(marbled). 

DISTRIBUTION. — Widespread in the Indo-Pa- 
cific, from Polynesia to east Africa and the Red 
Sea (Klausewitz 1969). It is usually collected 
over sand or sand and rock bottoms using ich- 
thyocides from tidepool depths to 37 m. 

REMARKS. — This species is unmistakable in 
its coloration, in having biserial vomerine denti- 
tion, and a notable fleshy ridge between its snout 
and the origin of its dorsal fin. Smaller fish (25 
cm) have a slightly larger head and shorter tail, 
and generally less dark coloration. A color pho- 
tograph of the head and anterior trunk of C. 
marmorata appears in Randall et al. (1990:44). 

Bleeker twice described Dalophis marmorata 
in 1853. The first (1853a:247) was merely a 
listing of its body depth and fin formula at the 
end of his description of Dalophis moluccensis. 
The latter, in his Bijdrage tot de kennis der Mu- 
raenoi'den en Symbranchoiden . . . (1853b:37), 
provided its complete description and compari- 
son to other species. On the advice of W. N. 
Eschmeyer, I recognize the earlier work as the 
original description in that its publication pre- 
ceded the latter and it satisfies Arts. 1 1 and 16 of 
the RJCN. 

A large collection of C. marmorata made on 
29 September 1957 by the George Vanderbilt 
Foundation (CAS 91808) in Palau provided an 
opportunity to examine the sexual condition and 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



203 



gut contents of 62 individuals (225-575 mm). 
The rotenone collection was made over a sandy 
bottom with areas of short eel grass. The depth 
was from 0-1.3 m, the tide was rising, and the 
area covered was approximately 2,800 m . The 
field notes suggest that "50% of the biota (was) 
sampled" and that "ca. 1 5 species of fishes (were) 
collected." Other species in that collection in- 
cluded 35 Schismorhynchus labial is and six Mu- 
raenichthys sibogae (family Ophichthidae), 24 
Synodus variegatus (Synodontidae), 53 Diplo- 
grammus goramensis (Callionymidae), 12 
Trachinotus bailloni (Carangidae), 4 Polydacty- 
lies sex/ilis (Polynemidae), and single specimens 
of congrids, labrids, and kraemeriids. Thirty-two 
of the 62 C. marmorata (mean = 423 mm, range 
= 340-575 mm) were females with developing 
ova (~ 0.5-1 mm in diameter), 22 (mean = 353, 
range = 255^440 mm) were male (several were 
ripe, two were spent), and the gonadal state of the 
remainder (mean = 254 mm, range = 225-300 
mm) was not determined. The condition of the 
specimens did not allow histological examina- 
tion of their gonads; however, on the basis of my 
examination, it appears that females mature at 
and attain a larger size than do males. At least 
half of the specimens examined had large con- 
centrations of calcareous algae within their gut 
cavity. Dissection demonstrated that the algae 
had previously been consumed by polychaete 
worms, which were consumed intact by the eels. 
No other identifiable gut contents were discov- 
ered. 

Material Examined. — BMNH 1867.11. 
28.260, 860 mm, Siboga, Sumatra, the holotype 
of Dalophis marmorata. MNHN 2126, 473 mm, 
Tahiti, the holotype of Callechelys Guichenoti. 
POLYNESIA — CAS 91804, 486 mm, Tahiti; 
CAS 13961, 512 mm, Moorea. Marshall Is- 
lands — CAS 43342, 437 mm; CAS 53746, 
8(213-439 mm); CAS 58951, 430 mm; BPBM 
29235, 3(160.5-226 mm), Enewetak. GUAM — 
BPBM 11822, 4(91-1 17 mm). CAROLINE IS- 
LANDS — CAS 91800, 356 mm; CAS 91806, 
459 mm, Yap. CAS 91801, 8(398-575 mm), 
Ifaluk Atoll. CAS 91805, 2(325^*52 mm), Kap- 
ingamirangi. FIJI — USNM 259668, 2(353-393 
mm). Palau — CAS 91803, 545 mm, Aulup- 
tagel Island. CAS 91802, 2(490-530 mm), Ba- 
belthaup Island. CAS 91799, 14(125-595 mm); 
CAS 91789, 4(160-392 mm), Kayangel Atoll. 
CAS 91807, 63(108^185 mm), Choi Island. 



CAS 91808, 62(225-575 mm), Agulpelu Reef. 
AUSTRALIA — AMS LA 34318-027, 280 mm, 
Queensland, Townshead Island. AMS LA 939, 
465 mm, Lord Howe Island. TIMOR SEA — CAS 
29137, 655 mm, Ashmore Reefs. MALDIVES — 
BPBM 34700, 316 mm. CHAGOS ARCHIPELAGO 
— USNM 3 13842, 565 mm, Diego Garcia Atoll. 
Aldabra Atoll — RUSI 5220 (x-ray only); 
RUSI 5223 (x-ray only). MOZAMBIQUE — RUSI 
5218 (x-ray only). RED SEA — USNM 313825, 
420 mm, Israel, Ras Burqua. 

Callechelys muraena Jordan and Evermann 
(Fig. 10; Tables 1,2,4) 

Callechelys muraena Jordan and Evermann 1 887:466 
(Snapper Banks, Florida, holotype USNM 37966). 

DIAGNOSIS. — Moderately elongate, body 
depth 27-38 in TL; head 11-14 in TL; tail 
2.5-2.7 in TL; maxillary teeth 7-11, dentary 
11-19; coloration pale, overlain with numerous 
eye-sized brown spots and mottling, median fins 
brown at bases, pale or yellow at margins; MVF 
H-83-142; total vertebrae 139-144 (n = 7). 

SIZE. — A small species; the largest reported 
by McCosker et al. (1989) was 590 mm. 

ETYMOLOGY. — From the Latin muraena 
(moray); treated as a noun in apposition. 

DISTRIBUTION. — A rare species, infrequently 
taken in the western Atlantic between North 
Carolina and Florida, and from the Gulf of Mex- 
ico off northern Florida, and from the Yucatan 
Peninsula. It has been captured by trawl and by 
dredge at depths of 27-1 15 m. 

REMARKS. — The leptocephalus of C. mu- 
raena was described and illustrated by Leiby 
(1989). 

Material Examined. — A complete listing 
of the material examined is in McCosker et al. 
(1989:309-311). 

Callechelys papulosa, new species 
(PI. 1; Figs. 11, 12b; Tables 1,2,4) 

DIAGNOSIS. — Extremely elongate, depth 61 
in TL; head 17 in TL; tail 2.7 in TL; maxillary 
teeth 5, dentary 7; body coloration black, except 
for white patches behind eye, on jaws and snout; 
numerous small pits on face; vertebral formula 
H-126-215. 



204 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 






Figure 10. Callechelys muraena, ANSP 151696, 232 mm (from McCosker et al. 1989, fig. 298). 



Counts and Measurements of Holotype 
(in mm). — Total length 551; head 32.5; trunk 
314.5; tail 204; predorsal distance 10.8; body 
depth at gill openings 9.0; body width at gill 
openings 6.0; body depth at anus 8.3; body width 
at anus 6.3; snout 3.4; tip of snout to rictus of jaw 
7.9; tip of snout to tip of lower jaw 2.6; eye 
diameter 1.4; interorbital distance 2.6; isthmus 
width 2.2. Vertebral formula H-126-215. Left 
lateral line pores 208, 12 in branchial region, 126 
before anal fin, the last within 3 mm of the tail 
tip. 

Description. — Body extremely elongate, its 
depth at gill openings 61 in TL, laterally com- 
pressed throughout its length, its width behind 
gill openings 92 in TL, tapering posteriorly to a 
hard finless point. Head and trunk 1.6 and head 
17 in TL. Snout acute, rounded at tip. Lower jaw 
included, closer to a line drawn from margin of 
eye than to base of anterior nostrils. Eye small, 
about equal in diameter to length of anterior 
nostril tubes, its center above middle of upper 
jaw. Margin of upper lip slit by posterior nostril 
which opens into mouth and appears as a short 
barbel beneath eye. A posterodorsally-directed 
shallow pocket in upper lip begins in lip behind 
posterior nostril and extends behind margin of 
eye. Tip and dorsal surface of snout and chin 
covered with microscopic hair-like prickles. 
Lower jaw, cheeks, nape, and dorsal surface of 
head and branchial basket covered with hundreds 
of small acne-like round pits (Fig. 11), about 
equal in size to cephalic pores. Tongue adnate. 
Branchial basket expanded. 

Cephalic (Fig. 11) and lateral line pores obvi- 
ous, those behind eye and along lower jaw diffi- 
cult to discern from skin pits. Five mandibular, 2 



preopercular, 1 + 3 supraorbital, 4 + 2 infraorbi- 
tal, and single interorbital and supratemporal 
pores. Twelve lateral line pores above branchial 
region, 126 before the anal fin, 208 total. 

Teeth small and conical (Fig. 12b). An anterior 
intermaxillary tooth (the largest), flanked by a 
single tooth, closely followed by a pair, all within 
lateral skin folds of the snout; this is followed by 
a gap, then a patch of 5 irregularly uniserial 
vomerine teeth. Jaw teeth uniserial and smaller, 
7 on each side of maxilla and 9 (with 2 gaps) on 
right and 1 1 on the left side of dentary. 

Body coloration based on photographs of holo- 
type (PI. 1) when fresh and specimen in iso- 
propanol. Trunk, tail, and anal fin black. Dorsal 
fin appears black in preservative, but when ele- 
vated the fin appears nearly transparent (except 
near tail tip), revealing the black underlying fin 
rays. Anus, isthmus, trailing margin of gill open- 
ing, axillary region, snout tip and chin pale white. 
Tip and lateral edges of lower jaw and patches 
behind eyes chalky white. A white spot on upper 
lip in advance of posterior nostril flap. 

SIZE. — Known only from the 55 1 mm holo- 
type, a mature male. 

ETYMOLOGY. — From the Latin papulosa, 
meaning blistered or pimpled, in reference to its 
facial condition. 

DISTRIBUTION. — Known from Papua New 
Guinea. 

REMARKS. — On the basis of its coloration 
and pock-marked physiognomy, C. papulosa 
would not be mistaken for any of its congeners. 
Its uniform black tail, trunk and head, except for 
the striking white patches behind its eyes, cheeks 
and chin, and surrounding the anus, are unique. 
As well, it has more vertebrae than any other 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



205 




206 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



Callechelys save C. lutea and C randalli. The 
presence of acne-like round facial pits is shared 
by C. bitaeniata, C. catostoma, and C randalli; 
however, in none is the condition so dramatically 
developed. Callechelys bitaeniata differs from 
C. papulosa in its coloration and vertebral 
number, and the latter two species differ from C 
papulosa in having comparatively shorter tails 
and in the development of their anterior inter- 
maxillary dentition. 

It seems likely that the extreme development 
of facial pits is associated with a sensory func- 
tion. That, and the restriction of the white colora- 
tion to the orbital, cheeks, snout and chin region, 
indicates that C. papulosa resides within a bur- 
row with only its snout and eyes protruding, 
awaiting passing prey. 

Material Examined. — Holotype: BPBM 
36290, 551 mm, PAPUA NEW GUINEA, D'Entre- 
casteaux Islands, Normandy Island, Buname 
Bay, west of Make-ia Point. Collected in 10 m 
using rotenone over a silty sand and sparse sea 
grass bottom by J. L. Earle on 17 December 
1993. 

Callechelys randalli, new species 
(Figs. 12a, 13; Tables 1,2,4) 

Diagnosis. — Extremely elongate, body 
depth 59-73 in TL; head 15- 18 in TL; tail 
3.5-3.6 in TL; teeth recurved, 2 or 4 large ante- 
rior intermaxillary fangs, maxillary 4, dentary 7; 
ethmoidal pore absent; skin of throat and vent 
area with prominent longitudinal grooves; body 
coloration white with a wide brown stripe from 
mid-head to tail tip; MVF H-146.5-222; total 
vertebrae 220-224 (n = 2). 

Counts and Measurements (in mm) of 
Holotype AND Paratype (in parentheses). — 
Total length 464 (219); head 26.2 (14.5); trunk 
309 (141.5); tail 129 (63); predorsal distance 13 
(6.5); body depth at gill openings ~7.8 (-3); body 
width at gill openings -5.3 (-2.6); body depth at 
anus -5.5 (-2.8); body width at anus -4.9 (-2); 
snout 4 (-2.3); tip of snout to rictus of jaw 8.1 
(-4.7); tip of snout to tip of lower jaw 3.4 (2.2); 
eye diameter 1.2 (-0.8); interorbital distance 2.2 
(-1.2); isthmus width 2.4 (-1 ). Vertebral formula 
H- 147-220 (H- 146-224). Left lateral line pores 
217, 9 in branchial region, 147 before anus, the 
last within 4.5 mm of the tail tip. 



DESCRIPTION. — Body extremely elongate, 
its depth at gill openings 59-73 in TL, laterally 
compressed throughout its length, its width be- 
hind gill openings 84-88 in TL, tapering poste- 
riorly to a hard finless point. Head and trunk 1 .4 
and head 15-18 in TL. Snout acute, rounded at 
tip. Lower jaw included, closer to a line drawn 
from base of anterior nostrils than to margin of 
eye. Eye moderate in size, its diameter about 
twice the length of anterior nostril tubes, its cen- 
ter above middle of upper jaw. Margin of upper 
lip slit by posterior nostril which opens into 
mouth. Lip smooth between anterior and poste- 
rior nostrils, without an apparent barbel. A pos- 
terodorsally-directed, shallow pocket in upper 
lip begins in lip behind posterior nostril and 
extends behind margin of eye. Nape, cheeks and 
dorsal surface of head and branchial basket cov- 
ered with numerous small acne-like round pits 
smaller in size than cephalic pores. Tongue ad- 
nate. Branchial basket expanded. 

Cephalic (Fig. 13) and lateral line pores obvi- 
ous, those above branchial basket difficult to 
discern. Five mandibular, 2 preopercular, 3 su- 
praorbital (ethmoidal pore absent), 4 + 2 infraor- 
bital, and single interorbital and supratemporal 
pores. Eleven lateral line pores above branchial 
region, 147 before the anal fin, 217 total. 

Teeth recurved, moderate in size for Calleche- 
lys (Fig. 12a). Two pairs (that of the holotype; the 
paratype has a pair of large teeth and a large space 
in the intermaxillary region from which a tooth 
may have been removed) of linearly-arranged 
large recurved anterior intermaxillary teeth 
within the snout groove, followed by a space, and 
a smaller tooth pair. Five widely-spaced teeth on 
each maxilla and 8 widely-spaced teeth on each 
dentary. 

Coloration of holotype in isopropanol white, 
overlain by a wide brown stripe from mid-head 
to tail tip, its width about 3/4 or more of the body 
depth; ventral surface and upper dorsal surfaces 
and base of dorsal fin white; dorsal fin margin 
dark brown; anal fin and extreme tail tip white. 
Snout, chin, and most of throat white. A brown 
band about equal to eye diameter lies behind eyes 
and across behind mid-orbit. An irregular brown 
freckling on cheeks and behind post-orbital band. 
Lateral line and cephalic pores not surrounded by 
white spots; a faint white spot lies above each 
lateral line pore from mid-trunk nearly to tail tip. 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



207 




Figure 12. Semi-diagrammatic representation of the dentition of the holotypes of A. Callechelys randalli n. sp., BPBM 
1844, 464 mm, and B. Callechelys papulosa n. sp., BPBM 36290, 551 mm. 



SIZE. — The largest known is the holotype, a 
464 mm male. 

ETYMOLOGY. — Named in honor of John E. 
Randall, friend and the collector of this and many 
of the specimens reported herein. 

DISTRIBUTION. — Known from the Marque- 
sas Islands. 

Remarks. — The new species is most closely 
related to, and could easily be mistaken for, the 
shallow- water Callechelys catostoma. The two 
share the unique synapomorphy of large, recur- 
ved anterior intermaxillary fangs and the absence 
of a pair of ethmoidal pores. Callechelys randalli 
differs in having more vertebrae (220-224 vs. 
192-205), a generally more slender body, a 
wider brown body stripe, and lacks a lip barbel. 



It is distinguishable from all other species of 
Callechelys in the unique specializations de- 
scribed above as well as in the characters men- 
tioned in the key. 

The large anterior intermaxillary fangs, a 
shared synapomorphy of C. randalli and C ca- 
tostoma, differ in number but not in condition 
between the two type specimens. The holotype 
has two pairs of fangs and the paratype has a pair 
and what appears to be the empty socket of a 
third. Variation in the number of these fangs 
occurs in C. catostoma as well (see Remarks 
concerning that species); however, its typical 
condition is to have two linearly-arranged fangs 
slightly adjacent to a fleshy flap along the 



208 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



midline, and atypically to have three or four 
teeth. 

It is not surprising that so few specimens of this 
or other species of Callechelys have been cap- 
tured in deep water poison stations. These eels 
are rarely seen above the sand surface (C. lutea 
is exceptional) and are very slow to react to 
rotenone. By the time that individuals have suc- 
cumbed to the ichthyocide, most collectors have 
far exceeded the time at depth safely allowed by 
conventional scuba diving. 

Material Examined. — Holotype: BPBM 
1 1844, 464 mm, Central PACIFIC, Marquesas 
Islands, Fatu Hiva, off point at north end of 
Hanauu Bay. Collected over sand in 35 m using 
rotenone by J. E. Randall, D. B. Cannoy and R. 
McNair on 21 April 1971. Paratype: CAS 
91839, 219 mm, collected with the holotype. 

Callechelys springeri (Ginsburg) 
(Fig. 14; Tables 1,2,4) 

Gordiichthys springeri Ginsburg 1951:484, fig. 16 

(from the stomach of a shark caught off Salerno, 

Florida, holotype USNM 1 2 1 604). 
Cryptopteiygiuni holochroma Ginsburg 1951:482, 

fig. 1 5 (off Cape Fear, North Carolina, holotype 

USNM 154994). 
Callechelys springeri, Rosenblatt and McCosker 

1970:494. (Robins et al. 1980:71, as first revisers, 

synonymized C. holochroma with C. springeri). 
Callechelys holochromus, McCosker and Rosenblatt 

1 972:22. 

DIAGNOSIS. — Elongate, body depth 45-53 in 
TL; head 14-18 in TL; tail 2.8-2.9 in TL; max- 
illary teeth 4-5, 9-10 on dentary; coloration 
cream, overlain with dark brown to black spots 
and blotches; MVF H- 108- 169; total vertebrae 
168-171 (n = 3). 

SIZE. — The largest reported specimen is a 
801 mm female (McCosker et al. 1989). 

ETYMOLOGY. — Named for Stewart Springer, 
who obtained the type specimen; treated as a 
noun in the genitive case. 

DISTRIBUTION. — Known from three speci- 
mens from off North Carolina and central Flor- 
ida, between depths of 22-36 m. 

REMARKS. — The three known specimens of 
C. springeri are in such poor condition that 
McCosker et al. (1989) cautiously recognized 
them as distinct from C. guineensis, an inhabitant 
of shallower water (see above). The vertebral 



counts and colorations differ slightly and there is 
a slight difference (although significant overlap) 
in the mean tail proportions. The leptocephalus 
of C. springeri is not known; those described by 
Leiby (1984:412) as '"Callechelys species cf. 
springeri" were tentatively reidentified by Leiby 
(1989) as Gordiichthys randalli. 

Material Examined. — USNM 121604, 
372 mm, from the stomach of a Carcharhinus 
milberti taken off Salerno, Florida, the holotype 
of Gordiichthys springeri. USNM 154994, 801 
mm, off Cape Fear, North Carolina, the holotype 
of Crvptopten'gium holochroma. FLORIDA AT- 
LANTIC: ANSP 151698, 367 mm, 27°42'N, 
80°04.5'W. 

Xestochilus, new genus 

Type species Callechelys nebulosus Smith 
1962:452. 

DIAGNOSIS. — Ophichthid eels, subfamily 
Ophichthinae, tribe Callechelyini (sensu 
McCosker 1 977) with an elongate body, its depth 
45-55 in TL, laterally compressed throughout its 
length; body and trunk longer than tail. Dorsal 
fin origin on nape, above supraoccipital, well 
before gill openings. Pectoral fins absent. Snout 
short and acute, rounded at tip, overhanging 
lower jaw; a short groove extends anteriorly to 
(but not reaching) level of anterior nostril bases 
(Fig. 1 b). Anterior nostrils tubular, posterior nos- 
tril opens entirely into mouth; upper lip entire, 
without a crease, split, or barbel at location of 
posterior nostril. Numerous papillae along pe- 
riphery of oral cavity. Gill openings low lateral, 
their aspect vertical, their length much greater 
than isthmus. Branchiostegal rays numerous, ca. 
1/4 not attached to hyoid; epihyal rays broadened 
at their bases; urohyal simple, not branched. 
Scapula present. Gill arches like those of all 
Callechelyini (McCosker 1977, table 3). Head 
pores reduced; three supraorbital pores, two pre- 
opercular pores, an ethmoidal pore and three 
pores in supratemporal canal. Teeth conical, 
slender, small and uniserial on jaws, those of 
intermaxillary followed by a single tooth on 
vomer. Body coloration striped and mottled. 

ETYMOLOGY. — From the Greek ^eaxoq 
(shaven), and x^i^oq (lip, neuter), in reference to 
its smooth upper lip. 

REMARKS. — Subsequent to my creation of 
the tribe Callechelyini (McCosker 1977), I have 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



209 




210 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



- z w # & w*» *'• ^.**^%.^ 




Figure 14. Callechelys springeri, ANSP 151698, 367 mm (from McCosker et al. 1989, fig. 301). 



had the opportunity to examine all of the known 
species. The poorly known Indian Ocean C. ne- 
bulosus has turned out to be much more widely 
distributed, and the opportunity to carefully ex- 
amine larger specimens has convinced me that it 
does not belong in Callechelys. Unfortunately, 
Leurophams lasiops is still known only from the 
small type specimen and I am thereby unable to 
stain, clear, dissect and adequately understand 
the anatomy of that taxon. 

The unique specializations of Xestochilus ne- 
bulosus include: the barely-grooved underside of 
its snout; the continuous upper lip; the state of its 
vomerine and intermaxillary dentition; and the 
presence of unattached branchiostegal rays. 
Other differences between it and other calleche- 
lyins are described in the preceding diagnoses 
and key. 

Xestochilus nebulosus (Smith) 
(PI. 1, Fig. lb, Table 4) 

Callechelys bitaeniatus nee Peters, Smith 1958:835. 
Callechelys nebulosus Smith 1962:452, pi. 65, fig. D 
(Pinda, Mozambique, holotype RUSI 1 15). 

DIAGNOSIS. — Elongate, body depth 46-55 in 
TL; head 13.7-17.7 in TL; tail 2.5-2.6 in TL; a 
low fleshy crest on forehead between interorbital 
and supratemporal pores; 2-3 stout teeth at inter- 
maxillary, not in aline, followed by a pair of stout 
teeth at anterior vomer; jaw teeth minute, 7-9 
maxillary, 12-14 dentary; coloration changes 
with size, in all sizes the jaw corner and preoper- 
cular pores are conspicuously brown or black, the 
median fins are pale except dorsal in trunk and 
anterior tail region which has a dark membrane; 
smaller specimens are tan to light olive with a 



complete brown band along the flanks from be- 
hind head to tail tip; in larger specimens (25 cm), 
the band breaks up in the anterior trunk region 
and brown spotting appears on the head, throat, 
and chest; largest specimens (35 cm) have nu- 
merous small spots on head and trunk region, the 
tail remaining as a brown band; MVF H-93-160; 
total vertebrae 155-165 (n = 19). 

SIZE. — The largest reported (Smith 1962) 
was 470 mm, the holotype, from Pinda, Mozam- 
bique. 

ETYMOLOGY. — From the Latin nebulosus 
(clouded). 

DISTRIBUTION. — Described from Mozam- 
bique and Aldabra, my examination of additional 
material of X. nebulosus has extended its range 
to include the Marshall Islands, the Marquesas 
Islands, Palau, Indonesia, and the Red Sea. It was 
collected over sand bottoms with rotenone be- 
tween depths of 2.5-42 m. 

REMARKS. — Small specimens of Xestochilus 
nebulosus could be mistaken for young C. bi- 
taeniata or C. catostoma. Upon careful examina- 
tion, Xestochilus is easily identifiable on the 
basis of the shortness of its snout groove (Fig. 
lb). The holotype of C. nebulosus has the verte- 
bral formula H-93- 161. 

The Agulpelu Reef, Palau, shallow water col- 
lection of 29 September 1957 (see Remarks un- 
der Callechelys marmorata) included numerous 
specimens of C catostoma and C marmorata 
along with X. nebulosus. 

Material Examined. — RUSI 115, 470 
mm, Pinda, Mozambique, the holotype of 
Callechelys nebulosus. Palau — CAS 92201, 
2(288-368 mm), Kayangel Atoll; CAS 92202, 
5(235-392 mm), Agulpelu Reef. MARSHALL IS- 
LANDS — CAS 53745, 7(244-326 mm); BPBM 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



211 



29234, 2(314-346 mm), Enewetak. MARQUE- 
SAS ISLANDS — BPBM 1 1914,2(151-224 mm), 
Tahuata; BPBM 12603, 235 mm, Nuku Hiva; 
BPBM 11846, 4(181-330 mm), Fatu Hiva. IN- 
DONESIA— USNM 313844, 5(175-338 mm), 
Ambon Island. RED SEA — SIO 71-165, 334 
mm. Eilat; USNM 273656, 300 mm, Eilat; SIO 
71-197, 2(283-355 mm, the smaller cleared and 
stained), Wesat. 

Slalus of Species Previously Referred 
to callechelys 

Described below are those ophichthids that 
have been described as species of, or have been 
subsequently referred to Callechelys but now 
belong in other genera (Table 4). 

Weber (1913) described Callechelys sibogae 
based on a 237 mm specimen from the south 
coast of Timor. Although having the general 
appearance of a species of Callechelys and de- 
scribed as lacking pectoral fins, its body propor- 
tions and general physiognomy are those of a 
bascanichthyin. Close examination of the holo- 
type (ZMA 109.034) discloses that it has small, 
rounded pectoral fins (ca. 0.7 mm in length) and 
is referable to Bascanichthys. 

Callechelys maculatus Chu, Wu and Jin (1981) 
was briefly described from two specimens 
(206-338 mm) from Pingtan Island (1I9°45'E, 
25°30'N) of the Peoples Republic of China. I 
have examined the 206 mm paratype (No. A 
01318). It has the general appearance of a 
callechelyin; however, on closer examination it 
is seen to possess a flattened snout (underside), 
vertebral formula 2-70-143, a dorsal fin origin 
behind the neurocranium, and other osteological 
features typical of a sphagebranchin. I provision- 
ally refer it to the genus Yirrkala pending a 
careful osteological analysis. 

Between 1885-1906, Jordan, and with various 
coauthors, referred species now recognized as 
Bascanichthys to Callechelys, including: 
Sphagebranchus scuticaris Goode and Bean 
(1880); S. teres Goode and Bean (1882); and 
Caecula bascanium Jordan (1885). Similarly, 
Deraniyagala (1931) placed Sphagebranchus 
longipinnis Kner and Steindachner (1866) and 
Ophichthys kirkii Giinther ( 1 870) in Callechelys . 
Storey (1939) referred all of the above-men- 



tioned species as well as several other species 
then recognized within Callechelys to Bas- 
canichthys, including: Ophichthys filaria Giin- 
ther (1872); Callechelys myersi Herre (1932); 
and Cfijiensis Seale (1935). 

ACKNOWLEDGMENTS 

Many individuals have generously assisted 
with this study, and I especially thank John E. 
Randall (BPBM) for providing many of the 
specimens and photographs crucial to this work, 
Peter H. J. Castle (VUW) for vertebral numbers 
of Indian Ocean specimens, and Eugenia Bohlke 
(ANSP) for sharing her vertebral and other data 
with me. As well, I thank the following: Roy 
Eisenhardt for computer and photographic assis- 
tance; David Catania for the preparation of radio- 
graphs; Molly Brown for the preparation of 
illustrations; Ed Robinson for photographs and 
advice on Hawaiian eels; Mysi Hoang for trans- 
lations; David Smith for advice and assistance; 
Alan Leviton for advice with this manuscript; 
William Eschmeyer for nomenclatorial advice; 
and Richard H. Rosenblatt and Carl Ferraris, Jr., 
for reading early drafts of this manuscript. Nu- 
merous curators have allowed me to examine 
specimens in their care, including: E. Bohlke 
(ANSP); J. Paxton, D. Hoese and M. 
McGrouther (Australian Museum, Sydney); A. 
Woolger and O. Crimmin (BMNH); A. Suzu- 
moto (BPBM); W. Eschmeyer. T. Iwamoto, D. 
Catania and J. Fong (CAS); W. D. Anderson 
(GMBL); L. Fishelson (Hebrew University); K. 
Hartel (MCZ); M. Bauchot and the late J. Blache 
(MNHN); M. Gomon (NMV); M. E. Anderson 
and the late M. M. Smith (Rhodes University); 
M. J. P. van Oijen (RMNH); H. L. Wu (Shanghai 
Fisheries University); R. H. Rosenblatt and H. J. 
Walker (SIO); B. W. Walker (then of UCLA); 
W. Bussing (UCR); W. A. Gosline (then of UH); 
C. R. Robins (then of UMML); the staff of the 
National Museum of Natural History (NMNH, 
USNM); I. J. H. Isbriicker and Han Nijssen 
(ZMA); and H. J. Paepke (ZMB). This project 
was assisted in part with grants from the David 
and Lucile Packard Foundation, the Elser Foun- 
dation, and during my tenure as a visiting fellow 
at the Australian Museum (Sydney). 



212 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



Literature Cited 

Allen, G. R. and D. R. Robertson. 1994. Fishes of 
the tropical eastern Pacific. Univ. Hawaii Press, 
Honolulu. 332 pp. 

Blache, J., M. L. Bauchot. and L. Saldanh a. 1 979. 
Designation d'un neotype pour Callechelys 
guineensis (Osorio, 1894), et validation de l'espece 
(Pisces, Anguilliformes, Ophichthidae). Cybium, 
Ser. 3, 3(7):97-98. 

Blache, J. and J. Cadenat. 1971. Contribution a la 
connaissance des Poissons anguilliformes de la cote 
occidentale d'Afrique. Dixieme note: les genre 
Myrichthys, Bascanichthys et Callechelys (Fam. des 
Ophichthidae). Bull, de l'l. F. A. N., Ser. A 
33(1):158-201. 

BLEEKER, P. 1 853a. Derde bijdrage tot de kennis der 
ichthyologische fauna van Ceram. Nat. Tijdschr. 
Ned. Ind. 5:233-248. 

. 1853b. Bijdrage tot de kennis der Mu- 

raenoiden en Symbrancho'iden van den Indischen 
Archipel. Verh. Bat. Gen. 25: 1-76. 



. 1864. Atlas Ichthyologique des Indes Orien- 
tates Neerlandaises, publie sous les auspices du 
gouvernement colonial. Tome 4, Murenes, Syn- 
branchies, Leptocephales. Leiden. 132 pp. 



. 1 865a. Description des deux especes inedites 

des genres Callechelys et Pisoodonophis. Ned. 
Tijdschr. Dierk. 2:213-216. 

. 1865b. Atlas Ichthyologique des Indes Ori- 



entals Neerlandaises, publie sous les auspices du 
gouvernement colonial. Tome 4, Murenes, Syn- 
branchies, Leptocephales. Leiden, plates 193-204. 

Bloch,M.E. and J. G.Schneider. 1801. M. E. Blo- 
chii, systema ichthyologiae iconibus CX illustra- 
tum. Post obitum auctoris opus inchoatum absolvit, 
correxit, interpolavit. Sander, Berlin. 584 pp. 

Boeseman, M. 1983. Forward. Pp. 1-12 in Atlas 
Ichthyologique des Indes Orientales Neerlandaises. 
Smithsonian Institution, Washington, D. C. 

BOhlke, E. B. 1 982. Vertebral formulae of type speci- 
mens of eels (Pisces: Anguilliformes). Proc. Acad. 
Nat. Sci. Philadelphia 134:31^19. 

BOhlke, J. E. 1967. The descriptions of three new eels 
from the tropical West Atlantic. Proc. Acad. Nat. 
Sci. Philadelphia 1 18(4):91-108. 



BOhlke, J. E. and J. C. Briggs. 1954. Callechelys 
cliffi, a new ophichthid eel from the Gulf of Califor- 
nia. Stanford Ichthyol. Bull. 4(4):275-278. 

BOhlke, J. E. and C. C. G. Chaplin. 1968. Fishes of 
the Bahamas and adjacent tropical waters. Living- 
ston Pub. Co., Wynnewood, Pennsylvania. 771 pp. 

Cadenat, J. 1954. Note d'ichtyologie Ouest-Afri- 
caine. VI. Sur quelques especes d'Apodes du genre 
Caecula. Bull, de lTnstitut. Francais d'Afrique 
Noire, Ser. A 16(l):238-244. 



. 1956. Note d'ichtyologie Ouest-Africaine. 

XVI. — Description d'une espece nouvelle 
d'Ophichthyidae: Caecula pantherina. Bull, de 
Flnstitut. Francais d'Afrique Noire, Ser. A 
18(4): 1267-1271. 

. 1961. Notes d'ichtyologie Ouest-Africaine. 



XXXIV. Liste complementaire des poissons de mar 
. . . de IT. F. A. N. a Goree. Bull, de lTnstitut. 
Francais d'Afrique Noire, Ser. A 23(1 ):23 1—245. 

Cervigon, F. 1 973. Los peces marinos de Venezuela. 
Complemento III. Contributiones Cientificas, Uni- 
versidad de Oriente, No. 4. 70 pp. 

Chu, Y. T. H, H. Wu, and X. Jin. 1981. Four new 
species of the families Ophichthyidae and 
Neenchelidae. J. Fish. China 5(l):21-27. 

Deraniyagala, P. E. P. 1931. Further notes on the 
anguilliform fishes of Ceylon. Spolia Zeylanica 
16(2): 131-1 37. 

Dingerkus, G. and L. D. Uhler. 1977. Enzyme 
clearing of alcian blue stained whole small verte- 
brates for demonstration of cartilage. Stain Tech. 
52:229-232. 

Fowler, H. W. 1928. The fishes of Oceania. Bernice 
P. Bishop Mus. Mem. 10. 540 pp. 

1932. The fishes obtained by the Pinchot 



South Seas Expedition of 1 929, with descriptions of 
one new genus and three new species. Proc. U. S. 
Natl. Mus. 80(6): 1-16. 

. 1938. The fishes of the George Vanderbilt 



South Pacific Expedition. 1937. Zool. results, part 
3. Monog. Acad. Nat. Sci. Philadelphia no. 2. 
349 pp. 

Ginsburg, I. 1 95 1 . The eels of the northern Gulf coast 
of the United States and some related species. Texas 
J. Sci. 3(3):43 1^185. 



MCCOSKER: SNAKE-EEL GENUS CALLECHELYS 



213 



Goode, G. B. and T. H. Bean. 1880. Catalogue of a 
collection of fishes obtained in the Gulf of Mexico, 
by Dr. J. w. Velie, with descriptions of seven new 
species. Proc. U. S. Natl. Mus. 2(98):333-345. 

1 882. Descriptions of twenty-five new spe- 



cies of fish from the Southern United States, and 
three new genera, Letharchus, foglossus, and Chri- 
odorus. Proc. U. S. Natl. Mus. 5(297):4 12-437. 

Gosline, W. A. 1951. The osteology and classifica- 
tion of the ophichthid eels of the Hawaiian Islands. 
Pacific Sci. 5(4):298-320. 

GONTHER, A. 1870. Catalogue of the fishes in the 
British Museum. Vol. 8. Catalogue of the Physos- 
tomi, containing the families Gymnotidae ... in the 
British Museum. British Museum, London, xxv + 
549 pp. 



. 1 872. On some new species of reptiles and 

fishes collected by J. Brenchley, Esq. Ann. Mag. 
Nat. Hist., Ser. 4, 10(60):4 18^26. 

. 1910. Andrew Garrett's fische der Sudsee. J. 



Mus. Godeffroy 9(17):389-514. 

Herre, A. W. C. T. 1932. Five new Philippine fishes. 
Copeial932(3):139-142. 

Jordan, D. S. 1885. A catalogue of the fishes known 
to inhabit the waters of north America, north of the 
Tropic of Cancer, with notes on the species discov- 
ered in 1883 and 1884. Rept. U. S. Comm. Fish, for 
1884. 185 pp. 

Jordan, D. S. and B. M. Davis. 1891 (1892). A 
preliminary review of the apodal fishes or eels in- 
habiting the waters of America and Europe. Rep. 
U.S. Comm. Fish and Fisheries for 1888, 
16(9):58 1-677. 

Jordan, D. S. and B. W. Evermann. 1887. Descrip- 
tion of six new species of fishes from the Gulf of 
Mexico, with notes on other species. Proc. U. S. 
Natl. Mus. 9(586):466-476. 

Jordan, D. S. and A. Seale. 1906. The fishes of 
Samoa. Bull. Bur. Fisheries (1905) 25:173-488. 

Kanazawa, R. H. 1952. More new species and new 
records of fishes from Bermuda. Fieldiana, Zool- 
ogy, 34(7):71-1 00. 

Kaup, J.J. 1856a. Uebersicht der Aale. Archiv fur 
Natur. 22(1 ):4 1-77. 



. 1856b. Catalogue of apodal fish, in the col- 
lection of the British Museum. British Museum, 
London, viii + 163 pp. 

Klausewitz, W. 1969. Fische aus dem Roten Meer. 
X. Callechelys marmoratus (Bleeker), ein Ne- 
unachweis fur das Rote Meer (Pisces, Apodes, 
Ophichthidae). Senckenbergiana Biol. 

50(l/2):39^0. 

Klunzinger, C. B. 1871. Synopsis der Fische des 
Rothen Meeres, II Theil. Verh. Zool.-Bot. Ges. 
Wien 21:441-688. 

Kner, R. and F. Steindachner. 1866. Neue Fische 
aus dem Museum der Herren Joh. Ces. Godeffroy 
und Sohn in Hamburg. Sitzungber. Akad. Wiss. 
Wien. 54:356-395. 

Leiby, M. M. 1984. Leptocephalus larvae of the tribe 
Callechelyini (Anguilliformes, Ophichthidae, 
Ophichthinae) in the western North Atlantic. Bull. 
Marine Sci. 34:398-423. 

. 1989. Family Ophichthidae. Pp. 764-897 in 

Fishes of the Western North Atlantic, part nine, vol. 
2. E. B. Bohlke, ed. Yale Univ., New Haven. 

Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. 
Dawson. 1985. Standards in herpetology and ich- 
thyology: part I. Standard symbolic codes for insti- 
tutional resources collections in herpetology and 
ichthyology. Copeia 1985:802-832. 

McCosker, J. E. 1970. A review of the eel genera 
Leptenchelys and Muraenichthys, with the descrip- 
tion of a new genus, Schismorhynchus, and a new 
species, Muraenichthys chilensis. Pacific Sci. 
24:505-516. 

. 1974. A revision of the ophichthid eel genus 

Letharchus. Copeia 1974(3):6 19-629. 

. 1977. The osteology, classification, and rela- 



tionships of the eel family Ophichthidae. Proc. 
Calif. Acad. Sci., Ser. 4, 41(1): 1-123. 

. 1979. The snake eels (Pisces, Ophichthidae) 



of the Hawaiian Islands, with the description of two 
new species. Proc. Calif. Acad. Sci., Ser. 4, 
42(2):57-67. 

McCosker J. E., E. B. Bohlke, and J. E. BOhlke. 
1989. Family Ophichthidae. Pp. 254-412 in E. 
Bohlke, ed., Fishes of the Western North Atlantic, 
part nine, vol. 1 . Yale Univ., New Haven. 



214 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 7 



McCosker, J. E. and P. H. J. Castle. 1986. Family 
Ophichthidae. Pp. 176-186 in The Sea Fishes of 
Southern Africa. M. Smith and P. Heemstra, eds. 
Macmillan Pub., Johannesburg. 

McCosker, J. E. andR. H. Rosenblatt. 1972. East- 
ern Pacific snake-eels of the genus Callechelys 
(Apodes: Ophichthidae). Trans. San Diego Soc. 
Nat. Hist. 17(2): 15-24. 

. 1995. Ophichthidae. Pp. 1326-1341, vol. 3, 



in Guia FAO para la identification de especies para 
los fines de la pesca. Pacifico centro-oriental. 
W. Fischer et al., eds. FAO, Rome. 

Myers, G. S. and C. B. Wade. 1941. Four new gen- 
era and ten new species of eels from the Pacific coast 
of tropical America. Allan Hancock Pacific Exped. 
9(4):65-lll. 

OSBURN, R. C. AND J. T. NICHOLS. 1916. Shore fishes 
collected by the "Albatross" expedition in Lower 
California with descriptions of new species. Bull. 
Amer. Mus. Nat. Hist. 35:139-181. 

OSORIO, B. 1894. Estudos ichthyologicos acerca da 
fauna dos dominios portuguezes da Africa. 3 e nota: 
Peixes maritimos das ilhas de S. Thome, do Principe 
e ilheo das Rolas. J. Sci. Math., Phisicas e Nat. da 
Acad. Real das Sci. de Lisboa, Ser. 2, 3(9): 1 73-1 82. 

Peters, W. C. H. 1878. Uebereineneuemerkwurdige 
Art von fliegenden Fischen, Exocoetus cirriger, aus 
China, und einen neuen Muraeniden, Ophichtkys 
bitaeniatus, aus Mombas. Monatsb. Akad. Wiss. 
Berlin: 555-556. 

Randall, J. E. 1996. Shore fishes of Hawaii. Natural 
World Press, Vida, Oregon. 216 pp. 

Randall, J. E., G. R. Allen, and R. C. Steene. 
1990. Fishes of the Great Barrier Reef and Coral 
Sea. Univ. Hawaii Press, Honolulu. 507 pp. 

Randall, J. Eand A. Wheeler. 1991. Reidentifica- 
tion of seven tropical Pacific fishes collected and 
observed by the Forsters during the voyage of /VMS 
Resolution, 1772-75. Copeia 1991(3):760-767. 

Robins, C. R., R. M. Bailey, C. E. Bond, J. R. 
Brooker, E. A. Lachner, R. N. Lea, and W. B. 
Scott. 1 980. A list of common and scientific names 
of fishes from the United States and Canada (fourth 
edition). Amer. Fisher. Soc, Spec. Pub. 12. I 74 pp. 

Rosenblatt, R. H. and J. E. McCosker. 1970. A 
key to the genera of the ophichthid eels, with de- 



scriptions of two new genera and three new species 
from the eastern Pacific. Pacific Sci. 24:494-505. 

Schultz, L. P. 1943. Fishes of the Phoenix and Sa- 
moan Islands collected in 1939 during the expedi- 
tion of the U. S. S. "Bushnell." Bull. U. S. Natl. 
Mus. No. 180.316 pp. 

Schultz, L. P. and Collaborators. 1953. Fishes of 
the Marshall and Marianas Islands. Families from 
Asymmetrontidae through Siganidae. Bull. U. S. 
Natl. Mus. No. 202(1): 1-685. 

Seale, A. 1935. The Templeton Crocker Expedition 
to western Polynesian and Melanesian islands, 
1933. Proc. Calif. Acad. Sci. (Ser. 4) 

21(27):337-378. 

Smith, J. L. B. 1958. The fishes of Aldabra. Part IX. 
(With a new eel from east Africa). Ann. Mag. Nat. 
Hist., Ser. 12, 10:833-842. 

. 1962. Sand-dwelling eels of the western In- 
dian Ocean and the Red Sea. Rhodes Univ. Ichthyol. 
Bull. 24:447-466. 

Snyder, J. O. 1 904. A catalogue of the shore fishes 
collected by the steamer "Albatross" about the Ha- 
waiian Islands in 1902. U. S. Bur. Fish. Bull. 
22:513-538. 

Storey, M. H. 1939. Contributions toward a revision 
of the ophichthyid eels. 1 . The genera Callechelys 
and Bascanichthys, with descriptions of new spe- 
cies and notes on Myrichthys. Stanford Ichthyol. 
Bull. l(3):61-84. 

Weber, M. 1913. Die fische der Siboga-Expedition. 
Siboga-Expeditie 32(57): 1-7 10. 

Wheeler, A. 1981. The Forsters' fishes. Pp. 781-803 
in The journal of the H. M. S. Resolution 
1772-1775. James Cook. Genesis Publications, 
Duildford, England. 

Whitehead, P. J. P. 1978. The Forster collection of 
zoological drawings in the British Museum (Natural 
History). Bull. Brit. Mus. (Nat. Hist), Hist. Ser. 
6:25-47. 

Winterbottom, R., A. R. Emery, and E. Holm. 
1989. An annotated checklist of the fishes of the 
Chagos Archipelago, Central Indian Ocean. Life 
Sci. Contrib. 145.226 pp. 



© CALIFORNIA ACADEMY OF SCIENCES, 1998 

Golden Gate Park 

San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 8, pp. 217-256, 11 figs., 1 table. 



April 27, 1998 



POLLEN MORPHOLOGY OF MEXICAN ACANTHACEAE: 
DIVERSITY AND SYSTEMATIC SIGNIFICANCE 



By 

Thomas F. Daniel 

Department of Botany, California Academy of Sciences 
Golden Gate Park, San Francisco, California 941 18 



APR 2 S 1998 



Diversity of pollen morphology is described for all 39 genera of Mexican Acanthaceae based on 
scanning electron microscopic study of 324 of the approximately 400 species of the family that are 
native or naturalized in the country. Although there is substantial variation in pollen morphology 
among the genera, species within most genera tend to be relatively conservative with respect to 
sculpturing elements such as the type, number, and position of apertures. Palynological diversity 
within the most speciose genus of Mexican Acanthaceae, Justicia, is as great as, or greater than, 
that recorded for the genus on a worldwide basis. Where possible or appropriate, pollen 
characters are used to assess the relative merits of two systematic classifications with respect to 
Mexican Acanthaceae. Some limitations of scanning electron microscopy for the characterization 
of pollen are noted. 

Received October 21, 1997. Accepted December 23, 1997. 



The remarkable diversity of pollen morphol- 
ogy in the Acanthaceae has been noted by virtu- 
ally all those who have studied palynological 
aspects of the family. The early palynological 
observations of Acanthaceae by Radlkofer 
(1883) were extended by Lindau (1893, 1895), 
who recognized 1 1 pollen types in the family. 
Lindau ( \$n$) based his widely used subfamilial 
classification of the family, in large part, on 
pollen morphology as encompassed by these 
types. The inadequacies of Lindau's pollen types 
for characterizing the observed palynological di- 
versity in the family have been noted by sub- 
sequent workers (e.g., Bhaduri 1944; 
Bremekamp 1948; Scotland 1990a, 1991). More 
recent palynological studies that have considered 
a large number of genera of Acanthaceae (Raj 
1961, 1973; Scotland 1990a), by necessity, have 
been based on only a few species per genus. 
Some limitations of broad, superficial paly- 
nological surveys of the family have been noted 



(e.g., Figueiredo and Keith-Lucas 1996; Immel- 
man 1987) and are further discussed below. Sev- 
eral comprehensive studies of pollen at the 
generic level have been completed recently (e.g., 
Figueiredo and Keith-Lucas 1996;Furness 1989, 
1990, 1991, 1996;Hilsenbeck 1990), sometimes 
revealing considerable palynological diversity 
within a genus. 

The present study attempts a comprehensive 
survey of all acanthaceous genera within a tax- 
onomically diverse geographic region, Mexico. 
With about 400 species in 39 genera, Mexico 
comprises a center of diversity for the family 
with considerable endemism at both the generic 
and specific levels. Pollen of 324 species repre- 
senting all 39 genera of Mexican Acanthaceae 
has been studied in order to: include palynologi- 
cal data in a forthcoming treatment of the family 
in Mexico; assist in classifying Mexican genera, 
particularly those endemic to the country or re- 
gion; assess the usefulness of pollen observations 



[217] 



218 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



based solely on relatively simple SEM prepara- 
tions in a regional geographic context; and add 
to the overall knowledge of acanthaceous pollen. 

In other regional studies of Acanthaceae pol- 
len, Rizzini ( 1 947) surveyed pollen of many Bra- 
zilian genera and utilized pollen characters in 
distinguishing and classifying genera there; 
Petriella (1968) described and figured pollen of 
30 species in 15 genera of Argentinean Acan- 
thaceae and provided a key to genera based ex- 
clusively on palynological characters; and 
Muller, Schuller, Straka, and Friedrich (1989) 
described 26 different pollen morphological 
types among Acanthaceae in Madagascar and 
several islands of the Indian Ocean. The ap- 
proach adopted here deemphasizes pollen types. 
The distinctions between types is often rather 
subjective (see under Justicia below) and a thor- 
ough study of pollen structure should be under- 
taken prior to their delimitation. Instead, 
comparative pollen descriptions are provided for 
each genus of Mexican Acanthaceae based on 
extensive sampling of species. Observed patterns 
and morphological correlations within and be- 
tween suprageneric taxa are noted. 

The subfamilial classification of Mexican 
Acanthaceae utilized here (Table 1) represents a 
compromise between that of Lindau (1895) and 
that of Bremekamp (1965). It maintains all of the 
taxa included by Lindau (1895) in the Acan- 
thaceae but follows Bremekamp's (1965) ar- 
rangement of taxa with retinaculate capsules. A 
revised classification of Mexican Acanthaceae is 
currently being prepared based on morphologi- 
cal, cytological, palynological. and molecular 
data. 

Methods 

Except where noted, pollen was observed and 
studied solely using scanning electron micros- 
copy (SEM). Only morphological attributes 
(e.g., shape, size, aperture number and arrange- 
ment, surface sculpturing) are considered here. 
These attributes are the easiest to observe with 
SEM and have been used most frequently in 
previous palynological studies of Acanthaceae. 
Knowledge of the structure of grains is neces- 
sary, of course, for a complete characterization 
of pollen. Studies of pollen structure are much 
needed and undoubtedly will provide additional 
systematic information. 



In all cases, pollen grains were removed indi- 
vidually from anthers on herbarium specimens 
with insect pins. They were mounted directly 
onto SEM stubs without any prior treatment such 
as acetolysis. Walker and Doyle (1975) noted 
that method and length of pollen preparation can 
affect pollen size and can cause the shape of 
grains to vary considerably as well. Hanks and 
Fairbrothers (1970) discussed specific effects of 
various preparation techniques on pollen of Fa- 
gaceae. The simple procedure noted above was 
used because SEM images of untreated pollen 
have been shown to portray more accurately the 
natural shape and dimensions of the grains 
(Daniel 1983a: 13) and the procedure does not 
remove the apertural surfaces which can contain 
important systematic data. Differences in size 
and/or shape between acetolyzed and unace- 
tolyzed pollen of Acanthaceae have been noted 
by Daniel (1983a, 1986a) and Valsaladevi and 
Mathew (1989). Immelman (1987) reported that 
no significant differences were found between 
acetolyzed and unacetolyzed pollen of two spe- 
cies of Justicia from southern Africa. Balkwill 
(1985) found that untreated pollen of Acan- 
thaceae provided better results with SEM than 
variously treated pollen. 

Photographs of appropriate grains repre- 
senting each species were made to document the 
pollen characteristics observed. Measurements, 
such as lengths of polar (P) and equatorial (E) 
axes, were taken directly from the photographs. 
The pollen figures depicted here were made by 
scanning the photographic negatives into com- 
puter files, adjusting brightness and contrast lev- 
els, cleaning backgrounds, and configuring the 
individual images into plates. ~ 

Data from all species within a 6 ~nus were 
pooled to formulate the generic descriptions be- 
low. In these descriptions, the name of the genus 
is followed by the number of species exam- 
ined/the number of species of the genus in Mex- 
ico/the number of species in the genus. Pollen 
figures are cited at the end of the descriptions. 
Generic discussions (palynological variation, 
systematic considerations, etc.) follow the de- 
scriptions and include references to previously 
published images of pollen of Mexican species. 
All species and collections sampled are listed in 
the Appendix. Most of the specimens repre- 
senting these collections are deposited at ASU. 
CAS. or MICH. 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



219 



Terminology in the descriptions generally fol- 
lows that of Walker and Doyle (1975). To the 
sculpturing elements described by them, the pre- 
fix "micro-" is sometimes added to characterize 
very small sculpturing elements (especially 
when a clear dichotomy in size of sculpturing 
elements is present). The term "bireticulate" is 
used for pollen with a reticulate exine in which 
the lumina appear foveolate to microreticulate. 
In most instances, this microreticulum would 
appear to result from the visibility of the col- 
lumelae (or collumelar branches), in side view, 
below the muri. For compound apertures, indica- 
tion that the width of the colpi barely exceeds the 
diameter of the ora means that it is less than twice 
the diameter of the ora. Elongate and presumably 
harmomegathic furrowlike indentations in the 
exine between compound apertures (i.e., in the 
interapertural regions or mesocolpia) are referred 
to as pseudocolpi. The terminology used to de- 
scribe the apertural regions in Justicia is pre- 
sented and discussed under that genus. 

All currently recognized genera of Mexican 
Acanthaceae were studied. Two other genera, 
Pachystachys Nees and Geissomeria Lindl., each 
based on a single collection from the nineteenth 
century, have been treated as occurring in Mex- 
ico. They are not included here. The sole species 
of Pachystachys known from Mexico was ex- 
cluded by Wasshausen (1986) pending further 
collections to establish its generic affinities. The 
identity of the only species of Geissomeria de- 
scribed from Mexico is currently under study. 

Palynological Descriptions and 
Discussions 

Anisacanthus 7/8/ca. 18. Pollen euprolate to 
perprolate (P:E = 1.4-2.4), 3-colporate, 6- 
pseudocolpate, P = 52-97 urn, E = 35^46 um; 
polar outline subtriangular to triangular; colpi 
fusiform, nearly as long as P, 8-9 (am wide at 
equator, not to barely exceeding diameter of ora 
in width, surfaces microverrucate to verrucate; 
pseudocolpi 2 per mesocolpium, shorter than or 
nearly as long as colpi, sometimes fusing near 
one or both poles thereby forming pseudocolpal 
arcs or ellipses in mesocolpia, surfaces microver- 
rucate to verrucate; interapertural surfaces (ex- 
clusive of pseudocolpi) rugulate to reticulate 
(reticulum homobrochate) to bireticulate. Fig. 8f. 



TABLE 1. Subfamilial classification of Mex- 
cian Acanthaceae. 

Subfamily Thunbergioideae 

Thunbergia 
Subfamily Mendoncioideae 

Mendoncia 
Subfamily Nelsonioideae 

Elytraria, Nelsonia, Staurogyne 
Subfamily Acanthoideae 
Tribe Aphelandreae 

Aphelandra, Holographis, Stenan- 
drium 
Subfamily Ruellioideae 
Tribe Trichanthereae 

Bravaisia, Sanchezia 
Tribe Louteridieae 
Louteridium 
Tribe Lepidagathideae 

Lepidagathis 
Tribe Ruellieae 

Subtribe Barleriinae 

Barleria, Lophostachys 
Subtribe Blechinae 

Blechum 
Subtribe Hygrophilinae 

Hygrophila 
Subtribe Petalidiinae 

Dyschoriste 
Subtribe Ruelliinae 
Ruellia 
Tribe Justicieae 

Subtribe Justiciinae 

Justicia 
Subtribe Isoglossinae (as "Rhytiglossi- 
nae" in Bremekamp, 1965) 
Stenostephanus 
Subtribe Odontoneminae 
Anisacanthus, Aphanosperma, 
Carlowrightia, Chalarothyrsus, 
Chileranthemum, Dicliptera, Gypsa- 
canthus, Henrya, Hoverdenia, Hy- 
poestes, Mexacanthus, Mirandea, 
Odontonema, Pseuderanthemum, 
Schaueria, Spathacanthus, Tetra- 
merium, Yeatesia 
Genera Incertae Sedis 
Poikilacanthus 



220 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



This American genus has centers of distribu- 
tion in Mexico and South America. Pollen of 
Mexican Anisacanthus is typical of that usually 
encountered in the Odontoneminae. Most of the 
palynological variation noted among species of 
Mexican Anisacanthus pertains to shape and size 
of the grains. 

Aphanosperma 1/1/1. Pollen euprolate toper- 
prolate (P:E = 1.5-2.3), 3-colporate, 6-pseudo- 
colpate, P = 39-50 urn, E = 21-27 urn; polar 
outline subtriangular, poles sometimes with a 
3-armed aperturelike indentation in the exine, 
arms aligned with colpi; colpi fusiform, nearly as 
long to as long as P, 2-6 urn wide at equator, not 
to barely exceeding diameter of ora in width, 
surfaces micro verrucate to verrucate; pseudo- 
colpi 2 per mesocolpium, equaling to longer than 
colpi, sometimes fusing near one or both poles 
thereby forming pseudocolpal arcs or ellipses in 
mesocolpia, surfaces psilate; interapertural sur- 
faces (exclusive of pseudocolpi) reticulate, 
reticulum homobrochate. Figs. 7d, e; 8e. 

Aphanosperma is a unispecific genus, endemic 
to northwestern Mexico, with flowers similar to 
those of some species of Carlowrightia, but with 
capsules unique in the Odontoneminae. Its pollen 
is typical for Odontoneminae except for the pres- 
ence of polar, aperturelike indentations in plants 
from one region of its range (see additional fig- 
ures in Daniel 1988a, 1997). Plants from south- 
em peninsular Baja California have this polar 
feature whereas plants from mainland Mexico do 
not. Wet preparations of pollen of Aphano- 
sperma have been reported as having P = 29-39 
urn, E = 25-34 urn, and P:E = 1.1-1.4 (Daniel 
1988a). 

Aphelandra 12/12/ca. 175. Pollen subprolate 
to perprolate (P:E = 1.3-2.4), 3-colpate, P = 
36-66 urn, E = 21-44 urn; polar outline circular 
to subtriangular, poles sometimes with a 3-armed 
aperturelike indentation in the exine, arms alter- 
nate with colpi; colpi fusiform or linear or hour- 
glass shaped, as long or nearly as long as P, 1-6 
urn wide at equator, sometimes fused at poles 
(i.e., syntricolpate), sometimes bifurcate near 
poles, sometimes with a ± prominent and swol- 
len margin, surfaces psilate to verrucate to gem- 
mate to rugulate; interapertural surfaces 
foveolate to fossulate to gemmate-pilate to rugu- 
late to reticulate, reticulum homobrochate to het- 
erobrochate with larger lumina positioned 
equatorially (especially near colpi). Fig. 2a, c. 



Aphelandra is neotropical in distribution. 
Wasshausen (1975) surveyed pollen of 60 spe- 
cies of Aphelandra and noted some palynologi- 
cal diversity among them. He further noted that 
the pollen types he recognized do not correlate 
well with macromorphological features used to 
distinguish the species. Daniel (1991a) described 
and figured palynological features of the Mexi- 
can species. Pollen of Mexican Aphelandra is 
generally consistent with that found elsewhere in 
the genus (McDade 1984; Wasshausen 1975, 
1996) and is similar to that of other Mexican 
Aphelandreae. Pollen of a single Mexican collec- 
tion from Oaxaca that otherwise greatly resem- 
bles A. gigantiflora is unique among species of 
the genus studied to date. It has a pseudocolpal 
ellipse in each mesocolpium (see figures in 
Daniel 1991a) and lacks the polar, aperturelike 
indentations of A. gigantiflora. The taxonomic 
status of this collection remains to be determined. 

Barleria 1/1/80-250. Pollen spherical to sub- 
prolate (P:E = 1.0-1.3), 3-colporate to 3-colpoi- 
dorate, P = 74-109 urn, E = 73-87 urn; polar 
outline subtriangular; colpi sometimes ± indis- 
tinct, sublinear to subfusiform, ca. 0.4-0.7 times 
as long as P, 6-17 urn wide at equator, not to 
barely exceeding or ca. 2 times diameter of ora 
in width, surfaces subpsilate to verrucate; in- 
terapertural surfaces coarsely reticulate, reticu- 
lum ± heterobrochate, muri psilate, lumina 
psilate to verrucate to baculate, those adjacent to 
colpi largest. Fig. 5e, f. 

Only one species, B. oenotheroides, of this 
primarily paleotropical genus occurs in the New 
World. Raj (1961) interpreted the apertures in 
Barleria as pores. I interpret them as compound 
apertures, as did Scotland (1990a). The infrafa- 
milial relationships of Barleria have been con- 
troversial since Bentham and Hooker's (1876) 
placement of the genus in the tribe Justicieae (cf. 
Bremekamp 1965; Lindau 1895; Natarajan 
1957). Raj (1961) and Daniel (1993a) concluded 
that its relatives include Lepidagathis and Lo- 
phostachys, both of which have similar pollen 
(see Lophostachys). Scotland, Endress, and 
Lawrence (1994) noted that a similar, and un- 
usual among Acanthaceae, aestivation pattern of 
the corolla is common to these genera as well. 

Blechum 2/2/ca. 8. Pollen oblate spheroidal to 
euprolate (P:E = 0.9-1.4), 3-syncolporate, P = 
40-57 urn, E = 35-59 urn; polar outline subcir- 
cular to circular; colpi fusiform, as long as P, 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



221 



fused at poles, 5-10 urn wide at equator, not to 
barely exceeding diameter of ora in width, sur- 
faces verrucate to gemmate; interapertural sur- 
faces finely (i.e., longest axis of lumina ca. 
0.4-1.3 urn) to coarsely (i.e., longest axis of lu- 
mina ca. 5-14 urn) reticulate, reticulum homo- 
brochate, muri subpsilate to microverrucate, 
lumina psilate to verrucate to gemmate. Figs. 5h; 

Hg- 

This American genus of fewer than 10 species 
is represented in Mexico by B. pyramidatum and 
B. grandiflorum. Differences in exine sculptur- 
ing between these two species were discussed 
and figured by Daniel (1995a), who also noted 
pollen with intermediate sculpturing in a Central 
American species, B. costaricense Oerst. Pollen 
of another Central American species, B. pana- 
mense Lindau (Mass & Dressier 663) is also 
intermediate in sculpturing between them. Based 
on this limited sampling, it would also appear 
that grains of B. pyramidatum (P = 40^19 urn) 
are smaller than those of B. grandiflorum (P = 57 
urn). 

All species of the genus examined to date are 
syncolporate (Daniel 1995a; Erdtman 1952; Raj 
1961; Ramamoorthy and Hornelas U. 1988; 
Scotland 1 990a). Polar fusion of colpi is encoun- 
tered elsewhere among Mexican Acanthaceae 
(e.g., Aphelandra, Carlowrightia, Justicia, 
Pseuderanthemum) as an incidental or species- 
specific part of palynological variation within a 
genus. In Blechum, polar fusion of the colpi 
appears to be a fixed generic characteristic. 

Bremekamp (1965) treated Blechum as consti- 
tuting a subtribe within the Ruellieae. Based on 
several macromorphological characteristics, the 
genus is similar to Ruellia. Features of the cap- 
sules and pollen, however, readily separate these 
genera (Daniel 1995a). 

Bravaisia 3/3/3. Pollen globose-oblong to 
globose-elongate (longer apertural axis:in- 
terapertural axis = 1.5-2.1), 2-colporate, 
polypseudocolpate, sculptural features of one 
face ± 90° out of phase with those of opposite 
face, longer apertural axis = 50-77 urn, 
shorter apertural axis = 48-70 urn, interaper- 
tural axis = 34-56 urn, apertural face subcir- 
cular to circular in outline, interapertural face 
elliptic to broadly elliptic in outline; colpi ± 
linear to fusiform, nearly as long as axes of 
apertural faces, 3-7 urn wide near ora, sur- 
faces verrucate; pseudocolpi 10-12 per aper- 



tural face, of vary ing lengths, some fusing near 
periphery of grain, oriented in± same direction 
as colpi (i.e., perpendicular to one another on 
opposite aperturate faces), some appearing to 
encircle pollen in interapertural view (i.e, divid- 
ing the 2 identical but skewed halves of the 
grain), surfaces verrucate; exine between 
colpi divided into bands by pseudocolpi, ± 
prominently protruding (liplike) on both sides 
of colpi at ora, bands psilate with a single row 
of foveolae down the center, longest axis of 
foveolae 0.2-2.7 urn. Fig. 3b, c. 

The distribution of this neotropical genus is 
centered in southern Mexico and northern Cen- 
tral America. Pollen of all species (see figures in 
Daniel 1988b) is similar except that grains of the 
larger flowered species, B. grandiflora, are about 
1.5 times longer (i.e., longer apertural axis) than 
those of the other two species. In the two samples 
of this species that were studied, most of the 
pollen was collapsed (see Daniel 1988b). Be- 
cause of the unusual symmetry of the exine 
sculpturing, there are no clear distinctions be- 
tween polar and equatorial axes in isolated pollen 
grains (i.e., what would appear to be the polar 
axis on one apertural face would appear to be the 
equatorial axis on the opposite face). Vasanthy 
and Pocock (1986) studied pollen of two species 
of Bravaisia and described its unusual form as 
"striate and rotationally symmetric." Daniel 
(1988b) referred to this type of pollen as "loxodi- 
colporate." All Trichanthereae share this pollen 
type. Vasanthy and Pocock (1986) discussed 
possible origins for this type of pollen and noted 
similarities to anomalous grains of the Old World 
genus Strobilanthes Blume (as Nilgirianthus 
Bremek.) in the Strobilantheae. 

Carlowrightia 23/23/24. Pollen prolate 
spheroidal to perprolate (P:E = 1.1-2.2), 3-col- 
porate, 6-pseudocolpate, P = 32-5 1 (-73) um, E 
= 18-39 um; polar outline subtriangular to trian- 
gular; colpi fusiform, nearly as long as P, 2-8 um 
wide at equator, not to barely exceeding diameter 
of ora in width, rarely fusing at cne or both poles 
(3-syncolporate), surfaces microverrucate to ver- 
rucate; pseudocolpi 2 per mesocolpium, shorter 
than or as long as colpi, sometimes fusing near 
one or both poles thereby forming pseudocolpal 
arcs or ellipses in mesocolpia, surfaces subpsilate 
to microverrucate; interapertural surfaces (exclu- 
sive of pseudocolpi) rugulate to reticulate 



222 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



(reticulum homobrochate) to bireticulate. Fig. 
8c. 

All but one species of this New World genus 
occur in Mexico. Pollen of Carlowrightia (see 
figures in Daniel 1981, 1983a, 1997) is similar to 
that of most other Mexican Odontoneminae. 
Most of the palynological variation noted within 
the genus results from differences in size and 
shape of grains. Daniel (1983a) noted that in wet 
preparations of Carlowrightia pollen P = 30-63 
Hin, E = 20-43 urn, and P:E = 1.1-2.0. 

Chalarothyrsus 1/1/1. Pollen euprolate (P:E 
= 1.8-1.9), 3-colporate, 6-pseudocolpate, P = 
54-59 um, E = 30-34 urn; polar outline subtri- 
angular to subcircular; colpi fusiform, nearly as 
long as P, 5 um wide at equator, not to barely 
exceeding diameter of ora in width, surfaces 
microverrucate to verrucate; pseudocolpi 2 per 
mesocolpium, as long as colpi, sometimes fusing 
near one or both poles thereby forming pseudo- 
colpal arcs or ellipses in mesocolpia, surfaces not 
seen; interapertural surfaces (exclusive of 
pseudocolpi) reticulate, reticulum homobro- 
chate. Fig. 8a. 

Chalarothyrsus is endemic to Mexico. The 
genus was placed in the Asystasieae by Lindau 
(1904), the "typical representatives" of which 
were included in the Odontoneminae by Bre- 
mekamp (1965). Its pollen is similar to that usu- 
ally encountered in that subtribe. The presence of 
four fertile stamens in Chalarothyrsus and 
Spathacanthus (also treated by Lindau in the 
Asystasieae) contrasts with other Mexican mem- 
bers of Bremekamp's Odontoneminae which 
have only two fertile stamens. 

Chileranthemum 3/3/3. Pollen oblate 
spheroidal to perprolate (P:E = 0.9-2.2), 3-col- 
porate, 6-pseudocolpate, P = 31-58 um, E = 
26-40 um; polar outline subcircular (with 
prominent constrictions at pseudocolpi) to circu- 
lar; colpi fusiform, nearly as long as P, 1-9 um 
wide at equator, not to barely exceeding diameter 
of ora in width, surfaces microverrucate to ver- 
rucate; pseudocolpi 2 per mesocolpium, shorter 
than or as long as colpi, often fusing near poles 
thereby forming pseudocolpal ellipses in meso- 
colpia, surfaces subpsilate to microverrucate; in- 
terapertural surfaces (exclusive of pseudocolpi) 
(subrugulate to) reticulate, reticulum homobro- 
chate. Fig. 7i. 

This genus is known only from Mexico. The 
questionable generic status of Chileranthemum 



and several of its relatives in the Odontoneminae 
was discussed by Daniel (1995b). Pollen of Chil- 
eranthemum (see figures in Daniel 1993b) is 
similar to that of most other Mexican Odon- 
toneminae and therefore does not assist in delim- 
iting this genus. 

Dicliptera 13/ca. 18/ca. 150. Pollen euprolate 
to perprolate (P:E = 1 .4-2.3), sometimes slightly 
hourglass shaped due to a slight equatorial con- 
striction, 3-colporate, 6-pseudocolpate, P = 
38-76 um, E = 22-34 um; polar outline subtri- 
angular; colpi fusiform, nearly as long as P, 2-6 
um wide at equator, not to barely exceeding 
diameter of ora in width, surfaces subpsilate to 
microverrucate; pseudocolpi 2 per mesocolpium, 
nearly to as long as colpi, sometime fusing near 
one or both poles thereby forming pseudocolpal 
arcs or ellipses in mesocolpia, surfaces subpsilate 
to microverrucate; interapertural surfaces (exclu- 
sive of pseudocolpi) reticulate (reticulum ± ho- 
mobrochate) to bireticulate. Fig. 7a. 

Dicliptera is pan tropical in distribution. The 
apparent difference in base chromosome number 
between New and Old World species of the genus 
(Daniel and Chuang 1993; Daniel et al. 1990) is 
not reflected by differences in pollen. Pollen of 
those species of Dicliptera examined (e.g., see 
figures in Daniel 1997) resembles that reported 
for Old World species (Balkwill 1985; Raj 1961; 
Scotland 1990a) as well as that of most other 
Mexican Odontoneminae. 

Dyschoriste 15/ca. 20/ca. 75. Pollen subpro- 
late to perprolate (P:E = 1.2-2.5), 3-colporate, 
irregularly polypseudocolpate, P = (38-) 43-85 
um, E = 26-38 um; polar outline circular; colpi 
fusiform, ca. 1/3-2/3 as long as P, 1-4 um wide 
at equator, not to barely exceeding diameter of 
ora, sometimes oblique (due to spiral/twisting of 
exine), surfaces not seen; pseudocolpi 4-12 per 
mesocolpium, mostly longer than colpi, often 
unequal in length, sometimes spiral/twisted, sur- 
faces subpsilate to microverrucate; exine be- 
tween colpi divided into bands by pseudocolpi, 
± prominently protruding (liplike) on both sides 
of colpi at ora, bands densely microverrucate to 
microechinate and often microfoveolate to mi- 
cro fossulate. Figs. 4a-c; 1 le. 

Dyschoriste occurs in various parts of the 
world. Furness (1995) studied pollen of primar- 
ily African species of Dyschoriste and recog- 
nized a single basic pollen type in the genus with 
numerous subtypes based on the number of 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



223 



pseudocolpi present. Among Mexican taxa, vari- 
ation was noted in grain shape and size, number 
of pseudocolpi (the number often varying within 
a collection or on individual grains), and density 
of the supratectal sculpturing elements. Other- 
wise, pollen of Mexican species (see figures in 
Daniel 1990a, 1996, 1997) is relatively uniform 
and is entirely consistent with that described 
from other parts of the world (Furness 1995). 
Furness (1995) noted palynological similarities 
between Dyschoriste and the African genus 
Chaetacanthus Nees. Among other Mexican Ru- 
ellieae, pollen of Dyschoriste is most similar to 
that of Hygrophila, but differs by its three (vs. 
four) colpi, shorter colpi, and supratectal sculp- 
turing. 

Elytraria 4/4/ca. 15. Pollen euprolate to per- 
prolate (P:E= 1.5-2.0), 3-colpate,P= 42-52 urn, 
E = 25-31 urn; polar outline subtriangular to 
circular; colpi narrowly linear or fusiform, nearly 
as long as P, 1-7 urn wide at equator, surfaces 
subpsilate (at least partially) to irregularly mi- 
croechinate or verrucate; interapertural surfaces 
finely (i.e., longest axis of lumina ca. 0.3-1.9 
urn) reticulate, reticulum ± homobrochate, muri 
microverrucate to microbaculate to microechi- 
nate. Figs, lg, h; 1 lb. 

The 15 or so species of this genus are distrib- 
uted in the tropics and subtropics of both Old and 
New Worlds. Scotland (1990a) noted colpate 
pollen in two New World species (including E. 
imbricata) and one Old World species and colpo- 
rate pollen in an Old World species. Roubik and 
Moreno P. (1991) noted colporate pollen in E. 
imbricata. Endoapertures (ora) were not ob- 
served in SEM preparations of pollen of Mexican 
Elytraria. 

The taxonomic affinities of the Nelsonioideae 
are not fully resolved. They have been treated as 
part of the Scrophulariaceae (e.g., Bremekamp 
1965) and as a distinct family (Sreemadhavan 
1977). Pollen of Mexican Elytraria (see figure in 
Daniel 1997) differs from that of other Mexican 
Nelsonioideae by its larger size and minute su- 
pratectal gemmae or echinae. 

Gypsacanthus 1/1/1. Pollen subprolate (P:E = 
1.2), 3-colporate, 6-pseudocolpate, P = 28-31 
|im, E = 24-26 urn; polar outline circular; colpi 
fusiform, nearly as long as P, 6 urn wide at equa- 
tor, barely exceeding diameter of ora in width, 
surfaces microverrucate to verrucate to gem- 
mate; pseudocolpi 2 per mesocolpium, shorter 



than colpi, sometimes fusing near one or both 
poles thereby forming pseudocolpal arcs or ellip- 
ses in mesocolpia, surfaces microverrucate; in- 
terapertural surfaces (exclusive of pseudocolpi) 
reticulate (reticulum homobrochate) to bireticu- 
late. Fig. 7g. 

Gypsacanthus is endemic to arid regions in 
southern Mexico. Pollen of Gypsacanthus stud- 
ied and figured by Lott et al. (1984) is euprolate 
(P:E = 1.7), somewhat larger (P = 46 um), and 
triangular in polar outline, but otherwise resem- 
bles that described here. The genus was included 
in subtribe Odontoneminae (Lott et al. 1984) in 
which this type of pollen is common. 

Henrya 2/2/2. Pollen euprolate to perprolate 
(P:E = 1.7-3.0), sometimes slightly hourglass 
shaped due to a slight equatorial constriction, 
3-colporate, 6-pseudocolpate, P = 50-71 um, E 
= 25-32 um; polar outline triangular; colpi ellip- 
tical, nearly as long as P, 8-9 um wide at equator, 
2.0-2.7 times diameter of ora in width, surfaces 
densely verrucate to gemmate, verrucae or gem- 
mae sometimes microverrucate; pseudocolpi 2 
per mesocolpium, shorter to longer than colpi, 
surfaces not seen; interapertural surfaces (exclu- 
sive of pseudocolpi) reticulate (reticulum homo- 
brochate) to bireticulate. Fig. 8b. 

Daniel (1990b) recognized two species in this 
genus of North and Central America; and he 
reported P = 59-83 um, E = 20-34 um, and P:E 
= 1.2-3.4 in wet preparations. The exceptionally 
broad colpi are unique among New World Odon- 
toneminae and are thus apomorphic for Henrya 
(see figures in Daniel 1 990b, 1 997). The basis for 
Gibson's (1974) description of pollen in Henrya 
as 4-porate remains unknown. 

Holographis 15/16/16. Pollen euprolate to 
perprolate (P:E = 1.4-2.3), 3-colpate (or 3- 
colpoidate), P = 32-65 um, E = 18^4 \im; polar 
outline subtriangular to triangular; colpi (or 
colpoids) nearly as long as P, linear to ± hour- 
glass shaped or sometimes bifurcating near poles 
(arms of bifurcations sometimes fused with ad- 
jacent arms of other colpi near one or both poles), 
1-8 um wide at equator, surfaces psilate to ± 
verrucate; interapertural surfaces psilate to ver- 
rucate to foveolate to rugulate. Fig. 2b, d. 

Holographis is endemic to Mexico where it has 
radiated among deserts and other dry regions. 
Pollen of the genus (see figures in Daniel 1983b, 
1986b, 1988c, 1997) resembles that of other 
Mexican Aphelandreae. 



224 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



Hoverdenia 1/1/1. Pollen euprolate (P:E = 
1.4-1.6), 3-colporate, 6-pseudocolpate, P = 
67-75 urn, E = 43^8 urn; polar outline triangu- 
lar; colpi nearly as long as P, 12-14 urn wide at 
equator, barely exceeding (i.e., 1.5-1.8 times) 
diameter of ora in width, ± elliptical, surfaces 
microverrucate to verrucate (often irregularly so 
with psilate regions); pseudocolpi 2 per meso- 
colpium, shorter than or as long as colpi, surfaces 
microverrucate to verrucate; interapertural sur- 
faces (exclusive of pseudocolpi) psilate to foveo- 
late, surfaces between colpi and adjacent 
pseudocolpi often ± broken up or appearing de- 
graded. Fig. 7c. 

This unispecific genus is endemic to north- 
eastern Mexico. Its pollen is generally similar to 
that of other genera of Odontoneminae, except 
for the psilate to foveolate (vs. reticulate) in- 
terapertural surfaces and the regions between the 
colpi and the adjacent pseudocolpi which often 
appear eroded. 

Hygrophila 1/2/80-100. Pollen spherical to 
prolate spheroidal (P:E = 1.0-1.1), P - 27-37 
urn, E = 27-33 urn, 4-colporate, irregularly 
polypseudocolpate; polar outline subsquare; 
colpi fusiform, ca. 1/2-2/3 as long as P, 2-7 (im 
wide at equator, not to barely exceeding diameter 
of ora in width, surfaces microverrucate to ver- 
rucate; pseudocolpi 3-6 per mesocolpium, ± un- 
equal in length, mostly longer than colpi, 
surfaces microverrucate (often with psilate re- 
gions); exine between colpi divided into bands 
by pseudocolpi, bands rugulate. Figs. 4d, e; 1 If. 

This pantropical genus is represented in Mex- 
ico by a native species, H. costata, and an intro- 
duced species, H. polysperma (Roxb.) T. 
Anderson. Furness (1994) described pollen of 34 
species of Hygrophila and recognized numerous 
pollen types and subtypes based on sculptural 
patterns, number and pattern of the pseudocolpi, 
and form of the endoapertures. Pollen of H. 
costata would appear to conform to her "//. 
brasiliensis-tyvQ," "subtype b." At least one spe- 
cies (H guianensis Nees) studied by Furness 
(1994), the pollen of which conforms to this 
subtype, was treated by Daniel (1995c) as con- 
specific with H. costata. Furness (1994) noted 
that pollen of Hygrophila and Brillantaisia P. 
Beauv., an Old World genus of about 1 1 species 
with pollen nearly identical to that of Hygro- 
phila, is unique in the Ruellieae. Elsewhere 
among Mexican Acanthaceae, 4-aperturate pol- 



len has been found only in a few species of 
Justicia. 

Hypoestes 1/1/70+. Pollen euprolate (P:E = 
1.7-1.9), 3-colporate, 6-pseudocolpate, P = 
44-50 urn, E = 25-26 urn; polar outline triangu- 
lar; colpi subfusiform, nearly as long as P, 1-2 
urn wide at equator, not to barely exceeding 
diameter of ora in width, surfaces microverru- 
cate; pseudocolpi 2 per mesocolpium, shorter 
than or nearly as long as colpi, surfaces micro- 
verrucate; interapertural surfaces (exclusive of 
pseudocolpi) reticulate (reticulum ± homobro- 
chate) to bireticulate. Fig. Hi. 

This paleotropical genus is represented in 
Mexico by the naturalized Malagasy endemic H. 
phyllostachya. Pollen of the species resembles 
that of native Mexican Odontoneminae as well 
as other Old World species of this genus (Balk- 
will and Norris 1985; Muller et al. 1989; Raj 
1961). 

Justicia 80/100+/ca. 600. Pollen 2-4-porate to 
2-4-colporate, if 2-aperturate then globose-ellip- 
tic to globose-elongate (rarely with an equatorial 
constriction) (P:E in apertural view = 1.2-2.3; 
P:E in interapertural view = 1.3—4.1; longer 
E:shorter E = 1.1-2.0) and P = 23-87 urn and E 
(apertural view) = 16—42 \im and E (interapertu- 
ral view) = 17-38 \im, if 3- or 4-aperturate then 
subprolate to perprolate (P:E = 1.2-2.7) and P = 
27-90 urn and E = 18-54 urn; polar outline el- 
liptic to broadly elliptic to subrectangular (2- 
aperturate) or triangular to subcircular 
(3-aperturate) or subcircular to square (4-apertu- 
rate); trema area (see discussion below) present 
or absent, containing peninsulae and/or 2-6 rows 
of ± discrete insulae, with an equal number of 
rows on each side of aperture(s) or (rarely) with 
more rows on one side than the other, insulae (2-) 
4-10 (-14) per row, 1.3-8.0 urn in diameter, 
subcircular to elliptic to ± irregular in shape, 
interinsular (and/or interpeninsular) surface of 
trema area subpsilate to microverrucate to verru- 
cate to verrucate-foveolate to microreticulate; 
colpi present (although often not prominent) or 
absent, usually fusiform, 2/5 as long to as long as 
P, 1.9-5.8 urn wide at equator, not to barely 
exceeding diameter of ora, rarely fusing at one or 
both poles (syncolpate), surfaces subpsilate to 
microverrucate to verrucate to verrucate-rugu- 
late; pseudocolpi absent or present, if present 
then 4 (in 2-aperturate grains) or 6 (in 3-apertu- 
rate grains) or 8 (in 4-aperturate grains), shorter 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



225 



to longer than colpi in length, sometimes fusing 
near one or both poles thereby forming pseudo- 
colpal arcs or ellipses in mesocolpia, surfaces 
subpsilate to microverrucate to verrucate; in- 
terapertural surfaces (exclusive of trema areas, if 
present) microreticulate and studded with psilate 
echinae (i.e., J. angiistiflora) to psilate-foveo- 
late-microechinate (i.e., J. medranoi) to subpsi- 
late to rugulate (with surfaces between rugulae 
verrucate in some J. masiaca) to reticulate 
(reticulum homobrochate to heterobrochate with 
lumina near poles smaller than lumina near equa- 
tor) to bireticulate, lumina of reticulum psilate to 
verrucate to gemmate. Figs. 9a-i; lOa-i. 

This is the largest genus of Acanthaceae with 
estimates of up to 600 species worldwide. Sev- 
eral genera with species in Mexico (e.g., Chae- 
tothylax Nees, Ixtlania M. E. Jones, Neohallia 
Hemsl., Siphonoglossa Oerst., Tabascinia Baill.) 
have been included in Justicia recently, based, at 
least in part, on the pollen evidence (Daniel 
1990a, 1995a; Graham 1988). Palynologically, 
Justicia is by far the most diverse genus of Mexi- 
can Acanthaceae (see figures in Acosta C. 1989; 
AcostaC. and Daniel 1993; Daniel 1990a, 1990c, 
1993b, 1995a, 1995d, 1997; Henrickson and 
Hiriart 1988; Hilsenbeck 1990; Hilsenbeck and 
Marshall 1983; Meagher 1974; Ramamoorthy 
1989; Wasshausen 1981; Wasshausen and 
Daniel 1995). Graham (1988) recognized 10 ma- 
jor types of pollen in the genus on a worldwide 
basis. Mexican species have been studied that 
resemble her types 1 and 3-9. Pollen resembling 
her type 2 might be present among the 12 species 
with 3-aperturate pollen; however, the distin- 
guishing feature of this pollen type (sexine thick- 
ness) was not determined in the grains studied. 
At least two species have pollen that greatly 
resembles her type 10 except for the number of 
apertures (three in J. longii and four in J. 
masiaca). Pollen of several species does not con- 
form to any of the types recognized by Graham 
(1988). For example, eight species studied have 
4-aperturate pollen and none of the types recog- 
nized by Graham include grains with four aper- 
tures. Considering the characters used by 
Graham for recognizing pollen types, at least six 
could be recognized among these eight species. 

Among the 80 species studied, eight have 4- 
aperturate pollen, 16 have 3-aperturate pollen, 
and 56 have 2-aperturate pollen. Among the lat- 
ter, two species have pseudocolpi and 54 have 



trema areas. For palynological descriptions of 
Justicia, trema areas are here defined as regions 
of thinner exine than found elsewhere on the 
grain, each containing a simple or compound 
aperture flanked by longitudinally arranged rows 
of more or less discrete insulae (which when not 
discrete from adjacent thicker portions of exine 
are termed peninsulae). The insulae vary greatly 
in size and number. They are usually arranged 
into ± distinct longitudinal rows or, rarely, appear 
to be randomly disposed (e.g., J. aurea). A single 
row on each side of the aperture (i.e., two rows 
per trema area) is the most common pattern 
among Mexican species. The number of rows of 
insulae is not always consistent within a species. 
For example, in J. candicans the number of rows 
within a trema area varies from four to six. Sculp- 
turing of the insulae resembles that found outside 
of the trema area. The trema areas rarely fuse 
across the poles (e.g., J. valvata, with connec- 
tions between two of the four opposing pairs of 
trema areas) or join with adjacent trema areas 
near the equator (e.g., J. jitotolana, with sculp- 
turing outside of the trema area restricted to the 
vicinity of the poles). Trema areas sometimes 
fuse near the poles to adjacent trema areas (e.g., 
J. borrerae, J. tabascina, J. valvata). Fusion of 
this type is reminiscent of fusion seen in pseudo- 
colpi elsewhere in the family. In fact, pollen with 
pseudocolpi and pollen with trema areas appear 
to represent ends of a continuum rather than 
completely distinct pollen types (see also 
Meagher 1974 who concurs and Hilsenbeck 
1990 who disagrees). The differences between 
them are a matter of degree and all intermediate 
stages between the extremes are evident among 
pollen of Justicia in Mexico. For example, col- 
porate pollen with pseudocolpi is found in J. 
ixtlania; pollen in which the region of thick exine 
between the colpi and pseudocolpi is barely bro- 
ken up into subdiscrete insulae is found in some 
J. breviflora, some J. mirandae, and in J. sp. 
{Cowan & Magaha 3 J 38); and pollen with a well 
delimited trema area is found in most species 
(e.g., J. candicans). Sometimes, even within a 
species, a similar trend can be observed. Daniel 
(1995d) noted and illustrated variation in size 
and surface sculpturing in pollen of J. masiaca in 
which grains vary from having discrete, inter- 
nally gemmate insulae in a trema area to having 
what appear to be fused insulae flanked by 



226 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



pseudocolpi to lacking either a trema area or 
pseudocolpi. 

Tricolporate, 6-pseudocolpate pollen similar 
to that generally encountered in subtribe Odon- 
toneminae is also found in Justicia of the Justici- 
inae (see also Henrickson and Hiriart 1988). 
Bremekamp's (1965) classification has the ad- 
vantage of uniting these taxa within a single tribe, 
Justicieae. The relative distinction between pol- 
len with pseudocolpi and pollen with trema areas 
is noted above. Although superficial pollen char- 
acters neither provide evidence formonophylesis 
of Justicia nor distinguish the genus from other 
genera of Justicieae, the considerable diversity of 
palynological characters has been used for in- 
trageneric classification (Graham 1988) and can 
be useful in species delimitation and recognition 
(e.g., Daniel 1995a, 1995d). Knowledge of pol- 
len morphology in Justicia can reveal potential 
taxonomic problems as well. For example, Gra- 
ham (1988) indicated that/, liebmanii V. A. W. 
Graham has type 9 pollen. Daniel (1995c) treated 
this species as conspecific with J. spicigera. Pol- 
len of other specimens of J. spicigera have pollen 
resembling Graham's type 7, suggesting either 
taxonomic imprecision or the presence of differ- 
ent pollen types in a single taxon. 

Some patterns of variation similar to those 
noted here are evident among southern African 
species of Justicia (Immelman 1987). 

Lepidagathis 1/1/ca. 100. Pollen subprolate to 
euprolate (P:E = 1 .2- 1 .8), 3-colporate, P - 21-36 
um, E = 16-20 urn; polar outline subcircular to 
circular; colpi fusiform, ca. 2/3 to nearly as long 
as P, 2-3 um wide at equator, not to barely 
exceeding diameter of ora in width, surfaces 
verrucate; interapertural surfaces reticulate, 
reticulum heterobrochate, muri psilate, lumina 
verrucate to pi late, some of those flanking (but 
not immediately adjacent to) colpi larger than 
those at poles and near center of mesocolpia. 
Fig. 5i. 

Lepidagathis is mostly paleotropical in distri- 
bution. Like its putative relatives Lophostachys 
and Barleria, its subfamilial position has never 
been stable. The sole Mexican species is some- 
times treated in Teliostachya Nees (see Daniel 
1995a for discussion of these genera). 

Lophostachys 3/3/20. Pollen subprolate to 
euprolate (P:E - 1 .2-1 .4), 3-colporate, P - 64-8 1 
um, E = 46-60 urn; polar outline subcircular; 
colpi ± linear to fusiform, ca. 1/2-2/3 as long as 



P, 2-6 um wide at equator, not to barely exceed- 
ing diameter of ora in width, surfaces psilate to 
verrucate; interapertural surfaces coarsely reticu- 
late, reticulum heterobrochate, muri ± psilate, 
lumina verrucate to gemmate to baculate to 
pilate, those near poles smallest and those flank- 
ing colpi largest. Fig. 5d, g. 

Lophostachys is neotropical in distribution 
with three endemic species in Mexico. Daniel 
(1993a) illustrated the pollen of each. Among 
Mexican and Central American Acanthaceae 
pollen with similar surface sculpturing is known 
in Ruellia (Ruelliinae), Lepidagathis (Lepida- 
gathidinae), and Barleria (Barleriinae), each of 
which would be included in a different subtribe 
utilizing Bremekamp's (1965) classification. 
Benoist (1911) concluded that Lophostachys is 
not distinct from Lepidagathis. While his conclu- 
sion has not been generally accepted, the putative 
distinctions between these genera deserve addi- 
tional study. 

Louteridium 7/8/10. Pollen spherical (P:E = 
1.0), pantoforate, diameter = 98-140 um; foram- 
ina more than 50, circular, 7-14 [im in diameter, 
surfaces microverrucate to microbaculate to mi- 
cropilate; interapertural surfaces gemmate to 
baculate, regions between gemmae or baculae 
psilate to microverrucate to micropilate to finely 
rugulate. Figs. 3a; 1 Id. 

This genus is restricted to Mexico and Central 
America. Its pollen is unique in the family and 
apomorphic for the Louterideae. 

Mendoncia 2/2/50-60. Pollen spherical (P:E 
= 1.0), 5-colpate (see discussion), diameter 
41-56 um; polar outline subpentagonal to sub- 
circular; colpi narrowly linear, ca. 1/4-1/3 as 
long as pollen diameter, 1-2 um wide at equator, 
surfaces irregularly verrucate, margins ± pro- 
truding (liplike); interapertural surfaces finely 
rugulate, rugulae microverrucate and forming a 
dense, anastomosing patchwork over the surface. 
Figs, le, f; 11a. 

Mendoncia occurs in the tropics of the New 
World, Africa, and Madagascar. Pollen of both 
Mexican species was figured by Daniel (1992). 
Pollen of the genus from regions outside of Mex- 
ico has been described as 3-6-colporate (Erdt- 
man 1952; Mulleret al. 1989; Raj 1961;Roubik 
and Moreno P. 1991; Scotland 1990a). En- 
doapertures (i.e., ora) were not observed in SEM 
preparations of Mexican species. 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



227 



The taxonomic status of Mendoncia and its 
relatives is not resolved. The subfamily has been 
treated as a separate family (e.g., Bremekamp 
1965; Cronquist 1981) and as a tribe within the 
Thunbergioideae (e.g., Takhtajan 1997). Scot- 
land (1990a) noted that the tectal ornamentation 
and the morphology of the compound aperture 
(not described) are apparently unique features of 
Mendoncia pollen. As noted by Scotland 
(1990a), the uniqueness of its pollen does not 
help to resolve the taxonomic position of Men- 
doncia. 

Mexacanthus 1/1/1. Pollen globose-oblong to 
globose-elongate (P:E = 1.6-1.8 in colpal view, 
3.2-3.4 in intercolpal view; longer E:shorter E = 
1.7-2.0), 2-colporate, 4-pseudocolpate, P = 
48-66 urn, E = 27-39 urn (colpal view) and 
20-23 (intercolpal view); polar outline subrec- 
tangular to elliptic; colpi fusiform, nearly as long 
as P, 7-8 um wide at equator, barely exceeding 
diameter of ora, surfaces microverrucate to ver- 
rucate; pseudocolpi 2 per mesocolpium, shorter 
than or nearly as long as colpi, surfaces micro- 
verrucate to verrucate; interapertural surfaces 
(exclusive of pseudocolpi) reticulate, reticulum 
homobrochate. Fig. 8d. 

Mexacanthus is endemic to western Mexico. 
Its Mexican relatives in the Odontoneminae (e.g., 
Carlowrightia, Anisacanthus, Miranded) all 
have 3-colporate pollen. The secondarily bilat- 
eral (i.e., 2-colporate) pollen of Mexacanthus 
(see figures in Daniel 1981) would appear to be 
apomorphic for the genus in the subtribe. 

Mirandea 4/4/4. Pollen spherical to perprolate 
(P:E = 1.0-2.0), 3-colporate, 6-pseudocolpate, P 
= 33-66 um, E = 25-44 um; polar outline circu- 
lar to triangular; colpi fusiform, nearly as long as 
P, 2-8 um wide at equator, not to barely exceed- 
ing diameter of ora in width, surfaces microver- 
rucate to verrucate; pseudocolpi 2 per 
mesocolpium, shorter than or as long as colpi, 
sometimes fusing near one or both poles thereby 
forming pseudocolpal arcs or ellipses in meso- 
colpia, surfaces psilate to microverrucate; in- 
terapertural surfaces (exclusive of pseudocolpi) 
microverrucate- foveolate to bireticulate. Fig. 8h. 

Mirandea is endemic to Mexico. Its pollen (see 
figures in Daniel 1978, 1986b) is similar to that 
of most other Mexican Odontoneminae. Miran- 
dea huastecensis differs from M. andradenia and 
M. sylvatica by its pollen with interapertural 
surfaces microverrucate and foveolate (vs. 



bireticulate). One collection of M. grisea (Daniel 
852) has microverrucate sculpturing whereas an- 
other (Daniel 331) has foveolate-reticulate 
sculpturing. 

Nelsonia 1/1/1+. Pollen euprolate (P:E = 
1.6-1.9), 3-colpate, P = 34-39 um, E = 18-21 
um; polar outline subtriangular to triangular; 
colpi nearly as long as P, 1-2 um wide at equator, 
narrowly linear, surfaces not seen; interapertural 
surfaces reticulate, reticulum ± heterobrochate, 
larger lumina irregularly dispersed. Figs, li; lie. 

The number of species recognizable in this 
pantropical genus is controversial (cf. Barker 
1986; Hossain 1984). Scotland (1990a) de- 
scribed pollen of Nelsonia as 3-colporate; 
Muller, Schuller, Straka, and Friedrich (1989) 
and Erdtman (1952) described it as 3-colporoid 
(or 3-colporoidate); and Roubik and Moreno P. 
(1991) described it as 3- or 4-porate (although 
their figures reveal 3-colporate grains). En- 
doapertures (ora) were not seen in SEM prepara- 
tions examined by me. Among Mexican 
Nelsonioideae, exine sculpturing of Nelsonia 
pollen is more like that of Staurogyne than 
Elytraria. Pollen of Nelsonia is intermediate in 
size between these other genera. 

Odontonema 8/8/29. Pollen oblate spheroidal 
to spherical to euprolate (P:E = 0.9-1.8), 3(-4)- 
colporate, 6(-8)-pseudocolpate, P = 33-62 um, 
E = 30^4 um; polar outline circular to subtrian- 
gular; colpi fusiform, nearly as long as P, 2-11 
um wide at equator, not to barely exceeding 
diameter of ora in width, surfaces psilate to ver- 
rucate; pseudocolpi 2 per mesocolpium, shorter 
than or as long as colpi, sometimes fusing near 
one or both poles thereby forming pseudocolpal 
arcs or ellipses in mesocolpia, surfaces psilate to 
verrucate; interapertural surfaces (exclusive of 
pseudocolpi) foveolate to rugulate to reticulate 
(reticulum homobrochate) to bireticulate. Fig. 8i. 

Odontonema is entirely American in its distri- 
bution. Baum (1982) and Daniel (1995b) noted 
that pollen of most species of Odontonema (see 
figures in Daniel 1995b) resembles that of most 
other Odontoneminae. They also noted the ex- 
ceptional pollen of Odontonema cuspidatum 
which differs from that of all other species in the 
genus by having four colpi and eight pseudo- 
colpi. 

Poikilacanthus 3/3/10+ . Pollen spherical to 
euprolate (P:E = 1.0-1.4), 5 (-6?)-porate, P = 
53-65 um, E = 49-57 um; polar outline ± pen- 



228 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



tagonal (where seen); pores ± circular, 6-7 um in 
diameter, surfaces psilate to subverrucate; in- 
terapertural surfaces covered with subcircular to 
rectangular to polygonal insulae, insulae com- 
prising gemmate regions enclosed by thick, psi- 
late to foveolate muri, either discrete (except 
sometimes near poles) and ± evenly distributed 
over surface or sharing common endwalls and 
arranged in loops and bands (loops enclosing a 
band), sometimes becoming irregularly reticu- 
late with internal muri; surfaces between insulae 
(or between bands and/or loops) irregularly re- 
ticulate. Fig. 6e, f. 

This genus is entirely American in distribution. 
Two distinct types of pollen (see figures in 
Daniel 1991b; Ramamoorthy 1989; Scotland 
1992) are evident among Mexican species of the 
genus. Poikilacanthus macranthus has discrete 
insulae whereas P. capitatus and P. no- 
vogalicianus have similar "insulae" linked in 
loops and bands. The apertures of Poikilacanthus 
pollen might with equal justification be treated 
as compound (i.e., comprising a colpus and an 
os) in both pollen types. Scotland (1992) noted 
that pollen of Poikilacanthus is 6-aperturate. Pol- 
len of two Mexican species was observed to be 
5-aperturate. The number of apertures in pollen 
of P. capitatus could not be determined with 
certainty, but likely is six. 

Pollen of Poikilacanthus has been considered 
to be unique in the family (Raj 1961). Because 
pollen characters have traditionally been 
weighted heavily in infrafamilial classification 
of the Acanthaceae, the tribal position of the 
genus has sometimes been unclear. Based on 
macromorphological characters, especially the 
presence of a rugula in the corolla, Poikilacan- 
thus resembles genera of Bremekamp's Justicii- 
nae. In fact, pollen characters appear to be the 
only consistent means of distinguishing Poikila- 
canthus from Justicia. Pollen like that of Poikila- 
canthus is not known among the diverse pollen 
types so far recorded fox Justicia (Graham 1988; 
although see discussion of J. masiaca above un- 
der Justicia, the pollen of which shows some 
similarities in exine sculpturing to that of 
Poikilacanthus). Pollen somewhat similar to that 
of Poikilacanthus macranthus (i.e., with the sur- 
face covered by discrete insulae) is known in the 
unispecific American genus Megaskepasma Lin- 
dau (which appears to have six or more aper- 
tures). Studies of the latter genus that address its 



relationship to the former are desirable. Based on 
macromorphological and, to a lesser extent, pa- 
lynological lines of evidence, Poikilacanthus 
could be readily accommodated in the Justici- 
inae. 

Pseuderanthemum 9/10-15/40-60. Pollen 
oblate spheroidal to euprolate (P:E = 0.9-1.7), 
3-colporate, 6-pseudocolpate, P = 34-68 um, E 
= 34-46 um; polar outline circular to subtriangu- 
lar; colpi fusiform, nearly as long as P, 1-15 um 
wide at equator, not to barely exceeding diameter 
of ora in width, sometimes fused at poles (i.e., 
3-syncolporate), surfaces (subpsilate to) verru- 
cate; pseudocolpi 2 per mesocolpium, shorter 
than or as long as colpi, sometimes fusing near 
one or both poles thereby forming pseudocolpal 
arcs or ellipses in mesocolpia, surfaces (subpsi- 
late to) verrucate; interapertural surfaces (exclu- 
sive of pseudocolpi) (rugulate to) reticulate, 
reticulum homobrochate, lumina sometimes 
containing one or more pilae. Fig. 7f. 

Pseuderanthemum is a pantropical genus 
much in need of revision. Pollen of those Mexi- 
can species examined (see figures in Daniel 
1993b) is typical of that generally found in other 
Mexican Odontoneminae. 

Ruellia 35/ca. 65/ca. 250. Pollen spherical to 
subspheroidal (including at least oblate spheroi- 
dal and prolate spheroidal, see discussion be- 
low), 3-porate (as observed in eight species, see 
discussion), longest diameter 45-99 urn; polar 
outline circular; pores 7-20 urn diameter, sur- 
faces psilate to verrucate; interapertural surfaces 
coarsely reticulate, reticulum ± homobrochate, 
muri sharply acute to rounded, psilate (to irregu- 
larly microverrucate to microechinate), some- 
times incomplete or irregular (especially near 
pores), lumina psilate to verrucate to gemmate, 
verrucae or gemmae usual ly ± evenly distributed, 
varying in number and size. Figs. 5a-c; 1 lh. 

Species of Ruellia occur worldwide and the 
genus is the second largest among Mexican 
Acanthaceae. Considering the diverse vegetative 
and floral characteristics among Mexican Ruel- 
lia, pollen of Mexican species (see figures in 
Daniel 1990a, 1990d, 1993b, 1996, 1997; 
Ramamoorthy and Hornelas U. 1988) is remark- 
ably uniform in general aspect. This uniformity 
is further evident on a worldwide basis (Furness 
and Grant 1996; Muller et al. 1989; Raj 1961). 
Most of the 3 1 species of Ruellia from Africa and 
Madagascar examined by Furness and Grant 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



229 



(1996) have pollen greatly resembling that of 
Mexican taxa. They noted deviations from this 
basic pattern in two species that were 5-porate, 
three species that have a region around the three 
pores resembling a very short colpus, and two 
species with a clavate (vs. coarsely reticulate) 
surface. They further noted that each of the 3 1 
Old World species studied has a unique combi- 
nation of pollen characters. Most of the observed 
variation in pollen of Mexican species of Ruellia 
involves grain size, form of the muri, and size and 
sculpturing of the lumina. Further study of Mexi- 
can species of Ruellia will be necessary to deter- 
mine whether their pollen provides a similar suite 
of diagnostic characters at the specific level. 

The particular form and sculpturing of Ruellia 
pollen can make thorough analysis of the grains 
based solely on SEM preparations difficult. Be- 
cause of their more or less spherical form and 
coarse reticulum (which often makes locating the 
apertures difficult), the polar axis and equatorial 
plane can be impossible to determine. These are 
discernible only when at least two apertures are 
in the field of view. In the limited number of 
grains where such was the case, the P:E varies 
from oblate spheroidal to prolate spheroidal. In 
some preparations, no apertures were observed. 
Thus, it could not be determined with certainty 
whether all Mexican species studied indeed have 
three apertures or whether they all lack regions 
adjacent to the apertures suggestive of colpi. 

Furness and Grant (1996) noted correlations 
among species with large (92-117 urn) pollen 
and presumed moth and bat pollination. The 
Mexican species examined here were grouped 
into seven classes based on differences in floral 
size, shape, and color that are presumably related 
to pollination syndromes. Although Mexican 
species with flowers that would appear to fit into 
bat (Faegri and van der Pijl 1979) and humming- 
bird (Grant and Grant 1968) syndromes have the 
largest pollen (up to 99 urn diameter), there is 
sufficient overlap among all seven classes so as 
to suggest that making correlations between pol- 
len size and pollinators among Mexican species 
would be premature at this time. 

Among Mexican Acanthaceae, pollen of Ruel- 
lia most closely resembles that of Barleria (of 
Bremekamp's Ruellieae: Barleriinae). Similari- 
ties between pollen of these genera were also 
noted by Furness and Grant (1996). Pollen simi- 
lar to that of Ruellia is also known elsewhere 



among flowering plants (e.g. Geraniaceae, Scot- 
land 1990b). 

Sanchezia 2/2/ca. 60. Pollen globose-oblong 
(longer apertural axis:interapertural axis = 1.9), 
2-colporate, polypseudocolpate, sculptural fea- 
tures of one face ± 90° out of phase with those of 
opposite face, longer apertural axis 80-88 urn, 
shorter apertural axis 61-65 urn, interapertural 
axis 44 |im, apertural face elliptic in outline, 
interapertural face more narrowly elliptic in out- 
line; colpi ± linear, shorter than longer axis of and 
longer than shorter axis of apertural face, 1 urn 
wide near ora, surfaces microverrucate; pseudo- 
colpi 12-14 per apertural face, of varying lengths 
(progressively larger away from colpi), longer 
ones fusing near periphery of grain, oriented in ± 
same direction as colpi (i.e., ± perpendicular to 
one another on opposite apertural faces), some 
appearing to encircle pollen in interapertural 
view, surfaces microverrucate to subverrucate; 
exine between colpi divided into bands by 
pseudocolpi, ± prominently protruding (liplike) 
on both sides of colpi at ora, bands psilate with a 
single row of foveae down the center, longest 
axis of foveae 1-4 urn. Fig. 3d, e. 

Several species of this neotropical genus are 
cultivated for ornament. One of them, S. parvi- 
bracteata, is possibly native or naturalized in 
southern Mexico (Daniel 1995c) and another, S. 
speciosa is cultivated and possibly naturalized. 
Pollen of Sanchezia occurring in Mexico, like 
that of other Trichanthereae, is "loxodicolporate" 
(see Bravaisia). Based on specimens examined, 
the longer apertural axis is longer in Sanchezia 
than in Bravaisia and the perforations in the 
bands of Sanchezia tend to be larger than those 
in Bravaisia. Vasanthy and Pocock (1986) noted 
gradations between radially symmetric and "ro- 
tationally symmetric" pollen in a different spe- 
cies of the genus (S. lampra Leonard & L. B. 
Sm.). 

Schaueria 1/1/10-15. Pollen prolate spheroi- 
dal to subprolate (P:E = 1.1-1.3), 3-colporate, 
6-pseudocolpate, P = 42-46 Jim, E = 36-39 urn; 
polar outline not seen; colpi fusiform, nearly as 
long as P, 10-13 urn wide at equator, barely 
exceeding diameter of ora in width, surfaces 
microverrucate to verrucate; pseudocolpi 2 per 
mesocolpium, shorter than or as long as colpi, 
sometimes fusing near one or both poles thereby 
forming pseudocolpal arcs or ellipses in meso- 
colpia, surfaces microverrucate to verrucate; in- 



230 



PROCEEDINGS OF THE CALIFORNIA ACADEMv OF SCIENCES 

Volume 50, No. 8 



terapertural surfaces (exclusive of pseudocolpi) 
reticulate (reticulum ± homobrochate) to bire- 
ticulate, reticulum sometimes irregularly frag- 
mented (sometimes into insulae) and surrounded 
by thin, verrucate regions. Fig. 7h. 

This neotropical genus of 10 to 15 species 
occurs in Mexico, Guatemala, and South Amer- 
ica. Its pollen (see figures in Daniel 1990a; Hil- 
senbeck and Marshall 1983) is generally like that 
of other Mexican Odontoneminae. Fragmenta- 
tion of that portion of the mesocolpium between 
the pseudocolpi, as seen in pollen from several 
collections, has not been observed in other Mexi- 
can Odontoneminae. Although not observed in 
Schaueria, similar fragmentation in that portion 
of the mesocolpium between the colpus and the 
adjacent pseudocolpi would result in pollen simi- 
lar to that found in certain species of Justicia. 

Spathacanthus 2/2/3. Pollen prolate spheroi- 
dal to euprolate, (P:E = 1.1-1.5), 3 (-4)-colpo- 
rate, 6 (-8)-pseudocolpate, P = 39-64 um long, 
E = 28-43 urn; polar outline subtriangular to 
subcircular (to square); colpi fusiform, ca. 1/2 to 
nearly as long as P, 10-15 um wide at equator, 
not to barely exceeding diameter of ora in width, 
surfaces microverrucate to verrucate; pseudo- 
colpi 2 per mesocolpium, as long as or longer 
than colpi, sometimes fused near one or both 
poles thereby forming pseudocolpal arcs or ellip- 
ses in mesocolpia, surfaces subpsilate to micro- 
verrucate; interapertural surfaces (exclusive of 
pseudocolpi) foveolate to fossulate to verrucate 
to rugulate. Fig. 8g. 

Spathacanthus occurs only in southern Mexico 
and Central America. Its pollen is generally like 
that of most other Mexican Odontoneminae (see 
Chalarothyrsus), except that the interapertural 
surfaces are not conspicuously reticulate. One 
collection of S. parviflorus (Skutch 961) has pol- 
len with four colpi whereas another (Matuda 
1696) has some grains with three colpi and others 
with four. Variation of aperture number within a 
species was encountered elsewhere among 
Mexican Acanthaceae only in Stenandrium 
dulce. Figure 8 shows pollen of a Costa Rican 
species, S. hoffmannii Lindau, that resembles 
pollen of Mexican species. 

Staurogyne 1/1/80-140. Pollen (see discus- 
sion) perprolate (P:E = 2.2-2.3) or suboblate to 
spherical (P:E = 0.8-1.0), 3-colporate or 3-col- 
pate to 3-colporoidate, P = 28-30 |um and E = 
12-13 um or P = 15-17 urn and E = 17-18 um; 



polar outline subcircular to subtriangular; colpi 
narrowly linear or fusiform, nearly as long as P, 
1 um or 3-4 jim wide at equator, surfaces verru- 
cate (where seen); interapertural surfaces reticu- 
late, reticulum ± homobrochate. Fig. lj. 

Staurogyne is a pantropical genus with a ma- 
jority of species occurring in southeastern Asia. 
Pollen of the sole species known from North and 
Central America, S. agrestis, is described above. 
It appears to be similar to that reported elsewhere 
in the genus (Raj 1961; Scotland 1990a). Pollen 
of Lott et al. 3396 (see figure in Daniel and Lott 
1993) is spherical to oblate and conspicuously 
colporate whereas pollen of Lott et al. 3226 is 
perprolate and either colpate or colporoidate (i.e., 
endoapertures, if present, were not seen). Inter- 
estingly, these two collections were both made in 
the same general region of coastal Jalisco. Scot- 
land (1990a) indicated that pollen of species he 
studied were 3-colporate but often with the en- 
doapertures indistinct. Among Mexican Nel- 
sonioideae, the interapertural exine sculpturing 
of pollen of Staurogyne is more like that of 
Nelsonia than Elytraria. Based on the samples 
examined, S. agrestis has smaller pollen than 
other Mexican species of the subfamily. 

Stenandrium 9/9/ca. 65. Pollen (except some 
S. dulce, see below) euprolate to perprolate (P:E 
= 1.4—2.2), sometimes slightly hourglass shaped 
due to a slight equatorial constriction, 3 -colpate, 
P = 33-55 um, E = 18-29 jim; polar outline 
circular to triangular; colpi linear to hourglass 
shaped, nearly as long as P, 1-6 um wide at 
equator, sometimes bifurcating near poles (arms 
of bifurcations sometimes fusing with adjacent 
arms of other colpi near one or both poles), 
surfaces psilate to microverrucate to verrucate to 
subrugulate; interapertural surfaces psilate to 
foveolate to fossulate to rugulate. Pollen of some 
individuals of S. dulce: spherical (P:E = ca. 1.0), 
33-42 um in diameter, inaperturate, surfaces 
verrucate to gemmate, verrucae and gemmae 
usually of two distinct sizes. Fig. 2e-i. 

Stenandrium occurs in Africa, Madagascar, 
and the New World. Palacios C. and Quiroz G. 
(1994) described and figured pollen of seven of 
the Mexican species. The observations above 
include information from the two Mexican spe- 
cies, S. subcordatum and S. verticillatum, not 
examined by them. In general, my observations 
agree with those of Palacios C. and Quiroz G. 
(1994); however, I could not substantiate their 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



231 



descriptions of S. dulce as trichotomocolpate, S. 
mancnonense as dicolpate, and S. pilosulum as 
dicolpate to tricolpate. (In fact, some confusion 
exists in their paper concerning the number of 
apertures in the latter two taxa, cf. the table vs. 
the text.) Furness (1993) recognized four pollen 
types among Old World species of Stenandrium 
studied by her. Pollen of most Mexican species 
more closely resembles her pollen type I which 
is only found in taxa from Madagascar. The 
unusual inaperturate pollen of some individuals 
of S. dulce has been reported for the species from 
throughout its wide range (Ariza Espinar and 
Ferrucci 1982; Furness 1993; Heusser 1971; 
Marticorena P. 1968). I have also examined pol- 
len of this species from Florida (Standley 71), 
Argentina (Renvoize 3428), and Bolivia (David- 
son 3746); pollen from each of these collections 
resembles that described above for individuals 
with inaperturate grains. Lindau (1895) divided 
Stenandrium into two sections based solely on 
these two different types of pollen. 

Interestingly, among the Mexican collections 
of S. dulce studied, both 3-colpate and inapertu- 
rate pollen was encountered (but never both types 
from the same collection). Three collections 
(Breedlove & Almeda 45287, Chiang et al. 8198, 
and Hinton 6086) have the euprolate to perpro- 
late, 3-colpate pollen resembling other Mexican 
species of the genus whereas three other collec- 
tions (Angeles 2, Hernandez M. 5995, and 
Padilla 8) have the spherical, inaperturate grains. 
Further study is needed to determine whether the 
presence of two pollen types among Mexican 
collections treated as S. dulce is indicative of 
inadequate taxonomic refinement or a remark- 
able palynological dimorphism within a single 
species. The sections of Stenandrium as delim- 
ited by Lindau (1895) also would appear to re- 
quire some modification. Pollen similar to the 
spherical, inaperturate grains sometimes found in 
S. dulce also is known in the South American 
species (and apparent close relatives of S. dulce) 
S. trinerve Nees (Petriella 1968; Raj 1961) and 
S. pohlii Nees (Raj 1 96 1 ). In fact, Ezcurra ( 1 993) 
utilized this pollen difference in her key to spe- 
cies and included one variant of S. trinerve within 
S. dulce. 

Furness (1993) noted the palynological simi- 
larity between Stenandrium and Crossandra Sal- 
isb., an Old World genus of the Acantheae. 
Among New World Aphelandreae, pollen of 



Stenandrium resembles that of Holograph is and 
Aphelandra, the only other representatives of this 
tribe in Mexico. Even some of the same tenden- 
cies observed in Stenandrium (e.g., colpi bifur- 
cating near poles) are to be found in both of these 
other genera. 

Stenostephanus 15/15/ca. 65. Pollen globose- 
elliptic to globose-elongate (longer E:shorter E = 
1.0-2.8), 2-porate, diameter in apertural view = 
3 1-56 um, E in interapertural view = 16—48 urn; 
outline in apertural view circular to subcircular, 
outline in interapertural view narrowly elliptic to 
broadly elliptic to circular; pores 3-9 um diame- 
ter, verrucate to gemmate to echinate, each pore 
surrounded by a circular region 17-47 um in 
diameter, surface of circular regions subpsilate to 
microverrucate to microrugulate and also gem- 
mate to baculate to echinate, the two circular 
regions separated by a peripheral band 2-15 um 
wide, surface of peripheral band psilate to verru- 
cate to rugulate (and sometimes with a central 
row of gemmae to baculae to echinae). Fig. 6a-d. 

This neotropical genus is most diverse in the 
northern Andes of South America. Two distinc- 
tive forms of pollen (see additional figures in 
Daniel 1995a) were encountered among the 
Mexican species of Stenostephanus: globose- 
elongate (i.e., longer E:shorter E = 2.0-2.8 and 
the interapertural outline narrowly elliptic; 6 
spp.) and globose-elliptic (i.e., longer E:shorter 
E = 1 .0-1 .5 and the interapertural outline broadly 
elliptic to circular; 9 spp.). Whether this differ- 
ence can be correlated with other characters and 
thus be useful in the infrageneric classification of 
Stenostephanus is currently under study. Consid- 
erable variation in the width and sculpturing of 
the peripheral band was noted among species. 

Stenostephanus and its relatives were included 
by Bremekamp (1965) in subtribe Isoglossinae 
(as "Rhytiglossinae") of tribe Justicieae. Macro- 
morphologically, Stenostephanus is similar to 
the African genera Oreacanthus Benth. and 
Brachystephanus Nees. Pollen similar to both 
types found in Stenostephanus is also found in 
Brachystephanus and the globose-elongate pol- 
len of Stenostephanus is characteristic of Orea- 
canthus. Brachystephanus appears to encompass 
considerably more diversity in its pollen than 
seen among Mexican Stenostephanus (Figue- 
iredo and Keith-Lucas 1996). The taxonomic 
relationships of Stenostephanus and its New 
World relatives with similar pollen (e.g., Razisea 



232 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



Oerst.) to various other Old World genera with 
superficially similar pollen (e.g., Conocalyx 
Benoist, Forcipella Baill., Isoglossa Oerst., 
Leandriella Benoist, and Sphacanthus Benoist, 
as depicted by Muller et al. 1989) warrant further 
study. 

Tetramerium 20/21/28. Pollen subprolate to 
perprolate (P:E = 1.3-3.0), 3-colporate, 6- 
pseudocolpate, P = 29-79 (-120) urn, E = 14-33 
(-41) urn; polar outline circular to triangular; 
colpi fusiform, nearly to as long as P, 1-5 urn 
wide at equator, not to barely exceeding diameter 
of ora in width, surfaces micro verrucate to ver- 
rucate to subgemmate; pseudocolpi 2 per meso- 
colpium, nearly to as long as colpi, often fusing 
near poles thereby forming pseudocolpal ellipses 
in mesocolpia, surfaces psilate to microverru- 
cate; interapertural surfaces (exclusive of 
pseudocolpi) (rugulate to) reticulate (reticulum 
homobrochate) to bireticulate. Fig. 7b. 

Most of the species in this American genus 
occur in Mexico. Except for considerable vari- 
ation in size, pollen morphology is highly con- 
sistent throughout the genus and resembles that 
of other Mexican Odontoneminae (see figures in 
Daniel 1986a, 1997). 

Thunbergia 4/5/100-200. Pollen spherical to 
subprolate (P:E = 1.0-1.2), spiraperturate (= spi- 
rotreme), P = 49-74 um, E = 45-66 um; polar 
outline circular; apertures one or apparently sev- 
eral per grain, dividing grain into horizontal 
bands in equatorial view, 1—4 um wide, surfaces 
psilate to scabrate to verrucate; interapertural 
surfaces subpsilate (e.g., T. alata) to verrucate 
(e.g., T. erecta) to rugulate with a single line of 
prominent verrucae to baculae (e.g., T.fragrans). 
Fig. la-d. 

At least five species of this paleotropical genus 
are cultivated in Mexico and at least two of those 
are naturalized as well. The taxonomic position 
of the Thunbergioideae (i.e., whether treated as a 
subfamily of Acanthaceae or a separate family) 
remains to be resolved. Pollen of species studied 
here resembles that described from other collec- 
tions and other species of Thunbergia (Bre- 
mekamp 1955; Erdtman 1952; Natarajan 1957; 
Raj 1961, 1973; Roubik and Moreno P. 1991; 
Valsaladevi andMathew 1989). Linear-spiral ap- 
ertures (or their derivatives), as seen in pollen of 
Thunbergia, are synapomorphic for Pseudocalyx 
Radlk. and Thunbergia (but not Meyenia Nees) 
in the Thunbergioideae among Acanthaceae. 



Spiraperturate pollen is known outside the Acan- 
thaceae in genera representing several families 
(Sheta and Brack-Hanes 1984). 

Yeatesia 2/2/3. Pollen subprolate to euprolate 
(P:E = 1.3-1.4), 3-colporate, 6-pseudocolpate to 
6-pseudocolpoidate, P = 47-5 1 um, E = 36-42 
um; polar outline subtriangular; colpi fusiform, 
ca. 1/2 to nearly as long as P, 3-8 |im wide at 
equator, not to barely exceeding diameter of ora 
in width, surfaces micro verrucate to verrucate to 
gemmate; pseudocolpi 2 per mesocolpium, dis- 
tinct to subdistinct, mostly shorter than colpi, 
surfaces microverrucate to verrucate; interaper- 
tural surfaces (exclusive of pseudocolpi) reticu- 
late (reticulum homobrochate) to bireticulate, 
muri psilate or microverrucate. Fig. 7j. 

Yeatesia is endemic to North America. Its pol- 
len (see figures in Hilsenbeck 1989) is similar to 
that of other genera of Odontoneminae. Pollen of 
Y. platystegia differs from that of Y. mabryi by 
its reticulate (vs. bireticulate) sculpturing with 
microverrucate (vs. psilate) muri and poorly de- 
veloped (vs. well developed) pseudocolpi. Ace- 
tolyzed pollen of Y. platystegia (Hilsenbeck 
1989) shows even less distinct pseudocolpi and 
poorly developed colpi. In all other features, my 
observations agree with photographs of Yeatesia 
pollen provided by Hilsenbeck (1989). The only 
other species of the genus, Y. viridiflora (Nees) 
Small, which is endemic to the United States, has 
pollen similar to that of Y. platystegia but with 
even less distinct pseudocolpi. Species of the 
genus appear to exhibit decreasing development 
of the pseudocolpi in a northward geographic 
direction. 

Summary and Conclusions 

The tremendous diversity of pollen morphol- 
ogy in the family is amply reflected among the 
genera of Mexican Acanthaceae. Palynological 
diversity is evident in grain size (P varies from 
21 to 140 um) and shape (oblate to prolate); 
aperture number (two to more than fifty), shape 
(colpi or pores), structure (simple or compound), 
and position (equatorial to global); and exine 
sculpturing (e.g., psilate to fossulate to reticulate 
to gemmate). The diversity of acanthaceous pol- 
len in Brazil, India, and Madagascar would likely 
equal, or even surpass, that so far documented 
from Mexico if sufficiently detailed palynologi- 
cal studies of those countries, each also rich in 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



233 



Acanthaceae, were available. In the genera stud- 
ied here, pollen diversity is generally taxonomic 
rather than geographic in nature. For example, 
pollen of genera comprising both Mexican and 
Old World species (e.g., Dicliptera, Dyschoriste, 
Hygrophila, Justicia, Ruellia, and Stenandrium) 
is decidedly consistent within each genus. 

The morphological attributes considered in 
this study provide useful systematic data primar- 
ily at the generic and suprageneric levels. Most 
Mexican genera consist of species with pollen 
that is relatively homogeneous, especially in 
characteristics of the apertures. Therefore, pollen 
is often useful in the characterization of genera 
(e.g., Henrya, Louteridium, Ruellia, Poikilacan- 
thus). Although most genera have relatively ho- 
mogeneous pollen, others show heterogeneity 
among species to a greater (e.g., Justicia) or 
lesser (e.g., Odontonema) extent. The presence 
of one species with unusual pollen in an other- 
wise palynologically homogeneous genus was 
encountered several times (e.g., Stenandrium, 
Odontonema). One limitation of broad, superfi- 
cial surveys of pollen is that this latter type of 
variation, which can be crucial in determining 
character polarity for phylogenetic reconstruc- 
tion, is often missed. The presence of individual 
species with unique pollen within a genus em- 
phasizes the need to study pollen of all species of 
genera. A similar argument might be applied to 
species where extensive sampling of individuals 
should likewise be undertaken. For the most part 
in this study few samples from any single species 
were studied; therefore, variation at the species 
level usually was not ascertained. In most in- 
stances where more than one sample of a species 
was studied, little variation was observed. In 
some cases, however, where significant variation 
was seen, more extensive sampling was under- 
taken. Further studies will be needed to better 
understand the significance of palynological 
variation in species such as Aphanosperma si- 
naloensis, Spathacanthus parvi/lorus, and 
Stenandrium dulce. 

Other than the lack of extensive sampling 
within a species, a significant deficiency of this 
investigation is the lack of observations on pollen 
structure. Studies of the internal structure of the 
exine should help to resolve whether pollen of 
Mendoncia, Elytraria, and Nelsonia in Mexico 
have endoapertures as reported previously but 
which were not evident in the samples I ob- 



served. Also, the sculpturing patterns of in- 
terapertural surfaces of unacetolyzed pollen 
likely differs from that of acetolyzed grains in 
some instances (e.g., some surfaces here charac- 
terized as reticulate might be revealed to be 
bireticulate in acetolyzed pollen). 

Obvious correlations of pollen form to known 
pollinators were not established by my observa- 
tions (see Ruellia). In some instances a positive 
correlation between pollen size and flower size 
was noted (e.g., Braviasia and Justicia masiaca, 
cf. Daniel 1995d). Unusual features of pollen 
grains sometimes could be associated with tax- 
onomy, but other times could not. In several 
genera, the colpi often contain an elevated ridge 
surrounding the ora and tapering toward the 
poles. The ridge is usually covered with larger 
sculptural elements than the surrounding colpal 
surface. This colpal ridge is particularly evident 
in pollen of Gypsacanthus, Mexacanthus, and 
Schaueria; but it was also observed in some 
pollen of Anisacanthus, Carlowrightia, Tetra- 
merium, and Yeatesia. All of these genera are in 
subtribe Odontoneminae. Another unusual paly- 
nological feature, protruding (liplike) exine on 
each side of the ora, was seen in pollen of the 
taxonomically unrelated genera Dyschoriste and 
Bravaisia. A similar and very rare feature present 
in even more distantly related genera is the tri- 
partite, aperturelike indentation at the poles in 
some individuals of Aphanosperma and in some 
species of Aphelandra. These regions of thin 
exine are probably best considered as 
pseudoapertures, and like pseudocolpi, possibly 
have a harmomegathic function. 

Because pollen characters have been routinely 
used in suprageneric classification of Acan- 
thaceae, it is not surprising that there tend to be 
good correlations between pollen form and sys- 
tematic classification. As noted in the generic 
discussions above, several of the suprageneric 
taxa listed in Table 1 are supported by 
palynological as well as macromorphological 
characters. For example, each of the Mendon- 
cioideae, Trichanthereae, and Louteridieae has a 
unique pollen type in the family. Bremekamp's 
(1965) classification, although incomplete, of- 
fers some advantages with respect to that of 
Lindau (1895). Daniel, Chuang, and Baker 
(1990) noted that it incorporated some related 
taxa, previously treated in different tribes, into 
the same subtribe. It also unites within the Jus- 



234 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



ticieae two subtribes (Justiciinae and Odon- 
toneminae) that share a common pollen type (i.e., 
3-colporate, 6-pseudocolpate). Poikilacanthus , 
treated by Bremekamp (1965) as being of uncer- 
tain position in the Justicieae, could be accom- 
modated in subtribe Justiciinae. The unusual 
pollen of Stenostephanus and its relatives in sub- 
tribe Isoglossinae suggest that it may not be as 
closely related to the other subtribes in the Jus- 
ticieae as they are to one another. Bremekamp's 
classification would also appear to have some 
disadvantages with respect to that of Lindau. For 
example, Lindau ( 1 895) included Barleria, Lepi- 
dagathis, and Lophostachys within the same tribe 
(Barlerieae) whereas Bremekamp (1965) re- 
moved Lepidagathis to a different tribe. The 
palynological similarities of these genera, com- 
bined with certain macromorphological charac- 
teristics, suggest that they might be better treated 
in the same tribe. In fact, the generic distinctions 
between the latter two genera have been ques- 
tioned (Benoist 191 1). Also, Bremekamp's sub- 
tribe Odontoneminae brings together many 
Mexican genera with similar pollen (i.e., 3- 
colporate, 6-pseudocolpate or pollen derived 
from this type). Macromorphological and cyto- 
logical evidence, however, suggests at least three 
groupings among the genera here included in the 
subtribe (Daniel and Chuang 1993). 

Because pollen of Acanthaceae is diverse and 
because it is often rather homogeneous within a 
genus, palynological characters will continue to 
be used both in the characterization of genera and 
in assessing their phylogenetic relationships. De- 
spite more than a century of describing pollen of 
Acanthaceae, many genera remain palynologi- 
cally inadequately diagnosed and the pollen of 
most species has yet to be described. Descrip- 
tions of pollen, based on original observations, in 
monographic and floristic studies of Acan- 
thaceae are highly desirable. 

ACKNOWLEDGMENTS 

I am grateful to the curators of the herbaria 
(ASU, CAS, DS, ENCB, F, GH, K, LL, MEXU, 
MICH, MO, NY, POM, RS A, UC, US) that made 
specimens available for my study; to the Univer- 
sity of Michigan, Arizona State University, and 
the California Academy of Sciences for making 
SEM facilities available to me; to Darrell Ubick, 
Mary Ann Tenorio, and Lisa Borok for supervis- 



ing operation of SEM at the California Academy 
of Sciences; to Dong Lin for scanning and re- 
touching the photographic negatives; to Mary Jo 
Sutton for creating the composite plates from the 
electronically-stored images; and to Jim Doyle 
who first stimulated my interest in pollen. 

LITERATURE CITED 

ACOSTA C, S. 1989. Chaetothylax rzedowskii (Acan- 
thaceae), una especie nueva de Chiapas, Mexico. 
ActaBot. Mex. 5:5-11. 

Acosta C, S. and T. F. Daniel. 1 993. A new species 
of Justicia (Acanthaceae) from southern Mexico. 
Kew Bull. 48:1 19-123. 

Balkwill, K. 1985. Taxonomic studies in the tribe 
Justicieae of the family Acanthaceae. Unpublished 
Ph.D. thesis, Univ. of Natal, Pietermaritzburg, Re- 
public of South Africa. 

Balkwill, K. and F. G. Norris. 1985. Taxonomic 
studies in the Acanthaceae; the genus Hypoestes in 
southern Africa. S. African J. Bot. 51 : 133—144. 

Ariza Espinar, L. and S. Ferrucci. 1982. Las espe- 
cies de Stenandrium (Acanthaceae) del centro de 
Argentina. Kurtziana 15:63-80. 

Barker, R. M. 1986. A taxonomic revision of Aus- 
tralian Acanthaceae. J. Adelaide Bot. Gard. 
9:1-286. 

Baum, V. M. 1982. A revision of the genus Odon- 
tonema (Acanthaceae). Unpublished M.S. thesis, 
Univ. of Maryland, College Park. 

Benoist, R. 1911. Les genres Lepidagathis et Lo- 
phostachys sont-ils distincts? Notul. Syst. (Paris) 
2:139-144. 

Bentham, G. and J. D. Hooker. 1876. Genera plan- 
tarum, Vol. 2, part 2. Reeve and Co., London. 

Bhaduri, S. 1944. A contribution to the morphology 
of pollen grains of Acanthaceae and its bearing on 
taxonomy. J. Dept. Sci. Calcutta Univ. 1:25-58. 

Bremekamp, C. E. B. 1948. Notes on the Acan- 
thaceae of Java. Verhand. Konin. Nederl. Akad. 
Wetensch., Afd. Natuur. Tweede Sec. 45(2): 1-78. 



. 1955. The Thunbergia species of the Male- 

sian area. Verhand. Konin. Nederl. Akad. 



Wetensch., Afd. Natuur. Tweede Sec. 50(4): 1-90. 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



235 



-. 1965. Delimitation and subdivision of the 



Acanthaceae. Bull. Bot. Surv. India 7:21-30. 

Cronquist, A. 1981. An integrated system of classi- 
fication of flowering plants. Columbia Univ. Press, 
New York. 



. 1991b. A synopsis of Poikilacanthus (Acan- 
thaceae) in Mexico. Bull. Torrey Bot. Club 
1 18:451-458. 



. 1992. Acanthaceae: Mendoncioideae of 

Mexico. Acta Bot. Mex. 17:53-60. 



Daniel, T. F. 1978. A new Mirandea (Acanthaceae) 
from Nuevo Leon, Mexico. Syst. Bot. 3:428-433. 

. 1 98 1 . Mexacanthus, a new genus of Acan- 



thaceae from western Mexico. Syst. Bot. 
6:288-293. 

. 1983a. Carlowrightia (Acanthaceae). Flora 



Neotropica 34: 1-116. 



. 1983b. Systematics of Holographs (Acan- 
thaceae). J. Arnold Arbor. 64:129-160. 

. 1986a. Systematics of Tetramerium (Acan- 



thaceae). Syst. Bot. Monogr. 12:1-134. 



1 986b. New and reconsidered Mexican Acan- 



thaceae. II. Southw. Naturalist 31:169-175. 



-. 1 988a. Aphanosperma, a new genus of Acan- 



thaceae from Mexico with unusual diaspores. Amer. 
J. Bot. 75:545-550. 

. 1988b. A systematic study of Bravaisia DC. 



(Acanthaceae). Proc. Calif. Acad. Sci. 45:1 1 1-132. 
. 1988c. Three new species of Holographis 



(Acanthaceae) from Mexico. Proc. Calif. Acad. Sci. 
46:73-81. 

. 1990a. New and reconsidered Mexican Acan- 



thaceae. IV. Proc. Calif. Acad. Sci. 46:279-287. 



-. 1990b. Systematics of Henrya (Acantha- 



ceae). Contr. Univ. Michigan Herb. 17:99-131. 



1 990c. New and reconsidered Mexican Acan- 



thaceae. III. Contr. Univ. Michigan Herb. 
17:133-137. 

. 1 990d. New, reconsidered, and little-known 



Mexican species of Ruellia (Acanthaceae). Contr. 
Univ. Michigan Herb. 17:139-162. 

. 1991a. A revision of Aphelandra (Acan- 



thaceae) in Mexico. Proc. Calif. Acad. Sci. 

47:235-274. 



. 1993a. A synopsis of Lophostachys (Acan- 
thaceae) in Mexico and Central America. Selbyana 
14:64-70. 



1 993b. New and reconsidered Mexican Acan- 



thaceae. V. Contr. Univ. Michigan Herb. 
19:271-291. 

. 1 995a. New and reconsidered Mexican Acan- 



thaceae. VI. Chiapas. Proc. Calif. Acad. Sci. 
48:253-282. 

. 1995b. Revision of Odontonema (Acan- 



thaceae) in Mexico. Contr. Univ. Michigan Herb. 
20:147-171. 

. 1995c. Acanthaceae. Pp. 1-158 in Flora of 



Chiapas, Part 4, D. E. Breedlove, ed. California 
Academy of Sciences, San Francisco. 

. 1995d. Jiisticia masiaca (Acanthaceae), a 



new species from northwestern Mexico. Brittonia 
47:408^113. 

. 1 996. New and reconsidered Mexican Acan- 



thaceae. VII. Polibotanica2:l-9. 



1997. The Acanthaceae of California and the 



peninsula of Baja California. Proc. Calif. Acad. Sci. 
49:309-403. 

Daniel, T. F. and T. I. Chuang. 1993. Chromosome 
numbers of New World Acanthaceae. Syst. Bot. 
18:283-289. 

Daniel, T. F.,T. I. Chuang, and M. A.Baker. 1990. 
Chromosome numbers of American Acanthaceae. 
Syst. Bot. 15:13-25. 

Daniel, T. F. and E.J. Lott. 1993. Staurogyne 
agrestis (Acanthaceae), a genus and species new to 
Mexico. Sida 15:367-372. 

Erdtman, G. 1952. Pollen morphology and plant 
taxonomy. Almqvist and Wiksell, Stockholm. 

EZCURRA, C. 1 993. Acanthaceae. Pp. 278-359 in Flora 
de la Provincia de Jujuy (Repiiblica Argentina) 9, 



236 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



A. Cabrera, ed., Col. Cient. INTA 13, Buenos 
Aires. 

Faegri, K. and L. Van Der Pijl. 1979. The principles 
of pollination ecology, 3rd ed. Pergamon Press, 
Oxford. 

Figueiredo, E. and M. Keith-Lucas. 1996. Pollen 
morphology of Brachystephanus (Acanthaceae- 
Justicieae). Grana 35:65-73. 

Furness, C. A. 1989. The pollen morphology of 
Ecbolium and Megalochlamys (Acanthaceae). Kew 
Bull. 44:681-693. 

. 1990. Pollen morphology of Crossandra 



Salisbury and Crossandrella C. B. Clarke (Acan- 
thaceae: Acantheae). Grana 29:161-176. 

. 1 99 1 . The pollen morphology of Sclerochiton 



(Acanthaceae: Acantheae). Kew Bull. 46:51-59. 
. 1993. A pollen morphological survey of the 



Old World species of Stenandrium Nees (Acan- 
thaceae: Acantheae). Grana 32:1-1 1. 

. 1994. The pollen morphology of Hygrophila 



and Brillantaisia (Acanthaceae: Ruellieae). Acta 
Bot. Gallica 141:267-278. 

. 1995. A pollen morphological study of Dy- 



schoriste Nees and Chaetacanthus Nees (Acan- 
thaceae: Ruellieae). Rev. Palaeobot. Palynol. 
84:331-345. 

. 1996. Pollen morphology of Acanthopsis 



Harvey, Acanthus L. and Blepharis Jussieu (Acan- 
thaceae: Acantheae). Rev. Palaeobot. Palynol. 
92:253-268. 

Furness, C. A. and M. C. Grant. 1996. Pollen mor- 
phology of some Ruellia species (Acanthaceae) 
from Africa and Madagascar. Grana 35:231-239. 

Gibson, D.N. 1974. Acanthaceae. Pp. 328-461 in 
Flora of Guatemala, Fieldiana, Bot. 24 (part 10), 
P. C. Standley et al., eds. Field Museum of Natural 
History, Chicago. 

Graham, V. A. W. 1988. Delimitation and infra-ge- 
neric classification of Justicia (Acanthaceae). Kew 
Bull. 43:551-624. 

Grant, K. A. and V. Grant. 1968. Hummingbirds 
and their flowers. Columbia Univ. Press, New York. 



Hanks, S. and D. E. Fairbrothers. 1970. Effects of 
preparation technique on pollen prepared for SEM 
observations. Taxon 19:879-886. 

Henrickson, J. and P. Hiriart. 1988. New species 
and transfers into Justicia (Acanthaceae). Aliso 
12:45-58. 

Heusser, C. J. 1 97 1 . Pollen and spores of Chile. Univ. 
of Arizona Press, Tucson. 

Hilsenbeck, R. A. 1989. Taxonomy of Yeatesia 
(Acanthaceae). Syst. Bot. 14:427^438. 

. 1990. Pollen morphology and systematics of 



Siphonoglossa sensu lato (Acanthaceae). Amer. J. 
Bot. 77:27^40. 

Hilsenbeck, R. A. and D. L. Marshall. 1983. 
Schaueria calycobractea (Acanthaceae), a new spe- 
cies from Veracruz, Mexico. Brittonia 35:362-366. 

Hossain, A. B. M. E. 1984. Taxonomic notes on the 
Nelsonia canescens complex (Acanthaceae). Will- 
denowia 14:397^103. 

Immelman, K. L. 1987. A study of the genera of 
Justicia L. and Siphonoglossa Oerst. (Acanthaceae) 
in southern Africa. Unpublished Ph.D. thesis, Univ. 
of Natal, Pietermaritzburg, Republic of South Af- 
rica. 

LlNDAU, G. 1893. Beitrage zur Systematik der Acan- 
thaceen. Bot. Jahrb. Syst. 18:36-64. 



. 1895. Acanthaceae. Pp. 274-354 in Die 

nariirlichen Pflanzenfamilien 4 (3b), H. G. A. 
Engler and K. A. E. Prantl, eds Wilhelm Engel- 
mann, Leipzig. 

. 1904. Acanthaceae Americanae III. Bull. 



Herb. Boissier, Ser. 2, 4:313-328. 

LOTT, E. J., V. JARAMILLO L. AND J. RZEDOWSKI. 
1984. Un genero nuevo de la parte meridional de 
Mexico: Gypsacanthus (Acanthaceae, Justicieae, 
Odontoneminae). Bol. Soc. Bot. Mexico 46:47-5 1 . 

M articorena P., C. 1 968. Granos de polen de plantas 
Chilenas. 1. Gayana 17:1-66. 

McDade, L. 1984. Systematics and reproductive bi- 
ology of the Central American species of the 
Aphelandra pulcherrima complex (Acanthaceae). 
Ann. Missouri Bot. Gard. 71:104-165. 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



237 



Meagher, M. B. 1974. Biosystematic investigations 
in the genus Justicia (Acanthaceae). Unpublished 
Ph.D. thesis, Univ. of South Florida, Tampa. 

MULLER, J., M. SCHULLER, H. STRAKA AND B. FRIE- 

drich. 1989. Palynologia Madagassica et Mas- 
carenica. Fam. 182: Acanthaceae. Trop. Subtrop. 
Pflanzenwelt 67:138-187. 

Natarajan, A. T. 1957. Studies in the morphology of 
pollen — Tubiflorae. Phyton (Buenos Aires) 
8:21^12. 

Palacios C, R. and D. L. Quiroz G. 1994. Estudio 
palinologico de las especies mexicanas del genero 
Stenandriwn Nees (Acanthaceae). Phytologia 
76:433^140. 

Petrjella, B. 1968. El polen de las Acanthaceae 
Argentinas. Revista Mus. La Plata, Secc. Bot. 
11:51-68. 

Radlkofer, L. 1883. Ueber den systematischen 
Werth der Pollenbeschaffenheit bei den Acan- 
thaceen. Sitzungsber. Math.-Phys. CI. Konigl. 
Bayer. Akad. Wiss. Munchen 13:256-314. 

Raj, B. 1961. Pollen morphological studies in the 
Acanthaceae. Grana Palynol. 3:3-108. 

. 1973. Further contribution to the pollen mor- 
phology of the Acanthaceae. J. Palynol. 9:91-141. 

Ramamoorthy, T. P. 1989. Poikilacanthns capi- 
tatus: a new combination in Mexican Acanthaceae. 
Syst. Bot. 14:150-151. 

Ramamoorthy, T. P. and Y. Hornelas U. 1988. A 
new name and a new species in Mexican Ruellia 
(Acanthaceae). PI. Syst. Evol. 159:161-163. 

Rizzini, C. T. 1947. Estudos sobre as Acanthaceae. 
Bol. Mus. Nac. Rio de Janeiro, Bot. 8:1-38. 

Roubik,D. W. and J. E.Moreno P. 1991. Pollen and 
spores of Barro Colorado Island. Monogr. Syst. Bot. 
Missouri Bot. Gard. 36:1-270. 

Scotland, R. W. 1990a. Palynology and systematics 
of Acanthaceae. Unpublished Ph.D. thesis. Univ. of 
Reading, England. 

. 1 990b. Homology, pollen grains and micros- 
copy. Proc. Royal Microscop. Soc. 25:108. 



. 1991. A systematic analysis of pollen mor- 
phology of Acanthaceae genera with contorted co- 
rollas. Pp. 269-289 in Pollen and Spores, 
Systematics Association Special Volume 44, S. 
Blackmore and S. Barnes, eds. Clarendon Press, 
Oxford. 



1992. Pollen morphology and taxonomic 



characters in Acanthaceae. Syst. Bot. 17:337-340. 

Scotland, R. W., P. K. Endress, and T. J. 
Lawrence. 1994. Corolla ontogeny and aestivation 
in the Acanthaceae. Bot. J. Linn. Soc. 1 14:49-65. 

SHETA, M. H. and S. D. Brack-Hanes. 1984. Devel- 
opment of spiraperturate pollen in Thunbergia 
alata. Pollen and Spores 26:181-186. 

Sreemadhavan, C. P. 1977. Diagnosis of some new 
taxa and some new combinations in Bignoniales. 
Phytologia 37:412-416. 

Takhtajan, A. 1997. Diversity and classification of 
flowering plants. Columbia Univ. Press, New York. 

Vasanthy, G. and S. A. J. Pocock. 1986. Radial 
through rotated symmetry of striate pollen of Acan- 
thaceae. Can. J. Bot. 64:3050-3058. 

Valsaladevi, G. and P. M. Mathew. 1989. Pollen 
morphology of species of Thunbergia (Acan- 
thaceae) from South India. J. Palynol. 25:133-140. 

Walker, J. W. and J. A. Doyle. 1975. The bases of 
angiosperm phylogeny: palynology. Ann. Missouri 
Bot. Gard. 62:664-723. 

Wasshausen, D. C. 1975. The genus Aphelandra 
(Acanthaceae). Smithsonian Contr. Bot. 18:1-157. 

. 1 98 1 . New species of Justicia (Acanthaceae). 



Phytologia 49:65-68. 

. 1986. The systematics of the genus Pachy- 

stachys (Acanthaceae). Proc. Biol. Soc. Wash. 
99:160-185. 

. 1 996. New species and new combinations in 

Aphelandra (Acanthaceae) from Ecuador and adja- 
cent Peru. Nord. J. Bot. 16:389-407. 

Wasshausen, D. C. and T. F. Daniel. 1 995. Justicia. 
nevlingii (Acanthaceae), a new species from Mex- 
ico. Novon 5:1 14-117. 



238 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



Appendix 

Taxa and collections from which pollen was 
examined. Collections denoted with and asterisk 
(*) were made outside of Mexico. Taxa for which 
combinations have not been made in the genus in 
which they are included here are listed at the end 
of the genus (i.e., Justicia and Stenostephanus). 

Anisacanthus Nees 
A. andersonii T. F. Daniel: McCarten & Bittman 

2690. 
A.junceus (Torr.) Hemsl.: Daniel 479. 
A. linearis (Hagen) Henr. & E. J. Lott: Daniel 

538. 
A. puberulus (Torr.) Henr. & E. J. Lott: Johnston 

etal. 10571. 
A.pumilis (Dietr.) Nees: Lott & Magallanes 313. 
A. quadrifidus (Vahl) Nees: Daniel 256, Daniel 

1260. 
A. thurberi (Torr.) A. Gray: Daniel 118*. 

Aphanosperma T. F. Daniel 
A. sinaloensis (Leonard & Gentry) T. F. Daniel: 
Daniel 4070, Daniel et al. 6866, Gentry 7111, 
Van Devender et al. 93-1494. 

Aphelandra R. Br. 
A. aurantiaca (Scheidw.) Lindl.: Breedlove 

71287, Shapiro 262. 
A. gigant (flora Lindau: Breedlove 70900, Stan- 

dley 19771. 
A. guerrerensis Wassh.: Daniel 5376, Hinton 

1 1 199, Reko 6051. 
A. heydeana Donn. Sm.: Breedlove 65803. 
A. hintonii Wassh.: Hinton 16049. 
A. lineariloba Leonard: Daniel s.n.cv. 
A. madrensis Lindau: Daniel 5276, Langlasse 

806, Lott 865. 
A. scabra (Vahl) Sm.: Daniel 5328. 
A. schiedeana Schltdl. & Cham.: Ventura A. 

12241. 
A. speciosa Brandegee: Croat 40837. 
A. verticillataNees ex Hemsl.: Daniel et al. 3295, 

Hinton 13486, Moore 5503. 
A. wendtii T. F. Daniel: Breedlove 22121, Daniel 

8cWendt5813. 
A. sp.: A. Delgado S. 635. 



Barleria L. 
B. oenotheroides Dum. Cours.: Breedlove & 
Daniel 71 185, Daniel 5327, Daniel etal. 8412. 

Blechum P. Browne 
B. grandiflorum Oerst.: Breedlove 50544. 
B. pyramidatum (Lam.) Urb.: Daniel 2057, 
Daniel et al. 5454*. 

Bravaisia DC. 

B. berlandieriana (Nees) T. F. Daniel: Cabrera 
& Ibara 1149. 

B. grandiflora Donn. Sm.: Daniel & Bartho- 
lomew 4998. 

B. integerrima (Spreng.) Standi.: Daniel 2144. 

Carlowrightia A. Gray 

C. albiflora T. F. Daniel: Daniel 276. 

C. arizonica A. Gray: Daniel 1 73*, Daniel 203*, 
Daniel 983, Daniel 1157, Daniel et al. 6845. 

C.fuertensis T. F. Daniel: Rose et al. 13585. 

C. hapalocarpa B. L. Rob. & Greenm.: Daniel 
854. 

C. henricksoniiT . F. Daniel: Johnston & Crutch- 
field 5357. 

C. hintonii T. F. Daniel: Hinton 11326. 

C. huicholiana T. F. Daniel: Bauml & Voss 1896. 

C. lesueurii Henr. & T. F. Daniel: LeSueur 1254. 

C. linearifolia (Torr.) A. Gray: Daniel 565. 

C. mcvaughii T. F. Daniel: Feddema 173. 

C. mexicana Henr. & T. F. Daniel: Henrickson 
& Lee 16013. 

C. myriantha (Standi.) Standi.: Gaumer 1102. 

C. neesiana (Schauer ex Nees) T. F. Daniel: 
Daniel 417. 

C. ovata A. Gray: Pringle 695. 

C. parviflora (Buckley) Wassh.: Daniel 899. 

C. parvifolia Brandegee: Daniel 547. 

C. pectinata Brandegee: Daniel 936, Daniel 
6846. 

C. pringlei B.L. Rob. & Greenm.: Daniel 1291. 

C. purpurea T. F. Daniel: Daniel 351. 

C. serpyllifolia A. Gray: Daniel 221, Daniel 653. 

C. texana Henr. 8c T. F. Daniel: Daniel 108*. 

C. torreyana Wassh.: Daniel 97*. 

C. trichocarpa T. F. Daniel: Daniel 260. 

Chalarothyrsus Lindau 
C. amplexicaulis Lindau: Daniel & Bartholomew 
4842. 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



239 



Chileranthemum Oerst. 
C. lottae T. F. Daniel: Paray 2857, Paray 2858. 
C. pyramidatum (Lindau) T. F. Daniel: Breed- 
love 27655, Torres C. 121. 

C. trifidum Oerst.: Rosas R. 1020. 

Dicliptera Juss. 

D. acuminata (Ruiz & Pav.) Juss.: Ventura A. 
15758. 

D. anomala Leonard: Hernandez G. 840. 

D. haenkeana Nees: Daniel 5290. 

D. inequalis Greenm.: Daniel & Bartholomew 

4860. 
D. inutilis Leonard: Daniel & Bartholomew 

4877. 
D. nervata Greenm.: Daniel 4009. 
D. peduncularis Nees: Daniel 1262. 
D. resupinata (Vahl) Juss.: Daniel et al. 1947, 

Daniel et al. 2489, Daniel et al. 6865, Ross et 

al. 2061. 
D. sciadephora Donn. Sm.: Daniel & 

Bartholomew 5026. 
D. sexangularis (L.) Juss.: Palmer 93. 
D. sumichrasti Lindau: Tenorio L. et al. 5834. 
D. thlaspioides Nees: Breedlove 35983, Daniel 

1292. 
D. unguiculata Nees: Camp 2430. 

Dyschoriste Nees 

D. angustifolia (Hemsl.) Kuntze: Daniel 5310. 

D. decumbens (A. Gray) Kuntze: Jones 24437. 

D. capitata (Oerst.) Kuntze: Breedlove 26151. 

D. hirsutissima (Nees) Kuntze: Daniel & Butter- 
wick 3260. 

D. mcvaughii T. F. Daniel: Daniel & 
Bartholomew 4828, McVaugh 12187. 

D. microphylla (Cav.) Kuntze: Ventura A. 5714. 

D. novogaliciana T. F. Daniel: Daniel 2051, 
Daniel & Bartholomew 4779. 

D. ovata (Cav.) Kuntze: Breedlove & Strother 
46563. 

D. pringlei Greenm.: Villa C. & Chavez L. 755. 

D. purpusii Kobuski: Breedlove 69496. 

D. quadrangularis (Oerst.) Kuntze: Breedlove & 
Anderson 63705. 

D. saltuensis Fernald: Daniel & Bartholomew 



D. schiedeana (Nees) Kuntze: Orcutt 1654. 
D. xylopoda Kobuski: Wilbur & Wilbur 2137. 
D. sp.: Daniel & Bartholomew 4860.5. 



Ely tr aria Michx. 
E. bromoides Oerst. : Daniel 303, Daniel & Baker 

3683. 
E. imbricata (Vahl) Pers.: Daniel & Butterwick 

32 59gh, Porter 297. 
E. macrophylla Leonard: Carranza 2397, 

Dressier 2619. 
E. mexicana Fryxell & S. D. Koch: Daniel & 

Butterwick 3263gh. 

Gypsacanthus E. J. Lott, V. Jaram. & Rzed. 
G. nelsonii E. J. Lott, V. Jaram. & Rzed.: Trejo 
V 390. 

Henrya Nees ex Benth. 
H. insularis Nees ex Benth.: Daniel 2 116, Daniel 

& Bartholomew 473 1 , Daniel & Bartholomew 

4822, Sanders et al. 261 3gh. 
H. tuberculosperma T. F. Daniel: Daniel & 

Bartholomew 4868. 

Holographis Nees 
H. anisophylla T. F. Daniel: Lott 2359. 
H. argyrea (Lundell) T. F. Daniel: Daniel 1242. 
H. cuicatlanensis Salinas T.: Salinas T. 5274, 

Salinas T 6104, Salinas T. et al. 4779. 
H. ehrenbergiana Nees: Gonzalez 3581. 
H. ilicifolia Brandegee: Daniel 659. 
H. leticiana T. F. Daniel: Torres C. 637. 
H medusae T. F. Daniel: Daniel & Bartholomew 

4918. 
H. pallida Leonard & Gentry: Gentry 7022. 
H.parayana Miranda: Daniel 6203. 
H. peloria (Leonard) T. F. Daniel: Tellez V. & 

Miller 10628. 
H. pueblensis T. F. Daniel : Daniel & Baker 3771, 

Purpus 3346. 
H. tamaulipicaT. F. Daniel: Woodruff et al. 166. 
H. tolantongensis T. F. Daniel: Hernandez M. & 

Tenorio L. 6948. 
H virgata (Harv. ex Benth. & Hook.) T. F. 

Daniel: Breedlove 60981, Carter & Ferris 

3819, Carter & Sharsmith 4215, Carter et al. 

5724, Moran 3928. 
H websteri T. F. Daniel: Webster & Lynch 

17724. 

Hoverdenia Nees 
H speciosa Nees: Daniel & Baker 3739, Rze- 
dowski 36934. 



240 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



Hygrophila R. Br. 
H. costata Nees: Daniel & Butterwick 5884*, 
Daniel et al. 4316*, Daniel et al. 8163cv*, 
Wendt3109. 

Hypoestes Sol. ex R. Br. 
H. phyllostachya Baker. Breedlove & Daniel 
70764, Daniel s.n*. 

Justicia L. 

(including taxa for which nomenclatural com- 
binations have not been published) 
J. adenothyrsa (Lindau) T. F. Daniel: Daniel & 

Bartholomew 4981. 
J. alopecuroidea T. F. Daniel: Daniel & 

Bartholomew 4980. 
J. americana (L.) Vahl: Lieb & Roessling 645*. 
J. anagallis (Nees) Lindau: Ventura A. 7071. 
J. angustiflora D. N. Gibson: Reyes 1255. 
J. aurea Schltdl.: Breedlove 29496. 
J. austrocapensis T. F. Daniel: Moran 7373. 
J. bartlettii (Leonard) D. N. Gibson: Breedlove 

&Almeda 57832. 
J. borrerae (Hemsl.) T. F. Daniel: Breedlove 

35290. 
J. brandegeana Wassh. & L. B. Sm.: Daniel 291. 
J. breedlovei T. F. Daniel: Breedlove 56242. 
J. breviflora (Nees) Rusby: Breedlove 50689, 

Breedlove 68976, Daniel & Bartholomew 

5010, Lott & Magallanes 1860. 
J. californica (Benth.) D. N. Gibson: Daniel et 

al. 1542. 
J. campechiana Standi, ex Lundell: Cabrera C. 

9938, Miranda 5860, Miranda 6839. 
J. canbyi Greenm.: Garcia M. & Ramos 2909. 
J. candelariae (Oerst.) Leonard: Breedlove 

67008, Skutch 2009*. 
J. candicans (Nees) L. D. Benson: Daniel 8355, 

Moran 18787. 
J. carthagenensis Jacq.: Breedlove 7718, Breed- 
love 34000. 
J. caudata A. Gray: Breedlove 8c Daniel 70945. 
J. chol T. F. Daniel: Cowan & Magana 3138. 
J. clinopodium A. Gray ex Greenm.: Daniel etal. 

5873. 
J. colorifera V. A. W. Graham: Breedlove 

50321. 
J. comata (L.) Lam.: Breedlove 38648. 
J. decurvata Hilsenb.: Torrecillas 217. 
J. eburnea D. N. Gibson: Tun O. 1134. 



J.fimbriata (Nees) V. A. W. Graham: Breedlove 

49679, Daniel & Bartholomew 4992. 
J.fulvicoma Schltdl. & Cham.: Daniel & Baker 

3637. 
J. gonzalezii Henr. & Hiriart: Daniel & Baker 

3779. 
J. herpetacanthoides Leonard: Breedlove & 

Daniel 71004. 
J. hians (Brandegee) Brandegee: Carter et al. 

4164. 
J. inaequalis Benth.: Matuda 3990. 
J. insolita Brandegee: Carter 5711, Leon de la 

Luz 2334. 
J. ixtlania T. F. Daniel: Daniel 2070. 
J. jitotolana T. F. Daniel: Thome & Lathrop 

41662, Zwill 631. 
J. kanal T. F. Daniel: Breedlove & Bourell 

68262. 
J. linearis B.L. Rob. & Greenm.: Daniel & Baker 

3719. 
J. lindeniana (Nees) J. F. Macbr.: Breedlove 

49908. 
J. longii Hilsenb.: Butterwick & Hilly >ard 7384*, 

Kearney & Peebles 14969*. 
J. macrantha Benth.: MacDougall s.n.. 
J. madrensis T. F. Daniel: Breedlove 38656. 
J. masiaca T. F. Daniel: Daniel et al. 2546, Van 

Devender & Sanders 92-1058, Van Devender 

etal. 93-827, White 3590. 
J. medranoi Henr. & Hiriart: Daniel & Baker 

3742. 
J. mirandae T. F. Daniel: Breedlove 50163. 
J. multicaulis Donn. Sm.: Breedlove 49946. 
J. nevlingii Wassh. & T. F. Daniel: Dor antes 

2799, Marcks & Marcks 913, Vazquez et al. 

411. 
J. palmeri Rose: Carter 5442. 
J. pectoralis Jacq.: Breedlove 48559. 
J. phlebodes Leonard & Gentry: Van Devender 

93-483. 
J. pilosella (Nees) Hilsenb.: Breedlove 69439. 
J. pringlei B. L. Rob.: Breedlove & Strother 

46783. 
J. purpusii (Brandegee) D. N. Gibson: Moran 

6941. 
J. ramosa (Oerst.) V. A. W. Graham: Daniel & 

Baker 3778. 
J. rzedowskii (Acosta) T. F. Daniel: Ventura & 

Lopez 1074. 
J. salviiflora Kunth: Breedlove & Raven 20133. 
J. santelisiana Acosta & T. F. Daniel: Acosta C 

1276, Morton & Makrinius 2364. 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



241 



J. soliana Standi.: Laughlin 2565. 

J. spicigera Schltdl.: Avila B. 28, Breedlove 

9431. 
J. tabascina T. F. Daniel: Cowan 2860. 
J. teletheca T. F. Daniel: Breedlove 56314. 
J. tianguensis T. F. Daniel: Breedlove 7365. 
J. torresii T. F. Daniel: Torres C. 11472. 
J. turipachensis T. F. Daniel: Breedlove 31242. 
J. valvata T. F. Daniel: Contreras 9311, Doran- 

tes et al. 3538, Vasquez T. et al. 2582. 
J. warnockii B. L. Turner: Rowell 5131. 
J. zopilotensis Henr. & Hiriart: Daniel 1192. 
J. sp.: Breedlove 28247. 
J. sp.: Cowan & Magaha 3138. 
J. sp.: Martinez R. 403. 
J. sp.: Lott & Bullock 21 71. 
J. sp.: Lott & Magallanes 443. 
J. sp.: Lott et al. 3733. 
J. sp.: Martinez R. 775. 
J. sp.: Purpus 5850. 
J. sp.: Salinas T. et al. 6624-a. 
J. sp.: Torres B. 536. 
J. sp.: Van Devender 94-54. 
J. sp.: Wendt & Hernandez 5727. 
J. sp.: Wendt et al. 2994. 
Rhytiglossa latifolia Nees: Cowan 1715. 
Siphonoglossa mexicana Hilsenb.: Lott & 

Guadalupe A. 2656. 

Lepidagathis Willd. 
L. alopecuroidea (Vahl) R. Br. ex Griseb.: 
Daniel et al. 5461 cv*, Martinez S. 18315. 

Lophostachys Pohl 
L. chiapensis Acosta: Breedlove 30873, Breed- 
love 66152. 
L. soconuscana T. F. Daniel: Boege 1086. 
L. uxpanapensis Acosta: Hernandez G. 642. 

Louteridium S. Watson 
L. brevicalyx A. T. Richardson: Hinton 15843. 
L. donnell-smithii S. Watson: Breedlove 57578. 
L. mexicanum (Baill.) Standi.: Breedlove 30786, 

Breedlove & Daniel 70879gh. 
L. parayi Miranda: Breedlove 70889. 
L. purpusii Brandegee: Breedlove 31613. 
L. rzedowskii T. F. Daniel: Kruse 1380. 
L. tamaulipense A. T. Richardson: Hutchinson 

s.n. 



Mendoncia Veil, ex Vand. 
M. guatemalensis Standi. & Steyer M.: Hernan- 
dez G. 1140. 
M. retusa Turrill: Daniel s.n.*, Matuda 16603. 

Mexacanthus T. F. Daniel 
M. mcvaughii T. F. Daniel: Daniel 2107, 
McVaugh 23016. 

Mirandea Rzed. 
M. andradenia T. F. Daniel: Daniel & Baker 

3693. 
M. grisea Rzed.: Daniel 331, Daniel 852. 
M. huastecensis: Daniel 252, Daniel & Baker 

3616. 
M. sylvatica Acosta: Wendt et al. 4104. 

Nelsonia R. Br. 
N. canescens (Lam.) Spreng.: Daniel et al. 
5452*, Daniel et al. 5452cv*. 

Odontonema Nees 
O. albiflorum Leonard: Tellez & Villasenor 

6634. 
O. auriculatum (Rose) T. F. Daniel: Daniel & 

Bartholomew 4913. 
O. callistachyum (Schltdl. & Cham.) Kuntze: 

Daniel & Bartholomew 5014. 
O. cuspidatum (Nees) Kuntze: Breedlove & 

McClintock 23784, Dwyer 2076. 
O. glaberrimum (M. E. Jones) V. M. Baum: 

Breedlove 49628. 
O. glabrum Brandegee: Breedlove & Bourell 

67391. 
O. mortonii V. M. Baum: Alexander 444. 
O. tubaeforme (Bertol.) Kuntze: Daniel & 

Bartholomew 4991, Hoover 137. 

Poikilacanthus Lindau 

P. capitatus (Leonard) Ramamoorthy: Hinton 
8075. 

P. macranthus Lindau: Breedlove 26283, Breed- 
love 39887. 

P. novogalicianus T. F. Daniel: McVaugh 15759. 

Pseuderanthemum Radlk. 
P. alatum (Nees) Radlk.: Daniel & Baker 3713. 



242 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 



P. cuspidatum (Nees) Radlk.: Breedlove & 

Daniel 70880. 
P.fasciculatum (Oerst.) Leonard: Ton 903. 
P.floribundum T. F. Daniel: Daniel 5381. 
P. hispidulum (Nees) Radlk.: Daniel 6205. 
P. pihuamoense T. F. Daniel: Daniel et al. 6283. 
P. praecox (Benth.) Leonard: Daniel 2131, 

McVaugh 26541. 
P. standleyi Leonard: McVaugh 23445, Maya J. 

862. 
P. verapazense Donn. Sm.: Wendt et al. 3113. 
P. sp.: Daniel & Ton 6171. 
P. sp.: Camp 2219. 
P. sp.: Clarke 356. 
P. sp.: Daniel et al. 5875gh. 

Ruellia L. 
R. amoena Sesse & Moc.: Daniel & Butterwick 

3287. 
R. bourgaei Hemsl.: Rzedowski 23294. 
R. breedlovei T. F. Daniel: Daniel & Bar- 
tholomew 5025. 
R. californica (Rose) I. M. Johnst: Daniel et al. 

6844, Wiggins 11411. 
R. cedilloiRamamoorthy: Cedillo T. & Torres C 

1425. 
R. cordata Brandegee: Boyd & Ross 5934. 
R. eumorphantha Lindau: Daniel & Bar- 
tholomew 4927. 
R.foetida Willd.: Daniel 2081 gh. 
R. fruticosa Sesse & Moc.: Daniel 8c Jessup 

5776. 
R. geminiflora Kunth: Breedlove 9163. 
R. guerrerensis T. F. Daniel: Hinton 11296. 
R. harveyana Stapf: Hernandez G. 447. 
R. hirsutoglandulosa (Oerst.) Hemsl.: Daniel & 

Baker 3736. 
R. hookeriana (Nees) Hemsl.: Breedlove & 

Daniel 71060. 
R. intermedia Leonard: Jenkins et al. s.n.. 
R. inundata Kunth: Daniel & Butterwick 6952, 

Daniel et al. 3315. 
R. jaliscana Standi.: Daniel & Bartholomew 

4855. 
R. jussieuoides Schltdl. & Cham.: Breedlove & 

Almeda 57192. 
R. lactea Cav.: Tenorio L. & Romero de T. 6720. 
R. leucantha Brandegee: Daniel & Butterwick 

6940, Brandegee s.n.. 
R. matagalpae Lindau: Breedlove & Daniel 

70899. 



R. matudae Leonard: Breedlove & Almeda 

57020. 
R. maya T. F. Daniel: Daniel & Bartholomew 

5000. 
R. mcvaughii T. F. Daniel: Cowan 4746. 
R. megasphaera Lindau: Breedlove 56280. 
R. novogaliciana T. F. Daniel: Hinton 12931. 
R. nudijlora (Engelm. 8c A. Gray) Urb.: Wiggins 

& Rollins 136. 
R. paniculata L.: Breedlove & McClintock 

23447. 
R. parryi A. Gray: Daniel 247. 
R. pereducta Standi, ex Lundell: Davidse et al. 

20405. 
R. petiolaris (Nees) T. F. Daniel: Daniel & 

Bartholomew 4930. 
R. pringlei Fernald: Torres C. et al. 226. 
R. puberula Leonard: Salinas T. et al. 6898. 
R. speciosa (Nees) Lindau: Pringle 13155. 
R. stemonacanthoides (Oerst.) Hemsl.: Daniel & 

Bartholomew 4871. 

Sanchezia Ruiz & Pav. 
S. parvibracteata Sprague & Hutch.: Breedlove 

& Daniel 71315. 
S. speciosa Leonard: Rosas R. 1261. 

Schaueria Nees 
S. parviflora (Leonard) T. F. Daniel: Contreras 
11180, Herrera 72, Trigos 198. 

Spathacanthus Baill. 

S. hahnianus Baill.: Breedlove & Smith 21618, 
Hernandez G. 1777, Hernandez G. & Gon- 
zalez L. 1855, Wendt et al. 2769. 

S. parvijlorus Leonard: Matuda 1696, Ramirez- 
Marcial & Quintana-Ascencio 507, Skutch 
961*, Steyermark 46632*. 

Staurogyne Wallich 
S. agrestis Leonard: Lott et al. 3226, Lott et al. 
3396. 

Stenandrium Nees 
S. barbatum Torr. & A. Gray: Johnston et al. 

10589.5. 
S. chameranthemoideum Oerst.: Breedlove 

67020. 
S. dulce (Cav.) Nees: Angeles 2, Breedlove & 

Almeda 45287, Chiang et al. 8198, Hernandez 

M. 5995, Hinton 6086, Padilla 8, Standley 71*. 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



243 



S. manchonense T. F. Daniel: Hinton 10460. 
S. nanum (Standi.) T. F. Daniel: Cabrera 1470. 
S. pedunculatum (Donn. Sm.) Leonard: Bullock 

1366. 
S.pilosulum (Blake) T. F. Daniel: Moran 21965. 
S. subcordatum Standi.: Chater et al. 3. 
S. verticillatum Brandegee: Purpus 3939. 

Stenostephanus Nees 

(including taxa for which nomenclatural com- 
binations have not been published) 
S. breedlovei T. F. Daniel: Breedlove 49644. 
S. chiapensis T. F. Daniel: Breedlove 34374. 
S. glaber (Leonard) T. F. Daniel: Cloud 4. 
S. gracilis (Oerst.) T. F. Daniel: Breedlove 

71524. 
S. latilabris (D. N. Gibson) T. F. Daniel: Matuda 

2485. 
S. monolophus (Donn. Sm.) T. F. Daniel: Breed- 
love & Smith 22692. 
S. purpusii (Brandegee) T. F. Daniel: Purpus 

6824. 
S. silvaticus (Nees) T. F. Daniel: Zuill 772. 
S. tacanensis (Acosta & R. Fernandez) T. F. 

Daniel: Breedlove & Almeda 47714. 
S. sp.:Reko3724. 
S. sp.: MacDougall s.n. 
S. sp.: Hinton 10758. 
S. sp.: Breedlove 72407. 
S. sp.: Campos V 4736. 
S. sp.: Breedlove 36060. 
S. sp.: Breedlove 61946. 
Habracanthus haematodes Nees: Ventura A. 

20491. 
Habracanthus harleyi Wassh.: Reveal et al. 

4239. 



T. butterwickianum T. F. Daniel: Daniel & But- 

terwick 3267. 
T. crenatum T. F. Daniel: Morton & Makrinius 

2680. 
T. diffusum Rose: Daniel 1141, Daniel 2097. 
T. emilyanum T. F. Daniel: Daniel & Butterwick 

3258. 
T. fruticosum Brandegee: Daniel & Butterwick 

6869, Daniel et al. 2490gh. 
T. glandulosum Oerst.: Daniel 2104, Daniel 

2104gh. 
T. glutinosum Lindau: Daniel et al. 3342. 
T. guerrerense T. F. Daniel: Hinton 11000. 
T. langlassei Happ: Daniel et al. 3310gh. 
T. mcvaughii T. F. Daniel: Daniel & Butterwick 

3247. 
T. nemorum Brandegee: Ventura A. 12283. 
T. nervosum Nees: Carter & Moran 5324, Daniel 

11 7*, Daniel 3443. 
T. oaxacanum T. F. Daniel: Fryxell & Lott 3390. 
T. obovatum T. F. Daniel: MacDougall s.n. 
T. ochoterenae (Miranda) T. F. Daniel: Gonzalez 

Q. 3631. 
T. rubrum Happ: Daniel & Butterwick 3270, 

Daniel et al. 3301. 
T. rzedowskii T. F. Daniel: Daniel & Butterwick 

3254 
T. tenuissimum Rose: Daniel 3384. 
T. yaquianum T. F. Daniel: Daniel 3372. 

Thunbergia Retz. 
T. alata Bojer ex Sims: Daniel s.n*. 
T. erecta (Benth.) T. Anders.: Ton 6367. 
T. fragrans Roxb.: Daniel & Butterwick 

6641gh*. 
T. grandijlora Roxb.: Breedlove 20052. 



Tetramerium Nees 
T. abditum (Brandegee) T. F. Daniel: Daniel 
3 364, Gentry 1178. 



Yeatesia Small 
Y. mabryi Hilsenb.: Daniel & Baker 3698. 
Y. platystegia (Torr.) Hilsenb.: McCart 7445, 
Peterson 1313. 



244 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 

Figure Legends 

FIGURE 1 . Pollen of Thunbergioideae, Mendoncioideae, and Nelsonioideae. a. Thunbergiafragrans 
(Daniel & Butterwick 6641 gh), equatorial view; b. T alata (Daniel s.n.), equatorial view; c. T. alata 
(Daniel s.n.), polar view; d. T. erecta (Ton 6367), equatorial view; e. Mendoncia retusa (Matuda 
16603), equatorial view; f. M. giiatemalensis (Hernandez G. 1140), polar view; g. Elytraria mexicana 
(Daniel 8i Butterwick 3263 gh), polar view; h. E. imbricata (Porter 297), intercolpal view; i. Nelsonia 
canescens (Daniel et al. 5452), colpal view; j. Staurogyne agrestis (Lott 3226), colpal view. Scale for 
a-d = 43 urn; for e, f = 36 urn; for g, h = 29 urn; for i = 22 urn; for j = 14 urn. 

FIGURE 2. Pollen of Aphelandreae. a. Aphelandra gigantiflora (Breedlove 70900), colpal view; b. 
Holographis tamaulipica (Woodruff et al. 166), polar view; c. A. schiedeana (Ventura A. 12241), 
intercolpal view; d. H. virgata (Carter & Ferris 3819), intercolpal view; e. Stenandrium dulce 
(Breedlove & Almeda 45287), intercolpal view; f. S. verticillatum (Purpus 3939), colpal view; g. S. 
dulce (Hinton 6086), interapertural view; h. S. barbatum (Johnston et al. 10589.5), interapertural 
view; i. S. dulce (Standley 71). Scale for a, d-i = 22 urn; for b = 14 (im; for c = 29 urn. 

FIGURE 3. Pollen of Louteridieae and Trichanthereae. a. Louteridium mexicanum (Breedlove 
30786); b. Bravaisia integerrima (Daniel 2144), colpal view; c. B. grandiflora (Daniel & 
Bartholomew 4998), intercolpal view; d. Sanchezia parvibracteata (Breedlove & Daniel 71315), 
colpal view; e. S. speciosa (Rosas R. 1261), intercolpal view. Scale for a = 72 urn; for b = 29 urn; for 
c-e = 43 urn 

FIGURE 4. Pollen of Petalidiinae and Hygrophilinae. a. Dyschoriste pringlei (Villa C. & Chavez L. 
755), intercolpal view; b. D. pringlei (Villa C. & Chavez L. 755), polar view; c. D. decumbens (Jones 
24437), colpal view; d. Hygrophila costata (Wendt 3109), polar view; e. H. costata (Daniel et al. 
8163cv), colpal view. Scale for a-c = 29 um; for d, e = 22 urn. 

FIGURE 5. Pollen of Ruelliinae, Barleriinae, Blechinae, and Lepidagathideae. a. Ruellia inundata 
(Daniel & Butterwick 6952), apertural view; b. R. pringlei (Torres C. etal. 226), interapertural view; 
c. R. intermedia (Jenkins et al. s.n.), probable polar view; d. Lophostachys uxpanapensis (Hernandez 
G. 642), intercolpal view; e. Barleria oenotheroides (Daniel et al. 8412), apertural view; f. B. 
oenotheroides (Daniel 5327), polar view; g. L. uxpanapensis (Hernandez G. 642), colpal view; h. 
Blechum pyramidatum (Daniel et al. 5454), polar view; i. Lepidagathis alopecuroides (Martinez S. 
18315), colpal view. Scale for a-c = 43 um; d, g = 36 um; e, f = 62 um; h = 29 um; i = 1 1 urn 

FIGURE 6. Pollen of Isoglossinae and Poikilacanthus. a. Stenostephanus gracilis (Breedlove 71524), 
apertural view; b. S. gracilis (Breedlove 71524), interapertural view; c. S. chiapensis (Breedlove 
34374), apertural view; d. S. chiapensis (Breedlove 34373), interapertural view; e. Poikilacanthus 
novogalicianus (McVaugh 15759), oblique view; f. P.macranthus (Breedlove 39887), polar view. 
Scale for a-d = 29 |im; for e, f = 36 urn 

FIGURE 7. Pollen of Odontoneminae. a. Dicliptera resupinata (Daniel etal. 6865), intercolpal view; 

b. Tetramerium fruticosum (Daniel & Butterwick 6869), colpal view; c. Hoverdenia speciosa 
(Rzedowski 36934), colpal view. d. Aphanosperma sinaloensis (Van Devender et al. 93-1494), 
intercolpal view; e. A. sinaloensis (Daniel 4070), polar view; f. Pseuderanthemum floribundum 
(Daniel 5381), polar view; g. Gypsacanthus nelsonii (Trejo V. 390), colpal view; h. Schaueria 
parviflora (Trigos 198), colpal view; i. Chileranthemum pyramidatum (Breedlove 27655), colpal 
view; j. Yeatesia platystegia (Peterson 1313), intercolpal view. Scale for a, d, e = 22 urn; for b, g = 
14 um; for c, f = 36 um; for h-j = 29 um. 

FIGURE 8. Pollen of Odontoneminae. a. Chalarothyrsus amplexicaulis (Daniel & Bartholomew 
4842), intercolpal view; b. Henrya tuberculosperma (Daniel & Bartholomew 4868), intercolpal view; 

c. Carlowrightia arizonica (Daniel et al. 6845), colpal view; d. Mexacanthus mcvaughii (McVaugh 
23016), intercolpal view. e. Aphanosperma sinaloensis (Daniel et al. 6866), colpal and subpolar 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 245 



views; f. Anis acanthus pumilis (Lott & Magallanes 3 1 3), polar view; g. Spathacanthus hoffinannii 
(Skutch 325), polar view; h. Mirandea grisea (Daniel 852), polar view; i. Odontonema cuspidatum 
(Breedlove & McClintock 23784), polar view. Scale for a, i =29 um; for b = 23 urn; for c, e, f, h = 22 
urn; for d, g = 36 um. 

FIGURE 9. Pollen of Justiciinae. a. Justicia tabascina (Cowan 2860), apertural view; b. J. medranoi 
(Daniel & Baker 3742), apertural view; c. J. breedlovei (Breedlove 56242), interapertural view; d. J. 
ixtlania (Daniel 2070), apertural view; e. J. fulvicoma (Daniel & Baker 3637), apertural view; f. J. 
zopilotensis (Daniel 1192), polar view; g. J. angustiflora (Reyes 1255), apertural view; h. J. 
angustiflora (Reyes 1255), polar view; i. J. fulvicoma (Daniel & Baker 3637), polar view. Scale for 
a, b, d, e, g-i = 29 urn; for c = 22 urn; for f = 32 urn. 

FIGURE 10. Pollen of Justiciinae. a. Justicia masiaca (White 3590), apertural view, b. J. candicans 
(Moran 18787), apertural view; c. J.jitotolana (Thome & Lathrop 41662), interapertural view; d. J. 
aurea (Breedlove 29496), apertural view; e. J. soliana (Laughlin 2565), interapertural view; f. J. 
warnockii (Rowell 5131), apertural view; g. J. warnockii (Rowell 5131), polar view; h. J. clinopodium 
(Daniel et al. 5873), apertural view; i. J. breviflora (Breedlove 50689), apertural view. Scale for a, b, 
e, h = 29 urn; for c = 36 urn; for d = 43 (im; for f, g =14 urn; for i = 22 um. 

FIGURE 11. Exine surfaces, a. Mendoncia retusa (Matuda 16603); b. Elytraria bromoides (Daniel 
303); c. Nelsonia canescens (Daniel et al. 5452); d. Louteridium purpusii (Breedlove 31613); e. 
Dyschoriste capitata (Breedlove 26151); f. Hygrophila costata (Daniel et al. 8163); g. Blechum 
pyramidatum (Daniel et al. 5454); h. Ruellia novogaliciana (Hinton 12931); i. Hypoestes phyl- 
lostachya (Breedlove & Daniel 70764). Scale for a, c, i = 2.1 um; for b = 2.7 um; for d = 10.7 um; 
for e-g = 3.1 um; for h = 5.4 um. 



246 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 




FIGURE 1 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



247 




FIGURE 2 



248 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 




DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



249 




FIGURE 4 



250 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 




FIGURE 5 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



251 




252 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 




FIGURE 7 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



253 




FIGURE 8 



254 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 




FIGURE 9 



DANIEL: POLLEN MORPHOLOGY OF ACANTHACEAE 



255 




FIGURE 10 



256 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 8 










FIGURE 11 



CALIFORNIA ACADEMY OF SCIENCES, 1998 
Golden Gate Park 
San Francisco, California 941 18 



JUN 22 



PROCEEDINGS 
CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 9, pp. 257-265, 9 figs., 1 table. 



June 19, 1998 



A NEW SPECIES OF ACANTHODACTYLUS (SAURIA: 

LACERTIDAE) FROM QASR-E-SHIRIN, KERMANSHAH 

PROVINCE, WESTERN IRAN 



By 

Nasrullah Rastegar-Pouyani 

Department of Zoology, Gothenburg University, Box 463, 
SE 405 30 Gothenburg, Sweden 



A new species of Acanthodactylus is described based on material collected in 1995 from the 
lowlands south of Qasr-e-Shirin, near the Iranian-Iraqi border, Kermanshah province, western 
Iran, at an elevation of about 285 m. Its larger-size, 8 rows of ventral plates, 48-52 relatively large, 
imbricate, keeled scales across dorsum, and distinctive color pattern distinguish it from other 
species of Acanthodactylus. The new species, Acanthodactylus nilsoni, is related to the "/I. 
boskianus-schreiberi" group; it appears to be restricted in distribution to the low lands of western 
Kermanshah and probably eastern Iraq. 

Received November 26, 1997. Accepted February 3, 1998. 



The lacertid genus Acanthodactylus Wieg- 
man. 1834 consists of about 31 species distrib- 
uted from Spain and Portugal in southwestern 
Europe, in North Africa across the Sahara desert 
and its periphery east to the Red Sea. throughout 
most of Arabian Peninsula, north to Cyprus and 
the Syrian-Turkish border; and eastward through 
Iraq, west, south, and east Iran, southern Af- 
ghanistan, to Pakistan and northwest India. All 
species are relatively small, ranging in size from 
52-105 mm snout to vent. They are diurnal and 
mostly ground-dwellers ordinarily found in rela- 
tively flat, open sandy, dry habitats, which usu- 
ally have at least some vegetation, ranging from 
open woodland to the borders of quite severe 
deserts. All species appear to be active hunters 
and feed largely on small invertebrates. All are 
oviparous; clutch size varies from two to about 
seven eggs. The majority of forms seem to ma- 
ture within ayear of hatching although some, like 
A. ety thrums in Spain and Portugal, may take 
two seasons (Arnold 1983). 



Six species of Acanthodactylus, the new spe- 
cies included, occur in Iran, primarily in southern 
and eastern parts of the country. Acanthodactylus 
opheodurus Arnold, may also occur in lowland 
southwestern Iran (Anderson, in press). 

The new species comes from extreme western 
Kermanshah Province, which to the north and 
northeast is dominated by the Zagros Mountains 
and to the west and southwest by lowlands with 
elevations of 200-700 m. 

Species Description 

Acanthodactylus nilsoni sp. nov. 
(Figs. 1-6) 

HOLOTYPE AND TYPE LOCALITY. — Adult 
male. GNHM Re. ex. 5145, collected by the 
author on 9 July, 1995, in the lowland regions, 5 
km south of Qasr-e-Shirin (34°30'N, 45°33'E), 
about 7 km to the Iranian-Iraqi border, at 285 m 
elevation, Kermanshah Province, western Iran. 



[257] 



258 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 9 



PARATYPES. — Adult male, GNHM Re. ex. 
5 146, other information as for the holotype. 

DIAGNOSIS. — A medium-sized lacertid (in 
comparison with the other species of Acantho- 
dactylus), maximum SVL = 73.9 mm, TL = 165 
mm; 8 longitudinal, and 26 transverse rows of 
ventral plates; 48-52 rather large, imbricate, 
keeled scales across dorsum; toes covered with 
three rows of scales; 9/7 supralabials, four of 
them anterior to subocular, the latter does not 
border the mouth; temporals mainly keeled and 
more than 100 on each side; gular fold distinct; 
26-27 gular scales; 10-12 collars; 23-24 lamel- 
lae under the fourth toe which is moderately 
denticulated laterally; eyelids with weak pectina- 
tion. 

Description of the Holotype (preserved in 
70% ethyl alcohol, condition good). — Body 
slightly depressed; head length 1.57 times its 
width and 28% of snout-vent length; snout 
pointed but not much narrow, perinarial regions 
rather distinctly swollen; canthus strong and 
loreal region slightly concave; a distinct median 
depression running from frontonasal to anterior 
frontal; limbs not slender, hind leg about 2.29 
times head length; tail length about 2.23 times 
snout-vent length, somewhat dorsoventrally flat- 
tened proximally, becoming laterally com- 
pressed distally; rostral rather prominent; paired 
internasals meet behind rostral, their common 
suture about 1/4 length of frontonasal, which is 
hexagonal and broader than long, its greatest 
width being almost as long as the internarial 
distance measured at the center of the nostrils; 
prefrontals longer than broad with a common 
suture of about 3/5 of their length; frontal dis- 
tinctly shorter than the distance from its anterior 
border to tip of snout, its maximum width about 
half of its length, much narrowed posteriorly 
where its width is about 1/5 of its length; fron- 
toparietal about half of the length of the frontal 
with an extensive common suture, each fron- 
toparietal in contact with frontal, third and fourth 
supraoculars, parietal, and interparietal; parietals 
longer than broad , their outer borders somewhat 



sinuous, their posterior margin convex; inter- 
parietal small, lozenge-like, surrounded by parie- 
tals and frontoparietals, with a clear parietal 
foramen; no occipital scale; four undivided su- 
praoculars on each side, first in contact with 
frontal and frontonasal and small, second and 
third supraoculars large, subequal, both in con- 
tact with frontal and frontonasal, fourth su- 
praocular the smallest and in contact with 
frontoparietal, parietal, first supratemporal, and 
third supraocular; 6/6 supraciliaries, first largest, 
keeled, and broadly in contact with first su- 
praocular; 9/11 granules between supraciliaries 
and last three supraoculars; nostril bordered pos- 
teriorly by a single postnasal, anteriorly and dor- 
sally by internasal, interiorly by first supralabial; 
rostral not in contact with nostril, separated from 
it by internasal and first supralabial; two loreals, 
second much larger than first and strongly 
keeled, both loreals longer than high; 9/7 su- 
pralabials, 4/4 anterior to subocular, fourth the 
largest and in broad contact with subocular; 
subocular does not reach mouth, it is much longer 
than high, strongly keeled, its lower edge in 
broad contact with fourth, fifth, and sixth su- 
pralabials; 7/7 infralabials, sixth the largest; 
mental roundish anteriorly and in contact with 
first infralabial and first pair of submaxillary 
shields, which are in five pairs, the first pair the 
smallest and in contact with mental and first and 
second infralabials, scales of first two pairs of 
submaxillary shields entirely in contact, third 
pair partly separated by a long gular scale, last 
two pairs broadly separated, the last three pairs 
subequal in size; eyelids slightly serrated, scales 
in center of lower eyelid enlarged and translu- 
cent; temporal region with about 104-108 scales 
on each side, anterior and upper scales distinctly 
keeled, inferior ones smooth and swollen and 
distinctly larger than upper ones; two keeled 
supratemporals, the first more than three times 
length of second and in contact with parietal and 
fourth supralabial; tympanic shield vertically 
elongate, keeled, and moderately developed; 
tympanum a vertical slit, almost as long as orbit. 



-» 



Figures 1-6. ( I ) Acanthodactylus nilsoni, holotype. A. Dorsal region, B. Ventral region with only 8 longitudinal rows of 
plates. (2) Acanthodactylus nilsoni, paratype. (3) Comparison of the holotype of Acanthodactylus nilsoni (left) with an adult 
male of A. boskianus (right). (4) Acanthodactylus nilsoni, temporal and labial regions. (5) Acanthodactylus nilsoni, posterior 
ventral regions and toes fringes. (6) Acanthodactylus nilsoni, upper head region with four pairs of intact supraoculars. 



RASTEGAR-POUYANI: ACANTHODACTYL US NILSONI 



259 




260 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 9 



moderately serrated anterior margin formed by 
5/6 scales; postocular higher than long and in 
contact with anterior supratemporal, fourth su- 
praocular, and last supraciliary; 26-27 gular 
scales in straight line from symphysis of chin 
shields to midline of collar, anterior ones small 
and almost juxtaposed, posteriorly becoming 
larger, more imbricate and broader; a clear gular 
fold; collar moderately serrated, attached in the 
region of midline, curved and formed of 10-12 
scales; nape with raised and granular scales with 
small keels, increasingly in size posteriorly and 
becoming flat and broad with very obtusely 
pointed posterior margins; mid-dorsal scales im- 
bricate and keeled; scales on flanks smaller, 
slightly keeled, although increasing in size to- 
wards the ventrals; 48-50 dorsal scales in a trans- 
verse row at midbody, of which about 23-25 in 
the middle of the series are large; 15-16 large 
dorsal scales in transverse row between hin- 
dlimbs; ventral scales strongly imbricate, in 
regular, straight longitudinal and transverse se- 
ries, almost all broader than long, 8 in longest 
longitudinal series across midbody, 26 transverse 
rows between the collar and hindlimbs; preanal 
region with two strongly enlarged plates which 
are broader than long, the more posterior more so 
than the other, both are surrounded by a semicir- 
cle of 7 enlarged scales which, in turn, are bor- 
dered by smaller ones; anterior and dorsal 
surfaces of upper forelegs (humerus) covered 
with large, imbricate plates that extend onto an- 
terior and dorsal surfaces of lower limb (forearm) 
but become smaller and keeled posteriorly, rest 
of forelimb covered with granular scales; fingers 
weakly denticulate, first four surrounded by three 
rows of scales, the fifth by four; anterior surface 
of thigh with row of very large, imbricate scales, 
bordered above and below by ones that are 
smaller and decrease in size posteriorly, and on 
dorsal surface where they are slightly keeled, 
grade into granules that extend onto posterior 
surface; 22/22 femoral pores, the two series meet 
at midline and extend laterally to reach the knee; 
upper surface of tibia with large, flat, keeled, 
pointed, imbricate scales that become smaller 
distal ly but are larger, more pointed and carinate 
laterally; ventral surface of tibia covered by lon- 
gitudinal rows of strongly enlarged, smooth, im- 
bricate plates; toes covered by three rows of 
scales, the posterior forming a denticulation 
stronger than on the fingers, and strongest on 



proximal half of fourth toe; 23/24 unicarinate. 
keeled lamellae beneath fourth toe; upper caudal 
scales much larger than posterior dorsals, ob- 
tusely pointed and mucronate, with strong keels 
that form straight lines along tail; ventral caudals 
smooth, 25-26 scales in fifth whorl behind vent. 
Coloration. Upper surface of head olive 
brown; temporal region and upper surface of 
forelegs light brown; upper surface of hindlimbs 
with many large whitish spots bordered by dark 
brown circles; dorsum greyish brown, with 6-8 
interrupted longitudinal dark stripes, the dorso- 
lateral ones having a tendency to form a reticula- 
tion; vertebral region without dark stripe; upper 
surface of tail light brownish grey; ventral sur- 
faces whitish. 

Measurements (in mm). Snout-vent length = 
73.4, tail length = 143, length of forelimb = 26, 
length of hindlimb = 48.5, head length = 21.1, 
head width = 13.4, head depth = 1 1, tip of snout 
to forelimb = 29.5. 

DESCRIPTION OF THE PARATYPE. — Adult 
male, similar to the holotype, differing as fol- 
lows: dorsal scales slightly keeled, 50-52 across 
widest part of dorsum from which 18-20 scales 
in midline of series are large; ventral plates in 29 
transverse rows from collar to hindlimbs; head 
length 1.53 times its width and 25% of snout-vent 
length; hindleg about 2.35 times head length; tail 
length about 1.63 times snout-vent length; 8/8 
supralabials; temporal scales slightly keeled su- 
periorly, raised and smooth inferiorly; postocular 
separated from last supraciliary by 2-3 granules; 
tympanum slightly serrated anteriorly, bordered 
by 4 / 4 scales; 10-11 collar scales; 21-22 uni- 
carinate and keeled lamellae under fourth toe, 
which is not as strongly serrated as in the holo- 
type; 24/24 femoral pores; 23-24 scales forming 
fifth caudal whorl behind vent; preanal plate 
smaller. 

Coloration. Upper head region olive brown 
with many irregular dark brown small spots, 
temporal region and upper surface of forelimbs 
with numerous dark brown blotches; dorsum al- 
most uniformly dark greyish brown with traces 
of 7-8 light stripes on neck and a very pale 
vertebral line; proximal upper caudal region with 
four brown and three light stripes, rest of tail with 
two brownish bands, bordered by three light 
stripes dorsal ly and laterally; upper surface of 
hind legs brown, with many light, round ocelli; 
ventral surfaces whitish; rostral, mental, and to 



RASTEGAR-POUYANI: ACANTHODACTYLUS N/LSONJ 



261 



some extent, submaxillary scales yellowish 
brown. 

Measurements (in mm). Snout-vent length = 
71.6, tail length =116, forelimb = 24, hindlimb 
= 42, tip of snout to forelimb = 25.9, head length 
= 17.3, head width = 1 1.6, head depth = 9.6. 

Habitat. — Semi-arid and arid steppes pre- 
dominate in the lowland regions of western Iran, 
west of the Zagros Mountains and its western 
foothills. 

The new species of Acanthodactylus reported 
on here is from the lowland areas of Kermanshah 
Province, western Iran. The province is located 
on the western periphery of the Iranian Plateau 
(Fig. 7). The Zagros Mountains dominate the 
province's northern and northeastern areas. To 
the west, near the Iranian-Iraqi border, are low- 
lands with elevations of 200-700 m. The type 
locality of Acanthodactylus nilsoni is located on 
this lowland area (Fig. 8), at an elevation of 285 
m. This region is the northernmost part of a wide 
lowland and desert belt that extends to the south, 
southeast, and southwest. It is characterized by 
long, hot and dry summers (35°C-50°C) and 
short, mild winters. The habitat of A. nilsoni is 
characterized by a soft sandy and alluvial sub- 
strate covered by desert-adapted vegetation in- 
cluding Tamarix, Astragalus, Zygophyllum, 
Artemisia, Euphorbia, and Alhagi, as well as 
numerous species belonging to the family 
Graminaceae (Fig. 9). 

I collected this lizard on the sandy and stony 
hills of the area south of Qasr-e-Shirin.The holo- 
type was foraging among the luxuriant bushes. 
The paratype was collected about a half hour 
later; it was actively looking for insects. When 
alarmed, these lizards took refuge beneath 
bushes or darted inside small holes. They were 
shy and wary and difficult to capture. Among 
other lizards found in this area may be men- 
tioned, Laudakia nupta nupta. Uromastyx lori- 
catus, Trapelus ruderatus ruderatus, Ophisops 
elegans, Mabuya aurata, and Varanus griseus 
griseus. 

Taxonomic Account 

The genus Acanthodactylus is primarily Sa- 
haro-Sindian in distribution and only one spe- 
cies, A. erythrurus, reaches the Mediterranean 
and Atlantic coasts of Europe in southern Portu- 
gal and Spain (Salvador 1982). The center of 



origin and diversity for this genus is believed to 
be the Arabian peninsula and Mesopotamian 
lowlands and Syrian Desert (Anderson 1966, 
1968, in press; Arnold 1973, 1980, 1983). 
Boulenger (1885) regarded Acanthodactylus as 
the close relative of Latastia, which occurs in 
Africa, but Arnold (1989) places Acanthodacty- 
lus as the sister taxon of Mesalina plus Ophisops 
-Cabrita. 

Acanthodactylus has recently been revised by 
Salvador (1982) and Arnold (1983); they recog- 
nized 9 species groups. Of these groups, three are 
mainly North African, and only two species, A. 
cantoris and A. blanfordi, reach as far east as 
southern Afghanistan, Pakistan, and the deserts 
of the northern Indian peninsula (Anderson, in 
press; Clark 1990; Salvador 1982). Acanthodac- 
tylus does not penetrate the Iranian Plateau to any 
great extent (Anderson 1966, 1968, in press; 
Arnold 1983; Leviton et al. 1992; Salvador 
1982). 

Of the six species of this genus, the new one 
included, that occur in Iran, A. micropholis and 
A. grandis belong to monotypic groups, A. blan- 
fordi and A. schmidti belong to the "cantoris" 
group, A. boskianus and A. nilsoni (sp. nov.) 
belong to the "boskianus-schreiberr group. 
Based on some similarities it has with both A. 
boskianus and A. schreiberi (e.g., more or less 
intact supraoculars, three series of scales on the 
fingers, usually four upperlabials anterior to the 
center of eye, straight longitudinal rows of ven- 
trals, keeled upper caudals), the new species is 
related to the "A. boskianus-schreibeiT group. It 
differs from both A. boskianus and A. schreiberi 
in several species-specific characters (Table 1). 
For instance, it is easily distinguishable from A. 
boskianus in having only 8 longitudinal rows of 
ventrals (10-12 in A. boskianus), more scales 
across dorsum (48-52 vs 18-43), weakly keeled 
temporal scales as well as dorsals, more pointed 
snout, and more lamellae under the fourth toe 
(23-24 vs 10-23); from A. schreiberi it differs 
in having larger and usually less numerous dor- 
sals (48-52 vs 43-64), 8 longitudinal rows of 
ventrals ( 10 in A. schreiberi ), 25-26 gular scales 
in a single longitudinal row (31-46 in A. 
schreiberi), more pointed snout, distinct pectina- 
tion of the anterior edge of tympanum, and slight 
pectination of eyelids. 

Acanthodactylus boskianus is widely distrib- 
uted throughout much of North Africa, reaching 



262 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 9 




Figure 7. Location of Kermanshah Province on the Iranian Plateau. 



45 : 



IRAQ 




50 km 



Figure 8. The type locality of Acanthodactylus nilsoni, 5 km south of Qasr-e-Shirin, Kermanshah province, western Iran. 
(□) = Type locality, (■) = Qasr-e-Shirin. 



RASTEGAR-POUY Mil. ACANTHODACTYLUS NILSONI 



263 





: 



■ 







Figure 9. Habitat and type locality of Acanthodactylus nilsoni, 5 km south of Qasr-e-Shirin, Kermanshah Province, western 
Iran. 



as far south as northern Nigeria and Eritrea; 
eastward it extends into the Arabian peninsula, 
Israel, Jordan, Iraq, Syria, and adjoining Turkey 
as well as western Iran (Rastegar-Pouyani, in 
press). On the other hand. A. schreiberi has been 
reported only from Cyprus, Lebanon, and Israel 
(Salvador 1982) as well as one doubtful record 
from Iraq (Reed and Marx 1959). The new spe- 
cies may well have originated as an isolate of A. 
boskianus, as the latter has been reported from 
Iraq (Boulenger 1 9 1 9. 1 92 1 ; Khalaf 1 959; Salva- 
dor 1982). The presence of A. nilsoni in western 
Iran only a few kilometers from the Iraqi frontier 
makes it likely that it also occurs inside Iraqi 
territory, at least in the easternmost parts of that 
country. Further investigation and field work is 
needed to establish the position of this lizard in 



Iraq, and also in southern lowland areas of Ham 
and Khuzistan Provinces in Iran. 

ETYMOLOGY. — Acanthodactylus nilsoni is 
named in the honour of my supervisor Dr. Goran 
Nilson for his contribution to the study of the 
Iranian Plateau and Middle East herpetofauna. 

Material Examined. — Acanthodactylus 
nilsoni (n = 2): GNHM Re.ex. 5145-6, from 
Qasr-e-Shirin, Kermanshah Province, western 
Iran. Acanthodactylus boskianus (n = 3): CAS 
203484; GNHM. Re. ex. 5142^13, from 2 km 
northwest of Harsin (34°17'N, 47°24'E), Ker- 
manshah Province, western Iran (first record for 
Iran). Acanthodactylus boskianus asper (n = 4): 
GNHM. Re. ex. 3333, 3346, 4799, 4937, from 
Sudan, Morocco, Libya, and Gaza. 



264 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 9 



Table 1. Comparison of morphological characteristics in Acanthodactylus nilsoni, A. boskianus, and A. 
schreiberi. [Based on material examined, as well as literature (e.g., Salvador 1982)] 



Characters 




A. nilsoni 


A. boskianus 


A. schreiberi 


Scales across dorsum 




48-52 


generally 1 8^43 


43-64 


Plates across venter 




8 


10-12 


10 


Collar scales 




10-12 


6-11 


8-10 


Gular scales in 1 










longitudinal row 




25-26 


21-31 


31-36 


Lamellae under fourth toe 


23-24 


10-23 


20-23 


Temporal scales 




weakly keeled 


distinctly keeled 


usually smooth 


Dorsal scales 




weakly keeled 


distinctly keeled 


usually granular 


Dorsal scales size 




moderate 


usually large 


small 


Snout position 




distinctly pointed 


slightly pointed 


obtuse 


Anterior edge of tympanum 


pectinate 


pectinate 


smooth 


Eyelids 




pectinate 


slightly pectinate 


unpectinate 


Subocular touches the 


mouth 


- 


+/- 


- 



Acanthodactvlus boskianus euphraticus (n = 
8): BMNH 1946.8.4.83-90, from Ramadieh, 
Iraq (33°25'N, 43°17'E). Acanthodactylus 
schreiberi (n = 3): GNHM. Re. ex. 4646 (1-3), 
from Cyprus. 

Abbreviations. — BMNH = British Muse- 
um of Natural History, London, England. CAS = 
California Academy of Sciences, San Francisco, 
California, USA. GNHM Re. ex. = Gothenburg 
Natural History Museum, Reptilia exotica, 
Gothenburg, Sweden. 

ACKNOWLEDGMENTS 

I would like to thank the Razi University 
authorities (Kermanshah-Iran) as well as Abbas 
Shanazari for their cooperation during field work 
in western Iran. I wish to thank Goran Nilson and 
Gothenburg Natural History Museum for loan of 
Acanthodactylus boskianus and A. schreiberi 
specimens. I am grateful to Colin McCarthy, 
Department of Zoology, British Museum of 
Natural History for loan of syntypes of Acantho- 
dactylus boskianus euphraticus. 

I am also thankful to S. C. Anderson, Biologi- 
cal Science Department, University of the 
Pacific, Stockton, California, for all his recom- 
mendations, encouragements, and for sending 
informative literature, especially his unpublished 



manuscript concerning the Iranian lacertids as 
well as critically reviewing this paper. 

As well, my special thanks go to Alan E. Levi- 
ton. Department of Herpetology, California 
Academy of Sciences, for his kind cooperation 
and for critically reviewing the manuscript. 

Literature Cited 

Anderson, S. C. 1966. The turtles, lizards, and am- 
phisbaeniansof Iran. Ph.D. Thesis. Stanford Univer- 
sity, Stanford, CA. 660 pp. 

. 1968. Zoogeographical analysis of the lizard 



fauna of Iran. Pp. 305-371 in The Cambridge His- 
tory of Iran, Vol.1 : The Land of Iran, W. B. Fisher, 
ed. Cambridge Univ. Press, Cambridge, UK. 

— . In press. The Lizards of Iran. Society for the 



study of Amphibians and Reptiles, Oxford, OH. 

Arnold, E. N. 1973. Relationships of the Palearctic 
lizards assigned to the genera Lacerta, Algyroides 
and Psammodromus (Reptilia: Lacertidae). Bull. 
Brit. Mus. Nat. Hist., Zool. Ser. 25(8):289-366. 



— . 1980. The reptiles and amphibians of Dhofar, 
Southern Arabia. Jour, of Oman Studies, Spec. 
Rept. 2:295-307. 



RASTEGAR-POUY ANI: ACANTHODACTYLUS NILSONJ 



>65 



— . 1983. Osteology, genitalia and the relation- 
ships of Acanthodactylus (Reptilia: Lacertidae). 
Bull. Brit. Mus. Nat. Hist., Zool. Ser. 44(5): 
291-339. 

— . 1989. Towards a phylogeny and biogeogra- 



phy of the Lacertidae: Relationships within an Old- 
World family of lizards derived from morphology. 
Bull. Brit. Mus. Nat. Hist., Zool. Ser. 55(2): 
209-257. 

Boulenger, G. A. 1 885. Catalogue of the Lizards in 
British Museum, Vol. 1. Taylor and Francis, Lon- 
don, UK. xii + 436 pp. 

— . 1919. On a new variety of Acanthodactylus 



boskianus Daudin, from the Euphrates. Ann. Mag. 
Nat. Hist.. Ser. 9, Vol. 3. pp. 549-550. 

— . 1921. Monograph of the Lacertidae, Vol. 2. 



British Museum (Natural History), London, UK. 
viii + 451 pp. 



CLARK, R. 1990. A report on herpetological observa- 
tions in Afghanistan. Brit. Herp. Soc. Bull. 
33:20-24. 

Khalaf, KamelT. 1959. Reptiles of Iraq, with some 
notes on the Amphibians. Ar-Rabitta Press, Bagh- 
dad, Iraq, vii + 96 pp. 

Leviton, A. E., S. C. Anderson, K. Adler, and 
S. A. Minton. 1992. Handbook to Middle East 
Amphibians and Reptiles. Society for the study of 
Amphibians and Reptiles, Oxford, OH. vii + 252 pp. 

Rastegar-Pouyani, N. In press. First record of the 
lacertid Acanthodactylus boskianus (Sauria: Lacer- 
tidae) for Iran. Asiatic Herpetol. Res. 

Reed, C. A. and H. Marx. 1959. A herpetological 
collection from northeastern Iraq. Trans. Kansas 
Acad. Sci. 62:91-122. 

Salvador, A. 1982. A revision of the lizards of the 
genus Acanthodactylus (Sauria: Lacertidae). Bon- 
ner Zool. Monogr. No. 16. 167 pp. 



CALIFORNIA ACADEMY OF SCIENCES, 1998 
Golden Gate Park 
San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 10, pp. 267-277, 7 figs., 1 table. June 19, 1998 



A NEW SPECIES OF LACERTA (SAURIA: LACERTIDAE) FROM 

THE ZAGROS MOUNTAINS, ESFAHAN PROVINCE, 

WEST-CENTRAL IRAN 

By 

Nasrullah Rastegar-Pouyani 
and 

Goran Nilson 

Department of Zoology, Gothenburg University, Box 463, 
SE 405 30 Gothenburg, Sweden 



A new saxicolous species of the genus Lacerta is described based on material collected in 1996 
from the Zagros Mountains, 3 km northwest of Fereydun Shahr, Isfahan Province, west-central 
Iran. The combination of relatively small size, 10 longitudinal rows of ventral plates, 7 pairs of 
submaxillary shields, single postnasal, masseteric shield minute or absent, complete row of 
granules between supraciliaries and supraoculars, 58-61 smooth, round, granular scales across 
dorsum, strongly compressed one-third distal part of tail, and unique color pattern distinguish it 
from all other species of Lacerta. The new species, Lacerta zagrosica, is related to the Lacerta Part 
II (sensu Arnold). It is likely restricted in distribution to the central Zagros Mountains in Esfahan 
Province, at about 2450 m elevation. 

Received November 26, 1997. Accepted February 3, 1998. 

The lacertid genus Lacerta Linnaeus 1758 In Iran, most species of Lacerta occur in 

(sensu lato) encompasses about 31 species, cen- northern, northwestern, and western parts of the 

tering on the least xeric regions of the Mediter- country, and are closely related in distribution to 

ranean and extending through the northern and the Zagros and Alburz Mountains. An exception 

western mountain ranges of southwest Asia. is Lacerta mostoufi Baloutch, 1976 which ap- 

Only two species, L. vivipara and L. agilis, occur pears to be a disjunct taxon, adapted to highly 

outside of this region, extending the known range xeric habitats and restricted in distribution to the 

of the genus across northern Eurasia to about Kavir-e-Lut on the southeastern part of the Ira- 

63°N in eastern Siberia and east to Japan. In nian Plateau (31°12'N, 59°19'E) (Baloutch 

southwest Asia, the genus is confined to the 1976). So far, 13 species of Lacerta have been 

Mediterranean coast, Turkey, and in Iran to the recorded for Iran, including the new and distinc- 

Zagros, Alburz, and western Kopet Dagh Moun- tive species from the Zagros Mountains, Ferey- 

tains. In the southeast, it is found on the Arabian dun Shahr, Esfahan Province (Figs. 1, 2) 

peninsula in the mountains of Oman {L. jayakari described here. 
Boulenger, 1887, and L. cyanura Arnold, 1972) 1 
(Anderson, in press; Arnold 1973). 



'These are regarded by some as belonging to a distinct genus, Omanosaura. 

[267] 



268 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 10 



Species Description 

Lacerta zagrosica sp. nov. 
(Figs. 3-6) 

HOLOTYPE AND TYPE LOCALITY. — Adult 
male, GNHM Re. ex. 5149, collected by the 
senior author on 14 August 1996, from the Za- 
gros Mountains, at about 2450 m elevation, 3 km 
northwest of Fereydun Shahr (50°04'E, 
32°58'N), about 140 km northwest of Esfahan 
city, Esfahan province, west-central Iran. 

Paratype. — Adult female, GNHM Re. ex. 
5150. Other information as for the holotype. 

DIAGNOSIS. — A relatively small-sized lacer- 
tid (maximum SVL, 67.8 mm; TL, 114 mm.), 
assignable to Part II of the genus Lacerta (see 
Arnold 1973), and differentiated from all other 
species by possessing a combination of 1 longi- 
tudinal rows of ventral plates, 58-61 smooth, 
granular scales across dorsum, single postnasal, 
7 pairs of submaxillary shields, complete row of 
granules between supraciliaries and supraocu- 
lars, obtusely keeled subdigital lamellae, mas- 
seteric shield very small or absent, dorsum 
greenish or olive brown with numerous dark 
spots, venter blue with black spots and dots on 
ventro-lateral region. 

Description of Holotype. — Adult male 
(preserved in 75% ethyl alcohol), with depressed 
body and slender tail; head distinctly depressed, 
its length less than twice its width ( 1 .6), its depth 
about 60% of its width; head length about 28% 
of distance from snout to vent; snout pointed, 
about as long as postocular part of head; tail 1 .69 
times as long as head and body, slightly de- 
pressed at base, tapering towards tip and strongly 
compressed in distal one-third; frontal slightly 
shorter than its distance from tip of snout, about 
1 .75 to 1 .90 times as long as its maximum width, 
narrowed posteriorly and in contact with two 
prefrontals, two supraoculars on each side, and 
two frontoparietals; parietals each about twice as 
long as its maximum width; occipital markedly 
wider than interparietal and more than half as 
long as interparietal length, and broadly in con- 
tact with it; four supraoculars, first small, not in 
contact with frontal, separated from it by prefron- 
tal and second supraocular, second and third 
supraoculars large, subequal, the second in con- 
tact with prefrontal and frontal, the third in con- 
tact with frontal and frontoparietal, the fourth 



small, almost as large as the first; supraciliaries 
7/7, first the largest, separated from second and 
third supraoculars by a complete row of 14/13 
granules; lower eyelid with 6-7 large, and about 
12 smaller scales that form an opaque central 
disk; nostril bordered by internasal, a single post- 
nasal, first upper labial, and rostral, the latter 
separated from naris by a very narrow rim; two 
loreals present, anterior loreal about half the 
length of posterior one, and in contact with first, 
second, and third supralabials, posterior loreal in 
contact with third and fourth supralabials inferi- 
orly and with prefrontal and first supraocular 
superiorly; 5/5 supralabials anterior to subocular, 
3/3 posterior to it; subocular more than twice as 
wide as long, borders mouth; three supratempo- 
rals decreasing in length posteriorly, the first 
longer than the two posterior temporals taken 
together, all supratemporals partly situated on 
parietal table of skull; temporal region covered 
by small scales, 60-63 on each side, scales larger 
than dorsals, 6 in a vertical line between su- 
pratemporals and last supralabial; anterior tem- 
porals larger than posterior ones, masseteric 
shield minute, tympanic shield weakly devel- 
oped and elongate; lower labials 6/6; submaxil- 
lary shields 7/7, first three pairs in complete 
contact, but the other pairs entirely separated, 
fourth pair largest and in contact with fourth and 
fifth infralabials and separated from each other 
medially by 3-4 small scales, sixth pair almost 
as large as first and in contact with the fifth, an 
extra (seventh) shield on each side, elongate and 
in contact with fifth and sixth shields (Fig. 5); 
28-29 gular scales in a straight line between 
collar and symphysis of chin shields; weakly 
developed gular fold present; collar not serrated, 
made up of 1 scales, fifth the largest and broader 
than long. 

Dorsal scales granular, non-imbricate, smooth, 
oval and round, 58-61 across widest part of 
dorsum, three of which correspond to each ven- 
tral plate; ventral plates almost rectangular with 
very weak posterior imbrication, arranged in 27 
transverse and 10 longitudinal rows, the first 
longitudinal row from midline widest, the outer- 
most rows the smallest; some lateral scales bor- 
dering ventrals enlarged, up to 1/3 the length of 
these shields; 3 lateral granules correspond to one 
ventral shield; preanal plate strongly developed, 
broader than long, bordered by a semicircle of 
seven large scales; a patch of five transverse 



RASTEGAR-POUYANI AND NILSON: LACERTA ZAGROSICA 



269 




Figure 1. Location of Esfahan Province on the Iranian Plateau. 




Figure 2. The type locality of Lacerta zagrosica, Zagros Mountains, 3 km northwest of Fereydun Shahr, about 140 km 
northwest of Esfahan city, Esfahan Province, west-central Iran. 



270 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 10 




Figure 3. Lacerta zagrosica, holotype. A. Dorsal region, B. Anterior ventral region. 



postanal scales, contrasting strongly with all 
other parts of body in coloration, being yellowish 
cream; upper forelimb with series of rather large 
shields above and granular scales below; a series 
of strongly developed shields under lower fore- 
limb; hind limb with lateral and ventral series of 
strongly enlarged plates, which are not separated 
from each other by small scales; dorsal surface 
of thigh with granular scales, its anterior, and 
lateral surfaces covered by two series of enlarged 
plates; ventral plates separated from the femoral 
pores by two rows of somewhat enlarged, over- 
lapping scales; 20-21 femoral pores, the two 
series just failing to reach the knees distally, 
separated from each other anteriorly by five 
small scales; tibia with two series of large plates, 
the largest series on the ventral and the other 



series on the lateral surface, dorsal surface cov- 
ered by small, granular, weakly pointed scales, 
slightly larger than dorsals, 20-2 1 in a transverse 
row in mid-tibia; digits of forelimb similar to 
hindlimbs in subdigital scalation; digits I, II, and 
V of hindlimb with one series of lamellae be- 
neath, which are obtusely keeled towards the tip 
of digits, digits III— IV with two series, except the 
first phalanx, where there is only a single series; 
digits of forelimb have the same arrangement; all 
digits of both fore- and hindlimbs covered above 
by smooth plates, fourth digit the longest, third 
and fifth digits almost the same length, followed 
by the second, and first respectively; caudal 
scales nearly rectangular, much enlarged and 
elongate, forming distinct whorls, the proximal 
10 whorls about half as long as median whorls. 



RASTEGAR-POUYANI AND NILSON: LACERTA ZAGROSICA 



271 




FiGi'RE 4. Lacerta zagrosica, paratype. Top: Dorsal region. Bottom: Ventral region. 



272 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 10 




Figure 5. The comparison of submaxillary region in Lacerta zagrosica (right) and L. brandtii (left) 



distal whorls smaller in size , their scales being 
more keeled and slightly pointed; each individual 
scale of the median whorls corresponds to 4-5 
dorsal granular scales in length; dorsal and lateral 
caudal scales rather strongly keeled longitudi- 
nally, proximal ventral caudals smooth, distal 
ones to some extent keeled, 3 1 scales in the fifth 
caudal whorl behind vent; tail strongly com- 
pressed in distal one-third with more elongate 
and keeled scales, both dorsally and ventrally. 

Measurements (in mm). Snouth-vent length 
(SVL): 67.8, tail length (TL): 114, head length: 
18.8, head width: 1 1.5, head depth: 7, length of 
forelimb: 25, length of hindlimb: 38, tip of snout 
to forelimb: 28. 

Coloration (in life and immediately after 
death). Dorsal surface of head olive brown green 
with irregular dark spots and dots, ground color 
of dorsum green, with numerous dark spots on 
sides having tendency to form a reticulation, 
encompassing light green spaces; these dark dots 
and spots less numerous on the vertebral region, 
which looks almost uniformly light green; dorsal 
surface of tail also light green with irregular dark 



spots scattered throughout; upper surface of 
limbs reticulated, dark oval and round ocelli en- 
compassing light green spaces; all of ventral 
surfaces dark blue, mixed with black spots and 
dots, these black markings most numerous on the 
ventro-lateral and gular regions; postanal region, 
corresponding four or five transverse scale rows, 
strongly contrasting in color with all other parts 
of body, being yellowish cream; ventral surface 
of tail uniformly light turquoise blue. 

Description of the Paratype. — The pa- 
ratype, an adult female, is similar to the holotype 
differing as follows: 

Four supratemporals present on the right side; 
masseteric shield absent; tympanic shield 
smaller and more elongate; 11-12 temporal 
scales in a straight line between tympanum and 
orbit; 34-35 scales in fifth whorl of tail; preanal 
plate bordered posteriorly by semicircle of eight 
scales; number of submaxillary shields as holo- 
type, but arranged slightly differently; the fourth 
pair of submaxillaries separated medially by 3 
small scales; 58-59 scales across widest part of 
dorsum; supraciliaries 6/7; femoral pores 18-20; 



RASTEGAR-POUYANI AND NILSON: LACERTA ZAGROSICA 



273 





Figure 6 The comparison of temporal region in L. zagrosica (top) and L. brandtii (bottom). Note the absence of masseteric 
shield and the presence of a rudimentary tympanic shield in L. zagrosica. 



lamellae under fourth toe 27/26; collar scales 1 1 ; 
11/11 granules between supraciliaries and su- 
praoculars; ventral plates in 28 transverse rows. 
Coloration of paratype (in life and immedi- 
ately after death). Dorsal surface of head light 
olive brown, with several very small dark spots; 
dorsum olive brown with numerous light ocelli 
scattered throughout dorsolateral regions; as well 
as numerous dark spots and dots on almost all 
pails of dorsum having tendency to make reticu- 
lation; dorsal surface of limbs as dorsum in col- 
oration, but with larger light ocelli; upper surface 



of tail light bluish brown with a few small dark 
spots on the proximal half; venter more or less 
whitish blue with a few dark blue spots on the 
v en tro- lateral region; lower surfaces of limbs and 
tail light blue; pectoral and gular regions tur- 
quoise blue with numerous dark blue spots. 

Measurements (in mm). Snouth-vent length 
(SVL): 57.8, tail length (TL): 97. head length: 
14.6, head width: 8.9, head depth: 5.3. length of 
forelimb: 20, length of hindlimb: 33, tip of snout 
to forelimb: 23. 



274 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 10 



BlOTOPE AND ECOLOGY. —Lacerta zagrosica 
inhabits the same rocks as Laudakia caucasia 
caucasia, foraging on the rocks or under the rock 
crevices. The holotype was collected while bask- 
ing on a rock; it was easy to see because of its 
vivid and brilliant blue ventral and greenish dor- 
sal coloration. The paratype was collected about 
an hour later; it had taken refuge under a huge 
rock. It was not as brilliantly colored as the male 
(holotype). The area is a part of the central Zagros 
with an elevation of about 2450 m; it is a typical 
habitat for Laudakia caucasia caucasia (Fig. 7). 
The area was surveyed extensively without find- 
ing more specimens. Lacerta zagrosica appears 
to be a saxicolous species apparently restricted in 
distribution to the Zagros Mountains. 

Also, two specimens of Mabuya aurata were 
collected from the same area. 

Taxonomic Discussion 

Before Arnold's revisionary work on Lacerta 
(sensu lato) in 1973, at least 55 species were 
assigned to the genus. Of these, 49 species are 
limited to a relatively small part of the southwest- 
ern Palearctic region, but two, L. vivipara Jaquin 
and L. agilis Linnaeus, which do not penetrate 
into the Iranian Plateau, occur not only in this 
area but also range much more widely in Eurasia. 
Four members of the genus are also found in 
Africa south of the Sahara desert (Arnold 1973). 
In his review of the genus, Arnold (1973) re- 
moved the tropical and southern African species 
from Lacerta, and raised the subgenera Podarcis 
and Gallotia to generic rank. This author split the 
genus Lacerta (sensu stricto) into two groups. 
Lacerta Part I consists of large, robust species 
having strongly imbricate ventrals and serrated 
collars. These species usually inhabit areas of 
dense, shrubby vegetation. This group is com- 
posed of Lacerta agilis, L. lepida, L. schreiberi, 
L. viridis, L. pamphylica, L. trilineata, L. prin- 
ceps, L. strigata, and L. media. The latter three 
species occur in Iran. The members of this group 
are distributed in north Africa (Morocco, north- 
ern Algeria, and northwestern Tunisia), main- 
land Europe, Mediterranean Islands, and western 
Asia. 

Arnold's Lacerta part II consists of smaller 
species (usually less than 90 mm snout-vent 
length, L.jayakari with SVL = 165 mm being an 
exception). Almost all have brightly colored ven- 



ters, at least the breeding males. They occupy a 
wide range of ecological niches, and many are 
adapted to living on or around the rocks. This 
group consists of about 24-26 species distributed 
discontinuously over almost the whole range of 
Lacerta (northern Africa, most of Europe, west- 
ern and southwestern Asia) with the exception of 
L. vivipara, which is found over a very large area 
of Palearctic Eurasia. 

Mayer and Bischoff (1996) in their taxonomic 
revision of the genus Lacerta based on morpho- 
logical and karyological studies, excluded the 
subgenera Zootoca Wagler, 1830 (type species = 
Lacerta vivipara), Omanosaura Lutz, Bischoff 
and Mayer, 1986 (type species = Lacerta 
jayakari), and Timon Tschudi, 1836 (type spe- 
cies = Lacerta lepida) from the genus Lacerta 
and raised them to the generic level. These 
authors regarded Lacerta (sensu stricto) as the 
sister group of Timon based on morphological 
features. Mayer and Benyr (1994) placed Zoo- 
toca as the sister group of Lacerta (sensu stricto) 
based on the relationships of their serum albu- 
mins. 

We have assigned L. zagrosica to Arnold's 
Lacerta Part II because of the many characters 
it shares with the other members of this group 
(e.g., small size, non-imbricating and brightly- 
colored ventrals, non-serrated collar, and special 
adaptations for living on the rocks). The mem- 
bers of this group often have small or disjunct 
ranges. This is almost certainly a relict distribu- 
tion and indicates that the group has undergone 
considerable reduction of the area of its total 
range (Arnold 1973). This seems to be the case 
with L. zagrosica as it is apparently restricted in 
distribution to the central Zagros Mountains in 
west-central parts of the Iranian Plateau. 

With regard to the present distribution of this 
lizard, we propose two alternative scenarios; (1) 
either it is the relict of a widely distributed group 
of lacertids which invaded from the Mediterra- 
nean region and northwestern Iran towards the 
southern regions along the Zagros Mountains 
during unfavorable climatic conditions of the 
Quaternary, acquiring special adaptations for liv- 
ing on or around the rocks [based on some simi- 
larities which it has with the archaeolacertids of 
Caucasus (e.g., depressed head, smooth dorsals, 
slender and fragile tail), this scenario is favored], 
or (2) it is one of the northernmost isolated and 
fragmented populations of an ancient group of 



RASTEGAR-POUYANI AND NILSON: LACERTA ZAGROSICA 



275 




Figure 7. A view of the Zagros Mountains, 3 km northwest of Fereydun Shahr, Esfahan Province, west-central Iran, habitat 
and type locality of L. zagrosica. 



lacertids which once were continuously distrib- 
uted throughout the central, south-central, and 
southeastern parts of the Iranian Plateau. The 
presence of L. mostoufii Ba\oulch, 1976 in south- 
eastern Iran and the occurrence of two lacertids 
(L. jayakari, and L. cyanura) on the mountains 
of Oman [which, most likely, once were con- 
nected to Iran through the shallow Strait of Hor- 
tnoz (Anderson 1968:363)] favors this latter 
hypothesis. 

The degree of relationship of L. zagrosica 
with these southern taxa is yet to be established. 
However, it is similar to L. cyanura in having a 
depressed habitus, two series of obtusely keeled 
lamellae under some digits (III— IV), slender 
limbs and tail, a series of strongly developed 
plates under lower forelimb, and approximate 
absence of masseteric shield. In the latter two 
characters, L. zagrosica is also similar to L. 
jayakari, though there are significant differences 
in body size and meristic counts between the two 
taxa. 

In 1968, the Street Expedition to Iran collected 
seven specimens of Lacerta brandtii from the 



Zagros Mountains, 10 km north of Kuh Rang in 
Esfahan Province about 60 km south of the type 
locality of L. zagrosica. 

Otherwise, Lacerta brandtii has a disjunct 
range, mainly distributed in northwestern Iran, 
East Azarbaijan Province as well as contiguous 
regions of the Azarbaijan Republic (Anderson, in 
press). 

Geographically, the nearest species of Lacerta 
part II to L. zagrosica is L. brandtii and then L. 
cappadocica urmiana. The latter has already 
been collected from the Zagros Mountains, Ker- 
manshah Province, western Iran (Rastegar- 
Pouyani, in press). We have examined and 
compared several species of Lacerta part II with 
L. zagrosica (see Material Examined). 

Lacerta zagrosica is easily distinguishable 
from Lacerta chlorogaster, L. praticola, and L. 
mostoufii in having smooth dorsal scales and a 
non-serrated collar; from L. cappadocica ur- 
miana in the absence of transparent shields on 
the lower eyelid, having 7 pairs of submaxillary 
shields, only one postnasal, and in color pattern; 
from L. brandtii in having only one postnasal, 7 



276 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 10 



Table 1 . Comparison of morphological characteristics in Lacerta zagrosica, L. brandtii, and L. 
cappadocica urmiana. 



Characters 


Lacerta zagrosica 


Lacerta brandtii 


L. cappadocica urmiana 


Ventral plates 


10 


8 


8 


(longitudinal) 








Submaxillary shields 


7 pairs 


5 pairs 


5 pairs 


Dorsals at midbody 


58-61 


47-52 


52-69 


Gular scales (single 


28-30 


23-26 


32-35 


row) 








Femoral pores 


18-21 


16-20 


22-27 


Translucent window on 


absent 


absent 


present 


the lower eyelid 








Masseteric shield 


rudimentary or absent 


strongly developed 


absent 


Tympanic shield 


small and narrow 


large and elongate 


almost absent 


Number of postnasals 


only one 


almost always two 


two or rarely three 


Subdigital lamellae 


weakly ( obtusely )keeled 


smooth 


strongly keeled 


Dorsal pattern of male 


greenish blue 


pale brown, olive grey 


greenish with dark 
reticulations and blue 
ocelli 


Male's throat and 


entirely blue 


usually yellowish white 


light blue 


abdomen 








Habitat 


rocky areas 


mainly steppes and 
hillsides 


rocky areas 



pairs of submaxillary shields, 10 longitudinal 
rows of ventrals, more scale counts across dor- 
sum, approximate absence of masseteric shield, 
small tympanic shield, and in color pattern (Ta- 
ble 1); from L. valentini valentini, L. steineri, L. 
defilippii, and L. raddei in having 5 supralabials 
anterior to subocular, complete row of granules 
between supraoculars and supraciliaries, 7 pairs 
of submaxillary shields, presence of pterygoid 
teeth and significant differences in color pattern; 
and from the Lacerta part I (Arnold 1973) in 
having much smaller size, non-serrated collar, 
rectilinear, or nearly so, ventral plates as well as 
differences in color pattern. 

In the more southern regions, many of the 
allopatric species of Lacerta Part II are strongly 
differentiated, probably indicating a long-stand- 
ing separation, but in the north the species are 
more similar to each other and here the reduction 
in range presumably was more recent. Probably 
the glacial and interglacial ages of Pleistocene 
caused the expansions and contractions of the 
range. Now, many species are restricted to the 
relatively moist or highland regions. 

In conclusion, however distinct many of these 
species are, including our new taxon, the classi- 



fication of Palearctic Lacerta presents many 
difficulties (Arnold 1972, 1973, 1989a, b; 
Boulenger 1920; Eiselt 1995; Lutz and Mayer 
1985; Mayer and Benyr 1994; Mayer and Bis- 
choff 1996; Mayer and Tiedemann 1982). Many 
of the characters used in systematics exhibit great 
interspecific variation, making the delimitation 
of species boundaries difficult. Many forms also 
show considerable intra-populational variability 
especially in the color and color pattern. And 
because of the uncertainties in the systematics of 
the group, a useful interpretation of the group's 
historical biogeography eludes us. 

ETYMOLOGY. — Lacerta zagrosica is so 
named as it is certainly restricted in distribution 
to the Zagros Mountains of west-central Iran. 

Material Examined. — Lacerta zagrosica 
(n = 2): GNHM Re. ex. 5149-50, from the Za- 
gros Mountains, 3 km northwest of Fereydun 
Shahr, Esfahan Province, west-central Iran. 

Lacerta brandtii (n = 9): FMNH 170956-62, 
from 10 km north of Kuh Rang, Esfahan Prov- 
ince, west-central Iran. GNHM Re. ex. 2873—4, 
from Ardabil, East Azarbaijan (Ardabial) Prov- 
ince, northwestern Iran. 



RASTEGAR-POUYANI AND NILSON: LACERTA ZAGROSJCA 



111 



Lacerta cappadocica urmiana (n = 3): CAS 
203483, GNHM Re. ex. 168RJ69R (field num- 
bers) from the Zagros Mountains, 30 km north- 
east of Kermanshah city, Kermanshah Province, 
western Iran. 

Lacerta chlorogaster (n = 10): GNHM Re. ex. 
2439-2448, from Transcaucasia. 

Lacerta defilippii (n = 3): GNHM Re. ex. 4413 
(1-3), from the Lar Valley. Alburz Mountains, 
northern Iran. 

Abbreviations. — CAS = California Acad- 
emy of Sciences, San Francisco, California, 
USA. FMNH= Field Museum of Natural His- 
tory. Chicago, Illinois, USA. GNHM Re.ex. = 
Gothenburg Natural History Museum, Reptilia 
exotica, Gothenburg, Sweden. 

Acknowledgments 

We would like to thank: Harold K. Voris, 
curator of Herpetology, Field Museum of Natural 
Histoiy. Chicago, for loan of Lacerta brandtii 
specimens. S. C. Anderson, Biological Science 
Department, University of the Pacific, Stockton, 
California, for all of his recommendations, en- 
couragements, and for sending informative lit- 
erature, especially his unpublished manuscript 
concerning the Iranian lacertids as well as criti- 
cally reviewing this paper. Alan E. Leviton, De- 
partment of Herpetology, California Academy of 
Sciences, for his kind cooperation and for criti- 
cally reviewing the manuscript. 

Also, our special thanks go to the Razi Univer- 
sity authorities (Kermanshah-Iran) as well as 
Alireza Hashemi and Masoud Abrishamian for 
their unsparing help during field work in west- 
central Iran. 

Literature Cited 

Anderson, S. C. 1968. Zoogeographical analysis of 
the lizard fauna of Iran. Pp. 305-371 in The Cam- 
bridge History of Iran, Vol.1: The Land of Iran, 
W. B. Fisher, ed. Cambridge Univ. Press, Cam- 
bridge, UK. 

— . In press. The Lizards of Iran. Society for the 



study of Amphibians and Reptiles, Athens, OH. 

Arnold, E. N. 1972. Lizards with northern affinities 
from the mountains of Oman. Zool. Meded., Leiden 
47:111-128. 



— . 1973. Relationships of the Palearctic lizards 
assigned to the genera Lacerta, Algyroides and 
Psammodromus (Reptilia: Lacertidae). Bull. Brit. 
Mus. Nat. Hist.. Zool. Ser. 25(8):29 1-366. 

. 1989a. Systematics and adaptive radiation of 

Equatorial African lizards assigned to the genera 
Adolfus, Bedriagaia, Gastropholis, Ho/aspis and 
Lacerta (Reptilia: Lacertidae). J. Natur. Hist., Lon- 
don 23:525-555. 



— . 1 989b. Towards a phylogeny and biogeogra- 
phy of the Lacertidae: relationships within an Old- 
World family of lizards derived from morphology. 
Bull. Brit. Mus. Nat. Hist., Zool. Ser. 55(2): 
209-257. 

Baloutch, M. 1976. Une nouvelle espeee de Lacerta 
(Lacertilia, Lacertidae) du sud-est de Llran. Bull. 
Mus. Natio. Hist. Nat. Paris, Zool. 294(417): 
1379-1384. 

BOULENGER, G. A. 1920. Monograph of the Lacerti- 
dae, Vol. 1 . British Mus. Nat. Hist., London, x + 352 
pp. (see pp. 282-301). 

ElSELT, J. 1 995. Ein Beitrag zur Kenntnis der Archae- 
olacerten (sensu Mehely, 1 909) des Iran (Squamata: 
Sauria: Lacertidae). Heropetozoa 8 (1/2): 59-72. 

Lutz, D. and W. Mayer. 1985. Albumin evolution 
and its phylogenetic and taxonomic implications in 
several lacertid lizards. Amphibia-Reptilia, Leiden 
6:53-61. 

Mayer, W. andG. Benyr. 1994. Albumin evolution 
und phylogenese in der Familie Lacertidae (Rep- 
tilia: Sauria). Ann. Naturhist. Mus., Wien 
96B:62 1-648. 

Mayer, W. and W. Bischoff. 1996. Beitrage zur 
taxonomischen Revision der Gattung Lacerta ( Rep- 
tilia: Lacertidae), Teil 1: Zootoca, Omanosaura, 
Timon und Teira als eigenstandige Gattungen. Sala- 
mandra 32(3): 163-1 70. 

Mayer, W. and F. Tiedemann. 1982. Chemotax- 
onomical investigations in the collective genus Lac- 
erta (Lacertidae: Sauria) by means of protein 
electrophoresis. Amphibia-Reptilia, Wiesbaden 
2:349-355. 

RASTEGAR-POUYANI, N. In press. A preliminary con- 
tribution to the lizard fauna of Kermanshahan Prov- 
ince, western Iran. Proc. II Asian Herpetological 
Meeting, Ashghabad-Turkmenistan, 6-10 Septem- 
ber 1995. Russian Jour. Herpetol. 5(1). 



CALIFORNIA ACADEMY OF SCIENCES, 1998 
Golden Gate Park 
San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 1 1, pp. 279-293, 7 figs., 1 table. July 20, 1998 



TWO NEW DISCODORID NUDIBRANCHS FROM THE 

WESTERN PACIFIC WITH A REDESCRIPTION 

OF DORIS LUTEOLA KELAART, 1858 



By 

Terrence M. Gosliner 
and 



JUL 2 7 1998 



David W. Behrens 

Department of Invertebrate Zoology, California Academy of Sciences 
Golden Gate Park, San Francisco, California 941 18 



Two new species of discodorid cryptobranch dorid nudibranchs are described from the tropical 
Indo-Pacific. Hoplodoris estrelyado is found from several localities in the western and central 
Pacific: Vietnam, Western Australia, Indonesia, Philippines and the Marshall Islands. It is the 
only member of the genus in which all of the radular teeth are denticulate. Taringa halgerda is 
known from Papua New Guinea, the Philippine Islands and Indonesia. It is similar in appearance 
to T. luteola, but differs in its coloration and tubercles and aspects of its internal anatomy. Taringa 
luteola (Kelaart, 1858) is redescribed and its systematic placement is discussed. 

Placement of the tw o species described here is regarded as tentative. Neither Hoplodoris estrelyado, 
Taringa luteola nor Taringa halgerda are placed with certainty in the genera where they are here 
assigned. They possess characteristics that are divergent from other described members of the 
genera Hoplodoris and Taringa. A major review and revision of dorid genera is required. 
Phylogenetic analysis would clarify the relationships between and within taxa and would 
determine which taxa represent monophyletic groups. Systematic revisions would then be made 
to reflect these monphyletic units. 

Received December 24, 1997. Accepted February 9, 1998. 



The tropical Indo-Pacific is rich in its diversity The status of the systematics of the doridacean 

of opisthobranch gastropods (Gosliner 1992; nudibranchs, especially the Cryptobranch ia, is in 

Ghiselin 1992; Gosliner and Draheim 1996). An- need of much revision. The separation of genera 

nual field collections in the Philippine Islands and their organization into families has been 

over the past several years have yielded speci- undertaken on an ad hoc basis, without employ- 

mens of numerous new species of opistho- ing phylogenetic methods. Many presently rec- 

branchs. Several of these have been recently ognized genera are based on what are likely 

described (Gosliner and Behrens 1997, 1998) or plesiomorphic characteristics and represent para- 

their descriptions are presently in press. This phyletic taxa. Problems associated with taxo- 

paper describes two new species of cryptobranch nomic allocation of species are illustrated by the 

dorid nudibranchs and compares them to other difficulties presently encountered in placing the 

species that are similar in appearance. two species described here into well-circum- 
scribed genera. 

[279] 



280 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 1 



Species Descriptions 
Family DlSCODORIDIDAE 

Hoplodoris estrelyado sp. nov. 
(Figs. IA,2,3) 

Hoplodoris nodulosa (Angas): Debelius, 1996:253, 
two bottom photos; not (Angas, 1 864). 

Discodoris sp. Gosliner, Behrens and Williams, 
1996:159, tig. 558. 

TYPE Material. — Holotype: CASIZ 088 1 14, one 
specimen, Twin Rocks, SW Calumpan Peninsula, 
Batangas Province, Luzon, Philippine Islands, 18 m 
depth, 4 March 1993, T. M. Gosliner. Paratypes: 
CASIZ 085950, one specimen, dissected, Bus Stop 
Reef, Balayan Bay, Batangas Province, Luzon, Phil- 
ippine Islands, 8 m depth, 24 March 1993, T. M. 
Gosliner. CASIZ 105664, three specimens, Hamilo 
Bay - The Head, Batangas Province, Luzon, Philip- 
pine Islands, 13 m depth, 4 March 1995, T. M. 
Gosliner. CASIZ 088113, one specimen, DeviFs 
Point, Maricaban Island, Batangas Province, Luzon, 
Philippine Islands, 13 m depth, 23 May 1993, M. D. 
Miller. CASIZ 110459, one specimen. Cathedral 
Rock, Balayan Bay, Batangas Province, Luzon, Phil- 
ippine Islands, 13 m depth, 23 May 1993, M. D. 
Miller. 

ETYMOLOGY. — The trivial name estrelyado 
is the Tagalog word for "sunny side up" which is 
derived from the Spanish word "estrella," for 
star, indicating the eggs are formed into starlike 
patterns with the yolk shining upwards. This 
applies to the unique color pattern on the notal 
surface, giving the appearance of fried eggs, 
served sunny-side up. 

Distribution. — Thus far, this species is 
known from Vietnam and Western Australia 
(Debelius 1996), Indonesia, the Philippine Is- 
lands (present study), and the Marshall Islands. 

NATURAE HISTORY. — Hoplodoris estre- 
lyado is found on the outer edges of rock walls 
and reef fronts in 5-30 m of water. 

External Morphology. — The living ani- 
mals (Fig. lA)are 15-40 mm in length. The body 



is oval and flat, the notal surface covered with 
large tapering tubercles (Fig. 2A), which lack 
spicules that project from the tubercles. The col- 
oration of the living animals is complex and 
strikingly unique. The ground color of the notal 
surface is translucent tan with dark brown and 
white specks. An oval pattern of tubercles resem- 
bling fried eggs, a white patch with yellow cen- 
ter, surrounds a medial group of dark brown 
tubercles. The size of the white, egg-like ring 
becomes smaller on the tubercles nearer the mar- 
gin. The tubercles forming the 6-8 largest fried- 
egg patterns are each encircled by 8 10 smaller 
white tubercles. A few tan colored tubercles are 
dispersed about the notal surface, and are particu- 
larly abundant near the center of the notum. The 
underside of the foot and mantle are speckled 
with dark brown. The anterior margin of the foot 
is bilabiate and notched. On either side of the 
head is a long, digitiform tentacle which also is 
covered with brown speckles. Six tripinnate 
branches form the branchial plume, which is tan 
with dark brown and white specks. The anus is 
situated within the gill circle. The perfoliate rhi- 
nophores bear about 13-19 lamellae and are dark 
reddish brown with white specks. 

Buccal Armature. — The buccal mass is 
large and muscular. At the anterior end of the 
muscular portion of the buccal mass is the thin, 
chitinous labial cuticle. It contains a pair of thick- 
ened triangular regions anteriorly that bear nu- 
merous irregularly tipped rodlets (Fig. 3A). The 
radular formula in one specimen is 26 x 65.0.65. 
The innermost lateral teeth (Fig 3B) are hamate 
with two minute, irregular denticles along the 
outer edge of the tooth. The radular teeth from 
the middle of the half-row (Fig. 3C) have a far 
more elongate cusp with up to 11 minute denti- 
cles on the outer margin. The teeth from the outer 
margin of the radula (Fig. 3D) have shorter, 
thicker cusps with 6-1 1 denticles along the outer 
margin. 

Reproductive System. — The reproductive 
system is triaulic (Fig. 2B). The ampulla is thick 
and tubular, narrowing somewhat before bifur- 



Figiri I. Living animals. A. Hoplodoris estrelyado sp. nov., specimen from Batangas Province, Philippine Islands, 
photograph by T. M. Gosliner. B. Taringa luleola Kelaart, 1858, Msimbati, Tanzania, photograph by T. M. Gosliner. 
C. Taringa halgerda sp. nov., specimen from Bus Stop Reef, Batangas Province, Philippine Islands, photograph by T. M. 
( i (is liner. 



GOSLINER AND BEHRENS: DISCODORID NUDIBRANCHS 



281 




282 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 1 





am 



ji^VfV(-;---^g.yJ--^V--v.'-->jv-J:;. l .;>-.-^ 



Figure 2. Hoplodoris estrelyado sp. nov. A. Living animal. B. Reproductive system, am = ampulla, be = bursa copulatrix, 
ej = ejaculatory duct, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina, vg = vestibular 
gland, scale = 1 .0 mm. C. Vestibular gland stylet, scale = 1 60 um. 



GOSLINER AND BEHRENS: DISCODORID NUDIBRANCHS 



283 




Figure 3. Hoplodoris estrelyado sp. nov. Scanning electron micrographs of buccal armature. A. Jaw rodlets, scale = 15 urn. 
B. Inner lateral teeth, scale = 43 urn. C. Lateral teeth from middle of the half-row, scale = 60 urn. D. Outer lateral teeth, scale 

= 43 urn. 



284 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 1 



eating into the oviduct and vas deferens. The 
short oviduct enters the female gland mass near 
the albumen gland. The thick proximal prostatic 
portion of the vas deferens is folded over itself 
once before it narrows into the long, narrow, 
highly convoluted, ejaculatory portion. The 
ejaculatory portion enters the short, wider penial 
bulb. The penis, which is adjacent to the slender 
vaginal duct at the common gonopore, is devoid 
of any armature, when stained and cleared. The 
female gland mass consists of the large mucous 
and membrane glands and a smaller albumen 
gland. The vagina is relatively long and straight 
with one partial loop. The proximal end of the 
vagina enters the large, spherical bursa copula- 
trix. Adjacent to the junction of the vagina and 
bursa is the thin duct which joins the thick- 
walled, club-shaped receptaculum seminis. This 
duct parallels the vagina for most of its length. 
The extremely short uterine duct also emerges 
from the receptaculum seminis and enters the 
female gland mass near the albumen gland. Near 
the exit of the mucous gland is a single, very large 
irregularly-shaped vestibular gland. The vestibu- 
lar duct has an opening separate from the 
gonopore. The vestibular gland is armed with a 
long tapering stylet (Fig. 2C), which is visible 
when the distal end of the gland is stained and 
cleared. The stylet is 800 um in length and is 
rounded at its proximal end. The stylet is not 
calcareous, as it was not dissolved when fixed in 
Bouin's fixative. 

DISCUSSION. - The genus Hoplodoris was 
described by Bergh (1880) with H. desmo- 
panpha designated as type species, by mono- 
typy. The anatomy of this species has not been 
completely described, although several aspects 
of its morphology have been depicted by Bergh 
(1880, 1905). The external morphology and 
color of the living animals were not described. 
The characters that Bergh (1905) used to distin- 
guish the genus were: presence of jaw rodlets. 
penial spines and a vestibular gland with a hol- 
low, cornicopiate spine. Burn (1969) considered 
Doris nodulosa Angas, 1864 as a species of 
Hoplodoris, based on the facts that it has penial 
spines and a vestibular gland with an armed 
stylet. Thompson (1975) confirmed the presence 
of a vestibular gland and penial spines, although 
neither author illustrated the reproductive struc- 
tures of this species. Miller (1991) redescribed 
the anatomy of Homiodoris novaezelandiae 



Bergh, 1904 and transferred this species to 
Hoplodoris on the basis that it has two distinct 
vestibular glands with hollow, conical stylets and 
a series of penial spines. These are the only three 
nominal species definitely placed in Hoplodoris. 

Thompson (1975 ), following his description of 
Australian specimens of Hoplodoris nodulosa, 
considered Carminodoris Bergh, 1889 as a jun- 
ior synonym of Hoplodoris, stating that Carmi- 
nodoris appears to be unnecessary since almost 
all of its characters can be embraced by the older 
Hoplodoris. Thompson noted that there were 
anatomical differences between the specimens of 
H. nodulosa he examined and those described by 
Kay and Young (1969) as Carminodoris nodu- 
losa. He attributed these differences to the fact 
that Kay and Young did not serial section their 
material. However, there are other anatomical 
differences between the Hawaiian and Australian 
material, such as the denticulation of the radular 
teeth, that suggest that they are not conspecific. 
It is likely that Thompson's material is conspeci- 
fic with Angas' Doris nodulosa since it is col- 
lected near the type locality, near Sydney, New 
South Wales. Kay and Young's material prob- 
ably represents an undescribed species. Absence 
of a stylet in the vestibular gland must be con- 
finned in the Hawaiian material. 

When the anatomy of the described species 
placed in Carminodoris and Hoplodoris is com- 
pared (Table 1 ), considerable variation between 
species is evident. All species have a nodular 
dorsal surface with numerous rounded tubercles. 
The radular teeth may be smooth or denticulate 
(Baba 1993). The penis and vagina may be lined 
with spines or may be unarmed. One or more 
vestibular glands may be present, which may be 
large or small, sessile or at the end of an elongate 
duct. The glands may be unarmed, armed with a 
curved, hollow spine or a solid, straight stylet. 
These differences are not correlated with each 
other and each species possesses a unique com- 
bination of characters. This suggests that 
Thompson's synonymy of Carminodoris with 
Hoplodoris may be justified and is here provi- 
sionally maintained. 

Hoplodoris estrelyado differs from all other 
described species of Hoplodoris and Carmino- 
doris in several regards. It is the only species in 
which all the teeth are denticulate. Only H. 
grandiflora (Pease, 1 860) and H. estrelyado have 
an unarmed penis (Kay and Young 1969; present 



GOSLINER AND BEHRENS: DISCODORID NUDIBRANCHS 



285 



■3 

o 

=: 



EC 



o 

X: 



C 

E ~ * 



=3 
£3 












































































i/l 












u 






aj 




13 




13 












<U 












15 






cS 




03 




a 












> 


G 

o 










DO 

c 






DO 

3 




DO 

C 




DO 

3 














i> 




u 




g 


o 




_o 


"-> 


c 


u 


S 


CJ 






13 






33 


'33 






53 


3 
— 


o- 


<L> 


3 


o 


3 


o 


3 
-J 


c^- 




'55 






o 


13 




-C 




c 






a 




3 




3 








13 






a. 


ca 




03 




o 






o 




C 




O 








EA 












































a3 






































x: 






































E 






C"-- 
































3 






































z 


■a 




O 

-a 








c^- 


•a 




-a 




■a 




^3 




T3 




o3 




13 




u 










13 




o 




13 




13 




O 




3 




E 




E 




T3 






E 




E 




E 




3 




E 




'So 
> 














c^- 
























03 




s 




E 






03 




3 




53 




53 




-3 






s 




c 








3 




C 




a 




c 




3 






3 




3 




03 






3 




3 




3 




3 




3 






















-a 








T3 






























<u 








o 












l/l 




TD 




T3 




— 




■a 


£ 




■a 




E 




~3 




T3 






U 




13 




U 




<u 




o 






V 




13 




'2 




E 




E 




E 




£ 


03 




E 




s 




E 




E 


















u- 


c 




u. 




3 








Uh 




Q. 




03 


■a 


03 


— 


03 


T3 


03 


3 


•a 


53 




3 


~3 


r5 




r3 


T3 


3 






03 




13 




o 






u 








o 








13 








3 




2 




3 






3 








3 








3 


£1 






5 




c 











"o 








c 








3 


13 




>^ 


> 


>. 


> 




> 


O- 




> 








^ 


C^- 




>1 


> 


"O 




13 


c 


~ 


s 


>, 


c 




>, 


s 






>. 


3 






— 


3 






J= 


c 


x: 





r- 


o 




IE 


c 


C 




c 


C 






x: 





o3 
> 




DO 


o 


DO 


u 


2 






Dp 

2 


u 


o 
x: 




Dp 

2 


o 








U 


x: 




jC 




-3 

1) 


x: 


-5 
o 






Xj 

o 

o 

1 


u 

3 

o 


x: 


Xj 




o 

C 




X3 

E 


Xj 



c 

E 

c/3 


Xj 




c3 




O 

o 




JD 


o 

o 


E 

t/3 




=8 


o 

c 


5 






E 




=a 








3 

03 




E 




— 


E 

c/3 


u 

s 






13 
c 


E 


E 

t/3 




i) 
c 
o 
c 




l-l 
13 

3 




E 




C£ 




"53 




E 




3 




c 


c 




"3 






C 




~~i 
















cs 




g 


J 








o 










su 










a 




o 




•S 


'a 








"a 




a 


_J 




3 






g 




a 




^ 




-2 


.,_, 




3 








_3 


, 


•^ 




.2 
G 


g 








"a 




a 


C3 


'S3 


l 

3 
=_ 






~s. 

3 


*3 

5 


e 

.a 




a 




-5 




00 




s 


_~ 


X 


"a 


*3 




s 


< 


s 


^3 

ts3 




Qu 

on 


U 




U 




u 




u 


u 




5: 




tt: 




li 




aj 





286 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 1 



study). Hoplodohs estrelyado and H. des- 
mopaiypha are the only two species that have a 
large spherical vestibular gland that is situated at 
the end of an elongate duct. It appears that H. 
estrelyado and Australian specimens of H. nodu- 
losa are unique in having an elongate, solid stylet 
associated with the vestibular gland. 

Confirmation of the generic placement of H. 
estrelyado must await further study of other 
members of the genus. Hoplodohs desmo- 
pcuypha, the type species of the genus, has not 
been recorded since its original and subsequent 
descriptions by Bergh ( 1880, 1905) and nothing 
is known about the color of the living animals. 
Similarly, the type species of Carminodoris, C. 
mauhtiana, is known only from the original de- 
scription (Bergh 1889) and its anatomy and the 
morphology of the living animal remain un- 
known. The absence of vestibular gland stylets 
in Hawaiian specimens identified as C. nodulosa 
and C. grandiflora require confirmation. Follow- 
ing the accumulation of this anatomical data, 
construction of aphylogenetic hypothesis testing 
the monophyly of these taxa must be undertaken. 

Members of other dorid genera (including As- 
ieronotus Ehrenberg, 1831, Sclerodoris Eliot, 
1903, Jorunna Bergh, 1876 and Kentrodoris 
Bergh, 1876) possess a vestibular gland. In As- 
teronotus the notum is ornamented with large, 
irregularly angled tubercles arranged in ridges 
rather than having densely arranged, rounded 
tubercles as in Hoplodoris (including Carmino- 
doris). Also, jaw rodlets are absent in Asterono- 
tus but are present in members of the latter genus. 
Species of Sclerodoris, Jorunna and Kentrodoris 
have the notum ornamented with spiculate 
caryopyllidia rather than rounded tubercles and 
lack jaw rodlets. In Sclerodoris the vestibular 
gland is sessile and lacks any armature while in 
Jorunna and Kentrodoris the gland has a long 
duct and a stylet as in Hoplodoris. Detailed com- 
parisons of this taxa with Hoplodoris must await 
a comprehensive parsimony-based phylogenetic 
study of the cryptobranch dorids to ascertain 
which similarities are due to common ancestry 
and which represent independent acquisition of 
these characters. 



Taringa luteola (Kelaart, 1858) 
(Figs. IB, 4, 5) 

Doris luteola Kelaart, 1 858: 1 03. 

Thordisa caudata Farran, 1905:340, pi. 2, figs. I 8, 19. 

Trippa luteola (Kelaart) Eliot, 1906:658, pi. 42, fig. 4. 

Material Examined. — CASIZ 099361, one 
specimen, dissected, 1 m depth, Manahuanja Island, 
Mtwara Region, Tanzania, 1 November 1994, T. M. 
Gosliner. CASIZ 099231, one specimen, 1 m depth, 
Manahuanja Island, Mtwara Region, Tanzania, 2 No- 
vember 1994, T. M. Gosliner. 

Distribution. — Thus far, this species is 
known only from the western Indian Ocean of 
Madagascar (Eliot 1906), Tanzania (present 
study) and Sri Lanka (Kelaart 1858; Farran 
1905). 

External Morphology. — Preserved ani- 
mals are 15-22 mm in length. The living animals 
(Fig. IB) are cream white with low, pointed 
tubercles. There is a yellow band along the entire 
margin of the mantle. An additional interrupted 
ring of yellow pigment is found in the central 
portion of the mantle extending from just poste- 
rior to the rhinophores to the region immediately 
anterior to the gills. The gills are white with 
brown to gray marginal pigment around the ulti- 
mate branches of the gill. The rhinophores are 
dark gray to black. The tubercles have a few 
irregular, digitate papillae extending from their 
dorsal surface (Fig. 4A). On either side of the 
head is a long, digitiform tentacle. The anterior 
end of the foot is bilabiate. The branchial plume, 
composed of six sparsely branched, bi- to tripin- 
nate gill branches, is large, nearly as wide as the 
body. A black line extends up the edge of each 
gill branch. The perfoliate rhinophores bear 
about 16 lamellae. 

Buccal Armature. — The buccal mass is 
large and muscular, with elongate curved sali- 
vary glands. At the anterior end of the muscular 
portion of the buccal mass is the thin chitinous 
labial cutical. It contains some thicker regions 
but is entirely devoid of chitinous rodlets. The 
radular formula is 28 x 37-39.1.37-39 in one 
specimen examined. The rachis (Fig. 5A) con- 
tains a row of thin, elongate rachidian plates. The 
inner lateral teeth are simply hamate and curved 
with two triangular denticles. The first 30 teeth 
(Fig. 5B) have 1-2 denticles while the next 13 
teeth lack any trace of a denticle. The outer 3-4 



GOSLINER AND BEHRENS: DISCODORID NUDIBRANCHS 



287 



teeth (Fig. 5C) are much shorter and have a finely 
fimbriate margin of elongate, bristly spines. 

Reproductive System. — The reproductive 
system is triaulic (Fig. 4B). The ampulla is thick, 
elongate and tubular, narrowing somewhat be- 
fore bifurcating into an oviduct and vas deferens. 
The short oviduct enters the female gland mass 
near the albumen gland. The proximal prostatic 
portion of the vas deferens is not much wider than 
the ejaculatory portion and is folded over itself 
once before it narrows slightly into the long, 
muscular ejaculatory portion. The ejaculatory 
portion consists of several loops. It narrows and 
then enters the moderately long penial bulb, 
which is devoid of armature but contains a thin 
cuticular lining near its apex (Fig. 4C). The penis 
is adjacent to the narrower vaginal duct at the 
common gonopore. The female gland mass con- 
sists of the large mucous gland and smaller mem- 
brane and albumen glands. The vagina is 
relatively long and curved. The proximal end of 
the vagina enters the base of the thin walled. 



spherical bursa copulatrix. Adjacent to this is a 
somewhat shorter thin duct to the pyriform re- 
ceptaculum seminis. The short uterine duct 
emerges from the base of the receptaculum and 
enters the female gland mass near the albumen 
gland. This species lacks any genital armature. 

DISCUSSION. — Doris luteola was originally 
described from Sri Lanka (Kelaart, 1858). Eliot 
( 1 906) described aspects of the anatomy of speci- 
mens from the eastern coast of Madagascar. Eliot 
considered Thordisa caudata Farran, 1905 as a 
junior synonym of Doris luteola, based on their 
similar color patterns and presence in Sri Lanka. 
Eliot transferred this species from Doris and 
Thordisa to Trippa on the basis that it has com- 
pound tubercles and ptylaine glands present on 
the buccal mass in addition to salivary glands. He 
also redescribed Trippa ornata Bergh 1880, type 
species of the genus Trippa, and strongly sug- 
gested that it is a synonym of Doris intecta 
Kelaart, 1859. Edmunds (1971) considered T. 
ornata as a junior synonym of Trippa intecta and 






Figure 4. Taringa luteola (Kelaart, 1858). A. Detail of tubercles, scale = 1 .0 mm. B. Reproductive system, am = ampulla, 
be = bursa copulatrix, ej = ejaculatory duct, fgm = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, 
v = vagina, scale = 1 .0 mm. C. Cuticular lining of vas deferens, c = cuticle, scale = 70 urn. 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 I 




Figure 5. Taringa luteola (Kelaart, 1858). Scanning electron micrographs of radula. A. Rachidian and inner lateral teeth, 
scale = 20 urn. B. Lateral teeth from middle of the half-row, scale = 30 urn. C. Outer lateral teeth, scale = 43 urn. 



GOSLINER AND BEHRENS: DISCODORID NUDIBRANCHS 



289 



described its anatomy in greater detail. We con- 
firmed this synonymy by re-examining the holo- 
type of T. ornata in the Zoologisk Museum, 
University of Copenhagen. This species differs 
from Taringa luteola in several important re- 
gards. The buccal mass of Trippa intecta has a 
small glandular portion (Edmunds 1971, fig. 8d) 
anterior to the insertion of the lateral muscula- 
ture. Such glands were not found in Taringa 
luteola. All of the raular teeth of Trippa intecta 
are simply hamate and devoid of auxiliary denti- 
cles or bristles. The radula of Taringa luteola has 
a row of vestigial rachidian teeth that are absent 
in Trippa intecta. Most of the lateral teeth have 
a single small denticle and the outer three laterals 
have fine, elongate bristles along their margin. 
The vaginal duct of Taringa luteola is far more 
elongate than that of Trippa intecta. 

Three other genera of dorids have radular teeth 
that are similar to those of Doris luteola. Species 
of Thordisa Bergh, 1877, Aporodoris Ihering, 
1886, and Taringa Marcus, 1955, all have elon- 
gate bristles on the outermost radular teeth. The 
latter two genera have denticles on most of the 
other lateral teeth, while denticles are absent in 
species of Thordisa. Marcus and Marcus (1967) 
stated that the two genera differ only by the 
presence of penial armature in Taringa and its 
absence in Aporodoris. Thompson and Brown 
( 198 1 ) considered Aporodoris as a junior syno- 
nym of Discodoris, based on a re-examination of 
the type material of Doris millegrana Alder and 
Hancock, 1854, type species of Aporodoris. 
Thompson and Brown united the genera despite 
the fact that most species of Discodoris, includ- 
ing the type species, Discodoris bolwliensis 
Bergh, 1877, lack denticles on the radular teeth. 
It appears that Aporodoris is more similar to 
Taringa than to Discodoris, but confirmation of 
its systematic placement requires detailed ana- 
tomical study of the penial papilla and sub- 
sequent phylogenetic analysis. The present 
species bears considerable resemblance to spe- 
cies of Taringa, in that the majority of radular 
teeth have denticles and the distal portion of the 
vas deferens has a cuticular lining. The absence 
of a distinctly thickened cuticular ring and the 
presence of a vestigial row of rachidian teeth 
distinguishes Do/75 luteola from other described 
members of Taringa. 



Taringa halgerda sp. nov. 
(Figs. 1C. 6. 7) 

Cadlinella sp. Debelius, 1996:21 7, top photo. 

Type Material. — Holotype: CASIZ 1 06469, one 
specimen. Bus Stop Reef, Balayan Bay, Batangas 
Province, Luzon, Philippine Islands, 13 m depth, 18 
April 1996, T. M. Gosliner. Paratypes: CASIZ 
110427, five specimens (one dissected). Bus Stop 
Reef, Balayan Bay, Batangas Province, Luzon, Phil- 
ippine Islands, 3 m depth, 23 April 1997, D. W. Behr- 
ens. CASIZ 083731, two specimens, Bus Stop Reef, 
Balayan Bay, Batangas Province, Luzon, Philippine 
Islands, 3 m depth, 21 February 1992, M. D. Miller. 

ETYMOLOGY. — The trivial name halgerda 
was chosen to indicate the overall, external, mor- 
phological similarity between this species and 
members of the genus Halgerda. 

DISTRIBUTION. — Thus far. this species is 
known only from the Philippine Islands (present 
study), from a photograph by Takamasa 
Tonozuka from Bali, Indonesia (Debelius 1996) 
and from a photograph by Jim Black from Milne 
Bay, Papua New Guinea. 

Natural History. — Taringa halgerda is 
found on the outer edges of rock walls and reef 
fronts in shallow water. The egg mass is a densely 
packed, open coiled ribbon, attached to the sub- 
strate along its edge. It has a characteristic pale 
blue color and is about 20 mm in diameter. 

External Morphology. — Preserved ani- 
mals are 22-33 mm in length. The living animals 
(Fig. 1C) are white with low, flat, yellow tuber- 
cles (Fig. 6A). The tubercles are digitate, with 
short, round papillae extending from their poste- 
rior surface (Fig. 6B). The largest are capitate 
with a narrow, short base. These are found me- 
dially on the dorsum. The anterior end of the foot 
is bilabiate and notched. On either side of the 
head is a long, digitifonn tentacle. The branchial 
plume, comprised of six sparsely branched, mul- 
tipinnate gill branches, is large, nearly twice the 
width of the body in diameter. The anus is situ- 
ated within the circle formed by the gill branches. 
The size and shape of the gill represent the basis 
for considering this species "Halgerda-Uke' in 
its appearance. A black line extends up the edge 
of each gill branch. The black, perfoliate rhino- 
phores bear about 17-18 lamellae. 

Buccal Armature. — The buccal mass is 
large and muscular. At the anterior end of the 



290 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 1 1 





B 





Figure 6. Taringa halgerda sp. nov. A. Living animals. B. Detail of tubercles, scale = 1 .0 mm. C. Reproductive system, 
am = ampulla, be = bursa copulatrix, ej = ejaculatory duct, fgm = female gland mass, p = penis, pr = prostate, rs = receptacu- 
lum seminis, v = vagina, scale = 1 .0 mm. D. Cuticular lining of vas deferens, c = cuticle, scale = 70 urn. 



GOSLINER AND BEHRENS: DISCODORID NUDIBRANCHS 



291 




FIGURE 7. Taringa halgerda sp. nov. Scanning electron micrographs of radula. A. Rachidian and innerteeth, scale = 60 pm. 
B. Lateral teeth from middle of the half-row, scale = 25 pm. C. Lateral teeth devoid of denticles, scale = 60 jam. D. Outer 
lateral teeth, scale = 43 pm. 



292 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. I I 



muscular portion of the buccal mass is the thin 
chitinous labial cutical. It contains some thicker 
regions but is entirely devoid of chitinous rodlets. 
The radular formula is 33 x 40. 1 .40 in one speci- 
men examined. The rachis (Fig. 7A) contains a 
row of thin, elongate rachidian plates The inner 
lateral teeth are simply hamate and curved with 
a single triangular denticle. The first 30 teeth 
(Fig. 7B) have a single denticle while the next 
5-10 teeth lack any trace of a denticle. The outer 
3 teeth (Fig. 7C) are much shorter and have a 
finely fimbriate margin of elongate, bristly 
spines. 

Reproductive system. — The reproductive 
system is triaulic (Fig. 6C). The ampulla is thick, 
elongate and tubular, narrowing somewhat be- 
fore bifurcating into an oviduct and vas deferens. 
The short oviduct enters the female gland mass 
near the albumen gland. The proximal prostatic 
portion of the vas deferens is not much wider than 
the ejaculatory portion and is folded over itself 
once before it narrows slightly into the long, 
muscular, ejaculatory portion. The ejaculatory 
portion is relatively short and contains one short 
loop. It narrows and then enters the short penial 
bulb, which is devoid of armature, but contains a 
thin cuticular lining near its apex (Fig. 6D). The 
penis is adjacent to the narrower vaginal duct at 
the common gonopore. The female gland mass 
consists of the large mucous gland and smaller 
membrane and albumen glands. The unarmed 
vagina is relatively long and curved. The proxi- 
mal end of the vagina enters the base of the thin 
walled, spherical bursa copulatrix. Adjacent to 
this is an equally long, thin duct which joins the 
bursa to the kidney-shaped receptaculum 
seminis. The short uterine duct emerges near this 
junction and enters the female gland mass near 
the albumen gland. This species lacks any genital 
armature. 

DISCUSSION. — Taringa halgerda is similar to 
T. luteola in most aspects of its anatomy. Both 
species have a similar color pattern, compound 
tubercles, a radula with a vestigial row of 
rachidian teeth, denticulate teeth and a simple, 
triaulic reproductive system. However, the two 
species differ in several regards. Taringa hal- 
gerda lacks the yellow marginal line and the 
incomplete interior yellow ring on the center of 
the mantle that are present in T. luteola. Rather. 
T. halgerda has yellow pigment restricted to the 
surface of the large tubercles in the central por- 



tion of the notum. The tubercles of T. halgerda 
are larger, more rounded and have more papillae 
extending from their apices than do those of T. 
luteola. The largest tubercles of T. halgerda are 
capitate while those of T. luteola are all conical. 
The gills of T. halgerda are proportionately 
larger than those of T. luteola. 

The radular teeth of T. halgerda have blunter 
cusps than do those of T. luteola. The teeth of T. 
luteola may have either one or two denticles 
while in T. halgerda the teeth have only a single 
denticle. More of the outer teeth lack denticles in 
T. halgerda than in T. luteola. 

The reproductive system of T. halgerda is 
similar in its morphology to that found in T. 
luteola. The primary difference between the two 
species is that the ejaculatory portion of the vas 
deferens is slightly more elongate and convo- 
luted in T. luteola. 

ACKNOWLEDGMENTS 

This work would not have been possible with- 
out the generous support of many individuals and 
institutions. Field work was made possible by 
financial support from Katharine Stewart, United 
Airlines and the Lindsay Field Research Fund of 
the California Academy of Sciences. Our collec- 
tions were made possible through the kind sup- 
port of our colleagues at the Bureau of Fisheries 
and Aquatic Resources of the Republic of the 
Philippines. Mike Miller first photographed and 
collected Hoplodoris estelyado and assisted 
greatly in documenting these species in the field. 
Dong Lin of the Photography Department of the 
California Academy of Sciences provided the 
prints and final production of scanning electron 
micrographs. Elizabeth Kools kindly provided 
technical support for producing this manuscript. 

LITERATURE CITED 

Baba, K. 1993. Two new species of Carminodohs 
(Nudibranchia: Dorididae) from Japan. Venus 
52(3):223-234. 

Bergh, R. 1 880. Malacologische Untersuchungen, 4. 
/// Reisen im Archipel der Philippinen. Zweiter 
Theil. Wissenschaftliche Resultate Suppl. 2, 
1 : 1-78, C. Semper, ed. Leipzig. 

— . 1889. Malacologische Untersuchungen, 3. //; 



Reisen im Archipel der Philippinen. Zweiter Theil 



GOSLINER AND BEHRENS: DISCODORID NUDIBRANCHS 



293 



Wissenschaftliche Resultate 16(2):8 15-872, C. 
Semper, ed. Leipzig. 

. 1 905. Malacologische Untersuchungen, 6. In 



Reisen im Arehipel der Philippine!!. Zweiter Theil. 
Wissenschaftliche Resultate 9:57-1 15. C. Semper, 
ed. Leipzig. 

Burn, R. 1969. A memorial report on the Tom Craw- 
ford Collection of Victorian Opisthobranchia. Jour- 
nal of the Malacological Society of Australia 
12:64-106. 

Debelius, H. 1 996. Nudibranchs and sea snails. Indo- 
Pacific field guide. IKAN — Unterwasserarchiv, 
Frankfurt. 321 pp. 

Edmunds, M. 1971. Opisthobranehiate Mollusca 
from Tanzania (Suborder: Doridacea). Zoological 
Journal of the Linnean Society 50(4):339-396. 

ELIOT, C. 1906. On the nudibranchs of southern India 
and Ceylon, with special reference to the drawings 
by Kelaart and the collections belonging to Alder 
and Hancock preserved in the Hancock Museum at 
Newcastle-on-Tyne. Proceedings of the Zoological 
Society of London for 1906:636-691. 

Farran, G. P. 1905. Report on the opisthobranehiate 
Mollusca collected by Professor Herdman, at Cey- 
lon, in 1902. Pp. 329-364 in Report to the govern- 
ment of Ceylon on the pearl oyster fisheries of the 
Gulf of Manaar 3, Suppl. Rept. No. 21. W. A. 
Herdman, ed. Royal Society, London. 

Ghiselin, M. T. 1992. How well known is the opis- 
thobranch fauna of Madang, Papua New Guinea? 
Proceedings of the Seventh International Coral Reef 
Symposium 2:697-701. 

GOSLINER, T. M. 1992. Biodiversity of tropical opis- 
thobranch gastropod faunas. Proceedings of the 
Seventh International Coral Reef Symposium 
2:702-709. 

GOSLINER, T. M. andD. W. BEHRENS. 1997. Descrip- 
tion of four new species of phanerobranch dorids 



(Mollusca: Nudibranchia) from the Indo-Pacific, 
with a redescription of Gymnodoris aurita (Gould, 
1852). Proceedings of the California Academy of 
Sciences 49(9):287-308. 

. 1998. Five new species of Chromodoris 



(Mollusca: Nudibranchia: Chromodorididae) from 
the tropical Indo-Pacific Ocean. Proceedings of the 
California Academy of Sciences 50(5): 139- 1 65. 

GOSLINER, T. M., D. W. BEHRENS, AND G. C. WIL- 
LIAMS. 1 996. Coral reef animals of the Indo-Pacific. 
Sea Challengers, Monterey, California. 314 pp. 

GOSLINER, T. M.andR. Draheim. 1996. Indo-Pacific 
opisthobranch gastropod biogeography: how do we 
know what we don't know? American Malacologi- 
cal Bulletin 12(l/2):37-43. 

Kay, E. A. and D. K. Young. 1969. The Doridacea 
(Opisthobranchia; Mollusca) of the Hawaiian Is- 
lands. Pacific Science 23(2): 1 72-23 1 . 

Kelaart, E. F. 1 858. Description of new and little 
known species of Ceylon nudibranchiate molluscs 
and zoophytes. Journal of the Royal Asiatic Society, 
Ceylon Branch, Colombo 3( 1 ):84-139. 

Marcus, Ev. and Er. Marcus. 1967. Tropical 
American opisthobranchs. Studies in Tropical 
Oceanography Miami 6:3-137. 

Miller, M. C. 1991. On the identity of the dorid 
nudibranch Homoiodoris novaezelandiae Bergh, 
1904 (Gastropoda: Opisthobranchia). Journal of 
Natural History 25:293-304. 

Thompson, T. E. 1975. Dorid nudibranchs from east- 
ern Australia (Gastropoda, Opisthobranchia). Jour- 
nal of the Zoological Society of London 

176:177-517. 

Thompson, T. E. and G. H.Brown. 1981. Allocation 
of the nudibranch Doris millegrana Alder and Han- 
cock, 1854 to the genus Discodoris Bergh, 1877, 
after re-examination of the type material. Zoologi- 
cal Journal of the Linnean Society 72(3):263-266. 



CALIFORNIA ACADEMY OF SCIENCES. 1998 
Golden Gate Park 
San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 12, pp. 295-306, 7 figs. 



July 20, 1998 



THE GENUS PECTENODORIS (NUDIBRANCHIA: 

CHROMODORIDIDAE) FROM THE INDO-PACIFIC, WITH THE 

DESCRIPTION OF A NEW SPECIES 



By 

Rebecca F. Johnson 
and 



JUL 2 7 1998 



Terrence M. Gosliner 

Department of Invertebrate Zoology, California Academy of Sciences 
Golden Gate Park, San Francisco, California 941 18 



The genus Pectenodoris is reviewed. The genus contains only Pectenodoris tiilineata and a new 
species, Pectenodoris aurora, described here. New records of P. tiilineata from the Philippines and 
Papua New Guinea and more information about the variation present in P. tiilineata are 
presented. The two species overlap in much of their ranges, throughout the western margins of 
the Pacific, and are similar in color pattern. They are part of different larger groups of similarly 
colored, sympatric chromodorids discussed here. 

Received December 24, 1997. Accepted March 11, 1998. 



Rudman (1984) described the monotypic ge- 
nus Pectenodoris. He hypothesized that many 
unknown, small chromodorids would be found 
to belong to this genus. One such small species 
has recently been collected and is described here 
as Pectenodoris aurora. 

Increased interest in colorful chromodorid 
nudibranchs has improved collections and 
knowledge of the fauna of the tropical Indo-Pa- 
cific. Traditionally, chromodorid nudibranchs 
have been described based on their external mor- 
phology, color pattern, radular morphology and 
reproductive anatomy. The arrangement and size 
of defensive mantle glands has previously been 
used to characterize genera, but not to distinguish 
between congeneric species (Rudman 1984, 
1986). Mantle gland configuration was found to 
be important in separating closely related species 
of the chromodorid genus Hypselodoris (Gos- 
liner and Johnson, in press). The variation pre- 



sent in the mantle glands of the two Pectenodoris 
species is discussed here. 

There are over three hundred species in the 
family Chromodorididae from the Indo-Pacific 
tropics (Gosliner and Draheim 1996). These 
large numbers can make it difficult to study 
relationships between species within large gen- 
era. One way of dealing with the enormity of the 
group has been the designation of color groups, 
groups of similarly colored species, that may or 
may not be closely related (Rudman 1982, 
1983a, 1985, 1986). Pectenodoris tiilineata had 
been included, by Rudman, in the "Noumea pur- 
purea color group,"' based on its resemblance to 
Noumea purpurea Baba, 1949, Noumea varians 
(Pease, 1871). Durvilledoris pusilla (Bergh, 
1874) and Durvilledoris similaris Rudman, 
1986. Once these distantly related species are 
placed into color groups, they can be identified, 
compared and discussed more easily (Rudman 



[295] 



2% 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 12 



1986). Pectenodoris aurora does not fit into this 
particular color group, but will be shown to be 
closer in color pattern to another group discussed 
by Rudman(1986). 

Species Descriptions 

Pectenodoris trilineata (Adams and Reeve, 

1850) 

(Figs. 1A, 2-4) 

Goniodoris trilineata Adams and Reeve, 1 850:68, pi. 

1 7, fig.4. 
Chromodoris virgata Bergh, 1 905: 1 60-1 62, pi. 4, fig. 

11; pi. 14, fig. 47. 
Pectenodoris trilineata (Adams and Reeve) Rudman, 

1984:159-163, figs 29-31. 
Pectenodoris trilineata (Adams and Reeve) Rudman, 

1986:319-320, figs 1C, 5. 

Material Examined. — CASIZ 105638, one 
specimen, dissected, Bethlehem, Maricaban Island, 
Batangas Province, Luzon Island, Philippines, 22 m 
depth, 26 February 1995, T. M. Gosliner. CASIZ 
083704, three specimens, one dissected, Devil's Point, 
SW side of Maricaban Island, Batangas Province, 
Luzon Island, Philippines, 8 m depth, 19 February 
1992, T. M. Gosliner. CASIZ 083711, three speci- 
mens, Kirby's Rock, NW side of Maricaban Island, 
Batangas, Luzon Island, Philippines, 23 m depth, 18 
February 1992, T. M. Gosliner. CASIZ 068813, one 
specimen, the Quarry, near Bunn village, 1 km south 
of Cape Croiselles, north of Madang, Papua New 
Guinea, 23 m depth, 8 August 1989, T. M. Gosliner. 
CASIZ 082924, one specimen, Rasch Passage, north 
coast near Madang Lagoon, Papua New Guinea, 9 m 
depth, 1 7 November 1990, T. M. Gosliner. 

DISTRIBUTION. — Pectenodoris trilineata is 
known from the tropical western margins of the 
Pacific, bounded in the north by the Tropic of 
Cancer and the south by the Tropic of Capricorn. 
The northern-most limit of its distribution is the 
South China Sea (Adams and Reeve 1850), and 
the most southerly record off. trilineata is from 
Heron Island, Capricornia Group, Great Barrier 
Reef. Australia (Rudman 1984, 1986). Bergh 
(1905) described this species as Chromodoris 
virgata, from the waters off of Sumbawa Island, 
Indonesia. Five new records of this species from 
Papua New Guinea and the Philippines are re- 
ported here. Pectenodoris trilineata is known 
from 8 to 23 m depth. 



External Morphology. — The variation 
observed in the nine newly collected specimens 
is consistent with that described by Rudman 
(1984, 1986). The animal from Devil's Point in 
the Philippines, CASIZ 083704, is pictured (Fig. 
1 A) as a comparison to P. aurora, the new mem- 
ber of the genus. All but two specimens exam- 
ined here were found to have glands around the 
entire mantle, opening ventrally. but all animals 
had at least one, and up to three, enlarged poste- 
rior glands. When more than one gland was en- 
larged, those glands immediately adjacent and to 
the right of the central posterior gland were 
larger. The two aforementioned specimens dif- 
fered only in the lack of all glands except the 
posterior ones (Fig. 2). 

Alimentary Canal. — In all specimens ex- 
amined, the length of the two portions of the 
buccal mass differed only slightly, while the 
width of the oral tube was about three times that 
of the muscular portion of the buccal mass. The 
jaws are small and delicate and very difficult to 
prepare for scanning electron microscopy, owing 
to the extreme reduction of the rodlets. The mor- 
phology of the jaw rodlets varies, from long and 
undivided, to others which are short and have 
multiple cusps (Fig. 3A). The radulae examined 
here display the same comb-like teeth described 
by Rudman (1984). The only notable difference 
is the lack of a vestigial, plate-like rachidian tooth 
(Figs. 3B-D). The two specimens examined here 
had radular formulae of 22 x 6.0.6 and 29 * 
15.0.15. This is similar to that reported pre- 
viously by Rudman (1984) for P. trilineata. 

REPRODUCTIVE System. — The arrangement 
of organs is triaulic (Fig. 4). The ampulla is wide 
and curved. It divides into a very long, convo- 
luted prostate and an extremely short oviduct, 
which enters the female gland mass. The prostate 
narrows slightly into the long ejaculatory por- 
tion, which terminates in an elongate, narrow 
muscular penial bulb. The vaginal duct is narrow 
and elongate. The large, curved, pyriform recep- 
taculum seminis is only slightly smaller than the 
ovoid bursa copulatrix. A wide duct connects the 
receptaculum seminis directly with the bursa 
copulatrix. The uterine duct emerges from the 
middle of the vagina, is long and narrow, and 
enters the female gland mass below the entrance 
of the oviduct to the mass. The female gland mass 
is large and completely developed. A small. 



JOHNSON AND GOSLINER: PECTENODORIS 



297 




Figure 1 . Living animals. A. Pectenodoris trilineata (Adams and Reeve, 1 850), from Batangas, Luzon, Philippines (CASIZ 
083704), photograph by T. M. Gosliner. B. Paratype of Pectenodoris aurora n. sp., from Bethlehem. Maricaban Island, 
Batangas, Luzon, Philippines (CASIZ 1 10443), photograph by T. M. Gosliner. 



298 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 12 




Figure 2. Pectenodohs trilineata (Adams and Reeve, 1 850), distribution of mantle glands. A. CAS1Z 105638, Philippines. 
B. CASIZ 083704, Philippines. C. CASIZ 068813, Papua New Guinea. D. CASIZ 083704, Philippines. E. CASIZ 083704, 
Philippines. 



lobate vestibular gland is present near the distal 
end of the female gland mass. 

DISCUSSION. — The defensive mantle glands 
of ' Pectenodoris trilineata are mentioned by Rud- 
man (1984, 1986) as a series of large single 
mantle glands, opening ventrally. This descrip- 
tion is consistent with present material, but some 
additional variation is noted (Fig. 2). 

In this species, the ratio of the glandular oral 
tube length to the muscular portion of the buccal 
mass length has been described as four to one 
(Rudman 1984, 1986). In the specimens exam- 
ined here, the length of the oral tube and the 
buccal mass was very similar. A difference in the 
width of the two portions was more apparent. 



This difference in length may be an artifact of the 
degree of extension or contraction of the buccal 
mass at the time of preservation. The shape and 
configuration of the radular teeth was found to be 
the same as described by Rudman (1984), the 
only differences being the lack of a vestigial 
rachidian plate-like tooth and the presence of 
more lateral teeth in some specimens. 

Rudman ( 1984) did not find a vestibular gland 
in his specimens, but mentioned a small glandu- 
lar structure that might have been an undevel- 
oped gland. In all of the specimens we examined 
a small, lobate vestibular gland was present 
(Fig. 4). 



JOHNSON AND GOSLINER: PECTENODORIS 



299 




Figure 3. Pectenodoris trilineata (Adams and Reeve, 1850), scanning electron micrographs. A. Jaw rodlets, (CASIZ 
105638), scale = 10 \xm. B. Inner lateral teeth, (CASIZ 083704) scale = 7.5 ^m. C. Lateral teeth (CASIZ 105638), scale = 20 
Urn. D. Lateral teeth, (CASIZ 105638), scale = 15 ^m. 



300 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 12 




Figure 4. Pectenodohs trilineata (Adams and Reeve, 1850), reproductive system (CAS1Z 083704). Abbreviations: am : 
ampulla, be = bursa copulatrix, ej = ejaculatory portion of vas deferens, fg = female gland mass, p = penis, pr = prostate, is 
receptaculum seminis, u = uterine duct, v = vagina, vg = vestibular gland. 



Pectenodoris aurora sp. nov. 
(Figs. IB, 5-7) 

Type Material. — Holotype: CASIZ 074702, one 
specimen, dissected, Seragaki Beach, ENE of Maeki- 
zaki, Ryukyu Islands, Okinawa, Japan, 58 m depth, 1 
September 1989, R. F. Bolland. Paratypes: CASIZ 
097631, two specimens, Batuangus Point, Lembeh 
Strait, north Sulawesi, Indonesia, 28 October 1993, 
Pauline Fiene-Sevems. CASIZ 110443, one speci- 
men, dissected, Bethlehem, Maricaban Island, Batan- 
gas Province, Luzon Island, Philippines, 22 m depth, 
21 April 1997, T. M. Gosliner. CASIZ I 1 1059, one 
specimen, Horseshoe Cliffs, 1 km WNW of Onna 
Village, Ryukyu Islands, Okinawa, Japan, 1 m depth, 
24 December 1994, R. F. Bolland. 

ETYMOLOGY. — Pectenodoris aurora is 
named for Aurora, the Roman goddess of the 



morning. This species is marked with all of the 
beautiful colors of a great sunrise. 

DISTRIBUTION. — The geographical range of 
P. aurora is limited to the western edge of the 
Pacific. This species is known only from the 
Indo-Pacific tropics of the Philippines and Indo- 
nesia and sub-tropical, Okinawa, Japan. It ranges 
from 10 to 58 m depth. 

External Morphology. — The living ani- 
mals are small, 6-8 mm in length (Fig. IB). The 
body shape is oval with a wide mantle overlap. 
The mantle, body and foot are all light pink in 
color. This color darkens to maroon on the edges 
of the mantle and the posterior tip of the foot. 
Three parallel whitish yellow, continuous or in- 
terrupted longitudinal bands run from the rhino- 
phores to the gills. These bands are each outlined 
by a thin opaque white line. The central and 



JOHNSON AND GOSLINER: PECTENODOR/S 



301 




Figure 5. Pectenodoris aurora n. sp., distribution of mantle glands. A. CASIZ 097631, Indonesia. B. CAS1Z 1 10443, 
Philippines. C. CASIZ 074702, Japan. D. CASIZ 097631, Indonesia. E. CASIZ 1 1059, Japan. 



longest of these bands completely encircles the 
gill cavity. The lighter pink areas between the 
bands are dotted with 5-7 dark pink to purple 
spots and occasional opaque white spots. The 
remainder of the notum lateral to the outer bands 
is marked with the same dark pink to purple and 
opaque white spots. The short posterior end of 
the foot is divided by a thin, medial white line. 
The rhinophores each have 8-9 lamellae. They 
have a dark orange base and apex with a single, 
transverse white band through the middle. The 
6 9 simply pinnate gills have the same dark 
orange base and apex with a central white band 
as do the rhinophores. 



The large mantle glands all open ventrally 
(Fig. 5). They are visible along the mantle edge 
in the living animal. All of the specimens exam- 
ined have 2^4 greatly enlarged posterior glands. 
There are 16-29 glands around the remainder of 
the mantle. There is some evidence that glands 
have been lost and regenerated in response to 
predation. It appears that more glands may be 
formed where the animal has been bitten or at- 
tacked. Areas of attack are visible in a number of 
specimens and these animals have more glands 
than their conspecifics, that lack damage along 
the mantle margin. 



302 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 12 




Figure 6. Pectenodoris aurora n. sp., scanning election micrographs. A. Jaw rodlets (CASIZ 074702), scale = 7.5 urn. B. 
Entire radula (CASIZ 074702), scale = 150 urn. C. Lateral teeth (CASIZ 097631), scale = 15 urn. D. Lateral teeth (CASIZ 
09763 1 ), scale = 1 urn. E. Lateral teeth (CASIZ 074702), scale = 1 5 urn. 



Alimentary Canal. — The oral tube is 
about three times the width of the muscular por- 
tion of the buccal mass. The jaws of this species 
are small, fragile and difficult to prepare for 
microscopy. The jaw rodlets are very small, 
fairly wide and appear to have multiple cusps 



(Fig. 6A). The radular formula of a specimen 
from Okinawa is 28 x 16.0.16 (Fig. 6B). It was 
not possible to determine the number of rows in 
the specimen from Indonesia, but there are thir- 
teen teeth per row. There is some variation in the 
number of denticles per tooth in the specimens 



JOHNSON AND GOSLINER: PECTENODORIS 



303 




Figure 7. Pectenodoris aurora n. sp., reproductive system (CASIZ 110443). Abbreviations: am = ampulla, be = bursa 
copulatrix, ej = ejaculatory portion of vas deferens, fg = female gland mass, p = penis, pr = prostate, rs = receptaculum seminis, 
u = uterine duct, v = vagina, vg = vestibular gland. 



examined here. One specimen from Okinawa, 
Japan has ten denticles on the inner lateral teeth 
and five to six denticles on the middle lateral 
teeth (Fig. 6E), whereas the specimen from the 
Philippines has only five to six denticles per 
tooth, in all teeth observed (Figs. 6C, D). 

Reproductive System. — The arrangement 
of organs is triaulic (Fig. 7). The ampulla is long 
and straight. It divides into a short, simply looped 
prostate and an extremely short, wide oviduct, 
which enters the female gland mass. The prostate 
narrows slightly into the very long, convoluted 
ejaculatory portion, which terminates in a short, 
bulbous, muscular penial bulb. The vaginal duct 
is narrow and elongate. The large, curved, pyri- 



form receptaculum seminis is longer than the 
ovoid bursa copulatrix. A wide duct connects the 
receptaculum and the bursa copulatrix at the top 
of the vaginal duct. The uterine duct emerges 
from the proximal third of the vagina, is long and 
narrow, and enters the female gland mass below 
the entrance of the oviduct to the mass. The 
female gland mass is large and completely devel- 
oped. A large, highly ramified vestibular gland is 
present near the distal end of the gland mass. 

Discussion 

Rudman (1984) created the genus Pecteno- 
doris based on the differences found in the radu- 



304 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 12 



lar morphology and mantle gland configuration 
of P. trilineata^ when compared to members of 
other chromodorid genera. Pectenodohs has 
been characterized by unique comb-like radular 
teeth; very fragile jaw rodlets; large, ventral ly 
opening defensive mantle glands and the lack of 
a vestibular gland (Rudman 1984, 1986). 

Both P. trilineata and/ 3 , aurora are very small, 
less than 10 mm in life. They are both purple to 
pink in color, although P. trilineata is darker. 
Both species have a series of three parallel, lon- 
gitudinal lines on the mantle. Pectenodohs 
aurora has yellow bands bordered by white. The 
bands in P. trilineata are smaller and are either 
solid yellow or yellow bordered by white, as in 
P. aurora. This three-lined color pattern was 
most likely present in the common ancestor of/ 3 . 
aurora and P. trilineata. It is due to descent, 
rather than convergence. Only P. aurora has 
purple and white spots on the mantle and orange 
and white banded rhinophores and gills. The only 
markings other than the bands present in P. tril- 
ineata are small maroon blotches at the base of 
the rhinophores and a thin white marginal line 
around the mantle. 

Pectenodoris aurora has radular teeth that are 
very similar in form to those of P. trilineata. The 
two species have similar radular formulae. Pec- 
tenodoris trilineata has a formula of 20-29 x 
6-15.0.6-15, while P. aurora has a formula of 
28 x 13-16.0.13-16. The number of denticles 
per inner tooth of both species is variable. The 
inner teeth have either 5-6 or 10-11 denticles, 
while the middle teeth have 5-6 denticles. Rud- 
man (1984) mentioned the presence of a 
rachidian plate in some specimens of P. tril- 
ineata. This type of tooth was not seen in any of 
the specimens of/ 3 , aurora or any specimens of 
P. trilineata examined here. 

The distribution of defensive glands around 
the mantle of these species is similar and exhibits 
a similar degree of variability. Rudman (1984, 
1986) described a general pattern for the genus 
as a series of single, large mantle glands opening 
ventrally, as he described for/ 3 , trilineata. In the 
five specimens of P. trilineata examined here, 
two distinct patterns were observed (Fig. 2). In 
both cases there are 1-3 greatly enlarged poste- 
rior glands, but two of the specimens have no 
other glands besides those in the posterior. The 
other three specimens have a series of equally 
spaced glands around the rest of the mantle. 



Pectenodoris aurora has 2—4 greatly enlarged 
posterior glands and a series of 16-19 smaller 
glands around the rest of the mantle (Fig. 5). The 
placement of these glands varies. One specimen 
lacks glands along one side, while others are 
missing glands at the anterior end. In every case, 
in both species, the central-most posterior gland 
is enlarged. One or more of the posterior glands 
adjacent to the postereomedial one are always 
enlarged, except for one P. trilineata that has 
only one enlarged gland. The presence or ab- 
sence of lateral, antero-lateral and anterior glands 
is variable, but all members of the genus have 
enlarged posterior glands. The ventrally open- 
ings of all of these glands is unique to members 
of the genus. 

The differences in the reproductive system of 
these two species include the size of the vestibu- 
lar gland (larger in P. aurora), the insertion point 
of the receptaculum seminis into the bursa copu- 
latrix, the size of the penis and the length of the 
prostate. Rudman (1984, 1986) reported the lack 
of a vestibular gland in P. trilineata, but men- 
tioned a minute glandular pouch at the genital 
opening. Here we found a small, but lobate and 
slightly ramified vestibular gland present in P. 
trilineata. The vestibular gland in P. aurora was 
much larger and much more ramified than that of 
P. trilineata. In P. trilineata, the receptaculum 
seminis enters directly into the bursa copulatrix, 
whereas in P. aurora the receptaculum seminis 
enters at the base of the bursa copulatrix at the 
vaginal duct. The penis in P. aurora is wide, 
muscular and short. In P. trilineata it is elongate, 
narrow and only slightly muscular. In addition, 
P. aurora has a long, highly-convoluted muscu- 
lar portion, while P. trilineata has a shorter sim- 
ple loop. Reproductive characteristics, like those 
mentioned above, have been shown to be impor- 
tant in distinguishing closely related species in 
other chromodorid genera, such as Hypselodoris 
(Gosliner and Johnson, in press). 

Pectenodoris trilineata has been reported from 
the western margins of the Pacific, from the 
South China Sea to the Great Barrier Reef. Pec- 
tenodoris aurora is also found along the western 
margins of the Pacific, but only as far north as 
Okinawa, Japan and as far south as the Lembeh 
Strait, Indonesia. The range of P. aurora is com- 
pletely circumscribed by that of P. trilineata. 
These species overlap within the bathymetric 
range, 8-23 m, although one specimen of P. 



JOHNSON AND GOSLINER: PECTENODOR1S 



305 



aurora was found at a depth of 58 m. In the 
Philippines, both species have been collected 
from Bethlehem. Maricaban Island, Batangas 
Province, Luzon at 22 m depth, within a few 
meters of each other. 

In an attempt to better understand the species 
included in the very large family Chromdoridi- 
dae, the members have been divided into groups 
of species that are not necessarily closely related, 
but share a common color pattern (Rudman, 
1982, 1983a, 1985, 1986). This method has 
helped us to more closely examine the different 
species and to bring attention to the possible 
importance of these color patterns in studying the 
evolution and biogeography of the family. It has 
been suggested that the sympatric members of 
each color group represent a case of Mullerian 
mimicry, that similar color patterns are a result 
of convergence and not common ancestry (Rud- 
man, 1991). Sympatry of these similarly colored 
species supports the idea of mimicry as a defense 
against predation. Pectenodoris trilineata was 
placed in the '"Noumea purpurea color group" by 
Rudman (1986) based on its resemblance to 
Noumea purpurea, Noumea varians, Dunille- 
doris similaris and Dur\illedoris pusilla. The 
basic pattern of these species is a pinkish purple 
background color with a median white line. All 
of the members of this color group have ranges 
that overlap with that of P. trilineata. Pecteno- 
doris aurora shares its color pattern with mem- 
bers of the "Chromodoris decora color group." 
This group consists of Chromodoris decora 
(Pease, 1860), Thorunna australis (Risbec, 
1928), Hypselodoris maculosa (Pease, 1871), 
Noumea alboannulata Rudman, 1986, and the 
juvenile stage of a Hexabranchus sanguineus 
(Ruppell and Leuckart, 1 830 3 1 ) (this species is 
not in the family Chromodorididae). All of these 
species are pinkish or yellowish pink with longi- 
tudinal white lines, purple and white spots and 
often orange and white banded rhinophores. 
Noumea alboannulata is the only member of this 
color group that has not been recorded in at least 
part of the range of P. aurora. However, a speci- 
men has recently been collected from Batangas, 
Philippines, within the range of P. aurora (M. 
Miller, pers. comm.) 

As mentioned above, color groups represent 
potential mimicry complexes (Rudman 1991). 
These groups are based only on the biological 
reality of perception of visual predators. The 



more variation that is present in the color patterns 
in each group the weaker the selective advantage 
of a mimicry complex becomes. There is quite a 
bit of variation present in each of these color 
groups, and potential overlap between the groups 
depending on which set of patterns are chosen to 
separate these groups. Different portions of color 
patterns in another chromodorid genus Hypselo- 
doris were found to be the result of both conver- 
gence and common ancestry (Gosliner and 
Johnson, in press). 

Further work in determining the phylogenetic 
relationships of all genera in the family Chromo- 
dorididae is necessary. The only way to properly 
address questions of mimicry, convergence and 
evolution of color patterns by descent is with a 
testable hypothesis of the relationships between 
species, a phylogeny. 

ACKNOWLEDGMENTS 

Robert Bolland, Mike Miller, and Pauline 
Fiene-Severns kindly provided specimens and 
data for the species studied here. Field work was 
supported by Katharine Stewart, The Christen- 
sen Research Institute, the In-house Research 
fund of the California Academy of Sciences and 
United Airlines. Elizabeth Kools provided valu- 
able technical assistance in the preparation of the 
manuscript and Dong Lin printed the final elec- 
tron micrographs. 

Literature Cited 

Adams, A and L. Reeve. 1 850. Mollusca, Part 3. The 
Zoology of the Voyage of the H.M.S. Samarang 
under the command of Capt. Sir Edward Belcher 
during the years 1843^46. Reeve, Benham and 
Reeve, London. Pp. 45-87, pis. 1 8-24. 

Baba, K. 1949. Opisthobranchia of Sagami Bay. 
Iwanami Shoten, Tokyo. Pp. 1-194, pis. 1-50. 

Bergh.L. S. R. 1874. NeueNacktschneckenderSud- 
see. 2. Journal de Museum Godeffroy 2:91-1 16. 

. 1905. Die Opisthobranchiata der Siboga Ex- 



pedition. Siboga Expeditie Report 50:1-248. 

Gosliner, T. M.andR. Draheim. 1996. Indo-Pacific 
opisthobranch gastropod biogeography: how do we 
know what we don't know? American Malacologi- 
cal Bulletin 12:37-43. 



306 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 12 



GOSLINER, T. M. AND R. F. Johnson. In press. Phy- 
logeny of Hypselodoris (Nudibranchia: Chromo- 
dorididae) with a review of the monophyletic clade 
of Indo-Pacific species, including descriptions of 
twelve new species. Zoological Journal of the Lin- 
nean Society. 

Pease, W. H. 1860. Descriptions of new species of 
Mollusca from the Sandwich Islands. Proceedings 
of the Zoological Society of London 28: 1 8-37. 



1871 



Descriptions of new species of nudi- 
branchiate Mollusca inhabiting Polynesia. No. 2. 
American Journal of Conchology 7: 1 1-19, pis 3-9. 

RlSBEC, J. 1928. Contribution a 1 'etude des nudi- 
branches Neo-Caledoniens. Faune des colonies 
Francaises 2:1-328. 

RUDMAN, W. B. 1982. The Chromodorididae (Opis- 
thobranchia: Mollusca) of the Indo-West Pacific: 
Chromodoris quadricolor, C lineolata and Hyp- 
selodoris nigrolineata colour groups. Zoological 
Journal of the Linnean Society 76:183-241. 

— . 1983a. The Chromodorididae (Opisthobran- 
chia: Mollusca) of the Indo-West Pacific: Chromo- 



doris splendidia, C. aspersa and Hypselodoris 
placida colour groups. Zoological Journal of the 
Linnean Society 78:105-173. 

. 1984. The Chromodorididae (Opisthobran- 



chia: Mollusca) of the Indo-West Pacific: a review 
of the genera. Zoological Journal of the Linnean 
Society 81:1 15-273. 

. 1985. The Chromodorididae (Opisthobran- 



chia: Mollusca) of the Indo-West Pacific: Chromo- 
doris aureomarginata, C verrieri and C. fidelis 
colour groups. Zoological Journal of the Linnean 
Society 83:241-299. 

. 1986. The Chromodorididae (Opisthobran- 



chia: Mollusca) of the Indo-West Pacific: Noumea 
purpurea and Chromodoris decora colour groups. 
Zoological Journal of the Linnean Society 
86:309-353. 

ROppell, E. and F. S. Leuckart. 1 830 or 1 83 1 . Neue 
wirbellose Thiere des Rothen Meeres. Atlas zu der 
Reise im nordlichen Afrika von Eduard Ruppell. Pp. 
1-47, pis. 1-12. 



CALIFORNIA ACADEMY OF SCIENCES, 1998 
Golden Gate Park 
San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 13, pp. 307-314, 5 figs. 



July 20, 1998 



A NEW SPECIES OF DORIOPSILLA (MOLLUSCA, 

NUDIBRANCHIA, DENDRODORIDIDAE) FROM THE PACIFIC 

COAST OF NORTH AMERICA 



By 

Angel Valdes 
and 

David W. Behrens 

Department of Invertebrate Zoology, California Academy of Sciences 
Golden Gate Park, San Francisco, California 941 18 



A new species of the genus Doriopsilla, D. spaldingi, is described on the basis of four specimens 
collected from La Jolla, California. This new species is characterized externally by an iridescent 
blue band around the mantle margin. Internally, D. spaldingi differs from other eastern Pacific 
yellow species in lacking a pyloric gland, having a very long vagina, an elongate, almost tubular, 
prostate and in the penial hooks morphology. Additional information on the distribution and 
natural history of this species is provided. Thus far D. spaldingi is only known from southern 
California and the northwestern extreme of Mexico. 

Received February 4, 1998. Accepted March 1 1, 1998. 



Until recently, seven nominal species from the 
family Dendrodorididae were known along the 
eastern Pacific coast (Behrens 1991). However, 
Gosliner, Schaefer, and Millen (in press) have 
recently reviewed the systematic status of the 
yellow species of Doriopsilla from this area, and 
they concluded that only two species are valid. 

The present paper describes a third yellow 
species of the genus based on material recently 
collected from the California coast, and depos- 
ited in the Department of Invertebrate Zoology 
of the California Academy of Sciences (CASIZ). 

Species Description 

Doriopsilla spaldingi sp. nov. 
(Figs. 1-5) 

Type Material. — Holotype: CASIZ 1 12492, La 
Jolla Submarine Canyon, San DiegoCounty, Califor- 



nia, April 1996, one specimen 48 mm preserved 
length, collected by Michael D. Miller. Paratypes: 
CASIZ 112493, La Jolla Submarine Canyon, San 
Diego County, California, 10 August 1997, 56 m 
depth, three specimens 23-31 mm preserved length, 
two of them dissected, collected by George E. Spald- 
ing. 

External Morphology. — Living animals 
reach up to 85 mm in length. The general color 
of the living animals is creamy white to yellow- 
orange (Fig. 1), the hyponotum being nearly 
translucent cream. The mantle margin is edged 
by an iridescent blue band. The rhinophores and 
gill are yellow, paler in the cream specimens. 

The body is oval (Figs. 1 , 2), high, stiffened by 
a subepidermal network of strong spicules over 
the entire body surface. The dorsum is covered 
by a number of low, simple conical tubercles, 
stiffened with spicules. Tubercles medial on the 



[307] 



308 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 13 



dorsum are larger, decreasing in size toward the 
borders of the mantle. The mantle margin is wide 
and slightly undulate compared to species of 
Dendrodoris. The rhinophores are perfoliate 
with 20 lamellae. The gill is composed of 4-5 
bipinnate leaves. The anus is eccentric to the left. 

Ventrally (Fig. 3), the border of the mantle is 
edged by an iridescent blue band. In some speci- 
mens similar pigmentation exists on the edge of 
the foot, as well. The oral tentacles are small, 
separate and grooved laterally. The anterior bor- 
der of the foot is notched. 

Internal Anatomy. — The buccal bulb is 
oval (Figs. 4A; 5A, B), covered by minute, rather 
undifferentiated oral glands on its proximal por- 
tion (Fig. 4B). The tubular esophagus leads from 
the buccal bulb. At this point two retractor mus- 
cles insert onto the posterior of the bulb. The 
esophagus is very long and convoluted (Fig. 5B). 
Posteriorly, it broadens into a short muscular 
portion, which has two retractor muscles at- 
tached (Figs 4C, 5B). The intestine runs posteri- 
orly in the usual position and lacks any pyloric 
gland (Fig. 5C). 

The ampulla is large and elongate (Figs. 5D, 
E). It divides into a short oviduct, which enters 
the female gland, and the prostate. The prostate 
is elongated, almost tubular, but slightly flat- 
tened. From its distal end. the prostate leads into 
an elongated and convoluted deferent duct which 
is almost as wide as the prostate. The penis, when 
everted, is very long and contains 16 rows of 
penial hooks. The penial hooks are approxi- 
mately 70-80 um wide at the base and up to 
40^45 um in length (Fig. 4D). The vagina is very 
long and convoluted. At its proximal end is a 
large, thin-walled, spherical bursa copulatrix. 
The seminal receptacle is small, having a short 
duct which joins the vagina at the point where it 
connects the bursa copulatrix. The uterine duct 
also emerges from this point. 

The buccal ganglia lay just behind the central 
nervous system. 

The circulatory system consists of a large heart 
(Fig. 5A), joined by the aorta with a flattened 
blood gland, placed behind the central nervous 
system. 

ETYMOLOGY. — The name spaldingi was cho- 
sen to recognize George E. Spalding III of Solana 
Beach, California, who originally discovered this 
species. 



DISTRIBUTION. — Thus far this species is 
known only from La Jolla Submarine Canyon 
(present study) and Point Loma (photographs by 
Mike Miller, Herb Gruenhagen and Bob Bayer) 
in San Diego County. This species is also present 
in San Miguel Island, the Santa Barbara Channel 
Islands and South Coronado Island, Baja Califor- 
nia, Mexico (photographs by Marc Chamber- 
lain). 

Natural History. — This species has al- 
ways been found on sandy to rocky substrate, at 
depths from 27 to over 60 m. Careful temperature 
records kept with numerous specimens collected 
indicate that in most instances this species prefers 
water of 15°C or lower (G. Spalding, pers. 
comm.), explaining its deep water occurrence. 
Several specimens have been observed under 
warmer conditions of surface downwelling (G. 
Spalding, pers. comm.). 

The egg mass is a typical, flat ribbon coil 
attached to the substrate on its side (J. Lance, 
pers. comm.), rather than on one edge, as re- 
ported by Gos liner et al. (in press) for Doriopsilla 
albopunctata. The egg mass of D. spaldingi is 
similar to that of the new species these authors 
describe, which is also classified as type B (Todd 
1983). The larvae exhibit direct or lecithotrophic 
development (J. Lance, pers. comm.). 

Discussion 

According to the recent diagnosis of the genus 
Doriopsilla published by Valdes and Ortea 
(1997), Doriopsilla spaldingi clearly belongs to 
this genus. The main generic diagnostic features 
of this species are the mantle stiffened by spic- 
ules, dorsum covered by tubercles, anus eccentric 
to the left, oral tentacles reduced, digestive sys- 
tem lacking ptyaline glands and penis eversible 
with numerous internal hooks. Valdes and Ortea 
(1997) indicated that Eliot (1906) was likely 
incorrect stating that in Doriopsilla the buccal 
ganglia lie immediately behind the rest of the 
central nervous system. However, the present 
description, as well as that by Gosliner et al. (in 
press) for the species they have studied, contra- 
dicts Valdes and Ortea (1997) and supports 
Eliot's observations. 

The two other yellow species of Doriopsilla 
recognized by Gosliner et al. (in press) from the 
Pacific coast of North America are clearly distin- 



VALDES AND BEHRENS: DORIOPSILLA 



309 




Figure 1 . Living animals of Doriopsilla spaldingi sp. nov. A. Yellow-orange color phase specimen from La Jolla Canyon, 
California, 50 m depth. B. Creamy white color phase specimen from La Jolla Canyon, 50 m depth. Photographs by Mike Miller. 



310 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 13 






yJSf&fSpg&j- 



msm 






**fc 



^'#-=>?i.V!S 



'•??■: •=■*.• ••?■: 



' mm 



y&Zxr 



M 



a ft -J 5 • » . 



^^ 



«..-. o : .p- 



'*'• 



&K 



KH* r fc'. ics 



&^' 



PPM 






SaflpB 



»' 



qp 



iw$si\ ! r>?'G:i>.xx 



« 6. # 



»i 



%>"» 




••■» •;';■.•'. V s ,-#'0 




V ■ •£"':■ 



■ 



y*to 



itt: !*••.■•.;■•■<> .'V;. V** 






•'«&:•; i 



*■ :^ifi>-. ?■%*;■* 



£**& 



!$$&£&> 



&£ 



M 



fcX^; ^Ml 






Figure 2. Doriopsilla spaldingi sp. nov., dorsal view of a living animal from Point Loma, scale bar = 8 mm. 




Figure 3. Ventral view of the anterior border of the foot of Doriopsilla spaldingi sp. nov. (C ASIZ 1 1 2493 ). 



VALDES AND BEHRENS: DORIOPSILLA 



311 




Figure 4. Scanning electron micrographs of Doriopsilla spaldingi sp. nov. (CAS1Z 1 12493). A. Buccal bulb, scale bar = 
1.5 mm. B. Detail of the oral glands, scale bar = 100 um.C. Muscular esophageal region, scale bar = 1.5 mm. D. Penial hooks, 
scale bar= 60 urn. 



312 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 13 






am4 


\\y' : - 


■#M \ 


sr 




\l\ \ 








f- 





Figure 5. Anatomy of Doriopsilla spaldingi sp. nov. (CASIZ 1 12493), scale bar = 3 mm. A. Dorsal view of the anatomy. 
B. Anterior portion of the digestive system. C. Intestine. D. General view of the reproductive system. E. Dissected reproductive 
organs. Abbreviations: a = aorta, am = ampulla, b = buccal bulb, be = bursa copulatrix, bg = blood gland, d = digestive gland, 
dd = deferent duct, e = esophagus, f = female gland, h = heart, i = intestine, m = muscular esophageal region, ot = oral tube, 
pr = prostate, r = retractor muscle, rs = renal sac, sr = seminal receptacle, v = vagina. 



VALDES AND BEHRENS: DORIOPS1LLA 



313 



guished from Dohopsilla spaldingi for their ex- 
ternal morphology and anatomy. Doriopsilla al- 
bopunctata and the unnamed species have the 
dorsum ornamented with white spots, which are 
absent in D. spaldingi, and lack the iridescent 
blue line around the mantle margin of this latter 
species. No other described species of the family 
Dendrodorididae has a color pattern similar to D. 
spaldingi. 

Internally, Doriopsilla spaldingi is charac- 
terized by the presence of an elongate, almost 
tubular prostate, which is different from the wide 
and flattened prostates of other species of the 
genus (see Valdes and Ortea 1997; Gosliner et 
al., in press). In addition. D. spaldingi differs 
from D. albopunctata and the other Californian 
unnamed species in the short length of the semi- 
nal receptacle duct, in lacking a pyloric gland, in 
having a long vagina and in the penial hook's size 
and shape. According to Gosliner, Schaefer and 
Millen (in press), there are 6 rows of penial hooks 
in their unnamed species and 16 rows in D. 
albopunctata, as in D. spaldingi. They also re- 
port the penial hooks of both their species to be 
very elongated with a short base (about 25-35 
um wide and 50 urn long in D. albopunctata, and 
10-15 jam wide and 15-20 um long in their un- 
named species), whereas in D. spaldingi they are 
elongated but the base is longer than the cusp 
(70-80 um wide and 40^45 um long). 

The other three nominal species of Doriopsilla 
present on the Pacific coast of North America are 
Doriopsilla nigromaculata (Cockerell in Cock- 
erell and Eliot, 1905), Doriopsilla rowena Mar- 
cus and Marcus, 1967 and Doriopsilla janaina 
Marcus and Marcus, 1967. Doriopsilla rowena 
is probably a junior synonym of D. nigromacu- 
lata (see Behrens 199 1 ), but anatomical compari- 
sons between specimens from San Diego, 
California and Puerto Penasco, Mexico (the type 
localities of both nominal species) are needed to 
confirm this point. These three species differ 
from D. spaldingi in coloration and anatomy. 
Doriopsilla janaina is light red, orange or yel- 
low with white dots (Marcus and Marcus 1967a), 
and D. nigromaculata and D. rowena are yel- 
lowish to orange with brown and white spots 
(Marcus and Marcus 1967b; Behrens 1991). 
Both species lack any blue pigment. In addition. 



the prostate of D. rowena and D. janaina is rather 
flattened instead of tubular, as in D. spaldingi. 

ACKNOWLEDGMENTS 

The authors would like to recognize the gener- 
ous support of several individuals. Mike Miller. 
George Spalding, Marc Chamberlain, Herb 
Gruenhagen, Nick Galluzzi, Bob Bayer and Jim 
Lance provided specimens or photographic re- 
cords of the species here described. Mike Miller 
permitted us to use his photographs for prepara- 
tion of color plates. Darrell Ubick prepared the 
samples using the critic point technique for SEM 
examination, and Dong Lin printed the scanning 
electron micrographs. Terry Gosliner provided 
unpublished information on the yellow species 
of Doriopsilla from California, and reviewed the 
manuscript. 

This paper has been supported in part by the 
Ministerio de Educacion y Cultura of Spain 
(SEUI), through its postdoctoral fellowships pro- 
gram. 

R.ESUMEN 
Se describe una nueva especie del genero 
Doriopsilla, D. spaldingi, en base a cuatro ejem- 
plares recolectados en California. Esta nueva 
especie se caracteriza externamente por la pre- 
sencia de una banda azul iridiscente alrededor del 
borde del manto. Internamente, D. spaldingi se 
diferencia de otras especies amarillas del Paci- 
fico Este en que carece de glandula pilorica. en 
que posee una vagina muy larga, una prostata 
alargada, casi tubular, y en la morfologia de las 
espinas peneales. Se incluye ademas informacion 
adicional sobre la distribucion e historia natural 
de esta especie. Hasta el momento, D. spaldingi 
se conoce solamente en el Sur de California y del 
extremo noroeste de Mexico. 

Literature Cited 

Behrens, D. W. 1991. Pacific coast nudibranchs — 
a guide to the opisthobranch Alaska to Baja Califor- 
nia. Sea Challengers, Monterey, California. 1 07 pp. 

Eliot, C N. E. 1906. The genus Doriopsilla Bergh. 
Journal of Conchology 1 1:366-367. 



314 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 13 



GOSLINER, T. M., M. C. SCHAEFER AND S. V. MlLLEN. 
In press. A new species of Doriopsilla (Nudibran- 
chia: Dendrodorididae) from the Pacific coast of 
North America, including a comparison with 
Doriopsilla albopunctata (Cooper, 1863). Veliger. 

Marcus, Ev. and Er. Marcus. 1967a. American 
Opisthobranch Mollusks. Part 1 : Tropical American 
Opisthobranchs. Studies in Tropical Oceanography 
6:1-137. 

1967b. American Opisthobranch Mollusks. 



Todd, C. D. 1983. Reproductive and trophic ecology 
of nudibranch molluscs. Pp. 225-255 in The Mol- 
lusca, VI - Ecology. Russell-Hunter ed. Academic 
Press, New York. 

VALDES, A. AND J. ORTEA. 1 997. Review of the genus 
Doriopsilla Bergh, 1880 (Gastropoda: Nudibran- 
chia) in the Atlantic Ocean. Veliger 40:240-254. 



Part 2: Opisthobranchs from the Gulf of California. 
Studies in Tropical Oceanography 6:141-248. 



CALIFORNIA ACADEMY OF SCIENCES, 1998 
Golden Gate Park 
San Francisco, California 941 18 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Vol. 50, No. 14, pp. 315-345, 19 figs., 3 tables. 



September 10, 1998 



REVIEW OF SOUTH ASIAN SISORID CATFISH GENERA 

GAGATA AND NANGRA, WITH DESCRIPTIONS OF A NEW 

GENUS AND FIVE NEW SPECIES 



By 

Tvson R. Roberts 



- Hole Oceanographic institution 



SEP 2 3 1998 



Affiliated Researcher. Smithsonian Tropical Research Institute 

and Wooas H 



Carl J. Ferraris, Jr. 

Department of Ichthyology, California Academy of Sciences 
Golden Gate Park, San Francisco, California 941 18 



The sisorid catfish genera Gagata and Nangra are endemic to southern Asia. The greatest 
concentration of species of these genera occurs in the Ganges basin, including four species of 
Gagata, all named previously, and four of Nangra, three of which are newly described here. Gagata 
gagata (Hamilton), G. sexualis (Tilak), G. youssoufi (Ataur Rahman), Nangra bucculenta n. sp., 
V. carcharhinoides n. sp., and N. ornata n. sp. are known only from the Ganges basin, while Gagata 
cenia (Hamilton) and Nangra nangra (Hamilton) also occur in the Indus basin, along with the 
endemic Nangra robusta IMizra and Awan. Gagata itchkeea (Sykes) is know n only from peninsular 
India. Two species of Gagata from Myanmar, previously identified as Gangetic species are 
described as (7. melanopterus, n. sp., from the Irrawaddy, Rangoon, Sittang, and lower Salween 
basins, and G. gasawyuh, n. sp., from the Irrawaddy, Salween and Tenasserim basins. The 
Gangetic species previously reported as Gagata (or Nangra) viridescens (Hamilton) does not 
belong in either of these genera and is placed in a new genus, Gangra. Once this monotypic genus 
is recognized, Nangra and Gagata are readily diagnosed as natural groups of the Nangrina. Nangra 
punctata Day, 1877, is a junior synonym of Gangra viridescens. A neotype is designated for Nangra 
nangra to stabilize the name according to current usage. 

Received September 27, 1997. Accepted April 7, 1998. 



Asian catfishes of the family Sisoridae have 
undergone an impressive radiation and include 
some of the most highly modified catfishes found 
in tropical Asia. Even relatively generalized sis- 
orids have highly specialized osteological fea- 
tures. With the notable exception of Mahajan's 
(1963-1967) excellent series of papers on Sisor 
rabdophorus Hamilton, 1 822, and Tilak's (1963) 
comparative study, relatively little work has been 
done on sisorid osteology until quite recently. 
Consequently, many sisorid genera are poorly 
delimited and difficult to identify. The South 
Asian genera Gagata Bleeker, 1 858, and Nangra 



Day, 1877, have been associated in ichthyologi- 
cal works since 1911, when they appeared as the 
first two genera in a dichotomous key to sisorid 
genera (Regan 191 1). They are closely related, as 
confirmed by de Pinna (1996). despite pro- 
nounced differences in overall appearance. 
Herein, we present revised diagnoses of Gagata 
and Nangra, review the included species and 
describe five new species. We propose a new 
generic name, Gangra, for Pimclodus 
viridescens Hamilton. 1822. Contrary to pre- 
vious studies, that species does not appear to be 



315] 



316 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 



more closely related to species of either Gagata 
or Nangra. 

Gagata, Nangra, and Gangra share a number 
of derived characters for sisorids that indicate the 
formation of a natural group. De Pinna (1996) 
uncovered six characters that were diagnostic of 
a group consisting only of species of these gen- 
era. At least one species of each of the three 
genera also have a nearly complete bony encap- 
sulation of the swim bladder by the parapophyses 
of the 4th and 5th vertebral centra. Anterior and 
posterior medial cranial fontanels are present in 
all species of the three genera, but the posterior 
fontanel is greatly reduced in Gangra viridescens 
and Gagata itchkeea (Sykes, 1839). Gagata and 
Nangra (but not Gagata itchkeea) have paired 
cranial fontanels. In addition, these three genera 
share a number of characteristics that are summa- 
rized here. The dorsal fin usually has six soft rays 
(up to nine in some Nangra) and the dorsal-fin 
spine is not serrated. The pelvic fin has one 
unbranched and five branched rays. The caudal 
fin is more or less deeply forked, and has 17 
principal rays. 

This study resulted from our independent 
fieldwork and research on freshwater fishes of 
South and Southeast Asia. We each obtained 
specimens of Gagata during fieldwork in Myan- 
mar and initially identified them as G. gagata 
(Hamilton, 1822) and G.cenia (Hamilton, 1822). 
Comparison of Gangetic material of G. gagata 
and G. cenia with our specimens from Myanmar 
quickly revealed that the specimens from Myan- 
mar represented undescribed species. The genus 
Nangra was also well represented in recent col- 
lections made by the first author in India and 
Bangladesh, including Nangra nangra (Hamil- 
ton, 1 822) and two new species, bringing the total 
number of Nangra species to five (three endemic 
to the Ganges drainage, one to the Indus, and 
Nangra nangra reported from both). Also in- 
cluded in samples from India and Bangladesh 
were numerous specimens of a species identified 
by various authors as Gagata (or Nangra) 
viridescens, treated here as the sole repre- 
sentative of our new genus Gangra. 

Materials and Methods 

Materials examined are deposited in the fol- 
lowing institutions: AMS, Australian Museum, 
Sydney; AMNH, American Museum of Natural 



History, New York; BMNH, The Natural History 
Museum, London; CAS and CAS(SU), Califor- 
nia Academy of Sciences, San Francisco; MCZ, 
Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge; MZUSP, Museu de Zoolo- 
gia da Universidade de Sao Paulo, Sao Paulo; 
NRM, Swedish Natural History Museum, Stock- 
holm; RMNH, Rijksmuseum van Natuurlijke 
Historic Leiden; SRS/ZSL Southern Regional 
Station of the Zoological Survey of India, Ma- 
dras; UMMZ, University of Michigan Museum 
of Zoology, Ann Arbor; and USNM, National 
Museum of Natural History, Smithsonian Insti- 
tution, Washington. 

Specimen measurements are given as standard 
length. Specimens were prepared for skeletal 
examination following Taylor and Van Dyke 
(1985). Most observations of teeth and gill rakers 
were made from cleared and stained specimens. 

Gill raker counts include all elements on the 
outer edge of the first gill arch that have a bony 
core. Counts are given as a range for upper limb, 
separated by a plus (+) sign from those of the 
lower limb. Vertebral counts include four ele- 
ments for the Weberian complex (the fifth cen- 
trum is sutured to the complex centrum, but 
recognizable) and the hypural complex counted 
as a single element. The posterior-most abdomi- 
nal vertebra is defined here as the last vertebra 
with its hemal spine lying entirely anterior to the 
anterior-most pterygiophore of the anal fin. All 
vertebrae posterior to this are referred to as 
postabdominal, following Roberts (1983). We 
begin a count just after the Weberian apparatus, 
either with vertebra 5 (identified by its greatly 
enlarged parapophysis) or vertebra 6 (identified 
as the first rib-bearing vertebra). 

Species distributions are plotted on Lambert 
azimuthal equal area projections of South and 
Southeast Asia. Country boundaries are repre- 
sented by dotted lines; rivers by solid lines. Riv- 
ers of Southeast Asia have been omitted. Plotted 
distributions represent specimens actually exam- 
ined, along with unconfirmed literature reports 
for which the stated locality was sufficiently 
precise to place on the map. 

The extensive synonymies listed in Day 
(1877), Hora and Law (1941) and Misra ( 1976) 
are not repeated here. 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



317 



Key to Genera of the Nangrina 
Di Pinna, 1996 

la. Pectoral girdle with rugose coracoid proc- 
ess covered with thin skin; outer and inner 
mental barbels widely separated, origin of 
inner mental barbels anterior to origin of 
outer mental barbels; lateral cranial fonta- 
nel absent; live specimens with viridescent 

or silvery supraopercular mark 

Gangra new genus 

lb. Pectoral girdle with coracoid process not 
visible externally; outer and inner mental 
barbels close together, their origins in a 
transverse row; lateral cranial fontanel 
generally present (absent in G. itchkeea); 
supraopercular mark absent 2 

2a. Body terete; head depressed, eye dorso 
lateral; snout elongate, depressed; maxil- 
lary barbel extends at least to pelvic-fin 
base; palatal teeth present; premaxilla ex- 
panded, firmly attached to correspondingly- 
expanded ventral plate of mesethmoid; 
caudal-fin base usually with a vertically 
elongate elliptical mark Nangra Day, 1877 

2b. Body compressed; head compressed, eye 
lateral; snout compressed, short; maxillary 
barbel extending only to level of pectoral- 
fin base; palatal teeth absent; mesethmoid 
strongly projecting anteroventrally, without 
lateral processes or expanded ventral plate; 
caudal-fin base without elliptical mark . . 
Gagata Bleeker. 1858 

Systematic Accounts 
Gagata Bleeker, 1858 

Gagata Bleeker, 1 858:204, 206 (Type species: Pi- 
melodus gagata Buch. [= Hamilton, 1822], by ab- 
solute tautonymy). Gender masculine. 

Callomystax Giinther, 1864:218 (Type species: Pi- 
melndus gagata Hamilton, I 822; unneeded replace- 
ment name for Gagata Bleeker, 1858). Gender 
masculine. 

DIAGNOSIS. — Summaries of counts of verte- 
brae, fin rays, and gill rakers are presented in 
Tables 1-3. Head and body compressed; eyes 
large, lateral; mouth inferior, relatively small and 
narrow; mesethmoid bone highly modified, 
strongly curved downward in front of snout, with 
laterally compressed, ventromedial! projection to 



which ascending process of premaxilla attached 
only by soft tissue; premaxilla with dorsomedial 
ascending process; jaw teeth finely conical, in 
few rows, absent from upper jaw in some species; 
palate toothless; branchiostegal membrane 
broadly joined to isthmus; dorsal fin with 6 
branched rays; caudal fin with 17 principal rays; 
pectoral girdle with coracoid process covered 
with thick skin and not visible externally. 

Comments. — Although the species of Ga- 
gata are similar in overall appearance and share 
a number of derived characteristics that indicate 
that they form a natural group, they exhibit great 
variation in the structure of the bony capsules of 
the swim bladder. Tilak (1963) described and 
illustrated the complex, somewhat incomplete, 
capsule of Gagata gagata. In that species, the 
swim bladder has a large dorsal bony roof and an 
incompletely ossified floor with finger-like proc- 
esses, but is otherwise uncovered by bone. In 
Gagata itchkeea bony investment of the swim 
bladder consists only of a large dome-like roof 
over the dorsomedial half of the swim bladder. 
The other species for which we have examined 
cleared and stained specimens have complete 
bony capsules. That is, the lateral wing of the 
swim bladder is entirely enclosed in a bony tube 
except at its open distal end which abuts the 
membranous tympanum. De Pinna (1996. fig. 
24) provided an excellent illustration of the fully 
encapsulated swim bladder of Gagata gasawyuh 
n. sp.. under the name G. gagata. In the two 
species which do not exhibit fully encapsulated 
swim bladders. G. gagata and G. itchkeea, the 
swim bladder and the tympanum are proportion- 
ally much larger than in the other species of 
Gagata. 

The seven species of Gagata appear to repre- 
sent an extraordinary instance of progressive re- 
duction of jaw teeth and gill rakers, possibly 
related to suctorial, fine particle detritivory. 
Maximum development of teeth and rakers oc- 
curs in G. gagata and G. melanopterus n. sp. In 
these two species, the upper and lower jaws have 
fine, close-set teeth for their full extent and rela- 
tively numerous, long, gill rakers on all gill 
arches. The first gill arch has rakers on the lead- 
ing and trailing edge in G. gagata but only on the 
leading edge only in G. melanopterus; the second 
through fourth arches have rakers on leading and 
trailing edges; and the fifth arch has rakers on the 



318 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 



— r~- in 



r- m v> 



— lo m — 



r-\ ON 



v© v~> rn 



6c 



ir> . — i/~> 'rj- r<"> 



2 

&C 



in rj m — ■— ' 



^ n «i n 



■^j- rn rio^-i^(N'-^ | n^r 



oo — — 



Q 


a 

6C 


a 

to 


a 


a 


a 


tr 


6C 


60 


a 







•-: c to 



o o o ^ o 



-Si ~ ~ 



<£* .^ ^ ,-« 



a 


-5 
a 


5 


a 




& 


5c 


5 


a 


? 



?\ ~ C ^J 



O ^ O ^ ^ ^ ^ 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



319 



tC o 



— « <n m rn —i ^ 



Tt OO ^ rf 



OO fS <N — ' 



— '<N<NW->— . en "^ "^ ° C> ^ vO 



i vo 



g~ 






•O m Tt lO 



wo 



■a -*= 



§33 









5 ?§ 

i3 Q^ 



a Q ?3 -s: 

a =* S 3 Q SJ 






S 5 

- v 5 c 



be 



bo 



I 



a 



o 



— i (N W-i — i 



— (N — ' — 



8 3 8 

3 to <i O 

.2 -s: fii t~ ^s u -a 



22 



320 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 



leading edge. Except for the somewhat larger 
rakers on the leading edge of the first gill arch, 
rakers on the remaining arches are nearly of the 
same size and number. The least development of 
jaw teeth and gill rakers occurs in G. youssoufi. 
In this species the jaws are edentulous, except for 
a few tiny teeth near the symphysis of the lower 
jaw. The leading edge of the first gill arch has a 
few tiny gill rakers, but its trailing edge and the 
remaining gill arches are entirely without rakers. 
This reduction in jaw teeth and gill rakers is not 
accompanied by a corresponding decrease in 
pharyngeal teeth. Teeth in the upper and lower 
pharyngeal tooth plates are well developed 
throughout the genus. The condition of the jaw 
teeth and gill rakers is intermediate in G. cenia 
and G. gasawyuh n. sp. These species have no 
upper jaw teeth, reduced lower jaw dentition, and 
fewer gill rakers than in G. gagata and G. 
melanopterus . 

Gagata youssoufi (and two specimens identi- 
fied as G. sexualis by Tilak 1970) exhibit another 
interesting modification of the gill arches. A 
single row of 8 to 10 slender teeth is present on 
the fourth gill arch. These teeth, which are re- 
stricted to the middle of the lower limb of the 
fourth gill arch, oppose the upper pharyngeal 
tooth plate and appear to be associated directly 
with the ceratobranchial. De Pinna (1996) re- 
ported the presence of tooth plates on the fourth 
ceratobranchial of Bagarius, another genus of 
sisorid catfish. In the specimens of Bagarius 
reported on by de Pinna, as well as those exam- 
ined by us (CAS 96580), the dentition of the 
fourth ceratobranchial is different from that of 
Gagata youssoufi. The teeth in Bagarius are lo- 
cated in a cluster, in an elevated part of the bone 
that appears to be a tooth plate that is fused to the 
underlying ceratobranchial. In Gagata youssoufi 
the teeth appear to be ankylosed directly to the 
ceratobranchial element and the teeth form a 
single row. 

NOTE ON GENDER. - The gender of Gagata 
was not indicated by Eschmeyer and Bailey (in 
Eschmeyer 1990:154), and has not been clearly 
established before now. Gagata is a Bengali 
name presumably of masculine gender. The first 
use of a Latin species name in connection with 
Gagata in which gender is unequivocal is that of 
Bleeker ( 1 863). Bleeker proposed the name Ga- 
gata typus as a replacement for Gagata gagata 



(Hamilton), a combination that he used when he 
first proposed Gagata as a generic name (Bleeker 
1 858). As with Gagata gagata, virtually all of the 
species names originally placed by Bleeker in 
Gagata were names Hamilton (1822) adopted 
from Bengali. Three species names placed in 
Gagata that were Latinized: Gagata pusillus, G. 
anisurus and G indicus, were only tentatively 
included in the genus, and subsequently placed 
elsewhere. As with Gagata typus, these names 
are also masculine. We follow the precept that 
gender of names that are not of Greek or Latin 
origin is determined by a species name of un- 
equivocal gender (ICZN 1985, art. 30d), and treat 
Gagata as masculine following Bleeker (1863). 
However, Sundagagata Boeseman, 1966. is 
treated as feminine. 

Key to Species of Gagata 

la. Dorsum of body with 4 or 5 dark saddles; 
caudal fin with dark markings 4 

lb. Body without dark saddles, although faint 
saddles sometime present; caudal fin im- 
maculate, or with fine black margin 2 

2a. Pectoral, pelvic, and anal fins not black dis- 
tally; dorsal fin with filamentous extension 
Gagata dolichonema He, 1996. 

2b. Pectoral, pelvic, and anal fins black distally; 
dorsal fin without filamentous extension . . 3 

3a. Dorsal fin relatively large, extending at least 
to adipose-fin origin when depressed; pelvic 

fin reaches anal-fin origin 

G. gagata (Hamilton, 1822) 

3b. Dorsal fin relatively small, falling far short 
of adipose fin when depressed; pelvic fin 

does not reach anal-fin origin 

G. melanopterus, new species 

4a. Caudal fin with single dark submarginal lu- 
nate mark: head and body with four saddles 
G. gasawyuh, new species 

4b. Caudal fin with square or round spot on each 
lobe; head and body with five saddles .... 5 

5a. Snout tip acutely pointed in lateral profile, 

with distinct notch anteriorly 

G. cenia (Hamilton, 1822) 

5b. Snout tip broadly rounded in lateral profile, 
without notch 6 

6a. Dorsal fin with filamentous extension in 
males G. sexualis Tilak, 1970 

6b. Dorsal fin without filamentous extension . 7 



ROBERTS AND FERRARIS: GAG AT A AND NANGRA REVIEW 



321 



7a. Anal fin with 8 to 10 branched rays; body 

depth equal to head length 

G. itchkeea (Sykes, 1839) 

7b. Anal fin with 11 to 14 branched rays; body 

depth much less than head length 

G. youssoufi Ataur Rahman. 1976 

Gagata cenia (Hamilton. 1822) 
(Figs. 1.2) 

Pimelodus cenia Hamilton 1 822: 1 74, 376, pi. 3 1 , fig. 
57 (type locality: rivers in the northern part of 
Bengal). 

Gagata cenia, Day 1 877:492 (partim; specimens illus- 
trated by Day as G. cenia are here reidentified as G. 
gagata and G gasawyuh). Hora and Law 1 94 1 :2 1 , 
pi. 1, figs. 5, 6. Jayaram 1981:240 (partim, exclud- 
ing Burma), Ataur Rahman 1990:220, fig. 130 
(Bangladesh). 

Nangra viridescens, Day 1877:494, pi. 1 15, fig. 7. 

MATERIAL EXAMINED. — 1 59 specimens, 26-80 
mm. BANGLADESH. CAS 95540 (3, 41^8 mm), 
Tangail District, Gala Khal, 02-08-92 [sic]. CAS 
95541 (4, 33-41 mm), Ganges basin, Sylhet district, 
Kushiara River at Saidpur, 16 May 1996, T. R. 
Roberts. CAS 95542 (20, 40-55 mm), Ganges basin. 
North Central Region, Tangail district. 1992, Bangla- 
desh Govt. 

INDIA. BMNH 1858.8.15:116-117 (2, 41-46 
mm), Ganges, Waterhouse. BMNH 1870.7.12:4 (1, 
not measured), Orissa, F. Day. BMNH 1 889.2. 1 :261 3 
(1. 33 mm), Calcutta, F. Day. BMNH 1889.2.1:2623 
(1, 66 mm), Poonpore River, F. Day. BMNH 
1889.2.1:2624-2625 (9, 39-43 mm), Sind, F. Day. 
BMNH 1889.2.1:2619-2622 (4, 56-80 mm), R. 
Jumna, F. Day. BMNH 1889.2.1:2616-2618 (3, 
41-55 mm), Darjeeling, F. Day. BMNH 
I 889.2. 1 :2626-263 1 (5, 42-49 mm), Orissa, F. Day. 
BMNH 1954.5.20:19-20 (2, 51-52 mm), Hooghly 
River (West Bengal) at Tribence Ghat, 8-1 1 -53 [sic], 
A. K. Datta. CAS 54538 (1, 43 mm), Assam State, 
Tezpur, B. Prashad. CAS 95539 (78, 36-72 mm), 
Ganges River at Patna, April-May 1996, T. R. 
Roberts. CAS(SU) 34841 (1, 59 mm), Bengal State, 
Hugh [= Hooghly] River, 11 April 1937, A. Herre. 
MCZ 4258 (4of 9:48-51 mm), Uttar Pradesh, Alla- 
habad, F. Day. 

PAKISTAN. BMNH 1872.1.30:4 (10, 28-39 mm), 
Indus, Beavan. CAS 24248 ( 1 0, 26-35 mm), Sind, 325 
miles north of Karachi (i.e., 5 miles north of Sukkur), 
1 Nov. 1968, E. S. Herald and party. 

Diagnosis. — A small species of Gagata ma- 
turing at less than 70 mm and reaching to about 
100 mm. Dorsum of body with dark saddles ex- 



tending ventrally only to lateral line. Caudal fin 
with transverse black bar across peduncle and 
round or square black spot on middle of each 
lobe. Dorsal fin with black spot on distal part of 
anterior rays. Upper jaw toothless; lower jaw 
with few small conical teeth in pocket or depres- 
sion near symphysis. Fourth ceratobranchial 
without teeth. Snout tip acutely pointed in lateral 
profile, tip separated from rest of snout by dis- 
tinct notch. 

COUNTS. — Dorsal fin with spinelet. spine and 
6 (rarely 7) branched rays; pectoral fin with spine 
and 8 (rarely 9) rays; anal fin with 4 or 5 simple 
rays and 9 to 1 1 branched rays. Vertebral column 
with 18 to 20 abdominal and 16 to 18 post ab- 
dominal vertebrae, total 35 or 36. First gill arch 
with 5-7 tiny rakers, nearly indistinguishable 
from surrounding tissue. 

DISTRIBUTION. — Gagata cenia is distributed 
widely in the Ganges basin, the Indus River 
system, and the Mahanadi River (Fig. 3). We 
have made a cursory examination of specimens 
from the Indus River, most of which are in poor 
condition, and we were unable to find any differ- 
ences between them and the Gangetic specimens. 
Shrestha (1994) reports this species from Terai, 
Nepal. Therein, G. cenia is reported to reach 100 
mm, which is substantially larger than any speci- 
men examined by us. 

COMMENTS. — There has been considerable 
confusion in identification of G. cenia (Fig. 2). 
This is due. in part, to the relative rarity of Ham- 
ilton's published monograph on Gangetic fishes 
(Hamilton 1822) and to the absence of type speci- 
mens for his species. But the real difficulties lie 
in the work of his successor Francis Day, whose 
interpretations of Hamilton were sometimes in 
error. As an example, Day ( 1 877) published two 
figures of specimens he identified as Gagata 
cenia, one of which was said to be an adult and 
the other a juvenile. The adult is Gagata gagata. 
as previously pointed out by Hora and Law 
(1941). The juvenile appears to be based on a 
specimen of our new species G. gasawyuh (iden- 
tified by diagnostic color features including dis- 
position of five saddle-marks on head and body 
and distinctive lunate mark on caudal fin). The 
specimen illustrated as the juvenile of G. cenia 
reportedly was collected in Delhi. We have not 
been able to examine any specimens of G. cenia 
from that locality, but the caudal-fin color pattern 
is unlike that of any specimen of Gagata cenia 



322 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 




Figure I . Gagata cenia, 47 mm, CAS 95539; India, Ganges River at Patna. 




Figure 2. Gagata cenia, illustration from Hamilton ( 1822), plate 31, figure 57; originally 72 mm. 



that we have examined. In fact, that pattern is 
characteristic of specimens of Gagata gasawyuh ^ 
from Myanmar. and we believe that Day's speci- 
men is most likely from there. 

Gagata dolichonema He, 1996 

Gagata dolichonema He 1996:380, fig. I (type local- 
ity : Daojieba [24°4 l'N, 99° 1 0'E], Baoshan County, 
Yunnan Province, China). 

Material Examined. — None. 

DIAGNOSIS (after He). — A species of Gagata 
with a median longitudinal groove that extends 
to end of occipital process; maxillary barbel 
longer than head; dorsal-fin spine produced into 
long filament; pectoral-fin spine without fila- 
mentous projection; eye diameter shorter than 
snout length; dorsal part of thoracic region of 
body with four black stripes; and isthmus without 
finger-like projections. 



COUNTS. — Dorsal fin with spinelet, spine and 
6 branched rays; pectoral fin with spine and 8 
branched rays; anal fin with 3 simple and 11(10 
mentioned in English summary) branched rays 
(from He, 1996; vertebral formula and gill rakers 
counts not stated). 

DISTRIBUTION. — Known only from the type 
locality, which is in the upper reaches of the Nu 
(= Salween) River drainage (Fig. 3). 

COMMENTS. — Gagata dolichonema is one of 
three species now recognized from the Salween 
River drainage. The two new species described 
in this paper (G. gasawyuh and G. melanopterus) 
are found in lower parts of the Salween River and 
its tributaries (as part of much wider distributions 
for each of these species), whereas G 
dolichonema is known only from much further 
up river. Gagata gasawyuh is readily distin- 
guished from this species by its distinctive color 
pattern, the absence of a filamentous extension 
of the dorsal spine, and in having an orbital 
diameter that is greater than the length of the 
snout. Gagata melanopterus lacks a filamentous 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



323 





70 


80 




90 

> 


100 












O* Gagata cenia 












■ Gagata dolichonema 


30 - 


•J^7 








▲ Gagata gagata 
x ▼ Gagata gasawyuh 






A I Vf 






•'/ 7 1 r 


20 - 








*«&\ j; 


% ):t<fb 














v^> 




j^/___ 


10 - 




W\ 









- 10 



90 



Figure 3. Distribution of Gagata cenia, G dolichonema, G. gagata, and G gasawyuh. Open circles represent unverified 
literature records of Gagata cenia. Distribution of Gagata dolichonema is an approximation, based on stated type locality. 



extension of the dorsal fin and has a distinctive 
color pattern that is not mentioned in the descrip- 
tion of G. dolichonema. In addition to the above, 
both G. gasawyuh and G. melanopterus typically 
have nine branched pectoral-fin rays, rather than 
the eight listed for G. dolichonema. 

Gagata gagata (Hamilton, 1822) 
(Figs. 4, 5) 

Pimelodus gagata Hamilton 1 822: 1 97, 379, pi. 39 fig. 

65 (type locality: fresh water rivers and estuaries of 

Bengal). 
Gagata typits Bleeker 1 863:90 (replacement name for 

Pimelodus gagata Hamilton 1822). 
Gagata cenia, Day 1877 (partim; pi. 1 15, fig. 4 of 

specimen from Delhi); Jayaram 1 98 1 , fig. 1 29 (after 

Day 1877). 
Gagata gagata, Hora and Law 1 94 1 : 1 5, pi. 1 figs. 1-2 

(partim, Ganges and Brahmaputra rivers only); Ti- 

lak 1970:214 (key); Ataur Rahman 1990:218, fig. 

218 (Bangladesh: Meghna River, near Chandpur). 

Material Examined. — 9 specimens, 40-97 mm. 
BANGLADESH. CAS 95543 ( 1 , 40 mm, cleared and 



stained), Ganges basin, Khulna markets, May 1 996, 
T. R. Roberts. 

INDIA. BMNH 1880.2.2:95 (1, 78 mm), Jamu, F. 
Day. BMNH 1889.2.1:2611-2612 (2, 90-97 mm), 
Calcutta, F. Day. BMNH 1858.8.15:113-115 (3, 
85-97 mm), Ganges, Waterhouse. CAS 54541 ( 1, 95 
mm). West Bengal State, Manirampur, Pulta Water 
Works. MCZ 36074 (1, 88 mm), West Bengal, Cal- 
cutta, F. Day. 

DIAGNOSIS. — A large species of Gagata, 
achieving at least 100 mm. Head and body sil- 
very overall, without pattern. Fins, except caudal 
fin, black distally, with clear basal portions; cau- 
dal fin entirely clear. Dorsal-spine tip extending 
well past adipose-fin origin when adpressed. Pre- 
maxilla with four rows of teeth. 

COUNTS. — Dorsal fin with spinelet. spine and 
6 branched rays; pectoral fin with spine and 7 to 
9 (modally 9) rays; anal fin with 5 or 6 simple 
and 1 1 branched rays. Vertebral column with 18 
or 19 abdominal and 19 or 20 postabdominal 
vertebrae, total 38 or 39. First gill arch with 10 
to 13 long, slender rakers. 



324 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 




Figure 4. Gagata gagata, 97 mm, BMNH 1855.8.15:1 13; Ganges River. 




FIGURE 5. Gagata gagata, illustration from Hamilton ( 1822), plate 39, figure 65; originally 162 mm. 



DISTRIBUTION. — Gagata gagata is known 
only from the Ganges basin of India and Bangla- 
desh (Fig. 4). 

COMMENTS. — Identification of the material 
reported here as Gagata gagata is based on the 
description and figure published by Hamilton 
( 1822). reproduced here as Figure 5. Our speci- 
mens agree with the account and illustration of 
Hamilton, except on two points: 1) according to 
Hamilton, the species "grows to about a foot 
= approx. 300 mm] in length, and is pretty com- 



mon both in the fresh waters and estuaries of 
Bengal,'" and 2) the illustration shows a second 
dorsal-fin spine that appears proportionally 
much stouter than that of any of our specimens. 
We suspect the species is no longer as common 
as it was in the early 1800s, which may explain 
why we have not seen specimens, or reports of 
specimens, much larger than 100 mm. Hora and 
Law (1941) examined a specimen from Alla- 
habad that was 102 mm. They also reported on 
specimens up to 143 mm, but from an unknown 



ROBERTS AND FERRARIS: GAGATA AND A 'A NGRA REVIEW 



325 



locality. Among the specimens they examined 
was at least one specimen of Gagata melanop- 
terus, from Myanmar. Thus, the specimens with- 
out associated locality information might be 
either of the two species. The largest specimen 
known to us is one reported by Ataur Rahman 
(1990) of 193 mm from the Gangetic delta re- 
gion. As for the much stouter dorsal-fin spine 
figured in Hamilton (1822), the spine may be- 
come disproportionately stout in large individu- 
als or its stoutness might have been exaggerated 
by the illustrator. We have not been able to 
examine a specimen as large as that illustrated in 
Hamilton, so we cannot choose between these 
two explanations. There remains, of course, the 
possibility that the Gangetic specimens we iden- 
tified as G. gagata are not conspecific with Ham- 
ilton's G. gagata. 

Specimens from The Natural History Museum. 
London, registered under the number BMNH 
1858.8.5:1 13-1 17. are listed as types of Gagata 
gagata. We see no reason to consider these speci- 
mens types, as Hamilton clearly indicated that he 
did not save any specimens (see Hora, 1929, for 
discussion). In addition, the specimens that are 
identified by us as Gagata gagata (BMNH 
1858.8.5:113 115) were mixed together with 
two specimens of Gagata ccnia (now BMNH 
1858.8.5:1 16-1 17), another Hamilton species. It 
is unlikely that Hamilton would have lumped 
together these two readily distinguishable spe- 
cies. The designation of these specimens as types 
can be traced back at least as far as Gunther 
(1864:218), although without explanation of 
why type status was conferred upon them. 

Gagata gagata is most similar to G. melanop- 
terus, n. sp., from Myanmar, in coloration and in 
having the highest gill raker counts among Ga- 
gata species. It differs from that species in the 
length of the dorsal fin: that of G. gagata extend- 
ing posteriorly when depressed beyond adipose 
origin (vs. falling far short of adipose fin), and in 
having four rows of teeth on the premaxilla (vs. 
2 rows in G. melanopterus). The structure of 
bony capsule of swim bladder differs between 
these two species (see discussion in diagnosis of 
Gagata). For further distinctions between G. ga- 
gata and G. melanopterus, see the account of the 
latter species. 

The unusual coloration of Gagata gagata is 
nearly identical to that of G. melanopterus. All 
other species of Gagata have prominent dark 



saddles on the dorsum of the body, and bands or 
spots on the caudal fin. Faint saddles have been 
observed in some specimens of G. melanopterus. 
but not in G. gagata. 

Gagata gasawyuh new species 
(Figs. 6, 7) 

? Gagata cenia. Day 1877, pi. 1 15, tig. 5 (specimen 
purportedly from Delhi). 

Gagata cenia, Vinciguerra 1890:121 (partim; Ran- 
goon, Mandalay, Bhamo); Smith 1945:394 (Thai- 
land: Salween River, Ta Ta Fang) 

Gagata gagata, de Pinna 1996:7 et seq., figs. 3, 15, 
24, 27, 44, 45, 47 (anatomical discussion). 

Material Examined. — 107 specimens, 20-130 
mm. Holotype: CAS 95544, 88 mm, MYANMAR, 
Tenasserim River mainstream upstream from Kita (or 
Htee-tah), gill-nets on steep sand bank, 8-9 March 
1992,T. R. Roberts. 

Paratypes: MYANMAR. Irrawaddy basin. AMNH 
8358 (3, 77-97 mm), Chindwin River, April 1923, B. 
Brown. AMNH 13776 (1, 68 mm), Chindwin River, 
April 1923, B. Brown. CAS 88614 (3, 42-58 mm), 
Mandalay market, 21 April 1996, C.J. Ferraris, D. 
Catania and Myint Pe. CAS 88899 (4, 65-93 mm). 
Myitkyina market, 21 April 1996, C.J. Ferraris, D. 
Catania and Myint Pe. CAS 88906 (22, 20-54 mm), 
Nyaung-U market (near Bagan), 13 April 1996, C. J. 
Ferraris and D. Catania. NRM 26667 (3, 46-99 mm). 
Dhweli Kyaung River, February 1935, R. Malaise. 
CAS 95546 (16, 21-41 mm), Myit-tha River (Pinda 
River) at Pinda Village. Sagaing Division, 
23°H'01"N, 94°05'32"E, 12 November 1996, C.J. 
Ferraris, Myint Pe and village fishermen. CAS 95547 
(6, 53-73 mm), Mandalay Division, Nyaung-U mar- 
ket, 8 Nov. 1996, C. J. Ferraris and Myint Pe. CAS 
95548 (6, 69-102 mm), Mandalay markets, 13-25 
April 1993, T. R. Roberts. NRM 42002 (3, 75-130 
mm), Myitkyina market, 4-8 November 1997, C. J. 
Ferraris. USNM 345150 (2, 76-77 mm). Mandalay, 
1 885, L. Fea. Sittang basin. USNM 345 151(5, 47-85 
mm), Bago Division, Taungoo Market, 7 April 1996, 
C. J. Ferraris and D. Catania. Tenasserim basin. CAS 
95545 ( 1, 99 mm), (taken with holotype) Tenasserim 
River mainstream upstream from Kita or Htee-tah, 
gill-nets on steep sand bank, 8-9 March 1992, T. R. 
Roberts. CAS 95549 (2, 95-108 mm), Tenasserim 
River and tributaries between Kita (or Htee-tah) and 
Baowasung, March 1992, T. R. Roberts. CAS 95550 
(4, 55-64 mm), Tenasserim River mainstream, 3 hrs 
upstream from Hteetah ( 1 st fishing village), seine and 
gillnet in sandbank, 7-8 March 1992, T. R. Roberts. 

THAILAND. Salween basin. BMNH 1992.3.10:1 
( 1 , 89 mm), Salween mainstream, 1 988, T. R. Roberts. 
CAS 9555 1 (6, 58-85 mm), sandy beach and sheltered 



326 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 




Figure 6. Gagala gasawyuh new species, holotype, 
Hee-tah). 



mm, CAS 95544; Myanmar, Tenasserim River above Keta (or 



bay with shallow muddy backwater on Sal ween main- 
stream, about midway between Mae Sam Lap and 
Paleh, 20 March 1 989, T. R. Roberts. CAS 95552 ( 1 2, 
65-98 mm), Sal ween mainstream, 20 km upriverfrom 
Mae Sam Leap, 21 April 1989. T. R. Roberts. CAS 
95553 ( 1 , 73 mm), Salween mainstream, about 5 km 
upstream from mouth of Menam Moei, 18-19 March 
1 989, T. R. Roberts. CAS 95554 (1,58 mm), sheltered 
backwater of Salween mainstream 20 km upstream 
from Mae Sam Leap, 21 April 1989, T. R. Roberts. 
CAS 95555 (2, 75-85 mm) and UMMZ 233234 (2, 
81-87 mm), rocky side channel of Salween main- 
stream about 40 km upriver from Mae Sam Leap, 20 
April 1989,T. R. Roberts. 

DIAGNOSIS. — A large species (exceeding 100 
mm) distinguished from all other species of G ci- 
gala by two color features: four distinct saddles 
on dorsum of body, usually extending ventrally 
on the sides of the body well below the lateral 
line, and dark marks on caudal fin lobes forming 
a single, continuous lunate mark in the middle of 
the caudal fin. 

COUNTS. — Dorsal fin with spinelet, spine and 
6 branched rays; pectoral fin with spine and 9 
(rarely 8 or 10) rays; anal fin with 4 to 6 simple 
rays and 10 or 11 (rarely 9 or 12) branched rays. 
Vertebral column with 20 to 22 abdominal and 
1 7 or 18 postabdominal vertebrae, total 38 or 39. 
First gill arch with 3 + 7-9 long slender rakers. 

DESCRIPTION. — Body elongate, slender; 
compressed in cross section at abdomen, some- 
what more compressed posteriorly. Skin smooth, 
without tubercles or ridges. Vent located just 
anterior to tip of adpressed pelvic fin. Lateral line 
complete, terminating at hypural-plate margin; 
canal running along lateral myoseptum, with 



pores emanating from short, ventrally directed 
oblique branches. 

Dorsal-fin origin above posterior half of ad- 
pressed pectoral-fin spine, its posterior insertion 
at vertical through pelvic-fin origin; dorsal fin 
with short spinelet preceding first fin ray; first ray 
a stout, sharply pointed, smooth spine; spine 
laterally compressed, its anterior margin a sharp 
keel; fin margin nearly straight, each branched 
ray slightly shorter than preceding ray; last fin 
ray without membranous attachment to body; tip 
of adpressed dorsal fin not reaching adipose-fin 
origin. 

Interval between dorsal fin and adipose fin 
1 '/4-2 times length of dorsal-fin base; adipose-fin 
base approximately equal to that of anal fin; its 
origin slightly in advance of anal-fin origin. 

Caudal fin deeply forked, lobes pointed, sym- 
metrical; outer principal rays nearly three times 
length of middle rays; procurrent rays of upper 
and lower lobes symmetrical, not extending far 
anteriorly. 

Anal-fin base short, approximately equal to 
that of adipose-fin base; first branched ray long- 
est, fin margin slightly concave, last ray without 
membranous attachment to body. 

Pelvic-fin origin at vertical through posterior 
insertion of dorsal fin; fin margin obtusely 
pointed, first branched ray longest; adpressed fin 
reaches to vertical through adipose-fin origin, but 
not to anal-fin origin. 

Pectoral fin with a stout spine, with sharply- 
pointed tip; anterior margin smooth; posterior 
margin feebly serrated; serrae restricted to mid- 
dle of spine, their length less then interval be- 
tween successive serrae; fin margin concave 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



327 




Figure 7. Gagata gasawyuh new species, Tenasserim River (specimen not retained). 



anteriorly, then convex; first pectoral-fin ray 
markedly longer than remaining rays; adpressed 
spine extending past vertical through middle of 
dorsal fin. but not reaching its posterior insertion. 

Head compressed; snout bluntly rounded in 
profile, nearly flat ventrally; dorsal profile nearly 
straight from dorsal spine base to anterior margin 
of orbit; broadly convex anterior of eye; dorsal 
surface of head with broad medial groove, ex- 
tending from level of nasal barbel to base of 
occipital process; eye lateral, above middle of 
head; dorsal margin of orbit contributing to dor- 
sal profile of head; orbital diameter approxi- 
mately 1 Vi times interorbital width, and similarly 
larger than snout length; eye covered with thin 
skin, no free orbital margin. 

Mouth inferior, near snout tip; oral opening 
transverse, en'irely ventral to nares; upper lip 
with single series of tiny finger-like lobes; lower 
lip free lateral to inner mental-barbel origin, lip 
bilobed. lateral lobe longer; upper jaw edentu- 
lous, oral surface covered with fine parallel rows 
of plicae; lower jaw with single row of 2 to 5 
widely-separated conical teeth; teeth not discern- 
ible through plicae covering oral surface, except 
in skeletal preparations; palate edentulous. 

Barbels in four pairs; nasal barbel tiny, extend- 
ing from fleshy flap that divides anterior and 
posterior nares and not reaching to anterior mar- 
gin of orbit; maxillary barbel connected to snout 
by very narrow sheet of membranous tissue that 
attaches to head at corner of mouth; membrane 
nearly indistinct on barbel; maxillary barbel 
reaching pectoral fin origin, or slightly beyond; 
mental barbels originate in transverse row, just 
posterior to lower jaw; for most of their length, 
mental barbels rest in shallow groove in isthmus 
skin; outer mental barbel extends to opercular 



margin, ventral to pectoral spine; inner mental 
barbel shorter. In addition, one pair of tiny, 
rounded, fleshy barbel-like flaps present between 
lower-jaw symphysis and inner mental-barbel 
base. 

Gill openings wide, branchiostegal mem- 
branes attached to isthmus at vertical through 
posterior margin of orbit. Isthmus with deep, 
longitudinal groove that meets paired oblique 
grooves just posterior to base of inner mental 
barbel. 

COLORATION. — Body covered with fine dark 
pigment, concentrated above lateral line; pig- 
ment absent from head and body ventral to level 
of pectoral-fin spine and pigment generally ab- 
sent from sides of body below lateral line; pig- 
ment forming series of distinct saddles across 
head and body, as follows: 1) head at eyes, ap- 
proximately as wide as orbital length and extend- 
ing to below lower margin of orbit; 2) nape, 
across occipital process and extending to pecto- 
ral-fin base; 3) posterior half of dorsal-fin base, 
extending obliquely to slightly above middle of 
adpressed pelvic fin; 4) anterior extent of adipose 
fin, extending obliquely towards, but not reach- 
ing, middle of anal-fin base; and 5) caudal pedun- 
cle, naiTower than preceding saddles and 
extending ventrally to below lateral line. Indis- 
tinct saddle sometimes present across dorsal-fin 
origin. Barbels without pigmentation. Dorsal fin 
with fine dark line along anterior margin of spine, 
and broad marginal band across margin; adipose 
fin with fine, sharply delimited terminal band; 
caudal fin with broad submarginal lunate band; 
anal fin with dusky spot near margin of middle 
rays, spot sometimes extending onto posterior 
rays; pectoral and pelvic fins with few. scattered. 



328 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 



tiny spots on dorsal surface, ventral surfaces 
clear. 

Distribution. — Gagata gasawyuh is 
known from the Irrawaddy, Salween, and Tenas- 
serim basins of Myanmar (Fig. 3). 

COMMENTS. — In all other Gagata with dark 
markings on the caudal fin, the marks consist of 
a square or round spot on the outer principal rays 
of each lobe and the inner rays are immaculate. 
In G. gasawyuh, each lobe has an elongate 
oblique mark that extends from the middle of the 
outer principal rays to the distal margin of the 
middle rays. 

The five saddle marks that extend over the 
dorsum of the head and body of Gagata gasaw- 
yuh are characteristic of this species. In other 
species of Gagata with distinct dark saddles, the 
saddles extend ventrally only to the lateral line 
and, instead of one distinct saddle across the 
dorsal-fin base, there is one at the dorsal-fin 
origin and one behind the posterior insertion of 
the dorsal fin. 

Gagata gasawyuh is substantially larger than 
any other species of Gagata characterized by 
distinct saddles. The largest specimen we exam- 
ined was 130 mm and specimens greater than 90 
mm are not uncommon, whereas the largest 
specimen of G. cenia, the next largest species, we 
examined was only 80 mm. Shrestha (1994), 
however, indicated that specimens of G. cenia in 
Nepal reach 100 mm. 

The extensive anatomical illustrations listed as 
Gagata gagata in de Pinna (1996) are clearly 
based on specimens of this species, the only 
species known from the Chindwin River basin. 



As noted in the comments in the account of 
Gagata, there is considerable variation in the 
form of the swim bladder encapsulation and the 
lack of upper jaw dentition, two of the characters 
mentioned in de Pinna's account. In both charac- 
ters, the descriptions and illustrations in de Pinna 
( 1996) fit G. gasawyuh and not G. gagata. 

ETYMOLOGY. — The species name is based on 
the Karen name "niya gasawyuh." Informants to 
the senior author indicate the derivation of the 
name as follows: niya, the Karen language prefix 
for fish; gasaw, elephant, and yuh, to respect or 
fear (meaning "the fish that elephants are afraid 
to step on"). 



Gagata itchkeea (Sykes, 
(Fig. 8) 



839) 



Phractocephalus itchkeea Sykes 1839:164 (type lo- 
cality: Dukhun [=Deccan]). Sykes, 1841:373, pi. 
67, fig. 1 (Beema River, near Pairgaon [= Bhima 
River (?), a large tributary of the Krishna River]). 

Gagata itchkea, Misra 1976:227 (partim, excluding 
Burma). 

Gagata itchkeea, Day 1877:492, pi. 115, fig. 1; Hora 
and Law 1941:18 (rivers of Deccan); Tilak 
1970:214 (key). 

Nangra itchkea, Jayaram 1979: 13 (review). 

Nangra itchkeea, Jayaram 1981:241 (key); Talwar 
andJhingran 1991:675. 

Nangra viridescens, Jayaram 1979, fig. 10b; Jayaram 
1 98 1 , fig. 131b; Talwar and Jhingran 1 99 1 , fig. 2 1 6. 

Material Examined. — 8 specimens, 32-54 mm. 
INDIA. CAS 62080 (7, 32-54 mm; 2, 36-38 mm, 
cleared and stained), Karnataka State, Tungabahdra 
River and Reservoir at Hospet and Hampi, 28 Jan. 





Figure 8. Gagata itchkeea, 52 mm, CAS 62080; India, Karnataka State, Tungabahdra River. 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



329 



20 




Figure 9. Distribution of Gagala itchkeea, G. melanopterus, G. sexualis, G. youssoufi, and Gangra viridescens. Open circles 
and squares represent unverified literature records. 



1985, Tyson Roberts. CAS(SU) 34S42 (I, 37mm), 
Maharashtra State, Deolali, Bombay, 15 Oct. 1935, 
A. G. L. Fraser. 

Diagnosis. — Apparently a small species, 
largest specimen examined 54 mm. Body deep, 
approximately equal to head length. Eye very 
large, greater than snout length. Snout blunt; 
mesethmoid greatly enlarged, more strongly pro- 
jecting ventrally than in any other species of 
Gagata. Tympanum large, its width equal to eye 
diameter; swim bladder chamber with greatly 
expanded dorsal bony roof but no ventral bony 
enclosure. Jaws with well-developed conical 
teeth in several irregular rows. Cranial roofing 
bones more superficial than in any other Nan- 
grina, and covered with thin skin; cranial surface 
rugose. Posterior cranial fontanel very small, less 
than one-third length of anterior fontanel; lateral 
fontanels absent; subtemporal fossa absent. 

COUNTS. — Dorsal fin with spinelet, spine and 
6 branched rays; pectoral fin with spine and 8 
(rarely 9) rays; anal fin with 4 or 5 simple and 8 
to 10 branched rays. Vertebral column with 17 to 
19 abdominal and 16 to 18 postabdominal verte- 



brae, total 34 or 35. First gill arch with 2 or 3 + 
8 to 1 1 short, slender gill rakers. 

DISTRIBUTION. — Gagata itchkeea is re- 
stricted to peninsular India (Fig. 9) and has been 
reported from the Narmada, Krishna and Cau- 
very basins (Hora and Law 1941). It has not been 
reported from the Ganges basin. Jayaram's 
(1981:241) statement that the species occurs in 
Myanmar was not supported by specimens, and 
we believe it to be in error. 

COMMENTS. — Inclusion of this species in 
Gagata is somewhat problematic. The species 
lacks the lateral cranial fontanel that otherwise 
characterizes Gagata, and it is the only species 
with almost no evidence of swim bladder ossifi- 
cation. On the other hand, G. itchkeea is quite 
similar in appearance to other species of Gagata, 
and shares characteristics such as the compressed 
head and large lateral eye that we consider diag- 
nostic for the genus. 

We have not had the opportunity to study a 
large number of specimens from the various river 
systems from which this species has been re- 
ported. Therefore we are not sure whether all the 
reports refer to this species, and we cannot rule 



330 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 



out the possibility that more than one species 
exists on peninsular India. 

Recent publications on Indian fishes have 
added some unfortunate confusion about the 
identity of this species. Jayaram (1979, 1981) and 
Talwar and Jhingran (1991) provide a perfectly 
recognizable illustration of Gagata itchkeea, but 
in each case, the figures are labeled Nangra 
viridescens. 

Gagata melanopterus new species 
(Fig. 10) 

Gagata cenia, Vinciguerra 1890:121 (partim, Man- 

dalay). 
Gagata gagata, Hora and Law 1941:15 (partim; 

Prome, Burma). 

Material Examined. — 518 specimens, 16-158 
mm. Holotype: USNM 348852 (99 mm), Myanmar, 
Yangon Division, Hlaing River, 16°53'41"N, 
96°05'28"E. 31 October 1997, C.J. Ferraris, Mya 
Than Tun, and local fishermen. 

Paratypes: MYANMAR. AMNH 223191 (15, 
38-69 mm); CAS 99694 (300, 16-84 mm); MZUSP 
uncat.(35, 16-77 mm); NRM 42001 (10, 65-82 mm); 
and USNM 348851 (100, 37-76 mm), taken with 
holotype. CAS 99695 ( 1 5, 45-94 mm), Yangon Divi- 
sion, Yangon River mouth at Mee Pya, 16°25.9'N, 
96°26'E, 29 October 1997, C. J. Ferraris et al. CAS 
91569 ( 1 , 83 mm), Mandalay markets, 23 April 1996, 
C. J. Ferraris, D. Catania and Myint Pe. CAS 91570 
( I, 64 mm), Nyaung-U fish market (near Bagan), 13 
April 1 996, C. J. Ferraris and D. Catania. CAS 95556 
( 1 , 86 mm) and CAS 95557 (5, 82-91 mm), Mandalay 
markets, 1 3-25 April 1 993, T. R. Roberts. CAS 95558 
(3, 41-50 mm), Yangon Division, Thanlyin market, 
28 November 1996, C. J. Ferraris and Myint Pe. CAS 
95559 (3, 46-64 mm), and USNM 344658 (22, 36-70 
mm), Yangon markets, 10-1 1 April 1996, C. J. Fer- 
raris, D. Catania and Myint Pe. NRM 26668 (4, 
75-158 mm), Moulmein, 1 1 November 1934, R. Mal- 
aise. NRM 14893 (1, 81 mm), Mandalay, September 
1935, Malaise. USNM 44755 ( 1, 86 mm), Mandalay, 
1 885, L. Fea. 

DIAGNOSIS. — Body silvery, either without 
marking or with faint saddles. Dorsal, anal, pec- 
toral, and pelvic fins blackened at least distally; 
caudal fin without markings. Dorsal-spine tip not 
reaching to adipose-fin origin when adpressed. 
Premaxilla with 2 rows of teeth. Gill rakers 16 or 
more on outer face of first arch. Adpressed anal 
fin fails to reach first lower procurrent caudal-fin 
rays. 



COUNTS. — Dorsal fin with spinelet, spine and 
6 branched rays; pectoral fin with spine and 9 
(rarely 8 or 10) rays; anal fin with 4 to 6 simple 
and 10 to 12 branched rays. Vertebral column 
with 19 or 20 abdominal and 20 or 21 postab- 
dominal vertebrae, total 39 to 41. First gill arch 
with 6 or 7 + 10 or 1 1 rakers. 

DESCRIPTION. — Body elongate; triangular in 
cross section at abdomen, highly compressed 
posteriorly. Skin smooth, without tubercles or 
ridges. Lateral line complete, midlateral, with 
irregularly distributed, short, ventrally-directed 
oblique branches; on caudal-fin base, lateral line 
curves dorsal ly, but does not extend onto fin rays. 

Dorsal-fin origin above posterior half of ad- 
pressed pectoral-fin spine, posterior insertion at 
vertical through pelvic-fin origin; dorsal fin with 
short spinelet preceding first fin ray; first ray a 
stout, sharply pointed, smooth spine; spine later- 
ally compressed, its anterior margin a sharp keel; 
fin margin straight, each branched ray slightly 
shorter than preceding ray; last fin ray without 
membranous attachment to body; tip of ad- 
pressed dorsal fin not reaching adipose-fin ori- 
gin. 

Interval between dorsal fin and adipose fin 
approximately 2 times length of dorsal-fin base; 
adipose-fin base approximately equal to that of 
anal fin; its origin in line with anal-fin origin. 

Caudal fin deeply forked, lobes pointed, sym- 
metrical; outer principal rays nearly three times 
length of middle rays; procurrent rays of upper 
and lower lobes symmetrical, not extending far 
anteriorly. 

Anal-fin base short, approximately equal to 
that of adipose-fin base; first branched ray long- 
est, fin margin concave, last ray without mem- 
branous attachment to body. 

Pelvic-fin origin at vertical through posterior 
insertion of dorsal fin; fin margin obtusely 
pointed, first branched ray longest; adpressed fin 
reaching past vertical through tip of adpressed 
dorsal fin, but not to anal-fin origin. 

Pectoral fin with a stout spine, sharply pointed 
at tip; anterior margin of spine keeled, smooth; 
posterior margin heavily serrated; serrations 
longest at middle of spine, absent on distal quar- 
ter; fin margin straight, first ray markedly longer 
than remaining rays; adpressed spine extending 
past vertical through posterior insertion of dorsal 
fin, and past pelvic-fin origin. 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



33: 




Figure 10. Gagata melanopterus new species, 99 mm, holotype, USNM 348852; Myanmar, Hlaing River, Yangon. 



Head compressed, triangular; snout bluntly 
pointed in lateral profile, nearly flat ventrally; 
dorsal profile of head nearly straight from snout 
tip to base of occipital process, slightly convex 
thereafter; head with broad mid-dorsal groove, 
extending from snout to tip of occipital process; 
eye lateral, slightly above middle of head; orbital 
diameter greater than interorbital width, and only 
slightly less than snout length; eye covered with 
thin skin, no free orbital margin. 

Mouth inferior, near snout tip; oral opening 
transverse, entirely ventral to nares; free lips 
absent; upper jaw with two rows of slender, 
needle-like teeth; lower jaw with three irregular 
rows of shorter conical teeth; palate edentulous. 

Barbels in four pairs; nasal barbel tiny, extend- 
ing from fleshy flap on anterior margin of poste- 
rior naris and not reaching to anterior margin of 
orbit; maxillary barbel connected to snout by 
broad sheet of membranous tissue that attaches 
to short groove at corner of mouth; membrane 
extends along medial surface of barbel, for at 
least half its length, becoming progressively 
more slender distally; maxillary barbel reaches at 
least to pectoral-fin origin; mental barbels origi- 
nate in transverse row, just posterior to lower 
jaw; outer mental barbel extends to opercular 
margin, ventral to pectoral spine; inner mental 
barbel shorter. 

Gill openings wide, branchiostegal mem- 
branes attached to isthmus at vertical through 
posterior margin of orbit. Vent located just ante- 
rior to tip of adpressed pelvic fin. 

COLORATION. — Body covered with fine dark 
pigment, somewhat more concentrated above lat- 
eral line; pigment absent from head and body 



ventral to level of pectoral-fin spine; pigment 
forms narrow dark saddle behind head; lateral 
process of dorsal-spine pterygiophore darkly 
pigmented; in some individuals, indistinct sad- 
dles cross dorsal fin and adipose-fin bases and 
extend below lateral line; barbels without pig- 
mentation; dorsal fin with dark pigment concen- 
trated on spine and distally on fin membrane, as 
broad marginal band; adipose fin with marginal 
band similar to that on dorsal fin; caudal fin with 
widely scattered pigment, nearly indistinct; in 
some individuals, caudal fin with very fine dark 
marginal band; pectoral fin heavily pigmented on 
dorsal and ventral surfaces of spine and fin, ex- 
cept for basal parts of posterior rays; pelvic fin 
heavily pigmented for distal half, or more, of fin; 
anal fin darkly pigmented on distal pail of all but 
posterior-most rays; width of anal-fin pigmenta- 
tion variable, limited to fine marginal band in 
some individuals but covering distal half of fin 
in others. 

Distribution. — Gagata melanopterus is 
known from the Irrawaddy, Rangoon, Sittang 
and lower Salween basins of Myanmar (Fig. 9). 

Comments. — Gagata melanopterus differs 
from G. gagata in having (a) a dorsal-fin spine 
that fails to reach adipose-fin origin (vs. de- 
pressed dorsal-fin spine extending to, and often 
well past, adipose-fin origin), (b) a smaller pelvic 
fin that fails to reach anal-fin origin (vs. reaching 
anal-fin origin), (c) the last anal-fin ray fails to 
reach first lower procurrent caudal-fin rays (vs. 
extending posteriorly to origin of lower procur- 
rent caudal-fin rays); (d) complete bony encap- 
sulation of the swim bladder (vs. incomplete). 



332 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 



and (e) two rows of teeth on the premaxilla (vs. 
four rows). 

This distinctive new species is known from 
lowland regions of Myanmar. It was most often 
encountered in lower reaches of rivers, or in 
markets near the coast. In one collection from the 
Hlaing River, it was the most abundant of the 
more than 40 species taken. Although the distri- 
bution of this species broadly overlaps that of 
Gagata gasawyuh in the Irrawaddy and Sittang 
rivers, G. melanopterus has not been taken in the 
Tenasserim basin, the Salween River above the 
vicinity of its mouth, the Irrawaddy River above 
Mandalay or the Chindwin River. 

ETYMOLOGY. — The species name "melanop- 
16018," a masculine adjective, is from the Greek, 
melas, black, and pteron, fin, wing, or feather. 

Gagata sexualis Tilak, 1970 

Gagata sexualis Tilak 1 970:207, figs. 1-6 (type local- 
ity: [India] North Koel River at Daltonganj [Cho- 
tanagpur]). 

Material Examined. — 3 specimens, 39-52 mm. 
INDIA: RMNH 26072 (2, 39^10 mm), Ganga River, 
H. Chapra, 28 June 1967. NRM 40591 (1, 52 mm), 
Dibrugarh market, Brahmaputra drainage, F. Fang and 
A. Roos, 20 January 1998. 

DIAGNOSIS. — A small Gagata with filamen- 
tous extension of dorsal-fin spine in males that 
reaches to adipose-fin base. Mouth located at 
level of anterior margin of orbit. Snout bluntly 
rounded in lateral profile without notch anteri- 
orly. Upper jaw toothless; lower jaw with single 
row of teeth (diagnosis after Tilak 1970). 

COUNTS. — Dorsal fin with spinelet, spine and 
6 branched rays; pectoral fin with spine and 8 
branched rays; anal fin with 4 or 5 simple rays, 
followed by 12 to 14 branched rays, posterior two 
rays very close together. Vertebral column with 
19 or 20 abdominal and 17 or 18 postabdominal 
vertebrae, total 36 to 38. Gill rakers not countable 
on examined specimens. 

Distribution. — Gagata sexualis is known 
only from the Ganges basin, including the 
Brahmaputra River (Fig. 9). 

Comments. — Gagata sexualis was de- 
scribed from a small series of specimens that 
were said to exhibit pronounced sexual dimor- 
phism in the shape of the head and the length of 
filamentous extensions of the dorsal and pectoral 



fins. In addition to the type series of seven speci- 
mens, two specimens from the Ganges River 
(RMNH 26072) were determined by Tilak 
(1970) to be conspecific with his new species. 
We have examined these two specimens and 
agree that they represent a distinct species of 
Gagata somewhat similar in appearance to Ga- 
gata cenia. The two species differ in a number of 
characters as outlined in Tilak (1970). In addi- 
tion, they differ in anal-fin ray count, a distinc- 
tion not mentioned in the original description of 
G. sexualis. Although the two RMNH specimens 
were said to be male and female, we were unable 
to see the sexually dimorphic characteristics in- 
dicated in the original description. The specimen 
from the Brahmaputra River system (NRM 
40591) exhibits pronounced dorsal and pectoral 
fin filaments that Tilak indicated to be sexually 
dimorphic characteristics of males. The gonads 
of NRM 40591 appear to be testicular, although 
histological examination was not conducted. 

The distinction between G. sexualis and G. 
youssoufi is more problematic and is discussed in 
detail in the account of the latter species. 



Gagata youssoufi Ataur Rahman, 
(Fig. 11)* 



1976 



Gagata youssoufi Ataur Rahman 1976:5, tig. 1 (type 
locality: Old Brahmaputra River near Mymensingh, 
Bangladesh); Ataur Rahman 1990:221, fig. 131 
(Bangladesh). 

Material Examined. — 120 specimens, 19-50 
mm. BANGLADESH. CAS 95563 (27, 19-41 mm), 
Tangail district, FAP 1 7 fisheries survey, 1992. 

INDIA. BMNH 1 889.2. 1:2615 (1, 50 mm), Assam, 
F. Day. CAS 55560 (1,35 mm), Assam State, Tezpur, 
B. Prashad. CAS 95561 (85, 36-48 mm, 10 cleared 
and stained), Ganges River at Patna, April-May 1 996, 
T. R. Roberts. MCZ 4258 (5 of 9, 41-48 mm), Uttar 
Pradesh, Allahabad, F. Day. 

DIAGNOSIS. — A small species of Gagata, to 
50 mm. Body with five dark saddles. Snout 
broadly rounded in lateral profile, without notch. 
Teeth absent from upper jaw, limited to small 
symphyseal patch on lower jaw. Gill rakers few 
in number and nearly indiscernible, except in 
skeleton preparations. Fourth gill arch with sin- 
gle row of teeth on lower limb. 

COUNTS. — Dorsal fin with spinelet, spine and 
6 branched rays; pectoral fin with spine and 8 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



333 




It"* 



Figure 1 1 . Gagata youssoufi, 38 mm, CAS 95561 ; India. Ganges River at Patna. 



rays (rarely 7 or 9); anal fin with 4 or 5 simple 
rays and 11 to 13 (rarely 14) branched rays. 
Vertebral column with 14 (rarely 15) abdominal 
and 18 or 19 (rarely 1 7) postabdominal vertebrae, 
total 37. rarely 38. First gill arch with 1 + 3 or 4 
tiny rakers. 

Distribution. — Gagata youssoufi is known 
only from the Ganges basin (Fig. 9). 

COMMENTS. — Gagata youssoufi closely re- 
sembles the poorly known G. sexualis, except 
that it lacks a filamentous extension of the dorsal 
fin, which is said to characterize the latter species 
and it does not show the pronounced sexual 
dimorphism figured in Tilak's description of G. 
sexualis. Our specimens from India and Bangla- 
desh typically have only 12 or 13 branched anal- 
fin rays, while two of the three specimens of G. 
sexualis examined by us have 14. Whether G. 
sexualis really is a sexually dimorphic species, 
and whether G. sexualis and G. youssoufi are 
different species are matters that we are unable 
to resolve with the material we examined and 
must be left for further study. Until then, we 
choose to recognize both species as valid. Should 
the names G. sexualis and G. youssoufi represent 
but a single species, the name G. sexualis has 
priority. 

A single specimen that was brought to our 
attention just before this paper went to press may 
shed some light on the status of the name Gagata 
youssoufi. The specimen (NRM 40591) was ob- 
tained at the Dibrugarh market, Assam, and said 
to be taken from the Brahmaputra River. The 
specimen appears to be an adult male Gagata 
sexualis, as it exhibits the elongate dorsal and 
pectoral filaments that characterize that males of 
that species. However, it has only 12 pectoral fin 
rays, which we found to be the typical number 
for Gagata youssoufi, but not that of G. sexualis. 
At 52 mm, the specimen is larger than any speci- 



men of Gagata youssoufi examined by us. It is 
possible that the elongate filaments are either 
exhibited only seasonally, or only in individuals 
larger than 50 mm. The original description of 
Gagata sexualis indicates the length of only one 
of the four males examined by Tilak (1970), 
which was that of the 55 mm holotype. Although 
we continue to reserve judgment until more ma- 
terial becomes available, it appears more likely 
that Gagata youssoufi may ultimately be a syno- 
nym of G. sexualis. 

Gangra new genus 

Type species: Pimelodus viridescens Hamilton 1822. 
Monotypic. 

DIAGNOSIS. — Summaries of counts of verte- 
brae, fin rays, and gill rakers are presented in 
Tables 1-3. Head, mouth and jaws broad, jaws 
with several rows of conical teeth. Mesethmoid 
not greatly expanded, its dorsal profile slightly 
convex, Y-shaped anteriorly. Palate toothless. 
Barbels short, slender, and round in cross section; 
nasal barbel very short, extending posteriorly 
only to end of posterior naris; maxillary and 
mental barbels not extending posteriorly beyond 
head; maxillary-barbel membrane absent or 
greatly reduced; maxillary bone inside maxillary 
barbel very short; outer and inner mental barbels 
evenly and widely separated. Paired lateral cra- 
nial fontanel absent. Branchiostegal membranes 
free from isthmus. 

COMMENTS. — Gangra viridescens (Hamil- 
ton) has historically been a problematic species 
from the standpoint of its identity and relation- 
ships. The large number of primitive sisorid char- 
acters exhibited by this taxon has caused 
researchers to place it variously with Nangra or 
Gagata by focusing on shared primitive charac- 



334 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 




Figi rf 1 2. Gangra viridescens, 56 mm, CAS 95570; India, Ganges River at Patna. 




W 



Figure 1 3. Gangra viridescens, illustration from Hamilton ( 1 822), plate I I , Figure 56; originally 56 mm. 



ters. It seems to us that the difficulty in assigning 
G. viridescens unambiguously to either of those 
two genera was the primary reason for the use of 
a single generic name for all of the species dis- 
cussed here (e.g., Misra 1976; Burgess 1989; 
Ataur Rahman 1990). Although we concur with 
de Pinna's (1996) conclusion that all of these 
species form a natural group, we think that treat- 
ing them as a part of a single genus obscures the 
diversity of form exhibited by this group and 
overshadows the clear species groups of Nangra 
and Gagata. By placing Gangra viridescens into 
a separate genus, both of these other genera be- 
come readily diagnosable entities. 

ETYMOLOGY. — Gangra is a hybrid word 
coined from Gagata and Nangra. Gender mascu- 
line. 

Gangra viridescens (Hamilton, 1 822) new com- 
bination 
(Figs. 12, 13) 



Pimelodus viridescens Hamilton 1822:173, 376, pi. 

1 1 , fig. 56 (type locality: rivers of the northern parts 

of Bengal). 
Nangra punctata Day 1877:494, pi. I 15, fig. 8 (type 

locality: Sone River at Bheer Bhoom, Bengal). 
Gagata viridescens, Hora and Law 1 94 1 :24, pi. 1 , figs. 

7, 8; Tilak 1970:214 (key); Misra 1976:229; Ataur 

Rahman 1 990:2 1 9 (Bangladesh); Shrestha 1994:58, 

fig. 90 (Nepal). 
Gagata (Nangra) viridescens, Mahajan 1967:298 et 

seq., figs. 3, 12, 14 (osteology of axial skeleton, 

caudal fin). 
Nangra viridescens, Jayaram 1979:14 (review); 

Jayaram 1981:241 (key); Shrestha 1994:142 (Ne- 
pal). 

Material Examined. — 59 specimens, 30-72 
mm. BANGLADESH. CAS 95571 (28, 32-55 mm), 
Tangail district, FAP 17 fisheries survey, 1992; CAS 
95572 (2, 30-31 mm), Tangail District, N. Dhalesh- 
wari River, FAP 17 fisheries survey, 9 Oct. 1992. 

INDIA. AMS B.7566 ( 1 , 49 mm, syntype of Nan- 
gra punctata), Sone River, F. Day. BMNH unreg. (1, 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



335 



37 mm), Bengal, Tirpoot, Gordon Dalgleish. BMNH 
1871.4.17:2 (4, not measured), N. E. Bengal, Jerdon. 
CAS 95570 (23, 44-72 mm), Ganges River at Patna, 
April-May 1996, T. R. Roberts. 

Diagnosis. — Same as for Gangra. 

COUNTS. — Dorsal fin with spinelet. spine and 
6 branched rays; pectoral fin with spine and 8 
branched rays; anal fin with 4 to 6 simple rays 
and 7 to 9 branched rays (typically 5 + 8). Verte- 
bral column with 18 or 19 abdominal and 15 or 
16 postabdominal vertebrae, total 33 or 34. First 
gill arch with 5 or 6 rakers. 

COLORATION. — Gangra viridescens is pale 
greenish or grayish overall, with a vivid, almost 
viridescent, horizontal silvery mark in the mem- 
brane connecting the opercle to the side of the 
cranium. These supraopercular marks are obvi- 
ous, viewed from the side or especially when 
viewed from above. Similar marks do not occur 
in any other sisorid. 

DISTRIBUTION. — Gangra viridescens occurs 
throughout lowland parts of the Ganges- 
Brahmaputra system (Fig. 9). 

NangraDay. 1877 

Nangra Day 1 877:493 (Type species: Pimelodus nan- 
gra Hamilton, 1 822, by absolute tautonymy). Gen- 
der feminine. 

Diagnosis. — Summaries of counts of verte- 
brae, fin rays, and gill rakers are presented in 
Tables 1-3. Body elongate, slender. Head de- 
pressed; snout slightly to considerably spatulate, 
with ventral portion flat and more or less mark- 
edly projecting anterior to jaws. Eyes small, dor- 
solateral. Mesethmoid a large flattened plate; 
premaxilla immovably fixed to mesethmoid. 
Jaws well toothed. Palatal teeth present on bone 
tentatively identified as endopterygoid; tooth 
patch single, elongate and widely separated 
across midline; teeth few in number but present 
in all specimens of Nangra examined by us for 
this character. Branchiostegal membranes free 
from isthmus. Barbels very long, maxillary bar- 
bel extending at least to pectoral spine tip and 
usually beyond pelvic fins; nasal barbel extend- 
ing at least to eye. Maxillary barbel membrane 
well developed. Maxillary bone inside maxillary 



barbel very long, extending posteriorly nearly to 
end of head or beyond. 

NOTE ON GENDER. — Although Nangra. like 
Gagata, presumably is masculine in Bengali. 
Day and other ichthyologists have treated Nan- 
gra as feminine (e.g., Nangra punctata Day, 
1877; Nangra robusta Mirza and Awan, 1973). 
The feminine gender of Nangra is recognized by 
Eschmeyer and Bailey (in Eschmeyer 1990:258). 

COMMENTS. — The composition of Nangra, 
as used here, differs from all previous uses. We 
believe that the combination of depressed body, 
elongated spatulate snout, and toothed palate are 
indications of a natural group of sisorids. The 
snout is supported by a broadly expanded plate 
on the ventral surface of the mesethmoid comua. 
a feature not otherwise seen in sisorids. Simi- 
larly, palatal teeth are typically absent in sisorids, 
although they have been reported in Glypto- 
thorax cons (as Euclyptosternon cons) by Gtin- 
ther( 1864) and Regan (1911). 

Key to Species of Nangra 
la. Nasal barbel extends to margin of head or 
beyond; dorsal fin typically with 8 (rarely 

7 or 9) branched rays 

Nangra nangra (Hamilton, 1822) 

lb. Nasal barbel extends no further than poste- 
rior margin of orbit; dorsal fin with 6 or 7 

branched rays 2 

2a. Dorsal fin and pectoral fin with long fila- 
ment extending from first ray 3 

2b. Dorsal fin and pectoral fin with little or no 

filamentous extension 4 

3a. Pectoral fin with 10 or 11 branched rays, 

dorsal fin with 7 branched rays 

. . Nangra robusta Mirza and Awan, 1973 
3b. Pectoral fin with 8 branched rays, dorsal 

fin with 6 branched rays 

.... Nangra carcharhinoides, new species 
4a. Dorsal fin with 7 branched rays and dark 
semicircular spot on base of middle rays, 
maxillary barbel extends to anal-fin origin 

Nangra ornata, new species 

4b. Dorsal fin with 6 branched rays and with- 
out dark markings; maxillary barbel ex- 
tends only to tip of adpressed pectoral-fin 

spine 

Nangra bucculenta, new species 



336 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 





Figure 14. Nangra bucculenta new species, holotype, 28 mm, CAS 95564; Bangladesh, Tangail District. 



Nangra bucculenta new species 
(Fig. 14) 

Holotype: CAS 95564 (28 mm), Ganges River 
delta, Tangail District, North Central Region, Bangla- 
desh, FAP 17 fisheries survey, 1992. 

Paratypes: CAS 95565 (10, 15-34 mm; 3, 20-23 
mm, eleared and stained), same collection data as 
holotype. 

DIAGNOSIS. — Possibly a very small species, 
largest known specimen only 34 mm. Snout 
moderately projecting; cheeks relatively ex- 
panded, head terete. Anterior naris very near 
snout tip (distance between naris and snout tip 
less than eye diameter). Nasal barbel extending 
to eye; maxillary barbel extending only to end of 
pectoral fin; maxillary-barbel membrane broadly 
attached to head only on cheek; maxillary bone 
extending posteriorly to midway between eye 
and end of head. Vent opening directly to exte- 
rior; vent and genital papilla located between 
anterior third of pelvic fins. Dorsal and pectoral 
fins without filamentous extensions. 

COUNTS. — Dorsal fin with spinelet, spine and 
6 branched rays; pectoral fin with spine and 8 
(rarely 7) branched rays; pelvic fin with 1 un- 
branched and 5 branched rays; anal-fin rays 4 or 
5 simple and 7 or 8 branched rays, total 11 to 13. 
Vertebrae 18 or 19 abdominal and 15 or 16 
postabdominal, total 34. First gill arch 2 + 4 or 5 
rakers. 

Description. — Body elongate, slender; 
round in cross section at abdomen, progressively 
compressed posteriorly. 

Dorsal-fin origin above middle of adpressed 
pectoral fin, posterior insertion above pelvic-fin 
origin; fin with short spinelet preceding first fin 
ray; first ray a smooth spine for most of its length, 
flexible distally and not prolonged into filament; 
fin margin straight, each ray slightly longer than 
following ray; last fin ray without membranous 
attachment to body. 



Interval between dorsal fin and adipose fin 
greater than length of dorsal-fin base; adipose fin 
approximately equal in size to anal fin; its origin 
above, or just anterior to anal-fin origin. 

Caudal fin deeply forked, lobes pointed, lower 
lobe longer and broader than upper lobe; outer 
principal rays more than twice length of middle 
rays; procurrent rays of upper and lower lobes 
symmetrical, not extending far anteriorly. 

Anal-fin base short, approximately equal to 
that of adipose-fin base; first branched ray long- 
est, fin margin slightly concave, last fin ray with- 
out membranous attachment to body. 

Pelvic-fin origin at vertical through posterior 
insertion of dorsal fin; fin margin slightly con- 
vex, middle rays longest; adpressed fin not reach- 
ing anal-fin origin. 

Pectoral fin with stout spine; spine smooth 
anteriorly, but with 10 to 12 stout serrations on 
posterior margin; serrations progressively longer 
distally; spine with short filament extending 
from tip; fin margin straight or slightly convex, 
first ray longest; adpressed fin extending to ver- 
tical from base of second branched dorsal-fin ray. 

Head bluntly conical, somewhat depressed 
dorsal ly and nearly flat ventral ly; dorsal surface 
of head with broad medial groove, extending 
from snout to posterior extent of supraoccipital; 
groove narrow and shallow on occipital process; 
eye dorsolateral; eye diameter slightly more than 
one-half interorbital width or snout length; eye 
covered with thin skin, no free orbital margin. 

Mouth inferior, wide, located near snout tip; 
oral opening entirely anterior to eye, curved 
slightly; upper lip absent; lower lip continuous 
with skin of underside of head, except at corners 
of mouth. Jaws with several rows of tiny teeth, 
teeth nearly indistinguishable from surrounding 
tissue; palatal tooth patch small, oblong, with 
few irregularly-placed teeth. 

Barbels in four pairs. Nasal barbel extends no 
further posteriorly than anterior margin of orbit. 
Maxillary barbel attached to head by broad sheet 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



337 



30 - 




Figure 15. Distribution of species of Nangra. Open squares represent unverified literature records for Nangra nangra. 
Distribution of Nangra robusta based on type locality. 



of membranous tissue that extends to level of 
outer mental-barbel origin; membrane extends 
along medial surface of barbel, progressively 
more slender towards barbel tip; maxillary barbel 
reaches to level of second or third branched dor- 
sal-fin ray. Mental barbels originate just poste- 
rior to lower jaw; outer mental barbel reaches 
pectoral fin-spine base; inner mental barbel 
somewhat shorter. 

Gill openings wide, branchiostegal mem- 
branes narrowly attached to isthmus. Vent situ- 
ated between middle of adpressed pelvic fins. 
Head and body covered with scattered, elongate, 
ridged or keeled tubercles; tubercles abundant on 
dorsal surface of head and body, especially on 
cheeks and ventral surface of snout; ventral sur- 
face of abdomen smooth. 

COLORATION. — Head and body above lateral 
line covered with scattered dark flecks, concen- 
trated in three patches middorsally: 1 ) at dorsal- 
fin origin, 2) posterior to posterior insertion of 
dorsal fin, 3) along adipose- fin base; elongate 
dark spot covers dorsal half of opercle; pigmen- 
tation absent below lateral line and on lower half 



of head; barbels, pectoral fin, pelvic fin, and anal 
fin without pigmentation; dorsal fin with pig- 
ment on middle of first four branched fin rays, 
remaining rays and interradial membranes clear; 
adipose fin with scattered pigmentation basally; 
caudal-fin base with small dark triangular spot 
ending across bases of middle rays; some speci- 
mens with faint dark spot on middle of lower 
caudal-fin lobe. 

Distribution. — Nangra bucculenta is 
known only from the Tangail District of central 
Bangladesh, a flood plain area between large 
tributaries in the Ganges delta, about 15 km east 
ofthe Jammuna mainstream (Fig. 15). Our speci- 
mens were obtained from sites that yielded large 
numbers ofthe following other sisorids: Glypto- 
thorax telchitta (Hamilton, 1822), Gagata cenia, 
G. youssoufi, Nangra nangra, Gangra viri- 
descens, and Sis or rabdop horns Hamilton, 1822. 

ETYMOLOGY. — The name "bucculenta," a 
feminine adjective, is from the Latin "bucculen- 
tus," meaning with expanded cheeks. 



338 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 




Figure 16. Nangra carcharhinoicies new species, holotype, 79 mm, CAS 95566; India, Ganges River at Patna. 



Nangra carcharhinoides new species 
(Fig. 16) 

? Nangra nangra Hamilton 1822 (partim; pi. 11, tig. 
63). 

Holotype: CAS 95566 (79 mm), INDIA: Ganges 
River at Patna, April-May 1996, T. R. Roberts. 

Paratypes: CAS 95567 (30, 60-92 mm), SRS/ZSI 
F.4847 (6, 65-87 mm), and UMMZ 233235 (2, 73-82 
mm), collected with the holotype. 

Diagnosis. — A relatively large species of 
Nangra, largest specimen 92 mm. Dorsal fin 
with 6 branched rays; dorsal and pectoral-fin 
spines with filamentous extensions. Snout 
strongly projecting. Nasal barbel reaching only 
to posterior margin of orbit; maxillary barbel 
extending to end of pelvic fin or to anal-fin 
origin; maxillary barbel membrane with narrow 
attachment to cheek and broad attachment to 
corner of mouth; mental barbels extending pos- 
teriorly to end of head. Vent opens into pouch or 
pseudovent, with broad membranous opening 
near pelvic fin tip. Head, body and caudal pedun- 
cle slightly more elongate than in other Nangra. 

COUNTS. — Dorsal fin with spinelet, spine and 
6 branched rays; pectoral fin with spine and 8 
rays; anal fin with 4 or 5 simple rays and 7 to 9 
branched rays. Vertebral column with 18 to 20 
abdominal and 17 or 18 postabdominal verte- 
brae, total 35 to 37. First gill arch with 1 or 2 + 5 
or 6 long, slender rakers. 

DESCRIPTION. — Body elongate and slender, 
round in cross section at abdomen, slightly com- 
pressed posteriorly. Skin of head and body cov- 
ered with scattered, elongate, ridged or keeled 
tubercles; tubercles more abundant on dorsal sur- 



face of head and body, absent on ventral surface 
of abdomen. Lateral line midlateral, complete. 

Dorsal-fin origin above distal fourth of ad- 
pressed pectoral-fin spine, posterior insertion 
posterior to vertical through pelvic-fin origin; fin 
with short spinelet preceding first fin ray; first ray 
a smooth spine continued distally as flexible ray, 
with slender filament extending beyond tip of 
first branched ray; fin margin slightly concave, 
each branched ray slightly longer than following 
ray; last fin ray without membranous attachment 
to body. 

Interval between dorsal fin and adipose fin 
approximately V/i times length of dorsal-fin 
base; adipose fin approximately equal in size to 
anal fin; adipose fin origin above anal-fin origin. 

Caudal fin deeply forked, lobes pointed; lower 
lobe slightly longer and broader than upper lobe; 
outer principal rays twice length of middle rays; 
procurrent rays of upper and lower lobes sym- 
metrical, not extending far anteriorly. 

Anal-fin base short, approximately equal to 
that of adipose-fin base; first branched ray long- 
est, fin margin concave, last fin ray without mem- 
branous attachment to body. 

Pelvic-fin origin at vertical through middle of 
dorsal fin; fin margin obtusely pointed, first 
branched ray longest; adpressed fin reaching ver- 
tical through tip of adpressed dorsal fin, but not 
to anal-fin origin. 

Pectoral fin with stout spine, spine tip blunt. 
Spine smooth anteriorly, with 12 or more stout 
serrations along posterior margin; serrations 
longest at middle of spine; spinous ray continued 
as long slender filament, filament length only 
slightly less than length of bony spine; fin margin 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



339 



concave anteriorly, convex for last two rays; first 
branched ray markedly longer than remaining 
rays; spinous portion of first ray extending to 
vertical from base of second branched dorsal-fin 
ray; filamentous extension reaching to pelvic-fin 
origin. 

Head triangular with rounded, conical snout 
tip; head somewhat depressed dorsal ly and 
nearly flat ventrally; dorsal surface of head with 
broad groove, extending from snout on to occipi- 
tal process, but not to tip of process; eye dorso- 
lateral; orbital diameter approximately equals 
interorbital width, but only one-half snout 
length; eye covered with thin skin, no free orbital 
margin; anterior naris relatively remote from 
snout tip, distance between naris and snout tip 
equals orbital diameter. 

Mouth inferior, near snout tip; oral opening 
entirely anterior to eye, curved slightly; upper lip 
absent; lower lip continuous with skin of under- 
side of head, except at corners of mouth. Jaws 
with three irregular rows of small conical teeth; 
palatal tooth patch tiny, remote from midline of 
palate, and consisting of 2 or 3 small, irregularly 
placed teeth. 

Barbels in four pairs. Nasal barbel extends 
posteriorly at least to middle of eye and some- 
times to posterior margin of orbit. Maxillary 
barbel attached to head by broad sheet of mem- 
branous tissue that extends to level of corner of 
mouth; along medial surface of barbel, mem- 
brane becomes progressively more slender dis- 
tal ly; maxillary barbel reaches at least to tip of 
adpressed pelvic fin, and often to anal-fin origin. 
Inner mental barbel reaches to pectoral-spine 
base. Outer mental barbel originates lateral and 
posterior to inner mental barbel and extends at 
least to middle of adpressed pectoral-fin spine. 

Gill openings wide, branchiostegal mem- 
branes narrowly attached at anterior extent of 
isthmus. Vent opens into pouch or pseudovent, 
opening near tip of adpressed pelvic fin. 

Coloration. — Head and body above lateral 
line covered with scattered dark flecks, concen- 
trated middorsally on body and dorsal to pectoral 
fin; pigmentation often extends below lateral 
line, but not reaching ventral surface of head or 
body; maxillary and mental barbels, pelvic fin, 
and anal fin without pigmentation; dorsal and 
caudal fins with pigment on branched fin rays but 
with clear interradial membranes; adipose fin 
with fine, scattered pigmentation; caudal-fin 



base dusky without distinct dark basal spot; rays 
of lower caudal-fin lobe more heavily pigmented 
than those of upper lobe; pectoral fin with scat- 
tered pigment on dorsal surface of spine and 
basal portion of fin, distal parts of interradial 
membranes and rays clear. 

Distribution. — Nangra carcharhinoides is 
known only from the Ganges near Patna (Fig. 
15). At this place the river is very broad, and 
during the dry season (i.e., at the time of collec- 
tion) the bottom has a vast extent of fine white 
sand and seemingly little else. All of the speci- 
mens were obtained from markets or local fish- 
ermen. Therefore, precise information on the 
habitat of this species is unavailable. 

COMMENTS. — It is possible that Hamilton's 
(1822) original account of N. nangra may have 
been based in part on N. carcharhinoides. Ham- 
ilton's figure (Fig. 17) shows only 6 instead of 
the 8 soft dorsal fin rays mentioned in his diag- 
nosis, and his statement that N. nangra attains 3 
or 4 inches suggests N. carcharhinoides. The 
nasal barbel is a bit too long for N. carcharhi- 
noides. but not long enough for N. nangra. The 
length of the maxillary barbel is too short for N. 
nangra, but it is the right length for N. carcharhi- 
noides. The figure does not show the dorsal and 
pectoral-fin filaments characteristic of N. car- 
charhinoides and seems to be of a less elongate 
fish. The diagnostic character of 8 dorsal fin rays 
is a prominent part of Hamilton's account of 
Nangra nangra, and we follow a long tradition 
of recognizing the only known species with 8 
dorsal-fin rays as N. nangra. 

The filamentous extensions of the dorsal and 
pectoral-fin spines are composed mainly of un- 
consolidated actinosts, as many as 20 for the 
dorsal-spine filament and 15 for the pectoral- 
spine filaments. 

The pseudovent is a unique pouch-like feature 
observed only in Nangra carcharhinoides. The 
pouch is a little less than an eye diameter in width 
and up to 2 times the eye diameter in length, with 
a broad, transverse and strongly concave or semi- 
circular membranous opening posteriorly. The 
structure is fully formed in 19 of 24 specimens 
examined for this character. In four specimens it 
was more or less collapsed or reduced, and in one 
specimen it was absent. Collapse or reduction 
may be partly due to drying of specimens before 
preservation, and the absence due to injury of the 
structure during life. In all specimens the vent 



340 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 




Figure 1 7. Nangra nangra, illustration from Hamilton ( 1 822), plate 1 1 , figure 63; originally 52 mm. 




Figure 18. Nangra nangra, neotype, 47 mm, CAS 96626; India, Ganges River at Patna. 



itself lies near the tip of the pelvic fin, thus farther 
posteriorly than in any other species of Nangra 
(possibly excepting N. robusta, in which pres- 
ence or absence of a pseudovent and the position 
of the vent are unknown). 

Etymology. — The name "carcharhinoides" 
(adjective) refers to the shark-like appearance of 
this species, especially with regards to the shape 
of the snout. 



Nangra buchanani Day 1 877:494, pi. 1 1 3, fig. 3. (type 
locality: Ganges, Jumna, and Indus rivers). 

Gagata nangra, Hora and Law 1941:26, pi. 1, figs. 
9-10 (Ganges, Kosi, Hooghly, Jumna, Indus); Tilak 
1970:214 (key); Ataur Rahman 1990:217, fig. 128 
(Bangladesh). 

Nangra nangra, Mirza and Awan 1973:149 (Indus 
River); Mirza 1980:26 (Punjab); Jayarain 
1981:241, fig. 131a (key); Talwar and Jhingran 
1991:676, fig. 215; Shrestha 1994:142 (Nepal). 



Nangra nangra (Hamilton, 1822) 
(Figs. 17, 18) 

Pimelodus nangra Hamilton 1822:193, 378, pi. 1 
fig. 63. (type locality: Kosi River). 



Material Examined. — 303 specimens, 20-55 
mm. BANGLADESH. CAS 95569 (85, 20-40 mm), 
Ganges basin, North Central Region, Tangail district, 
FAP 17 fisheries survey, 1992. UMMZ 208457 (2, 
23-37 mm), Meghna River just downstream from 
Gazaria, 27 Oct. 1977, W. J. Rainboth et al. 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



341 



INDIA. BMNH 1889.2.1:2337-2346 (17, 22-35 
mm), Allahabad, F. Day. CAS 96626 (1, 47 mm, 
neotype of Nangra nangra), CAS 95568 (190, 37-52 
mm), and SRS ZSI F.4846 (10, 37-55 mm), Ganges 
River at Patna, April-May 1996, T. R. Roberts. 

DIAGNOSIS. — A medium-sized species of 
Nangra, reaching 55 mm. Dorsal tin typically 
with 8 branched rays. Snout moderately project- 
ing; anterior naris origin close to snout tip (dis- 
tance between anterior naris and snout tip less 
than eye diameter); nasal barbel extending pos- 
teriorly at least to end of head, sometimes reach- 
ing to dorsal-fin origin; maxillary barbel 
extending past adipose-fin origin; maxillary 
bone long, extending posteriorly almost to end of 
head; maxillary barbel membrane small, with 
narrow attachment to cheek only; vent opening 
directly to exterior. Vent and genital papillae 
near middle of pelvic fins. 

COUNTS. — Dorsal fin with spinelet, spine and 
8 (rarely 7 or 9) branched rays; pectoral fin with 
spine and 8 or 9 (modally 8) rays; anal fin with 3 
or 4 simple rays and 9 to 1 1 (modally 9) branched 
rays. Vertebral column with 19 to 20 abdominal 
and 16 or 17 postabdominal vertebrae, total 35 to 
37. First gill arch with 7 to 9 rakers. 

DISTRIBUTION. — Nangra nangra is known 
from the Gangetic basin and the Indus River (Fig. 
15). Mirza (1980) reported that this species was 
rare in the Punjab of Pakistan. 

COMMFNTS. — The identity of this species is 
somewhat problematic in light of conflicting in- 
formation provided in Hamilton (1822), and the 
lack of type specimens. As mentioned earlier. 
Hamilton did not save any types, so we have tried 
to match Hamilton's description and illustration 
(Fig. 17) with one of the Gangetic species of 
Nangra. As mentioned in the account of Nangra 



carcharhinoides, discrepancies exist between 
Hamilton's illustration and text in the number of 
branched dorsal-fin rays. We choose to associate 
the name Pimelodus nangra Hamilton with the 
one species that has eight dorsal-fin rays, which 
is arguably an observation directly made by 
Hamilton. We cannot know if the lesser number 
of rays in the illustration is real, or a mistake by 
the illustrator that went undetected by Hamilton. 
The decision to retain the name N. nangra for the 
eight-rayed species maintains the usage that has 
remained stable at least as far back as Day ( 1 877), 
and continued by Hora and Law ( 194 1 ) and more 
recent Indian workers. To maintain this stability, 
we have selected a specimen of this species (CAS 
96626) as neotype for Nangra nangra. 

Day (1877) proposed Nangra buchanani as a 
substitute for Pimelodus nangra Hamilton. 1822, 
to avoid the tautonomy caused by his proposal to 
use Nangra as a generic name. It appears not to 
be a strict replacement name, and specimens 
from the several localities mentioned in the ac- 
count must be considered syntypes of N. bucha- 
nani. Although we have not been able to examine 
any of these syntypes, Day's (1877) illustration 
and description are clearly that of Nangra nan- 
gra. 

Nangra ornata new species 
(Fig. 19) 

Holotype: UMMZ 233236 (37 mm), BANGLA- 
DESH: Gowain River and Khal at Gowainghat, north- 
ern Sylhet province (Surma or Meghna watershed), 2 1 
February 1978, W.J. Rainboth and A. K. Ataur 
Rahman. 

Paratypes: UMMZ 208746 (5, 25-34 mm), col- 
lected with the holotype. 




Figure 19. Nangra ornata new species, holotype, 37 mm, UMMZ 233236; Bangladesh, Sylhet Province, Gowain River 
and Kahl at Gowainghat. 



342 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 



DIAGNOSIS. — A small species of Nangra, 
largest specimen only 37 mm. Eye relatively 
large, its diameter about equal to snout length; 
nasal barbel extends to just past posterior margin 
of orbit; maxillary barbel with broad membrane, 
barbel reaches to anal-fin origin; semicircular 
spot on middle of caudal-fin base, about equal to 
eye diameter; dark spot extending across bases 
of second through fifth branched dorsal-fin rays. 

COUNTS. — Dorsal fin with spinelet, spine and 
7 branched rays; pectoral fin with spine and 8 
branched rays; anal fin with 4 or 5 simple and 8 
or 9 branched, total 12-14 rays. Vertebral col- 
umn with 19 or 20 abdominal and 16 or 17 
postabdominal vertebrae, total 35 (rarely 36). 
First gill arch with 1 or 2 + 6 to 8 rakers, total 
7-10. 

DESCRIPTION. — Body elongate, slender; 
round in cross section at abdomen, progressively 
compressed posteriorly. 

Dorsal-fin origin above distal fourth of ad- 
pressed pectoral fin, posterior insertion above 
pelvic-fin origin; fin with short spinelet preced- 
ing first fin ray; first ray stiffened for basal half, 
flexible distally and not prolonged into filament; 
fin margin straight or slightly convex, each ray 
slightly shorter than preceding ray; last fin ray 
without membranous attachment to body. 

Interval between dorsal fin and adipose fin 
greater than length of dorsal-fin base; adipose fin 
approximately equal in size to anal fin; its origin 
above, or just anterior to anal-fin origin. 

Caudal fin deeply forked, lobes pointed; outer 
principal rays more than twice length of middle 
rays; procurrent rays of upper and lower lobes 
symmetrical, not extending far anteriorly. 

Anal-fin base short, approximately equal to 
that of adipose-fin base; first branched ray long- 
est, fin margin slightly concave, last fin ray with- 
out membranous attachment to body. 

Pelvic-fin origin at vertical through posterior 
insertion of dorsal fin; fin margin straight, rays 
of approximately equal length; adpressed fin not 
reaching anal fin origin. 

Pectoral fin with stout spine; spine smooth 
anteriorly, but with 7 to 9 stout serrations on 
posterior margin; serrations progressively longer 
distally; fin margin straight or slightly convex, 
first ray longest; adpressed fin extending to ver- 
tical from base of second branched dorsal-fin ray. 

Head bluntly conical, somewhat depressed 
dorsally and nearly flat ventrally; dorsal surface 



of head with broad median groove, extending 
from snout to tip of occipital process; eye dorso- 
lateral, large; eye diameter approximately equals 
interorbital width, and nearly equals snout 
length; eye covered with thin skin, no free orbital 
margin. 

Mouth inferior, near snout tip; oral opening 
entirely anterior to eye, curved slightly; upper lip 
absent; lower lip continuous with skin of under- 
side of head, except at comers of mouth. Jaws 
with several rows of tiny teeth, teeth nearly in- 
distinguishable from surrounding tissue; palatal 
tooth patch elongate, teeth irregularly placed. 

Barbels in four pairs. Nasal barbel extends 
posteriorly to just past posterior margin of orbit. 
Maxillary barbel with broad sheet of membra- 
nous tissue along medial margin that extends to 
level of comer of mouth; membrane progres- 
sively more slender towards barbel tip; maxillary 
barbel reaches to anal-fin origin. Inner mental 
barbel originates just posterior to lower jaw and 
reaches to middle of pectoral-fin spine; outer 
mental barbel originates lateral and posterior to 
inner mental barbel and extends to level of tip of 
adpressed pectoral-fin spine. 

Gill openings wide, branchiostegal mem- 
branes narrowly attached across isthmus. Vent 
situated between middle of adpressed pelvic fins. 
Skin of head and body covered with scattered, 
elongate, ridged or keeled tubercles; tubercles 
more abundant on dorsal surface of head and 
body, absent on ventral surface of abdomen. 

COLORATION. — Head and body above lateral 
line covered with scattered dark flecks, concen- 
trated middorsally on body and above pectoral 
fin; diffuse dark spot covers posterior extent of 
occipital process; pigmentation absent below lat- 
eral line and on lower half of head; barbels, pelvic 
fin, and anal fin without pigmentation; dorsal fin 
with pigment concentrated on lower 1/3 of sec- 
ond through fifth branched fin rays and, to a 
lesser extent, on intervening interradial mem- 
branes; remainder of fin with little or no pigmen- 
tation; basal half of adipose fin with scattered 
pigmentation; caudal-fin base with dark semicir- 
cular spot extending across bases of all principal 
rays, covering basal third of middle rays, but not 
extending onto caudal peduncle; sparsely scat- 
tered pigmentation reaches distally approxi- 
mately two-thirds length of caudal-fin rays, but 
absent from fin-ray tips and interradial mem- 
branes; pectoral fin with scattered pigment on 



ROBERTS AND FERRARIS: GAGATA AND NANGRA REVIEW 



343 



dorsal surface of spine and basal parts of rays, 
interradial membranes, distal parts of rays and 
ventral surface clear. 

Distribution. — Nangra ornata is known 
only from the type locality in the Surma or 
Meghna watershed in northeastern Bangladesh 
(Fig. 15). 

COMMENTS. — Nangra ornata is readily dis- 
tinguished by two prominent features. It has the 
largest eye of any Nangra, orbit length approxi- 
mately equal to that of the snout. In all other 
species, the eye diameter is more typically one- 
half to one-third the snout length. The dorsal fin 
of N ornata is unique in having a dark spot at the 
base of middle rays. 

ETYMOLOGY. — From ornatus, Latin for 
decorate or embellish, in reference to the bold 
spots on the caudal peduncle and the dorsal-fin 
base. 

Nangra robusta Mirza and Awan, 1973 

Nangra robusta Mirza and Awan 1973:145, fig. I 
(type locality: Indus River at Jinnah Barrage near 
Kalabagh, Pakistan). 

Material Examined. — None. 

DIAGNOSIS (after Mirza and Awan). — A rela- 
tively large species, attaining 100 mm. Dorsal 
and pectoral-fin spines with filamentous projec- 
tions. Snout strongly projecting. Nasal barbel 
extending to eye; maxillary barbel reaching to 
anal fin. 

COUNTS. — Dorsal fin with spinelet, spine and 
7 branched rays; pectoral fin with spine and 10 
or 1 1 branched rays; anal fin with 2 simple rays 
and 8 branched rays (counts from Mirza and 
Awan, gill rakers and vertebrae not reported). 

Distribution. — This species is reported 
from the Indus River of Pakistan (Fig. 15), where 
it is considered to be rare (Mirza, 1980). 

COMMENTS. — Nangra robusta is similar to 
our new species. Nangra carcharhinoides, both 
in overall appearance and in size. None of the 
other species of Nangra reach the 90 to 100 mm 
size attained by these two species. All specimens 
ofN carcharhinoides examined by us have only 
6 branched dorsal fin rays and 8 branched pecto- 
ral fin rays. These are consistently different from 
the 7 dorsal rays and 10 or 11 pectoral rays 
reported by Mirza and Awan (1983). These dif- 



ferences, in combination with the more slender 
body of N. carcharhinoides, and the disjunct 
distribution of the two forms, lead us to consider 
them as separate species. 

Acknowledgments 

The first author's (T. R. R.) Asian fieldwork 
has been supported by grants from the Smith- 
sonian Tropical Research Institute, the Institute 
for Biological Exploration, and the Committee 
for Research and Exploration of the National 
Geographic Society (grant 5 141-93). Specimens 
from the Fisheries Studies and Pilot Project 
(FAP) 17 fisheries survey of the Tangail district 
of Bangladesh were presented to T. R. R. 
through the kindness of M. A. Wahab and Alan 
Tollervey of the Department of Fisheries Biol- 
ogy and Limnology. Bangladesh Agricultural 
University, Mymensingh. 

The opportunity for the second author 
(C. J. F.) to conduct field work in Myanmar was 
provided by the Food and Agriculture Organiza- 
tion (FAO) of the United Nations, and the Union 
of Myanmar Department of Fisheries. Numerous 
people from each of these organizations provided 
support. Dora Blessish. Kent Carpenter, U Hla 
Win. U Nyi Nyi Lwin. U Myint Pe and U Mya 
Than Tun are recognized by C. J. F. for their 
special efforts. 

Research and writing was done in the Depart- 
ment of Ichthyology of the California Academy 
of Sciences, with space and access to equipment 
provided by William Eschmeyer and Tomio 
Iwamoto. Dong Lin took the black and white 
photographs. David Catania prepared radio- 
graphs and some cleared-and-stained specimens. 
Douglas Nelson provided the specimens from 
UMMZ. and Barbara Brown provided specimens 
and radiographs from AMNH. Specimens were 
examined at NRM and BMNH, with the assis- 
tance of Sven Kullander and Eric Ahlander. and 
Darrell Siebert and Oliver Crimmen, respec- 
tively. Fang Fang (NRM) allowed us to examine 
her recent collection of Brahmaputra River 
fishes. Support for travel to the museums in New 
York, London, and Stockholm was provided by 
the American Museum of Natural History, FAO, 
and the In-house research fund of CAS, respec- 
tively. This manuscript was improved by reviews 
of one or more drafts by John McCosker, Darrell 
Siebert. and Mario de Pinna. We thank these 
people and organizations for their assistance. 



344 



PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 

Volume 50, No. 14 



Literature Cited 

Ataur Rahman, A. K. 1976. A new species of the 
genus Gaga/a Bleeker from river [sic] of Bangla- 
desh. Bangladesh Jour. Biol. Sci. 5(l):4-8. 



. 1990. Freshwater fishes of Bangladesh. The 

Zoological Society of Bangladesh, Dhaka. Pp. 
1-364. 

BLEEKER, P. 1858. Ichthyologiae Archipelagi Indici 
Prodromus. Vol. 1, Siluri. Batavia. Pp. i-xii + 
1-370. 

— . 1863. Systema silurorum revisum. Nederl. 



Tijds. Dierk. 1:77-122. 

BOESEMAN, M. 1966. A new sisorid catfish from Java, 
Sundagagata robusta gen. et spec. nov. Proc. Kon- 
inkl. Nederl. Akad. Wetensch. (Amsterdam), ser. C 
69(2):242-247 + I table. 

Burgess, W. E. 1989. An atlas of freshwater and 
marine catfishes. T. F. H. Publ., Neptune City, New 
Jersey. Pp. 1-784. 

Day, F. 1877. The fishes of India, being a natural 
history of the fishes known to inhabit the seas and 
fresh waters of India, Burma, and Ceylon. Bernard 
Quarich, London. Part 3:369-552, pis. 79-138. 

ESCHMEYER, W. E. 1990. Catalog of the Genera of 
Recent Fishes. California Academy of Sciences, 
San Francisco. Pp. i-v + 1-697. 

GUNTHER, A. W. C. T. 1 864. Catalogue of the fishes 
of the British Museum. 5. Catalogue of the Physos- 
tomi, containing the families Siluridae, Characini- 
dae, Haplochitonidae, Sternoptychidae, Scopelidae, 
Stomiatidae in the collection of the British Museum. 
Trustees of the British Museum, London. 5:i-xxii +