The Biological Investigation

of Malpelo Island, Colombia

JEFFREY B. GRAHAM

EDITOR

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 176
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 17 6

The Biological Investigation

of Malpelo Island, Colombia

Jeffrey B. Graham

EDITOR

SMITHSONIAN INSTITUTION PRESS

City of Washington

1975
ABSTRACT

Graham, Jeffrey B., editor. The Biological Investigation of Malpelo Island,

Colombia. Smithsonian Contributions to Zoology, number 176, 98 pages, 35

figures, 1975.—The results of joint Smithsonian Institution and U. S. Navy terrestrial and marine investigations of Malpelo Island, Republic of Colombia are

reported in 15 papers in this volume. A new species of lizard (Phyllodactylus),

a new starfish (Tamaria), two new species of shrimp (Alpheus and Synalpheus),

and a new species of fish (Chriolepis) are described. The terrestrial ecology of

Malpelo and the behavior and natural history of the lizards Anolis agassizi and

Diploglossus millepunctatus are described and discussed. Genie variability in

A. agassizi has been investigated and karyotypes of A. agassizi and D. millepunctatus are reported. The ecology of the island's benthic marine communities is

detailed and papers listing and discussing zoogeographically interesting features

of the island's crustacean, starfish, and fish species are included. The geology of

Malpelo is briefly described and an improved map of the island is presented.

The importance of Malpelo Island in the understanding of biogeographic problems in the eastern tropical Pacific Ocean is reviewed.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5095. SERIES COVER

DESIGN: The coral Montastrea cavemosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Main entry under title:

The Biological investigation of Malpelo Island, Colombia.

(Smithsonian contributions to zoology, no. 176)

Supt. of Docs, no.: SI 1.27:176.

"The results of joint Smithsonian Institution and U.S. Navy terrestrial and marine investigations

. . . are reported in 15 papers in this volume." I. Natural history—Malpelo Island. 2. Marine

biology—Malpelo Island. 3. Malpelo Island. I. Graham, Jeffrey B., ed. II. Smithsonian

Institution. III. United States. Navy. IV. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 176. [DNLM: 1. Ecology. 2. Lizards. 3. Marine biology. W1SM454N

no. 176 1974/QL244 B6134 1974]

QL1.S54 no. 176 [QH121] 591'.08s [WQ.V%6V5\ 74-23%

For sale by (he Superintendent of Documents,, U.S. Government Printing Office

Washington, D.C. 20402 - Price $2.20 (paper cover)
Contents

Page

INTRODUCTION, by Jeffrey B. Graham 1

TERRESTRIAL BIOLOGY OF MALPELO ISLAND: A HISTORICAL REVIEW,

by George C. Gorman and Terence L. Chorba 9

RECONNAISSANCE AND MAPPING OF MALPELO ISLAND, by A. Ross Kiester

and Jeffrey A. Hoffman 13

FIELD OBSERVATIONS ON THE GEOLOGY OF MALPELO ISLAND,

by Jeffrey A. Stead 17

THE ECOSYSTEM ON MALPELO ISLAND, by Henk Wolda 21

NATURAL HISTORY, BEHAVIOR, AND ECOLOGY OF Anolis agassizi,

by A. Stanley Rand, George C. Gorman, and William M. Rand 27

NOTES ON THE NATURAL HISTORY OF Diploglossus millepunctatus

(Sauria: Anguidae), by A. Ross Kiester 39

A NEW GECKO FROM MALPELO ISLAND (SAURIA: GEKKONIDAE:

Phyllodactylus), by Raymond B. Huey 44

ELECTROPHORETIC ESTIMATES OF GENIC VARIATION IN, AND THE

RELATIONSHIPS OF, Anolis agassizi, by T. Preston Webster 47

NOTES ON THE CHROMOSOMES OF Anolis agassizi (SAURIA: IGUANIDAE)

AND Diploglossus millepunctatus (Sauria: Anguidae), by Brad Stamm

and George C. Gorman 52

SWBTIDAL COMMUNITIES OF MALPELO ISLAND, by Charles Birkeland,

David L. Meyer, James P. Stames, and Caryl L. Buford 55

THE MACRURAN DECAPOD CRUSTACEA OF MALPELO ISLAND,

by Lawrence G. Abele ; 69

ASTEROIDEA FROM MALPELO ISLAND WITH A DESCRIPTION OF A NEW

SPECIES OF THE GENUS Tamaria, by Maureen E. Downey 86

FISHES COLLECTED AT MALPELO ISLAND, by John E. McCosker and

Richard H. Rosenblatt 91

A NEW SPECIES OF GOBY FROM MALPELO ISLAND (TELEOSTEI:

GOBIIDAE: Chriolepis), by Lloyd Talbott Findley 94

i n

The Biological Investigation

of Malpelo Island, Colombia

Introduction

Jeffrey B. Graham

On 28 February 1972 a group of 17 scientists

representing the Smithsonian Tropical Research

Institute, the Republics of Panama and Colombia,

and several U.S. universities embarked on a six

day expedition to investigate the marine and terrestrial biota of Malpelo Island, Colombia, a small

isolated Pacific island 270 miles to the west of Colombia and south of Panama. The expedition, a

joint project of the U.S Navy and the Smithsonian

Tropical Research Institute, was undertaken to

gain a more comprehensive understanding of

ecological processes and the natural history of

plants and animals on Malpelo. The scientific

party consisted mostly of biologists who made collections, carried out field observations, and conducted experiments on the island and in the waters

around it. (The Malpelo scientific party consisted

of C. Birkeland, J. B. Graham, D. L. Meyer, A. S.

Rand, A. Rodaniche, W. L. Smith, J. P. Stames,

and H. Wolda, all of the Smithsonian Tropical

Research Institute (STRI); O. Arroyo, Colombian

Institute of Natural Resources (INDERENA); J. Barreto S., Universidad George Tadeo Lozana, Colombia; G. C. Gorman, University of California Los

Angeles (UCLA); J. A. Hoffman, Smithsonian Astrophysical Laboratory; A. R. Kiester, Museum

of Comparative Zoology; D. B. Macurda, University of Michigan; W. M. Rand, Massachusetts

Institute of Technology (MIT); T. L. Chorba,

Jeffrey B. Graham, Smithsonian Tropical Research Institute,

P. O. Box 2072, Balboa, Canal Zone.

U.S.S. York County; J. A. Stead, U.S. Navy Hydrographic Center. R. J. Kinney, USN, served as expedition physician. The geology of Malpelo was

studied and a more accurate map of the island was

made by an exploration team.

The scientists were transported from Panama to

Malpelo by the U.S.S York County (LST 1175)

commanded by LCDR Lauren Seeber. At Malpelo,

the scientists were supported by York County crewmen who did everything from prepare box lunches

to operate small boats and assist in the collection

and sort of specimens. While on station the York

County conducted sea trials and made numerous

depth soundings and positional sightings of the

island. This work, while helping to establish that

the peaks on Malpelo are some 125 meters higher

than previous maps indicated, will also improve

the hydrographic charts of this area.

The Smithsonian-U.S. Navy Expedition to Malpelo marked the fourth time that Smithsonian

scientists and their colleagues have been able to

use a U.S. Navy vessel for research purposes. In

March 1970 biological investigations of the Secas

Islands in the Gulf of Chiriqui, Panama, were conducted by scientists aboard the U.S.S. Traverse

County. Later in the same year the U.S.S. Walworth County supported marine studies at Coiba

Island, Panama. In 1971 the U.S.S. Terre Bonne

Parish carried a research team to Bocas del Toro,

Panama, for three days of field work. In the summer of 1973, while the volume on Malpelo was
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

being compiled, the fifth and most ambitious expedition was undertaken when the U.S.S. Spartanburg County took 25 scientists to Cocos Island,

Costa Rica, 550 miles southwest of Panama.

These expeditions have been fruitful. The largest and best developed coral reef formations to be

found thus far in the eastern tropical Pacific were

discovered and subsequently studied by Peter

Glynn and colleagues during the Gulf of Chiriqui

trip (Glynn, Stewart, and McCosker, 1972). Previous to these investigations it was thought that no

well-developed coral reefs occurred in the eastern

tropical Pacific. The coral-eating starfish Acanthaster planci was found in great abundance and

several species of Indo-Pacific corals were also discovered by Glynn and his co-workers during the

same expedition. Our understanding of the faunal

composition and zoogeography of eastern tropical

Pacific fishes has benefited significantly from Navysupported research in this region (Rosenblatt,

McCosker and RubinofF, 1972).

The use of Navy ships is a valuable addition to

the research capacity of the Smithsonian Tropical

Research Institute. Navy vessels allow many scientists to participate in the expeditions and it is

possible to visit localities beyond the range of small

research vessels. Needless to say, costs and the

logistical support problems are also vastly reduced.

ACKNOWLEDGMENTS.—It is a pleasure to acknowledge Admiral C. H. Griffiths, Commander of the

United States Navy, Fifteenth Naval District, Fort

Amador, Canal Zone, and his staff who made

arrangements for the Malpelo Expedition. We also

thank Captain Lauren Seeber and the officers and

men of the U.S.S. York County for supporting our

investigations. We express our appreciation to the

Foreign Ministry of the Republic of Colombia for

permitting us to visit Malpelo Island and to the

U.S. Army Southern Command, 352nd Aviation

Detachment, for providing an aerial reconnaissance

of the island. I would like to thank my colleagues:

L. G. Abele of Florida State University, D. Dexter

of San Diego State University, J. McCosker of the

California Academy of Sciences, and A. S. Rand,

I. RubinofF, and R. R. Warner, all of the Smithsonian Tropical Research Institute, for their advice and assistance in the preparation of this volume; I acknowledge the technical assistance of

I. Downs, A. Kourany, and V. Vergel of the Smithsonian Tropical Research Institute.

Island Biology and Its Biogeographical

Significance

The biological investigation of islands has

made a valuable contribution to the studies of

evolution, biogeography, and ecology. Islands, by

virtue of their differences in size, shape, climatic

conditions, and distances from continents, present

a stepped progression of natural experimental

areas (MacArthur, 1972). Darwin's ideas on the

natural variability and selection of species developed in part through his observations at the Galapagos Islands. Some biologists investigate islands

to test different aspects of modern evolutionary

theory. Islands, however, are intrinsically interesting because each is unique, and they afford biologists the chance to study the effects of isolation

on the resident species. Island populations may

have greatly expanded or otherwise modified

niches and may have decreased levels of genetic

variability or, if isolated for a long period, endemic

species may be formed (MacArthur and Wilson,

1967).

Malpelo Island (3°51'07"N, 81°35'40"W) is one

of several oceanic islands in the eastern tropical

Pacific. It is the only island on the Malpelo Ridge

(Figure 1) a solitary volcanic submarine ridge that

extends in a northeast-southwest direction with a

length of 150 miles and a width of 50 miles (Chase,

1968). An oceanic island, Malpelo occurs outside

the continental shelf and it is separated from the

mainland of Central America by depths greater

than 1800 fathoms. Malpelo has never been connected, even by shallow water, with any other

islands or the mainland.

Malpelo's distance from the mainland and the

depths of water between the island and the mainland are significant barriers to the colonization of

both terrestrial and shallow-water marine organisms. Colonization therefore must occur through

sea surface currents which would transport rafting

terrestrial organisms and the planktonic larvae and

subadults of marine forms to the island. Malpelo

is closer to the mainland than Cocos and the Galapagos islands (Figure 2) and it is reasonable to

assume it serves as a stepping stone for marine

forms enroute, via oceanic currents, to these

islands.

Consideration of the complex pattern of surface

circulation in the eastern tropical Pacific Ocean
NUMBER 176

(Wyrtki, 1965) provides insight as to how surface

currents may serve for transport from the mainland

to Malpelo and beyond to Cocos and the Galapagos islands. Only general features of surface

circulation in the area of these islands for the

months of January and July are presented (Figure

2) and the reader is referred to Wyrtki's paper for

details on monthly current vectors, velocities, and

the persistence of flow for the different areas in the

eastern tropical Pacific.

In the months from January to March the tradewinds cause a strong current flow (0.5 knots) out

of the Gulf of Panama towards Malpelo and the

Galapagos islands (Figure 2). A small anticyclonic

gyre that borders the mainland and encompasses

Cocos and Malpelo islands is also formed at this

time. If winds blow strongly for long periods the

condition known as "El Nino" occurs. This results

in the transport of a larger than normal amount

of warm surface water, which displaces the Peru

current to the southwest away from South America

and the Galapagos Islands. During £1 Nino the

Galapagos Islands may be completely bathed in

warm tropical waters.

The Equatorial Countercurrent (Figure 2) does

not penetrate very far into the eastern tropical

Pacific from January to April, but from May to

December this current is strong and reaches the

FIGURE 1.—Submarine topography of the central eastern tropical Pacific Ocean in the vicinity

of Malpelo, Cocos, and the Galapagos islands. The lines are 600, 1000, and 1200 fathoms. Data

are from Chase (1968).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

20*

10*

NORTH EQUATORIAL CURRENT

EQUATORIAL COUNTERCURRENT

SOUTH EQUATORIAL CURRENT

t t C

120' 110 IOO« 90' 80*

20*

70*

10 <

NORTH EQUATORIAL CURRENT

EQUATORIAL COUNTERCURRENT

SOUTH EQUATORIAL CURRENT

FIGURE 2.—Surface currents around the major oceanic island in the eastern tropical Pacific

Ocean. The islands indicated are Clipperton (CI), Cocos (C), Galapagos (G), and Malpelo (M).

Data are from Wyrtki (1965).
NUMBER 176

coast of Central America where it divides. The

major part of the Equatorial Countercurrent

flows northwest along Central America, while the

remainder flows south and then turns west as part

of the South Equatorial Current. From May to

December Clipperton, Cocos, and Malpelo islands

lie in the swath of the Equatorial Countercurrent,

and the Galapagos Islands occasionally receive

water from the southern part of this current from

August to October. The combination of strong

southern surface flow in the months from January

to May and the circulation pattern of the Equatorial Countercurrent probably account for the dispersal of mainland marine organisms to these

islands.

Another phenomenon that is of special interest

to marine zoogeographers is the presence and dispersal patterns of Indo-Pacific organisms (corals,

mollusks, crustaceans, and fishes) at the eastern

tropical Pacific islands and on the Central American mainland (Costa Rica and western Panama).

This has been recently reviewed for fishes by

Rosenblatt, McCosker, and Rubinoff (1972). Larvae of these animals are apparently transported

across 1000 miles of open sea by the Equatorial

Countercurrent, which, as was shown above, extends to the mainland of Central America and

sweeps across some oceanic islands for long periods

of the year. It is not presently known how many

species have successfully completed this transit or

whether they more frequently colonize oceanic islands or the mainland.

Summary of the Malpelo Expedition

The amount of information that was accumulated in the four full days of field work on Malpelo

is gratifying. Part of the expedition's success lay in

what was learned about the island by its earlier

explorers. Gorman and Chorba have traced the

history of Malpelo's exploration, which goes back

to 1542. Early visitors to Malpelo had the impression that the flow of energy to the terrestrial ecosystem had to be sustained by the sea and that this

was done mainly by birds. While acknowledging

that the success of the land crab (Gecarcinus) and

the large lizard (Diploglossus) definitely depends

on the activity of birds, Wolda has found that the

terrestrial community is much too large and complex for this to be the only pathway. He concludes

that lichens and mosses constitute the important

lower trophic levels for the land community.

Malpelo's endemic lizards were well studied on

this expedition. A new species of gecko (Phyllodactylns transversalis) was discovered and this is

described by Huey. This species, while similar in

many ways to Peruvian species has its closest affinity

to Mexican Phyllodactylus. The depth of biological knowledge about mainland and island Anolis,

together with their extensive research experience

with anoles enabled Rand, Gorman, and Rand to

ask specific questions about the ecology and behavior of Malpelo's A. agassizi. The Malpelo anole

is larger than most other anoles isolated on islands

and it reproduces seasonally. Its social behavior is

strikingly different from that of other Anolis; it

is not territorial, there is little aggression, and its

displays are few and simple. Anolis agassizi is an

active, curious lizard that is readily attracted to

things that appear in its habitat: scientists, their

equipment, and their fruit and candy as well.

Their curiosity and the remarkable tolerance of

individuals for each other, these authors believe,

enables A. agassizi to take advantage of resources

such as water and food that have variable distributions in time and space on Malpelo. Kiester describes Diploglossus millepunctatus as curious

and opportunistic and draws basically the same

conclusions about the ecological significance of

this behavior as were made for A. agassizi. Interestingly, Kiester postulates that D. millepunctatus,

because of its dependence on birds, may have its

reproductive seasonality synchronized with that

of the nesting seabirds. This awaits further study.

Webster measured a low degree of genie variability

in A. agassizi. From what is presently known this

seems to be a characteristic of old, solitary anole

populations. Stamm and Gorman found that the

karyotypes of A. agassizi and D. millepunctatus

are similar to the more primitive groups in these

two genera. Taken together these observations

might suggest that Malpelo's Anolis and Diploglossus have been in residence for a very long time.

But as Webster and Stamm and Gorman point out,

our present knowledge is too limited to say this

with certainty.

Kiester and Hoffman surveyed the island and,

with special climbing equipment, scaled its summits. Sightings from the peaks and from the bridge

of the York County enabled them to determine
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Left: Aerial view (4000^ of Malpelo Island. The northern end of the island is in

the foreground. Right: A York County landing craft approaching the west side of Malpelo

Island.

that the highest elevations of Malpelo are almost

125 meters higher than early maps indicated. Stead

conducted a geological survey of Malpelo. His

description of the island's weathering, together

with a compilation of the York County's depth

soundings around Malpelo enables us to appreciate the original size of this rock. Once 8 to 10

times its present size, Malpelo has been steadily

worn by the persistent pounding of the sea. When

we brought our small boats close to shore and

tried to land on the island we keenly sensed the

intensity of the struggle between Malpelo and the

sea (Figure 3).

Rough water and the island's steep slope made

getting from the boat to shore and returning a

major undertaking for the terrestrial party. Exploration ended prematurely for one scientist in

this group who severly injured his arm while trying

to get from the island into the boat. In spite of

these obstacles none in the terrestrial group elected

to spend their nights on Malpelo and, at days end,

all were glad to return to the haven of the York

County. The warship took on a curious appearance

at this time of day. Its main deck was littered with

biological specimens and collecting equipment and

was crowded with seamen who were rapidly transformed into eager assistants, audiences, or both.

The crewmen relished this and took great pleasure

in hearing about the discoveries and adventures

of the day. Accounts were passed rapidly to the

sailors on watch and expedition progress reports

were included in the ships daily bulletin. News of

the discovery of the new Phyllodactylus species at

Malpelo may have traveled faster than any other

sjiecies discovery in history as it was flashed the

same day to the ship's home port at Little Creek,

Virginia. Sorting, preservation, and labeling continued past dark while at the same time, in other

parts of the ship, other men worked, and scuba

tanks, thermos bottles, and lunch containers were

refilled for use the next day. When work was

finished for the day, showers, warm food, a recounting of the day's adventures, a soft bed, and last,

but by no means least, the evening movie fully

restored us for the next day.

Investigation of Malpelo's marine habitat was

also rewarding. The water was so rough, however,

that the small boats we had brought for the diving

teams were unsafe and diving operations had to

be staged from one of the York County's boats. We

initially tried to use the ship's landing crafts but

these had so much freeboard that they were not

practical. After struggling through a day's diving

from the landing craft, I related this difficulty to

Captain Seeber, who immediately offered us the

use of his launch. Thus it came to be that the

elegantly furnished Captain's Gig became our

diving boat. No doubt the launch has seen better

days both before and since we loaded it with divers,

their gear, and specimens. But it proved an invaluable asset at Malpelo. There were many sharks

around the island; they were seen at the surface

and on every dive. No particularly dangerous encounters occurred with sharks during our dives,
NUMBER 176

but their omnipresence was sobering and at times

their proximity was slightly unnerving.

Birkeland and colleagues completed their ambitious project of studying and characterizing the

subtidal benthic communities at Malpelo. The

smooth vertical subtidal walls are dominated by

large, sparsely distributed barnacles. Recruitment

success for transforming barnacle larvae is low but

once they are established, adults may live for a

long time. On the gently sloping southeastern edge

of Malpelo, Birkeland's group found extensive

coral development. Although they do not constitute a true coral reef, these corals are luxuriant

and depth zonation is marked for different species.

The occurrence of corals to as deep as 30 meters,

about the deepest known in the eastern tropical

Pacific, is attributable to the clear water around

Malpelo. A list of all the marine invertebrates

known from Malpelo is appended to the Birkeland

et al. paper. Abele records 43 species of decapod

Crustacea (18 species of macrurans) at Malpelo

and describes two new species of snapping shrimp.

Abele lists the distribution of each species and, by

drawing upon his other recently completed studies,

he makes an interesting observation about the

species diversity of decapod crustaceans at Malpelo,

Cocos Island, and several other eastern tropical

Pacific localities. The diversity of starfish at Malpelo is low and Downey lists only six species in

her paper. Three species occur throughout the

eastern tropical Pacific and a new species of Tamaria and a new subspecies of Narcissia are described for Malpelo. Until Downey's work, one

Malpelo starfish (Leiaster) was previously known

to occur only at the Hawaiian Islands.

Our knowledge of the Malpelo fish fauna is still

incomplete but McCosker and Rosenblatt report

that 70 species of fish are now known for the island.

In the specimens collected in the SmithsonianNavy Expedition, McCosker and Rosenblatt found

several new species, as well as a new genus of fish;

all awaiting description except the new species

of goby (Chriolepis) described by Findley. The

Abele, and McCosker and Rosenblatt papers demonstrate several interesting features that the decapod crustaceans and fishes of Malpelo have in

common. The majority of species in both groups

are Central American mainland forms; there are

also Indo-Pacific species and some endemics. Both

groups contain species that were previously

thought to be endemic at Cocos or the Galapagos

islands or were known only from these two localities. These findings strongly suggest that Melpelo is a stepping-stone for marine mainland

species enroute to more distant islands and, as the

surface current patterns indicate, for Indo-Pacific

species as well. Some of the terrestrial invertebrates

listed by Wolda appear to be most closely related

to Indo-Pacific groups, which prompts the conclusion that these were rafted from the old world

to Malpelo via the Equatorial Countercurrent.

The exchange of species between Malpelo, Cocos,

and the Galapagos Islands is implied by their similar fish and decapod faunas and by the marked

faunal similarity observed by Birkeland et al. between Malpelo and the Galapago Islands. Further

work with these faunas will clarify these relationships.

Although much was learned at Malpelo the

short time for study has left questions and the biota

of the island would doubtlessly be better understood if more sampling, observation, and experimentation could be conducted. It was agreed that

additional visits should be made, ideally at different times of the year so that the possible influence of seasons could be assesed. That there is

much to learn is evident when it is pointed out

that we still do not know how much rain annually

falls on Malpelo. Murphy (1945) suggested that

the island never receives rain, but Mapelo does

lie within the range of the convergence of the trade

winds (Intertropical Convergence Zone, Wyrtki,

1965), which suggests a seasonal rainfall. The island does have weathered, phosphatized rock

(McConnell, 1943), and seepage marks, dry

stream beds, and pools of water were found by the

Smithsonian-Navy expedition. All of these suggest

that rain is not uncommon but expeditions to the

island have practically all been during the months

of the dry season (December to March) and we

can only speculate about amounts of rain in the

rest of the year.

This volume, by reviewing and compiling Malpelo's literature and by reporting results to date

of the most thorough investigation of the island

sets the stage for future exploration. It describes

Malpelo but it cannot convey the impressive

scenario of that island. When viewed objectively

it is an isolated, inhospitable island, which by these
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

virtues has remained a natural laboratory for the

study of evolution.

When seen first hand, however, Malpelo is a

large grotesque rock that rises rudely, ominously

out of a monotonous sea. As we contemplated this

rock we reminded ourselves, and took pleasure in

the fact, that it was a sight that few people have

ever seen.

Literature Cited

Chase, R. E.

1968. Sea Floor Topography of the Central Eastern

Pacific Ocean. US. Bureau of Commercial Fisheries,

circular 291, 33 pages.

Glynn, P. W., R. H. Stewart, and J. E. McCosker

1972. Pacific Coral Reefs of Panama: Structure, Distribution, and Predators. Sonderdruck aus der Geologischen Rundschau, 61:483-519, 15 figures.

MacArthur, R. H.

1972. Geographical Ecology: Patterns in the Distribution

of Species. New York: Harper and Row.

MacArthur, R. H., and E. O. Wilson

1967. The Theory of Island Biogeography. Princeton:

Princeton University Press.

McConnell, D.

1943. Phosphatization at Malpelo Island, Colombia. Bulletin of the Geological Society of America, 54:707-

716, 2 plates.

Murphy, R. C.

1945. Island Contrasts. Natural History, 15:14-23.

Rosenblatt, R. H., J. E. McCosker, and I. Rubinoff

1972. Indo-West Pacific Fishes from the Gulf of Chiriqui,

Panama. Contributions in Science of the Los Angeles County Museum, 234:1-18, 3 figures.

Wyrtki, K.

1965. Surface Currents of the Eastern Tropical Pacific

Ocean. Bulletin of the Inter-American Tropical

Tuna Commission, 9:271-304.
Terrestrial Biology of Malpelo Island:

A Historical Review

George C. Gorman

and Terence L. Chorba

ABSTRACT

Records of the exploration of Malpelo Island date

back to 1542. Early biological investigations mainly

concerned the terrestrial fauna, particularly the

island's lizards. The complete list of Malpelo's

avifauna is presented. The etymology of Malpelo

is discussed.

That the biology of Malpelo would be poorly

known should surprise few people. The island

couples geographic isolation with hazardous accessibility. It also lacks spectacularly unique denizens

such as Komodo dragons or giant tortoises. Thus

interest in Malpelo is primarily restricted to the

dedicated naturalist who can appreciate that the

very isolation makes it a laboratory of evolution,

and who realizes that a species need not be the

largest or the strangest to be of genuine biological

interest. The joint Smithsonian-U.S. Navy Expedition to Malpelo was not the first to visit this unique

corner of the world; others have stopped on the

island, perhaps while enroute to the Galapagos, or

while collecting marine life off the coast of South

America. Our expedition probably was the first to

be dedicated to the Malpelo ecosystem per se, to

attempt to collect and identify as many of the

living forms as was feasible, to examine the behavior and ecology of at least some of the species,

George C. Gorman, Department of Biology, University of

California, Los Angeles, California 90024. Terence L. Chorba,

student in Final Honour School of Psychology, Philosophy,

and Physiology, Oxford University, and member of Brasenose

College, Oxford, England.

and to attempt to understand the flow of energy

in the system. The expedition will have been successful if, for no other reason, others are inspired

to visit this unique massive rock to pursue further

some of the interesting problems raised in these

pages.

The first record of a landing on Malpelo that

we know of took place in the late 18th century.

The Spanish Commodore, Alejandro Malaspina

(in Malaspina and Bustamente, 1885:551) quoting one of his officers wrote:

One of the shipmasters in the local trade, favoured by fine

weather and a smooth sea, once landed on a shelf of rock on

the north face of the island, and after climbing thirty steps

down hewn out by hand, came upon a large pool of rainwater, which was not overclean and had some bird feathers

in it; but ... . he used the opportunity to fill some casks.

This ... . is all that can be said about Malpelo.

At the time there was more that could have been

said, but Malapsina either was not familiar with

any previous mention of the island by the chroniclers, or did not consider what they said to be of

sufficient importance.

Pedro de Cieza de L£on (1881:92) wrote that

the Spanish colonial administrator in Peru, Cristobal Vaca de Castro, en route from Panama to

Buenaventura in 1542 "came to a rocky islet which

the sailors call Mai Pelo."

Literally, Malpelo means "bad hair." The origin

of this name is unknown but the island appears on

a parchment map of the world of 1550 (Desceliers,

1550) that is in the British Museum. The name on

the map is "ye mallabry." The term "malabrigo"

was frequently used by cartographers to designate

islands and bays. Literally, "malabrigo" means
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

shelterless. Such a name would be well deserved.

Another possibility, however, is that the word

"Malpelo" has roots in the Latin malveolns (literally "inhospitable" or "spiteful"), which in the

vernacular could have degenerated to "malbolo"

and at some point been interpreted as "malpelo."

At the time that Vaca de Castro sailed past

Malpelo, though perhaps unknown to the Crown,

the island was a territory of Spain by virtue of the

Papal Bull issued in 1493 by Pope Alexander VI,

and the Treaty of Tordesillas (1494) under which

Spain and Portugal divided the non-Christian

world in half. Subsequently, Malpelo passed automatically to Peru and later to Colombia without

dispute. Inaccessible, uninhabitable, lacking arable

land and suitable anchorage, it is not the kind of

real estate for which one battles furiously.

One of the first scientists to discuss life on Malpelo was C. H. Townsend (1895), who visited

the island in March 1891 on board the U.S. Fish

Commission Steamer Albatross. During the brief

visit, Townsend collected a Swallow-tailed Gull

(Creagrus furcattis) and noted that seabirds of

several species swarmed about the inaccessible

summit.

Townsend also collected the first reptiles with

the locality data "Malpelo Island." These lizards

were described as Anolis agassizi (Iguanidae) by

Stejneger (1900:163), who reported:

Mr. Charles H. Townsend, who collected these specimens in

Malpelo, informs me that they were running over the rocks

near the water. The island was too steep to afford a landing,

but the lizards were shot off or whisked off the face of the

cliffs, thus falling into the water, whence they were secured

by the collector.

Two years prior to the publication of Townsend's report, Faxon (1893) described the common

land crab of Malpelo as Gecarcinus malpilensis

(Gecarcinidae) from a single male specimen collected by the same expedition in which Townsend

collected the lizards. No notes other than a description were provided.

In November and December of 1927, Captain

A. G. Hancock made a trip to the Galapagos. J. R.

Slevin of the California Academy of Sciences was

along, and when the boat passed near Malpelo,

Slevin and a colleague were able to effect a landing

from a small skiff on the northeast corner of the

island. They collected 27 Anolis (a medium-sized

lizard) and 10 specimens of a large anguid lizard

which Slevin (1928) named Celestus hancocki. He

claimed that this lizard was most closely related to

Celestus millepunctatus O'Shaughnessy known

from a single specimen of unknown provenance,

but that it differed from C. millepunctatus in

several features of scutellation and limb development.

Slevin did not explore the island. All his specimens of Celestus were taken within a few feet of

the water's edge on a rocky ledge. When wounded,

he claimed that they would take to the water.

Stomach contents consisted of feathers and remains

of crabs.

The George Vanderbilt South Pacific Expedition

of 1937 stopped at Malpelo in February. Some field

notes by the Vanderbilt's collector Ronald Smith

were published by Fowler (1938).

Smith was confused in his taxonomy, since he

believed that there were five species of lizards

roaming about. But from his descriptions it is obvious that there were only the two known forms.

Sexual dimorphism, and perhaps distinctive size

classes led to his erroneous conclusion. Since we

can easily determine when Smith was referring to

Anolis, and when to Celestus, we might mention

a few of his ecological observations. He claimed

that the Anolis was found throughout the island,

all the way to the summits, but that the Celestus

was limited to the height of 150 meters. He also

reported a habitat difference. The anoles were

seen mostly amongst loose rocks and exposed surfaces, whereas the Celestus were found primarily

in small cave-like formations. The lizards were

described as fearless and quite bold.

The Vanderbilt expedition collected four specimens of Anolis and seven Celestus, and this served

as impetus for Dunn (1939) to reconsider the

lizard fauna of the island. He made two major

decisions at the generic level, one that has stood

the test of time, the other that has not. Dunn

pointed out that the genus Celestus could not be

maintained as distinct from Diploglossus. He assigned the large anguid to Diploglossus, the generic

name that it still bears. More interestingly, he

demonstrated that the previously described C. millepunctatus was not distinct from Slevin's C. hancocki. Thus the specimen described by O'Shaughnessy in the mid-nineteenth century was almost

certainly obtained from Malpelo, although no records of an expedition to the island have been
NUMBER 176 11

found. The second decision was to raise the anole

to a distinct genus (Mariguana). This decision

has not been accepted (Etheridge, 1960).

Bond and deSchauensee (1938) presented an

annotated checklist of the birds of Malpelo based

upon the observations and collections of the Vanderbilt expedition and incorporating data from

Townsend (1895). It is reprinted below in its

entirety.

ANNOTATED LIST OF THE BIRDS

OF MALPELO ISLAND

1. Sula dactylatra granti Rothschild. (Sulidae, Pelicaniformes) Masked Booby. Two adult males were collected

February 8, and an immature female February 9.

2. Fregata sp. (Fregatidae, Pelicaniformes). None collected

but Smith notes: "Man o'War Birds 50; small colony on

one of the outlying rocks on the southwest of the island."

3. Falco peregrinus anatum Bonaparte. (Falconidae, Falconiformes) Duck Hawk. Not collected but two birds

observed twice by Smith on Malpelo Island.

4. Actitis macularia (Linnaeus). (Scolopacidae, Charadriiformes) Spotted Sandpiper. Not collected but "ten seen

in company with tattlers."

5. Heteroscelus incanus (Gmelin). (Scolopacidae, Charadriiformes) Wandering Tattler. One male and two females,

in winter plumage, were collected February 8 and 9.

About twenty-five individuals were seen.

6. Creagrus furcatus (Neboux). (Laridae, Charadriiformes)

Swallow-tailed Gull. An adult male and female as well

as a nestling were collected February 8 and 9. Smith

records a colony of thirty birds on the island.

7. Anous stolidus ridgwayi Anthony. (Laridae, Charadriiformes) Noddy. An adult female, the only specimen

seen, was secured February 9. It was not in breeding

condition.

8. Anous minutus diamesus (Heller and Snodgrass). (Laridae, Charadriiformes) Black Noddy. An adult male collected February 9, was the only specimen seen. The bird

was not in breeding condition.

9. Hirundo rustica erthrogaster Boddaert. (Hirundinidae,

Passeriformes) Barn Swallow. An immature female was

secured February 8. Four birds were seen flying out of

a cave.

10. Progne sp. Martin. (Hirundinidae, Passeriformes). Recorded by Townsend but none secured. Smith did not see

any martins.

During March 1941, the schooner Askoy approached Malpelo and Dr. Robert Cushman

Murphy and another member of his party made a

landing. His popular article (Murphy, 1945) provides the best summary of the natural history of

the island. He estimated the booby population at

about 25,000; added an additional species to the

bird list of Malpelo (Red Billed Tropic Bird,

Phaethon aethereus); examined the stomach contents of a gull and found that it was eating

marine crabs; and, most important, noted the presence of a diversity of invertebrates including

spiders, pseudoscorpions, beetles, flies, ants, and

other insects. Samples of Malpelo rocks were obtained by Murphy and a study of phosphatization

at Malpelo was published by McConnell (1943).

The land crabs (Gecarcinns) impressed Murphy

(1945:16) who described them as

ghoulish in appearance . . . , fat and bloated looking creatures with shells of a ghostly white. They may be quite

harmless toward human beings, yet they seem to eye a visitor

with an intent that grows uncomfortable. If you sit long

enough the crabs will move up closer and closer, as if with

whetted appetites, and I have a feeling that they would

make life miserable for anyone who had to sleep ashore.

Garth (1948) provided important information

on this crab. He did not agree with Faxon (1893)

that it was an endemic species, but considered it

G. planatus Stimpson, a species widespread from

Baja California to Acapulco, that has successfully

colonized the Revillagigedo Islands and Clipperton, but that is notably absent from the Galapagos.

There have been additional landings on Malpelo, some documented by specimens. For example,

the San Diego Society of Natural History has

lizards collected in December 1931 and January

1933, by Cyrus Perkins. There is, however, little

additional published information about the ecology and natural history of the island or its inhabitants. What inspired the present expedition? Why

travel hundreds of miles to visit a barren rock?

The very harshness of the environment raises interesting questions. In addition to sea birds, and a

few nonmarine birds that are very strong flyers but

presumably not residents, only three common terrestrial residents, Anolis and Diploglossus (lizards)

and Gecarcinus (a crab) had been described and

identified. What species of plants (if any) and

invertebrates would be found? What are their biogeographic affinities? How does such a community

maintain itself?

Our working assumption was that this was a terrestrial ecosystem dependent upon primary production on the sea and not on the land itself.

Presumably the birds which fed upon marine life

provided food resources for small terrestrial inver-
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tebrates (parasites, carrion eaters, egg predators,

etc.) which in turn provided sustenance for the

lizards.

Several of us were specifically interested in the

Anolis lizards. The genus is comprised largely of

tropical arboreal species. They tend to be highly

territorial, defending fixed perch sites with elaborate displays. Here was an environment devoid of

the typical plant cover that we associate with

Anolis. Did they have any special physiological

adaptations that permitted life on warm, exposed

rocks? Was there any modification of the social

structure of populations in the absence of typical

perch sites that are associated with their territoriality?

These are the kinds of questions that were on

our minds when we first discerned the form of

Malpelo Island, gray itself against the gray of the

early dawn on the sea on 29 February 1972. Answers are what we sought when we braved the steep

swells and jumped off the landing craft to begin

serious exploration of the island and its life.

Literature Cited

Bond, J., and R. M. deShauensee

1938. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part II: The Birds of

Malpelo Island, Colombia. Proceedings of the Academy of Natural Science of Philadelphia, 90:155-157,

plates 9-11.

Cieza de Leon, Pedro de

1881. Guerras Civiles de Peru: Guerra de Chupas. Madrid: Publicado por Dom Pedro de Novo y Colson.

[English translation by Sir Clements Markham and

reprinted by the Hakluyt Society, London, 1918.]

Desceliers, P.

1550. Faicte a Arques . . . Ian 1550. Add. MSS 24065.

British Museum (Natural History), London.

Dunn, E. R.

1939. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part III: The Lizards

of Malpelo Island, Colombia. Notulae Naturae of

the Academy of Natural Sciences of Philadelphia,

4:1-3.

Etheridge, R. E.

1960. The Relationships of the Anoles (Reptilia: Sauria:

Iguanidae): An Interpretation Based on Skeletal

Morphology. Unpublished Ph.D. Dissertation, University of Michigan. Ann Arbor: University Microfilms Inc.

Faxon, W.

1893. Preliminary Descriptions of New Species of Crustacea. Part 6 in Reports on the Dredging Operations off the West Coast of Central America to the

Galapagos, to the West Coast of Mexico, and in

the Gulf of California, in Charge of Alexander

Agassiz, by the U.S. Fish Commissioner Steamer

"Albatross," during 1891, Lieutenant Commander

Z. L. Tanner, U.S.N., Commanding. Bulletin of the

Museum of Comparatix>e Zoology at Harvard College, 24:149-220.

Fowler, H. W.

1938. The Fishes of the George Vanderbilt South Pacific

Expedition, 1937. Monographs of the Academy of

Natural Sciences of Philadelphia, 2: V+1-349 pages,

10 plates. [Malpelo, pages 5-6.]

Garth, J. S.

1948. The Brachyura of the "Askoy" Expedition with

Remarks on Carcinological Collecting in the Panama Bight. Bulletin of the American Museum of

Natural History, 92(l):l-66, 4 figures.

Malaspina, A., and J. de Bustamante

1885. Viaje politico—Cientifico alrededor del Mundo por

las corbietas "Descubierta" y "Atrevida": desde

1789 a 1794. Madrid: Publicado por Dom Pedro

de Nova y Colson.

McConnell, D.

1943. Phosphatization at Malpelo Island, Colombia. Bulletin of the Geological Society of America, 54:707-

716.

Murphy, R. C.

1945. Island Contrasts. Natural History, 54:14-23.

Slevin, J. R.

1928. Description of a New Species of Lizard from Malpelo Island. Proceedings of the California Academy

of Science, fourth series, 16 (21):68I-684, plates 25,

26.

Stejneger, L.

1900. Description of Two New Lizards of the Genus

Anolis from Cocos and Malpelo Islands. Bulletin

of the Museum of Comparative Zoology, 36 (6):

161-164, 1 plate.

Townsend, C. H.

1895. Birds From Cocos and Malpelo Islands, with Notes

on Petrels Obtained at Sea. Bulletin of the Museum

of Comparative Zoology, 27:121-126, 1 plate.
Reconnaissance and Mapping

of Malpelo Island

A. Ross Kiester

and Jeffrey A. Hoffman

ABSTRACT

Daily observations of the exploration party are reported. Early maps of Malpelo show it to have one

peak with a maximum elevation of about 300 m.

The exploration party surveyed and mapped the

island and found that the island has twin peaks,

which are both approximately the same height and

about 150 m higher than previous maps indicate.

An improved map of Malpelo is given. It is believed that the Smithsonian exploration team is

the first to reach the island's summit.

Introduction

An overflight of Malpelo Island, arranged

through the courtesy of the U. S. Army, in preparation for the U. S. Navy-Smithsonian Expedition

to Malpelo showed serious defects in the official

map based on a 1954 aerial survey. The published

U. S. Navy Hydrographic Office map (No. 1685,

North Pacific Ocean, Isla del Coco, Malpelo Island

inset) showed a single major peak of 260 meters

(845 feet) on the island. Observers in the overflight

reported two distinct peaks of nearly the same

height, read as roughly 390 meters (1200 feet) on

the plane's altimeter. Initial observations from the

ship on arrival at Malpelo did not reveal which of

the peaks was higher. They were so close in height

that whichever peak was closest to the ship appeared the highest. A previous expedition had reA. Ross Kiester, Museum of Comparative Zoology, Harvard

University, Cambridge, Massachusetts 02138. Jeffrey A. Hoffman, Smithsonian Astrophysical Observatory, Harvard University, Cambridge, Massachusetts 02138.

ported reaching the summit of the rock, but had

not indicated which of the two summits they considered this to be. Indeed very little general

description of the topography of the island existed

in the literature, and no mention was made of the

existence of the two peaks. Because of this confusion, a general survey of the topography of the

island was one of the expedition's scientific

objectives.

In pursuit of this objective, two members of the

terrestrial party of the expedition extensively explored the entire island, while other members of

the party carried on intensive studies at one site.

In this report we give a general description of the

island as obtained by the survey party, and discuss

the mapping of the island.

Mapping Technique

As the party traversed the island, measurements

were made of the horizontal and vertical angles

between the various summits and landmarks. All

measurements were made with a marine sextant

and a Brunton compass. Further surveying was

done from the ship. Sextant measurement of the

angles between the various features of the island

can give the height of these if the distance to the

island is known. This method, however, limited

the survey to the various peaks and ridges visible

in profile from the ship. An accurate knowledge

of the location of each feature on the island was

necessary for the reduction of sextant sights. While

the ship was circling the island, azimuth bearings

were taken with a pelorus at regular intervals on

all of the main features, including the north and

13
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

south tips of the island. An attempt to use gunranging radar to fix the distance from the island

was frustrated by an uncertainty in determining

the radar reflection point on the island. Lacking

this distance measurement, the position of the ship

was fixed by the intersection of the azimuth bearings to the tips of the island and to the "South

Peak," which position was shown on the old map.

Further surveying had to accept the island outline

and scale of the published Hydrographic Office

map. But these are less likely to be in error than

the photographically determined contour lines.

The intersections of the azimuth bearings to each

of the features from the various ship positions located each feature on the map. Heights were

determined from vertical sextant measurements of

the angles between the features and the waterline,

and horizontal angle measurements of the width

of the island. An azimuth bearing to the South

Peak combined with the horizontal angle fixed

the ship's position during this phase of the surveying. The linear heights of the features were then

obtained by solving a simple proportional equation

and correcting for the height of the observer above

the waterline of the ship.

the southern end. Beside the peaks the most conspicuous feature of the island is a large arm extending out on the west side of the island and forming a small bay (Figure 4). The bay is narrow and

the cliffs surrounding it are very sheer, giving a

fjord-like appearance. Several sea caves and arches

were also seen on the circuit. Besides the main

island itself, there are several other rocks and sea

stacks in the vicinity. The two largest groups of

these rocks are clustered to the north and to the

south of the island.

A total of three days were spent exploring the

main island and one day was spent exploring the

largest of the offshore rocks.

After the circuit of the island, a landing site

was chosen on the south side of the end of the spur

which forms the fjord-like bay on the west side of

the island. This site became the base for most of

the exploration and the location of most of the

intensive studies of the terrestrial biology of the

island. This landing and study site was designated

Landing Site 1 (Figure 4). After landing the entire party and its equipment, the exploration team

started climbing up the spur. The vista up the

Itinerary of the Exploration Party

A preliminary idea of the topography of the

island was gained the first morning by making a

complete circuit of the island close into shore.

This survey also provided an opportunity to pick

out possible landing sites. The island is quite steep

along the sides and has a rather flat-topped appearance. It has three distinct peaks (Figure 4). At the

north end of the island is the smallest of the three

which was termed "North Peak." This summit is

sharply set off by nearly vertical walls all of the

way around it. The top, however, appears to be

relatively flat. This peak is separated from the next

by an east-west fault. This next peak south is cylindrical and butte-like, and is much higher than

the first. It appears to have equally as steep sides

and is also flat on top. This peak was referred to

as "Plug." To the south of it, separated by a huge

east-west fault which split the entire island is a

more rounded peak which appeared to be of the

same height as Plug. This summit was called

"South Peak." From it the island slopes clown to

the south until it reaches the rather sheer cliffs at

North Racks

5 Stools

.""Shoolt

FIGURE 4.—Reference map of Malpelo showing the various

landing sites (1-4) mentioned in the text, the routes taken

by the exploration party, and significant landmarks.
NUMBER 176 15

spur toward the saddle between Plug and South

Peak was a rocky slope completely devoid of vegetation. Climbing up the spur the party found several caves of various sizes, including one which

went through the spur from one side to the other.

The only vegetation that could be found was algae

which was growing inside of the caves and crevices

where some moisture was present. After some

scrambling, the saddle was reached. Here a cairn

with a bottle containing the names of several Colombians from two mapping and survey expeditions was found. In the saddle ttte lava was shot

through with holes of varying diameter, making

movement slow. From here the party ascended to

the summit of South Peak. Once on the summit

the party had a view down the entire south end of

the island. This view corresponded exactly to the

photograph taken from the summit by the Vanderbilt Expedition of 1937 (Bond and deSchauensee,

1938, plate 10B). Thus it was concluded that the

Vanderbilt Expedition had in fact climbed South

Peak.

As sighted from South Peak, the Plug had an

elevation of 1° to 1.5° horizontal, showing it to be

the true summit of the Malpelo rock massif. As

surveyed from the ship, Plug was determined to

be approximately 6 meters higher than the South

Peak. The root mean square errors for the measurements of the two peaks from various ship positions are about equal to the difference in height

of the two peaks. The 1° to 1.5° elevation of the

northern peak as sighted from the southern corresponds to a height difference of 6-9 meters, which

agrees with the difference as determined by sextant

sights from the ship.

From South Peak the party returned to the saddle and climbed up to the base of Plug. From this

point the climbing became more difficult. After

some searching, a chimney on the east side of Plug

was found that afforded a relatively easy, but nonetheless technical (class 5), climb up to the top.

No bottles or cairns were found on this summit

plateau. None of the previous parties climbing on

the island had mentioned the necessity of technical

rock climbing for the routes they used. Thus, the

Smithsonian party seems to have been the first to

reach the summit plateau of Plug.

As with the southern summit, Plug was devoid

of vegetation. However, both species of lizards

(Anolis and Diploglossus) were common up to the

top of the two peaks, as were the land crabs (Gecarcinus malpilensis). Blue-faced Boobies (Sula dactylatra) were also omnipresent. At the top of both

peaks a strong breeze was blowing continuously in

contrast to the still conditions lower down on the

leeward side of the island. From Plug the party

climbed back down to the saddle and thence down

to the landing site.

On the second day the party landed on the

northwest edge of the island (Landing Site 2, Figure 4). While two members of the party worked

near the shore, two others climbed towards North

Peak. Initially the peak was attempted by climbing

around the west side to its north face, which had

several large chimneys. Also in the area is a spectacular spire isolated from the main peak. The

route was frustrated because the chimneys were

seen to contain formidable chock-stones. On the

less steep slope to the northwest of the North

Peak were some of the few patches of grass to be

found on the island. The largest of these patches

was not more than several square meters in area.

Retracing their steps the party then climbed to

below the saddle between Plug and North Peak.

Here there was a gully which led up to the saddle.

In the gully several large pools of water were

observed. One of these was perhaps one-half meter

deep and was the largest body of standing water

seen on the island. The pools contained some

aquatic insects which unfortunately could not be

collected. Continuing up the gully into the fault

proved difficult and roped climbing was necessary

to scale a 25-meter lava wall. From the saddle it

was a relatively easy scramble up to North Peak.

Again both species of lizards and numbers of

boobies were present all the way up to the summit,

and the wind was blowing steadily in contrast to

the leeward side. From here the party climbed

down along the east side of Plug and over the

saddle south of Plug and down to Landing Site 1.

On the last day of exploration of the main island, a party of two landed on the east side of the

island (Landing Site 3, Figure 4). This landing

site was used by a Colombian expedition which

had placed a bronze plaque to mark the site. The

party made its way up to the saddle south of Plug

and then up to South Peak. From there they

climbed down to the southern end of the island.

The area southwest of South Peak is relatively

the flattest part of the island. The slope leading
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

from South Peak down to the end of the island

was a scree slope with small rocks cemented by

solidified lava. Lava bombs were strewn about

liberally, and all over the lava had solidified into

cubist architectural forms. The south end of the

island was extremely dry. Few lizards were seen on

the central dome of the south ridge, but both

species as well as the land crab were abundant on

the extreme south end. There was a small patch

of grass on the southernmost cliff. From the south

end the party hiked back up over South Peak and

then back down to Landing Site 1.

On the fourth day, the party visited the largest

of the offshore rocks (Landing Site 4, Figure 4).

This rock is in the South Rocks group and like the

other rocks it is quite steep. A landing was difficult

and roped climbing began from the tiny shelves

near the water. Near the top of the rock (45m =

130 ft) the slope decreased appreciably. Rather

surprisingly, a large patch of grass was found here.

This was the largest and lushest stand of vegetation found anywhere. Anolis were common in the

grass as well as on much of the top of the rock.

Diploglossus were also found, but not in large

0

FIGURE 5.—A revised map of Malpelo showing summit altitudes and land contours based on the surveying of the STRI

exploration party in March 1972.

numbers. No land crabs appeared to be on the

rock. Several small bushes and a fern were growing

at the top of the rock. The bushes frequently

showed large trunk masses with only small green

shoots. This was suggestive of large seasonal variations in rainfall.

A Red-footed Booby (Sula sula) was sighted

perched in the middle of the bushes. This species

habitually perches in trees in contrast to the Bluefaced Booby, and has not previously been reported

from Malpelo Island. Positive identification was

made when one of the members of the party was

able to climb down to within a few meters of the

bird before it flew, and secure several good photographs showing the characteristic red feet of this

species.

As with the landing, departure from the rock

was somewhat difficult due, in part, to the nearby

shoals.

Conclusion

The survey undertaken by the exploratory party

resulted in the map shown in Figure 5. As mentioned above, the outline of the island and the

scale are taken from the old Hydrographic Office

map, but the topography and contour lines are

the result of the new survey. It should be borne in

mind that the map may not be entirely accurate

due to the rugged nature of the terrain, and to

the limited time and equipment of the survey

party.

Future visitors to Malpelo should be able to

make substantial improvements in the map, particularly with regard to geological details. Any extensive on-site surveying, however, could not be accomplished without a good deal of technical climbing.

We hope that this map will be of some help to any

future visitors.

Finally we remark that the rather dry description given here in no way does justice to the sustained surrealism of this unique and unusual

island.

Literature Cited

Bond, J., and R. M. deSchauensee

1938. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part II: The Birds of

Malpelo Island, Colombia. Proceedings of the Academy of Natural Science of Philadelphia, 90:155-157,

plates 9-11.
Field Observations on

the Geology of Malpelo Island

Jeffrey A. Stead

ABSTRACT

Malpelo Island had a volcanic origin. Depth soundings around the island indicate that it was once

8 to 10 times larger than its present size. Marine

weathering has eroded the island and has formed

steep cliffs and sea caves along its sides. Several

types of igneous rocks and minerals occur on the

island. The distribution of major joints and faults

is shown.

Published accounts of the geology of Malpelo

Island include McConnell's (1943) analysis of

three rocks from the island and the field observations of Murphy (1945, also in McConnell, 1943).

The amygdaloid specimen assayed by McConnell

(1943) showed a small amount of phosphatization

(replacement of natural minerals by phosphate

compounds derived from guano) while this process

was very advanced in another rock (phosphate

rock). The specimen of augite andesite was only

slightly altered by phosphates. Murphy reported

that guano was not very concentrated on the island

and McConnell (1943) concluded that phosphatization occurs mainly in porous areas of the island

(along faults and fractures) where guano could be

carried and concentrated by rains.

Malpelo is volcanic in origin, the present island

is the remnant of a much larger structure. Most of

the island's perimeter consists of nearly vertical

rock faces which extend from 60 to 230 meters

above sea level (Figure 6).

Jeffrey A. Stead, Defense Mapping Agency, Hydrographic Center, Rodman Office, Box 2023, Rodman, Canal Zone. Present

address: Hahnemann Medical College, Philadelphia, Pennsylvania 19102.

Several types of igneous rocks are present on

Malpelo. Field identifications are as follows: dacite, trachyte, tuff, basalt, and andesite. Up to an

altitude of 210 to 240 meters, the island appears

to be mostly trachyte, with lesser amounts of

dacite and tuff. This lower zone is intruded by at

least two sets of dikes made up of basic rocks. The

top of the island is covered by an andesite cap.

Field identification of minerals is as follows:

Specimens of dacite collected near the landing site

contained amygdules of quartz and apatite. One

amygdule contained fine, hairlike, zeolite crystals

coated with hematite. Small amounts of chrysocolla

were found in a joint at an altitude of about 240

meters. Guano occurs in limited amounts on all

parts of the island.

Fresh water seepage was observed in several

places. Red streaks of iron oxide are deposited on

the rock faces where seepage has occurred. Some

of these streaks represent principal areas of runoff

following rainstorms rather than permanent seep

age areas.

Steep cliffs around the perimeter are probably

maintained by the combined effects of wave action,

which loosens and removes rock in low areas

(Steers, 1953), rain, and alternate heating and

cooling, which loosen the soft lavas (mostly

trachytes) above the normal splash zone until they

fall off along joints. The lava cap is a very resistant

rock, and probably does not fall until the rock

underneath it has been weathered away.

Sea caves (Figure 6) are formed by the scouring

action of waves on brecciated rocks around faults

(Guilcher, 1958). There are many small faults on

Malpelo, most of which intersect the waterline at

large (60°-90°) angles. The largest cave was

estimated to be 9 meters wide and 13 meters high

17
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(above sea level) and to have a 10-fathorn water

depth at its mouth. Some of the caves have a white

formation on their ceilings which resembles flowstone. This is probably composed of insoluble substances from guano, washed down along the faults

and redeposited on the ceilings of the caves.

Small amounts of soil, consisting of variable

mixtures of clay and animal feces, are found in

sheltered areas of the island. Soil is found nowhere

in large amounts, but increases in abundance with

increasing altitude, and rarely occurs below 90

meters. The absence of soil in the lower levels can

be attributed to steep slopes, rain, and to wave

action which, during severe storms, can splash

water to considerable heights (Johnson, 1919).

Two major sets of joints and faults were measured in the fjord area (Figure 7). Their average

measurements were as follows: strike 210°T, dip

79°NW; strike 110°T, dip 82°SW. The eastern

wall of the fjord is the scarp of a major fault, represented by a cliff over 60 meters high and half a

kilometer long. This fault, which is the most prominent feature of the fjord, strikes 2O5°T and dips

78°NW. The largest sea cave on the island is at

the apex of the fjord (Figure 7).

The U.S.S. York County made continuous depth

soundings around Malpedo Island. These soundings, plus those from a 1952 U.S. Navy sketch survey, are shown in Figure 7.

Bottom depth falls off greatly to 50-60 fathoms

on all sides of the island, but the bottom gradient

increases rapidly at 60-70 fathoms (Figure 7).

The area out to the 60-fathom curve may represent

a wave-cut plateau formed during the Pleistocene

(Shepard, 1963). If this is assumed, the 60-fathom

curve would indicate the minimum size of the original volcano. A line has been drawn (Figure 7) to

indicate a depression in the sea floor. This may be

a fault. Its trend, 218°T, is close enough to the

210±T average of one of the major joint sets on

the island to strongly indicate a structural relationship. The dashed line trends 136°T, and also may

indicate a fault, or it may mark the division between two previously existing cones.

FICI'RE 6.—The southwestern tip of Malpelo Island showing the lava cap, the steep cliffs, and

narrow caves that occur at the watcilinc all around the island.
NUMBER 176 19

FIGURE 7.—Map of Malpelo Island showing depth soundings from a U.S. Navy 1952 sketch

survey (boldface numbers), those from the T.S.S. York County (lightface numbers) and depth

profiles around the island. The location of two submarine faults (218°T and 136°T) are

indicated.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ACKNOWLEDGMENTS.—I thank D. Meyer of the

Smithsonian Tropical Research Institute, B. Macurda of the University of Michigan, and R. Stewart of the Panama Canal Company, Engineering

Division, for reading this manuscript and offering

suggestions. Macurda assisted me with field work

and Steward helped me interpret photographs of

the island.

Literature Cited

Guilcher, A.

1958. Coastal and Submarine Morphology. London:

Methuen and Co., Ltd.

Johnson, D. W.

1919. Shore Processes and Shoreline Development. New

York: John Wiley and Sons.

McConnell, D.

1943. Phosphatization at Malpelo Island, Colombia. Bulletin of the Geological Society of America, 54:707-

716, 2 plates.

Murphy, R. C.

1945. Island Contrasts. Natural History, 15:14-23.

Shepard, F. P.

1963. Submarine Geology. New York: Harper and Row,

Inc.

Steers, J. A.

1953. The Sea Coast. London: Collins Clear-Type Press.
The Ecosystem on Malpelo Island

Henk Wolda

ABSTRACT

A rather high percentage of predators was found

among the terrestrial invertebrates of Malpelo Island. Except for the birds and one of the lizards,

almost all of the animals depend on a purely terrestrial economy based on numerous algae, lichens,

and mosses.

Studies on the biota of isolated islands and

groups of islands have produced exciting results,

influencing and even initiating the theory of evolution. Almost all existing islands have now been

surveyed, at least to some extent, but until 1972

Malpelo remained a notable exception. Except for

the vertebrates no collections had been made, presumably because it is remote and almost inaccessible. The location of the island, however, suggests

that it may contain species that are highly interesting biogeographically, especially because introductions of species by man are very unlikely. Although

some landings have been made (Fowler, 1938;

Bond and deSchauensee, 1938), very few people

have ever set foot on the island.

A number of seabirds, a land crab, and two

species of lizards are known from Malpelo, but

statements in the literature about plants and invertebrates are not very encouraging. It is reported

that there is very little (Bond and deSchauensee,

1938) or no (Slevin, 1928) plant life that could

support a community of animals. In fact, Slevin

(1928) reports that he could not find "any life

whatever" except for the species mentioned above.

Such a situation seems a priori very unlikely. Even

very forbidding habitats, including bare rocks in

Henk Wolda, Smithsonian Tropical Research Institute, P. O.

Box 2072, Balboa, Canal Zone.

the Pacific (Clapp, 1972) are known to contain a

variety of species and all kinds of animals are able

to reach islands more remote than Malpelo (Carlquist, 1965). Moreover, one would expect at least

one of the lizards (Anolis agassizi) to feed predominantly on insects. Indeed, Dunn (1939) found

"two or three species of insects" in stomachs of

Anolis. Bond and deSchauensee (1938) report an

abundance of "small black gnats or flies" and a

chirping, as if by crickets. The most encouraging

statement is that of Murphy (1945), who, in the

stomachs of Anolis, found "remains of small crabs,

spiders, pseudoscorpions, beetles, flies and fly larvae, ants and other insects." The problem to be

><?

South

Rock

FIGURE 8.—Map of Malpelo Island with collecting sites.

21
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

investigated is what animals occur on the island

and how do they make a living. The near-absence

of plant life reported in the literature suggested

that the economy of Malpelo is entirely based on

the birds bringing in food from the ocean, their

droppings, and dead bodies.

In the first week of March 1972, I was able to

test this idea. My collections and observations were

made in localities indicated on Figure 8 and I

examined some material collected from the South

Rock.

The general impression one obtains from the

main island is that it is bare rock (Figure 9), with

no higher plants except for very few small patches

of a yet unidentified grass. On the South Rock this

grass was fairly abundant and here also a shrub

occurred, possibly a legume, and a fern, PityroFIGURE 9.—General view of site 1.

gramma dealbata (Presl) Tryon. On the main

island, however, lower forms of plant life were

abundant. On vertical rock faces moist with fresh

water and under ledges, algae, mosses, and lichens

could be found everywhere (Figures 10, 11). A few

pieces of rock were collected and eight species of

lichens were identified from them: Caloplaca

sp., Candelbria sp., ?A.spicilia sp., ?Diploschistes sp.,

Lecidca sp., Graphidaceae sp., fBasidea sp. and

Pyxine cf. glebosa Tuck. Considering the smallness

of the sample, the number of sj>ecies present may

be much larger than this.

This vegetation is the basis for quite a complex

community of animals. In crevices and, especially,

under rocks, a multitude of animals was discovered.

My collection probably does not cover all the

species present, but the fact that mine was about

as good as the anoles (see below) suggests that I

may not have done too badly. I did not make extractions from soil under the rocks and any microarthropods living there would have been missed.

The abundance and diversity of the invertebrates

obtained (Table 1) is rather impressive. At least

17 species of insects were found, belonging to 8

orders. The "black flies" mentioned by Bond and

deSchauensee (1938) represent a new genus of

Chloropidae and the crickets they heard belong

to the genus Hygronemobius, related to H. liura

described from British Guyana (Gurney, pers.

comm.). These two species were among the most

abundant insects, together with a little black ant

(Pheidole sp.) and staphylinid beetles. The two

species of snails were fairly common in clusters

under rocks at locality 1. There was an extraordinary variety of spiders, some eleven species of

Araneida, none of which were webbuilders.

Worms and isopods were abundant under rocks at

locality 2, far above the intertidal zone.

It is much too early to start speculating about

the origin of the species found, but there are indications that some of them may have come from the

Indo-Pacific. After a preliminary investigation of

the centipeds, Crabill (pers. comm.) suggests that

the closest relatives of some of these may not be

South American species. Representatives of the

opilionid family Assamidae seem to be mostly

living in the Indo-Malayan region (Brues, Melander, and Carpenter, 1954). The land crab and all

three species of lizards (including the new gecko
NUMBER 176 23

described by Huey, pp. 44-46 herein) are all found

on Malpelo only, but how many of the other

species are endemic is not yet known.

To depict the structure of the ecosystem on

Malpelo, i.e., to construct a foodweb, roles had to

be assigned to each of the species concerned. The

classification of these species as a herbivore, a predator, or a scavenger is based on what is known

about relatives living elsewhere and on some observations. The classification of the staphylinids

as herbivores, for instance, was made because they

were very abundant and found mostly among

mosses. The beetle, Enochrus, is classified as a predator, although the adults may be omnivorous. The

results are included in Table 1 and the tentative

foodweb is illustrated in Figure 12. The landcrab

Gecarcinus malpilcnsis, was very common and was

frequently observed nibbling on stones, presumably feeding on lichens and algae. Remains of insects were often found in their feces. Many of these

may have been scavenged or obtained while eating

the feces of lizards, but the crabs can also act as

predators. On one occasion I saw two crabs that

were holding a male Anolis, which was still very

much alive, although one of its hindlegs was already eaten. A third crab was eating the tail.

These crabs also feed on dead birds. They are

opportunistic omnivores, but probably mostly

scavengers.

I had the opportunity to examine the gut contents of 38 individuals of Anolis agossizi from the

main island, plus five from the South Rock. These

were kindly given to me by A. S. Rand. Virtually

all the items listed in Table 1 were observed, including some juvenile crabs. The only obvious exception were the earthworms, presumably because

they are living in soil under rocks and thus inaccessible. It is certainly not because they are distasteful. When I removed the rocks, several anoles came

in immediately and feasted on the worms. The

anoles from the South Rock also had a variety of

arthropods in their guts, but differed from those

from the main island in that they had their rectums filled with grass seed.

Both individuals of the gecko, Phyllodactylus,

had eaten crickets, a few ants (probably Strumigenys) and a lepidopteran larva, possibly Ereunetis. The number of species per individual was

much smaller than that observed in Anolis. Ants

(Pheidole sp.) and staphylinid beetles, both of

which made up the bulk of the diet of anoles, were

absent in the geckos. These differences could be

accidental, but are probably real, Anolis being

diurnal and Phyllodactylus nocturnal.

Nematodes, probably several species, were found

in 60 percent of the anoles, averaging three per

individual (range 1 to 14). The geckos had 11 and

12 nematodes, respectively.

The large lizard, Diploglossus millepunctatus,

has a quite different way of life. The feces that I

examined did not contain any arthropods at all.

They were observed feeding on fish regurgitated

"••& •

FIGURE 10.—Moist rock face and land crabs at site 1. Fici'RF. II.—Moist wall of empty creek bed at site 2.
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

by boobies (Sula dactylata granti), on fish leftovers

after the boobies had fed their young, and on fresh

bird droppings. Slevin (1928) and Dunn (1939)

found crab remains in Diptoglossus' stomachs and

Garth (1948) suggested that this lizard feeds on

both crabs and bird eggs. The latter may be true,

but I have no evidence to support this idea.

The foodweb (Figure 12) shows two interesting

features. The first relates to the rather large variety

of predators, some 24 species, including two lizards.

In contrast I found only nine species of herbivores

and eight scavengers. This suggests that among the

species living on Malpelo, an unusually high percentage are predators, many of which may be generalists rather than specialists. Two of the herbivores, the snails, seem to be eaten exclusively by

the lizards, not by any of the invertebrate predators. These predators are all eaten by Anolis, but

the bulk of their food consists of herbivores. More

exhaustive collections, made at different times of

the year, are needed to determine whether indeed

the predators make up such a large proportion of

the fauna. If this is so, it might have some bearing

on theoretical considerations which state that the

TABLE 1.—List of invertebrates found on Malpelo Island with estimate of their

food habit (H = Herbivore; P = Predator; S = Scavenger; Par. = Parasite; O =

Omnivore)

Oligochaeta

Nematoda

Gastropoda

Diplopoda

Geophilomorpha

Scolopendromorpha

Decapoda

Isopoda

Chelonethida

Araneida

••

Opilionida

Schizomida

Acarina

Thysanura

Collembola

Orthoptera

Hemiptera

Coleoptera

Diptera

Lepidoptera

"

Hymenoptera

Chilenophilidae

Cryptopidae

Gecarcinidae

Oecobiidae

Selenopidae

Agelenidae

Lycosidae

Salticidae

Segestriidae

Pholridae

Assamidae

Entomobryidae

"

Gryllidae

Enicocephalidae

Hydrophilidae

Staphylinidae

Scydtnaenidae

Ephydridae

"

Sciaridae

Chloropidae

?Pyralidae

Tineidae

Formicidae

1 (?)sp.

spp.

2 spp.

sp.

Ribautis sp.

Scolopocryptops sp.

Cryptops sp.

Gecarcinus malpilensis

sp.

sp.

sp.

Selenops sp.

spp.

spp.

about 3 spp.

PAriadna sp.

sp.

Paramitraceras sp.

sp.

sp.

sp.

Lepidocyrtus caprelsi

Lepidocyrtus sp.

Hygronemobius sp.

Systelloderes sp.

Enochrus sp.

sp.

PEuconnus sp.

Nostima sp.

Scatella sp.

Bradysia sp.

sp.

sp.

Ereunetis sp.

Odontomachus sp.

Pheidole sp.

Strumigenys sp.

S

Par.

H

H

P

P

P

O

S

P

P

P

P

P

P

P

P

P

P

Par.

H

S

S

H

P

P

H

P

S

s

s

H

H

S

P

H

P
NUMBER 176 25

number of predator species can never exceed the

number of prey species (cf. Levins, 1968), except

in some special circumstances (Stewart and Levins,

1973). If the phenomenon is real, it could be that

some of the necessarily simplified assumptions on

which such theories are based are violated. It could

be that predation by the lizards on the invertebrates, prey and predators alike, is so intense, at

least during some times of the year, that competition between the invertebrate predators is greatly

reduced, thus enabling more of them to coexist.

In that case, the problem boils down to what limits

the numbers of lizards. We found no evidence of

predation by birds on the lizards, nor do the

anoles seem to limit their density through behavioral means, as they are nonterritorial (see Rand,

Gorman, and Rand, pp. 27-38 herein). Their gut

contents did not suggest that they were experiencing food shortage, but in other seasons this could

be different.

The other interesting feature of the foodweb is

that my original idea, that the entire economy of

the island was based on the sea, may be wrong.

Only the birds and Diploglossus depend completely,

and the landcrab partly, on income from the

ocean. All the other species seem to depend on a

purely terrestrial economy. I would predict that if

the birds, which provide the link between the

TERRESTRIAL

GRASS

SOUTH ROCK

A

ALGAE

LICHENS

MOSSES

V

HERBIVORES

9spp.

V

V

PREDATORS

22spp.

A

FECES

CORPSES

SCAVENGERS

8spp.

V?

V

NEMATODES

ANOLIS

PHYLLODACTYLUS

A

V

GECARCINUS

A

BIRDS

V

FECES

FOOD REMAINS

CORPSES

DIPLOGLOSSUS

TICKS

MARINE

FIGURE 12.—Tentative structure of the foodweb on the island of Malpelo. The four question

marks refer to uncertainties in utilization of grass by the herbivores, crabs eating living insects,

Diploglossus taking living crabs, and the importance of bird droppings in providing nutrients

for the plants. For simplicity, some relationships are omitted. These include lizards and crabs

providing food for the scavengers and ticks being eaten by Anolis.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ocean and the land, disappeared, Diploglossus

would soon be extinct and the land crab would

reach lower densities, but that this would have

little, if any, effect on the rest of the fauna. The

only condition under which this would not be true

is when the weathering rock would not provide

enough nutrients for the plants to grow, so that

fertilization (e.g., phosphates) by the birds is of

vital importance.

ACKNOWLEDGMENTS.—I wish to express my gratitude to all those who identified the specimens:

Drs. M. J. Dibben and T. Esslinger of Duke University for the lichens, Dr. D. B. Lellinger of the

National Museum of Natural History, Smithsonian Institution, for the fern, Dr. L. Abele of the

Smithsonian Tropical Research Institute and Drs.

R. E. Crabill, W. B. Peck, D. L. Wray, A. B.

Gurney, J. L. Herring, P. J. Spangler, J. M. Kingsolver, W. W. Wirth, R. J. Gagne, C. W. Sabrosky,

D. C. Ferguson, D. M. Weisman, and Dr. D. R.

Smith, all of the Department of Agriculture, for

identification of the items listed in Table 1.

I also gratefully acknowledge Drs. N. G. Smith

and J. B. Graham of the Smithsonian Tropical

Research Institute and G. C. Gorman of the University of California at Los Angeles for critically

reading the manuscript.

Literature Cited

Bond, J., and R. M. deSchauensee

1938. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part II: The Birds of

Malpelo Island, Colombia. Proceedings of the Academy of Natural Sciences of Philadelphia, 90:155-

157, plates 9-11.

Brues, C. T., A. L. Melander, and F. M. Carpenter

1954. Classification of Insects. Bulletin of the Museum of

Comparative Zoology at Harvard College, 108:1-917.

Carlquist, S.

1965. Island Life. Garden City, New York: Natural History Press.

Clapp, R. B.

1972. The Natural History of Gardner Pinnacles, Northwestern Hawaiian Islands. Atoll Research Bulletin,

163:1-25,7 figures.

Dunn, E. R.

1939. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part III: The Lizards

of Malpelo Island, Colombia. Notulae Naturae of

the Academy of Natural Sciences of Philadelphia,

4:1-3.

Fowler, H. W.

1938. The Fishes of the George Vanderbilt South Pacific

Expedition, 1937. Monographs of the Academy of

Natural Sciences of Philadelphia, 2: v + 349 pages,

10 plates. [Malpelo, pages 5-6.]

Garth, J. S.

1948. The Brachyura of the "Askoy" Expedition with

Remarks on Carcinological Collecting in the Panama Bight. Bulletin of the American Museum of

Natural History, 92(1): 1-66, 4 figures.

Levins, R.

1968. Evolution in Changing Environments. Princeton:

Princeton University Press.

Murphy, R. C.

1945. Island Contrasts. Natural History, 54:14-23.

Slevin, J. R.

1928. Description of a New Species of Lizard from

Malpelo Island. Proceedings of the California

Academy of Sciences, fourth series, 16:681-684,

plates 25, 26.

Stewart, F. M., and R. Levins

1973. Partitioning of Resources and the Outcome of

Interspecific Competition: A Model and Some

General Considerations. The American Naturalist,

107:171-198.
Natural History, Behavior, and Ecology

of Anolis agassizi

A. Stanley Rand,

George C. Gorman,

and William M. Rand

ABSTRACT

Anolis agassizi is found throughout Malpelo Island, and in places is extremely abundant. In one

study area we estimated one lizard per two square

meters. Social behavior contrasts markedly with

other anoles in the apparent lack of territoriality,

infrequent aggressive encounters, and reduced display repertoire. There is a strong tendency for

social facilitation—the species is aptly described as

"curious." Home ranges are large, and lizards tend

to move long distances during the course of the

day. Resources such as food, water, and perch sites

may be shared by many individuals. There appear

to be no special adaptations to temperature extremes nor to desiccation. Reproduction is apparently highly seasonal, as evidenced by the complete

absence of small juveniles in museum collections

made in December and January, and in our collections of February and March. In our sample

about half the females were carrying eggs. The

species is larger than most solitary anoles (adult

males often > 100 mm) and markedly sexually

dimorphic for size (males larger). Adult males

fall into two discrete reproductive categories that

have morphological correlates. Some have black

heads and permanently erect nuchal crests—these

have active testes. Others lack the nuchal crest

and have spotted heads like the females—these

have completely regressed testes.

A. Stanley Rand, Smithsonian Tropical Research Institute,

P. O. Box 2072, Balboa, Canal Zone. George C. Gorman,

Department of Biology, University of California, Los Angeles, California 90024. William M. Rand, Department of

Nutrition, Massachusetts Institute of Technology, Cambridge,

Massachusetts 02139.

Introduction

Almost all islands in the Caribbean have been

colonized by one or more stocks of Anolis lizards.

Patterns of distribution, ecological relationships,

and behavior have been extensively studied in

these West Indian anoles by Ruibal (1961), Rand

(1964), Gorman (1968), Schoener (1969), Schoener and Schoener (1971), Lazell (1972), Williams

(1972) and many others. The Pacific waters adjacent to the tropical American landmass are relatively impoverished in islands, and only two are

inhabited by Anolis. One is Cocos Island, and the

second is Malpelo. Malpelo is more isolated than

any of the West Indies, which have been naturally

colonized by anoles, although strictly in terms of

distance it is within the limits known for colonizations by Caribbean Anolis (Williams, 1969; Rand,

1969). Malpelo is also more devoid of vegetation

than any West Indian island. Caribbean anoles

are mostly arboreal, in the strictest sense, the exceptions usually perch on other sorts of vegetation

(bushes, grass, cactus, etc.). There are a few anoles

associated with rocks, such as A. rimanim on boulders in Hispaniola and A. bartschi on cliffs around

cave entrances in Cuba; but none is from an environment so devoid of higher plants as is Malpelo

Island.

Previous to the expedition reported here, our

knowledge of Anolis agassizi was limited, but the

very nature of the habitat suggested that this

species might be rather different in behavior and

ecology from better known anoles of the Antilles.

The species was first described by Stejneger (1900:

161-163) with a detailed account of external mor27
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

phology. The paper included a superb watercolor

plate of an adult male. Stejnegar also provided

the few field observations available: "Mr. Charles

H. Townsend, who collected these specimens in

Malpelo, informs me that they were running over

the rocks near the water. The island was too steep

to afford a landing, but the lizards were shot off or

whisked off the face of the cliffs, thus falling into

the water, whence they were secured by the collector." Slevin, in 1927, landed briefly on Malpelo

and collected a series of A. agassizi, but he did not

explore the island.

Ronald Smith of the George Vanderbilt South

Pacific Expedition of 1937 did explore the island

and found anoles all the way to the summit. He

described them as fearless and quite bold (Fowler,

1938). Dunn (1939) utilized the Vanderbilt Expedition specimens to reconsider the status of the

lizards of Malpelo. Because of peculiarities, particularly a small throat fan, he erected a new,

monotypic genus for agassizi. This he called Mariguana. He reported that the agassizi stomachs contained small insects of two or three species. He

remarked that the unusual color pattern (see below) was similar to that of the other totally unrelated lizard species on the island, Diploglossus

millepunctatus.

Etheridge (1960) considered the relationships

of all available species of anoline lizards including

agassizi. He showed that this species fit squarely

into the latifrons species group of Anolis, and he

therefore did not recognize the validity of Mariguana. The latifrons series is an osteologically

primitive group of anoles otherwise restricted to

the southernmost of the Lesser Antilles and the

tropical American mainland from Costa Rica

southward. Anolis agassizi is virtually indistinguishable from the Lesser Antillean members of

the latifrons series on osteological characters alone,

but genetic similarity evidence (Webster, p. 50

herein) implies that it is not particularly close to

either the island or continental species examined.

Compared with West Indian anoles, the Malpelo

species was known to be unusual but not unique

in several aspects of its biology. Anolis agassizi is

larger than most anole species that occur on islands without congeners (Schoener, 1969:389),

but it is somewhat smaller than Anolis ferreus of

Marie Galante. Anolis agassizi has a very reduced

dewlap in both sexes, but not more reduced than

A. hendersoni of Hispaniola or A. bartschi of Cuba.

The color pattern of A. agassizi is unusual, consisting of minute light spots on a very dark ground

color, but many species of Anolis on small islands

have unique color patterns.

We were able to verify these attributes and to

discover additional peculiarities particularly in

the social behavior of A. agassizi during our visits

from 29 February to 3 March 1972.

ACKNOWLEDGMENTS.—We are grateful to Drs.

Thomas A. Jenssen of the Virginia Polytechnic

Institute and Ernest E. Williams of Harvard University for reading and commenting on the manuscript. Travel funds for G. C. Gorman were provided by a grant from the National Geographic

Society; partial support for A. S. Rand was provided by National Science Foundation grant No.

B019801.

Distribution and Numbers

Anolis agassizi were found everywhere on Malpelo from sea level to the highest peaks, and also

on the only one of the small rocks south of the island that was explored. They were rare on the

south plateau and uncommon on the black rocks

in the lowest 3 meters.

Anoles seemed less common on the large smooth

rock faces than on the more irregular rocky areas

which provided more hiding places and probably

more food and water. They certainly did not avoid

either the vertical cliffs or the more nearly horizontal edges. The concept of structural niche as

defined by perch dimensions (Rand, 1964) fails

completely in an environment such as Malpelo.

Home Range Size and Population Density

Our home range data are based on marked individuals. On the first afternoon we marked all of

the anoles that we could catch in an area of perhaps 20 m long by 10 m high (Figure 13). We

visited this area on three succeeding days and recorded the numbers of anoles seen, as well as their

activities.

We marked 29 anoles. Of these, we saw 21 again

during the next three days. Of the six large adult

males marked, five were resighted. The marked

lizards did not seem to prefer specific perches;

rather they moved back and forth across the rocks,
NUMBER 176 29

some moving 5 to 10 meters during a morning. One

lizard was seen about 150 meters away from where

it had been marked; however, another marked lizard was seen at virtually the same spot on three

successive days.

Anoles gathered from distances as great as 10 to

15 meters at concentrated resources, such as the

oranges and jellied candy ("Chuckles") that we

offered, and at water seeps. Home ranges are large

and, even for adult males, are widely overlapping

and seem little defended. The long distance moved

by one anole and the fact that several marked animals were not resighted suggests that there is a

floating population like those postulated for

other anoles (e.g., Philibosian, 1972).

During our observations, and when lizards congregated at sliced oranges we had set out (p. 31),

we made periodic counts of the numbers present

and the proportion of these that were marked.

Eleven counts showed 8 to 25 anoles present (an

FIGURE IS.—Part of the study area on Malpclo where Anolis

agassizi were marked and observed and the orange and

"Chuckles" experiments conducted.

average of 16 anoles) of which 0 to 31 percent

(average of 16 percent) were marked. If we were

seeing about the same proportion of unmarked as

of marked anoles, and assuming that of the 29

marked anoles, 20 remained in the area, then the

total number of anoles in our area would have

been about 120. With this small sample and these

many assumptions, we hesitate to calculate confidence intervals. We estimate, however, that anoles

were concentrating at the oranges from an area of

perhaps 500 to 1000 square meters. This would

indicate a population density of about one anole

per 5 to 10 square meters. Although not all areas

of the island are equally habitable by Anolis, there

must be at least 100,000 Anolis agassizi on the

island.

Measurements were taken from 45 animals,

which were preserved and deposited in the Museum of Comparative Zoology. This series contained

four juveniles whose sex could not be determined,

14 adult females (snout-vent length, SV x 77.1

[71-84] SD 4.6) and 27 males. Fourteen of the

males had enlarged testes, dark heads, and welldeveloped nuchal crests (SV x 97.6 [89-114] SD

7.3). The remaining 13 had small testes, femalelike coloration, and small, if any, nuchal crests

(SV x 84.2 [71-99] SD 7.9). (The latter males

are discussed on p. 32).

Weights of these animals just before preservation were found to be highly correlated with snoutvent length in each sex (r = 0.% for males, r =

0.85 for females). The lower correlation for females

is due primarily to their smaller size range. The

linear regression equations for weight (W) on

snout-vent length (SV) are: males, W = — 40 +

0.62 SV; females, W = -1 8 + 0.35 SV. Only a

small number (13.4 percent) of the lizards captured

had unbroken tails, and an additional 6.8 percent

were broken in capture. The sexes do not differ in

the percentage of tails broken (p. 32). In the 18

lizards with unbroken tails, the tail was slightly

more than twice SV (mean = 2.19; SD = 0.12).

The regression of tail length (T) on SV is T =

22 + 1.9 SV, with an r of 0.92.

Schoener (1969) has shown that Anolis species

which live on islands without congeners tend to

fall into a restricted size range. The largest third

of the males usually have a snout-vent length between 55 and 90 mm, and the females between 40

and 60 mm (for any given species, the female is
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

always significantly smaller). Anolis agassizi is an

exception. The largest third of the males averages

105.4 mm SV, the maximum is 114 mm. The comparable female average is 85.2 mm SV with a recorded maximum at 87 mm. Thus A. agassizi is

about the size of the Lesser Antillean giants A.

richardi and A. bimaculatus, both of which are

sympatric with smaller species.

Two hypotheses come to mind to explain the

larger than predicted sizes of A. agassizi: (1) There

was a congener on the island that is now extinct.

This is unknowable, but unlikely. The very remoteness of the island makes the probability of a

second colonization extremely low; and insertion

of a new colonist onto a small, already occupied

island is the exception rather than the rule for

West Indian Anolis, where competitive exclusion

seems to prevail (Williams, 1969; Gorman and

Boos, 1972). (2) The average prey size is larger on

Malpelo than in the Antilles. This is simply not

the case (see below).

Any number of additional hypotheses to explain

the body-size anomaly could be conjured up (e.g.,

selection to resist predation; intense sexual selection), but we have only our ignorance of the historical factors that have acted during the evolution

of this species with which to discuss such hypotheses. Thus we resign ourselves to the observation

that A. agassizi is large.

Food and Feeding

FIELD OBSERVATIONS AND ANALYSIS OF STOMACH

CONTENTS.—The stomachs of 43 anoles were examined and the contents identified by H. Wolda,

who collected invertebrates on Malpelo (see

Wolda's paper herein). Most of the species of invertebrates that he found on Malpelo, except

earthworms, were represented in the Anolis' guts.

Major groups taken by the anoles include: snails,

juvenile land crabs, isopods, millipedes, centipedes,

spiders, pseudoscorpions, ticks, and insects. The

insects included beetles, flies, true bugs, caterpillars,

ants, crickets, and Thysanurians. Many of these

animals are cryptic. They were found under rocks

and in crevices. Anoles certainly do enter crevices

and holes in the rocks where they probably catch

some prey.

The most common items in the stomachs were

ants. These occurred in all guts and in larger numbers than any other prey. The next most common

items were beetles, both larvae and adults, which

occurred in 72 percent of all guts. Together, beetles and ants made up the majority of the food

eaten and probably more than half of the biomass.

The food items were small, nothing longer

than 20 mm, few over 10 mm. The majority of the

items and probably of the volume was in the 5 mm

class.

This range of prey corresponds with our subjective impressions of what was available. We saw

almost no invertebrates longer than 20 mm except

earthworms, which were perhaps too well concealed

to be captured, and the large land crabs, which

were too big to be taken.

As previously pointed out, A. agassizi is larger

than might be predicted on the basis of Schoener's

(1969) study of single species anole communities.

The feeding data, however, do not support the hypothesis that A. agassizi is larger because it feeds

on unusually large prey. No precise comparisons

can be made, but the food of A. agassizi is certainly

smaller than that of the smaller mainland species

A. polylepis (Andrews, 1971), and on the same

order as that of the smaller A. roquet, which is

alone on the island of Martinique (Schoener and

Gorman, 1968).

Anoles made repeated efforts to catch a hookless

trout fly which we presented to them. They jumped

into the air for it when the artificial fly was dangled overhead. One suspects that they may gather

at a bird carcass to feed on the invertebrates which

also gather there. In this situation, they may leap

into the air to catch circling flies.

A small series of anoles was taken from one of

the south rocks, where grass and bushes grew much

more abundantly than on Malpelo itself. All of

these anoles had their lower intestines packed with

grass seeds which appeared to be undigested.

About 60 percent of the guts contained a few

(1 to 4) small, presumably parasitic, nematodes.

FEEDING EXPERIMENTS.—As pointed out above,

the anoles seemed particularly bold and curious,

and often came to us and even jumped on us.

Casual observation implied that they were particularly attracted to the color orange. Thus, they

seemed especially interested in a Kodak film package and in an orange screw-cap on a container of

suntan lotion. This led us to perform two very

simple field experiments.
NUMBER 176

The Orange Experiment: On several occasions

in and near the study area, we set a half of an

orange on a rock. Anoles from the surrounding

areas gathered at the orange to lick repeatedly at

the cut surface, and to bite, pull off, chew, and

swallow, bits of the pulp (Figure 14). They also

licked the peel, but did not persist in this. The

anoles were intially attracted by the bright color

of the orange, and persisted because of its taste.

We saw lizards come to the orange from as far

as 15 meters across the rock face. Five or ten individuals would feed at the orange itself at one time,

while another 10 to 20 remained within a 2 meter

radius of it. There was almost no aggression shown

at the orange though a male occasionally spread

his small throat fan or bobbed.

The anoles seemed attracted to the area of the

orange by the activities of the anoles already there,

as well as by the orange itself. On one occasion we

saw a succession of anoles drinking (1 to 4 at a

time) from a small wet spot on the rocks. Again we

had the impression that these lizards were attracted

by the activities of other individuals.

Anoles did not stay at the orange for more than

a few minutes though the same lizard might have

returned several times in the course of an hour or

two.

The "Chuckles" Experiment: To test the possibility that the anoles actually showed a color preference in making initial feeding choices, we used

the jellied candy called "Chuckles." A package of

c . \>

FIGURE 15.—Anolis agassizi making a choice between two

equal sized pieces of "Chuckles."

this candy contains five colors (flavors): red,

green, black, orange, and yellow. Comparisons of

different colored candy were made by setting out

pairs of equal sized bits of candy close together in

areas where we had not conditioned the lizards to

oranges. We noted which bit of candy was selected

by the first lizard to approach (Figure 15).

The results (Table 2) show that yellow and

orange are equivalent and that they both are

chosen more frequently than the other colors (p<

0.01). Of the remainder, red and green are about

equal, and both preferable to black, though the

difference is not significant (p>0.05). We hasten

to add that this appears to be a color preference

and not a taste preference, for once the less optimal candy was licked by the lizard, it was immediately eaten.

We cannot separate hue from brightness in

these color choices, but all candies presented were

conspicuous.

TABLE 2.—Candy color preferences of Anolis agassizi (numerator is the number of times "Color

selected" was chosen before "Other color in test

pair"; denominator is the total number of trials)

Color selected Other color in test pair

Yellow Red Green Black

FICURE 14.—Anolis agassizi gathered about an orange half.

Orange

Yellow

Red

Green

3/7 3/3

5/6

-

-

6/6

4/6

2/6

-

6/8

6/6

5/6

4/6
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The preference for orange and yellow seems

most peculiar because there are certainly no fruits

of this color on the island. The only objects that

we could find in this color range were the large

land crabs, and the anoles showed not the least

interest in these. One possible explanation is that

the anoles are attracted to and feed on the yolks of

broken seabird eggs. Many boobies nest on Malpelo and broken eggs may be a frequent occurrence

at certain times of year.

Predation

The most conspicuous potential anole predators

on the island are the large lizards, Diploglossus,

the land crabs and the birds. The seabirds could

certainly eat anoles, but Neal Smith (pers. comm.)

says this is unlikely. The peregrine falcon (Falcon

peregrinus) has been recorded on the island (Bond

and deSchauensee, 1938), but it is normally a bird

predator. The land crabs were everywhere. We

saw a small group tearing apart and eating a still

living anole (see p. 23). How it was caught is unknown for the crabs are relatively slow moving

animals.

The Diploglossus appear to depend largely on

the exuvia of boobies and on land crabs for food,

and they gather where an adult booby is feeding

its young. They do occasionally eat anoles, however, for a specimen in the American Museum of

Natural History collections had one in its stomach

(C. Meyers, pers. comm.). The Diploglossus could

not be induced to eat oranges, but one big male

repeatedly approached an orange that had attracted a large group of anoles, and the Anolis

hurriedly fled at each approach. Though this Diploglossus was much swifter than the land crabs,

the anoles were even faster and easily evaded him

by keeping about a meter ahead of him on the

rocks.

There is strong circumstantial evidence that

predation is important. Approximately 85 percent

of the anoles that we examined had their tails

broken and regenerated. Given the low level of

aggressiveness that we observed and the fact that

females showed as high a precentage of breakage

as males, the implication is that the breakage was

not caused by interaction with conspecifics, but

rather by predation attempts. Pianka (1970) has

argued for a correlation between frequency of

tail breakage and intensity of predation in lizards

of the western United States.

If, as it seems likely, crabs and Diploglossus are

the important predators of anoles, it is likely that

they catch more small than large ones. The color

patterns of Anolis are highly cryptic on the rocks,

suggesting selection by a visually hunting predator.

Adult males with their permanently erect nuchal

crests and black heads are very striking and are

more conspicuous than females and juveniles.

Adult males of many Antillean Anolis also are

more conspicuous than females, presumably because sexual and social selection favors conspicuousness in males and the larger size of the males

reduces their predation and consequently the selection against being conspicuous.

Temperature

An open question in our minds, before seeing

the anoles of Malpelo in the field, concerned temperature adaptations. This species lives on bare

rock near the equator. Would A. agassizi prefer

high body temperatures? Would it be able to resist

high environmental temperatures?

Our short-term, relatively simple studies indicate

conclusively that with respect to temperature A.

agassizi is a rather typical anoline lizard. Forty

body-temperature measurements of active animals

were taken in the field with a Schultheis thermometer. The mean temperature, representing pooled

samples of several mornings and early afternoons

was 30.6°C (24-38°C) with a standard deviation

of 1.8. We believe that this approximates the preferred body temperature as, quite obviously, the

lizards could easily have had higher body temperatures by sunning themselves briefly on exposed

rocks. Conversely, by spending more time in crevices, lower body temperatures could have been

maintained. The A. agassizi temperatures were

very similar to the Lesser Antillean species A. richardi and A. aeneus (Schoener and Gorman, 1968),

and comparable to, but slightly lower than, A.

homolechis on Cuba (Ruibal, 1961), a filtered

sun animal.

Anolis agassizi does not show a temperature

adaptation to particularly high or low temperatures. This is what one might expect from the

structure of the island, which receives a great deal

of insolation but also provides a great many hiding
NUMBER 176

places to escape the sun. The most extensive slopes

and cliffs orient north and south so that large parts

of the island are in total shade for either the

morning or the afternoon. This is probably the

most peculiar aspect of this anole's thermal ecology.

The A. agassizi were active during most of the day

in the shade and were seen basking only briefly

when the sun first reached them. Since we worked

on the west side, there were large areas where we

saw anoles active in the early morning and which

they abandoned fairly shortly after the sun reached

them (see p. 35).

The situation is quite unlike that which is usual

for a tree lizard who seldom needs to move more

than a few inches to get into the shade. An A.

agassizi must move several meters if it is to exploit

many of the rocks and cliffs. In the absence of flying predators it does not need to worry about keeping close to shelter.

Anoles that live in areas of high insolation tend

to have either enlarged scales or black peritoneal

pigment. Anolis agassizi has the latter. The walls

of the body cavity are pigmented, most heavily on

the posterior half. There is some pigment on the

heart and on the last 1 or 2 cm of the intestine. In

some examined A. agassizi the entire dorsal surface

of the lower intestine is pigmented.

To determine adaptations to high temperatures,

a simple heating experiment was performed on

animals brought to Panama from Malpelo. These

experiments were conducted immediately upon

our return, and the lizards could not have acclimated to the new climatic regime. Individuals

representing all size classes were tested. In these

experiments we suspended a bright lamp above a

cage that measured approximately 30 X 30 X 45

cm. The cages were constructed of wood with two

glass sides. Two parallel series of experiments were

run: in one, the lamp was placed at a height of

30 cm above the cage top; in the second, at 42 cm.

Each individual was dropped into the cage, the

light turned on, and the response observed for 3

minutes.

It has been observed that when an anole is

dropped into a novel situation its typical first response is to "freeze." As it warms up under these

experimental conditions, it starts moving about

with the onset of discomfort; finally it opens its

mouth and begins panting (Ruibal, 1961).

All lizards moved within 50 seconds with the

light source at 30 cm, and in less than two minutes

with the light source at 42 cm. The temperature of

onset of panting was not correlated with body size,

and only slightly with the distance between heat

source and cage. With the heat source at 30 cm,

ten lizards began panting at a mean body temperature (MBT) of 34.8°C; with the heat source at 42

cm, seven lizards began panting at MBT of 33.7°C.

We scored trials only when the lizard moved,

then panted, the reason being that occasional animals opened the mouth soon after being dropped

into the cage. In all cases where mouth opening

preceded movement, the time of the trial was very

short and the body temperature was well below

that of all other trials. To confirm that movement

was a response to the heat, a transparent heat filter

was inserted between the light and the cage. In

eight of 10 cases the lizard did not move at all for

more than 2.5 minutes (trial was discontinued at

3 minutes).

This temperature at which panting begins

(pooled mean, 34.2°C) is considerably lower than

published data for several Puerto Rican species of

Anolis (Heatwole et al., 1969), and is between data

for two Cuban species (Ruibal, 1961), A. allogus

of deep shade (30.3°C) and A. homolechis of filtered sun (36.2°C).

Quite clearly, A. agassizi is not specifically

adapted to high body temperatures. Because of the

north-south orientation of Malpelo and the great

contrast between sun and shade on each part of

the island with the passage of each day, some of

the social organization discussed below may be related to the lack of temperature adaptation. An

animal cannot afford to defend a restricted territory if it is unliveable for half of every day.

Water and Drinking

Many anoles that occur in very arid microhabitats (A. poncensis in Puerto Rico, A. auratus

in Panama) tend to have enlarged dorsal scales.

Anolis agassizi does not. This reinforces our impression that Malpelo is not as dry as the lack of

vegetation suggests. Lying within the doldrums and

being of some height, Malpelo probably receives

frequent rain during most of the year. Water probably does not stand very long on the exposed bare

surfaces of cliffs and rocks. But fresh water was

found spottily over most of the island during our
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

visit, even though it did not rain. The volcanic

rock of which the island is made is porous and

there are many small seeps, springs, and rock pools.

We saw anoles drinking together at a small seep

and it may be that much of their water comes from

such limited and shared sources.

Initial tests done by Paul Licht of the University of California at Berkeley on captive A. agassizi

suggest that they lose water under drying conditions much as typical Antillean anoles do, and

have no special adaptations either to prevent desiccation or to tolerate water loss. Several specimens

collected on Malpelo immediately excreted fluid

urine, implying no special need for water conservation.

Reproduction, Sex Ratio, and Size Class

Distribution

Our collection on Mapelo leads to the conclusion that the lizards must be highly seasonal (or at

least periodic) in their reproduction, or that we

were guilty of very biased sampling, or both (they

are not mutually exclusive). The smallest lizard

collected was 41 mm (SV). This is well above the

hatching size of 25 to 30 mm (p. 35). Thus, we

failed to find evidence of any really young lizards.

It is often the case that Anolis are more secretive

as young juveniles and special efforts are needed

to collect them, but we had enough people spending enough time in prime lizard habitat to make

us believe that hatchlings were truly scarce, if not

totally absent.

One of us (GCG) examined the two large museum series of A. agassizi: the Slevin specimens in the

California Academy of Sciences, collected in December 1927; and the Cyrus Perkins specimens in

the San Diego Society of Natural History, collected

in December 1931 and January 1933. Slevin (1928)

wrote of his trip to Malpelo, whereas no information is available on the two journeys made by

Perkins. Both these series then were made at approximately the same time of year (DecemberJanuary) and ours was made in the first days of

March. The smallest lizards in these collections

were 64 mm (San Diego) and 56 mm (California

Academy). Thus nothing is known about the time

of hatching or hatchling ecology in nature.

Studies on the reproductive cycles of tropical

Anolis (Licht and Gorman, 1970; Sexton et al.,

1971) indicated that within a population some females were likely to be reproductive (with an

oviducal egg or an enlarging follicle) throughout

the year, the percentage seemingly affected by rainfall. Thus in wet months 100 percent of the females might be scored reproductive, but in extensive dry seasons, reproductivity could approach

zero.

The reproductive condition of the females on

Malpelo also hints at seasonality. In our collection, five females had oviducal eggs, the smallest

female being 72 mm SV; two additional females

had enlarging follicles, these were both large specimens (86 and 87 mm). Without adequate samples

we cannot determine the size at sexual maturity.

Since our smallest reproductive was 72 mm, let us

consider this our cut-off point. There were six

females larger than 71 mm which were completely

nonreproductive. Thus, in early March, about 50

percent of the collected female A. agassizi (6 of 13)

were nonreproductive. In the museum samples

(December-January) six of eight females (75 percent) larger than 71 mm were nonreproductive.

To determine whether there is true seasonality or

whether there is merely a low level of reproductive activity parceled out throughout the year

would take further collections of A. agassizi.

The males presented the most aberrant and unexpected findings. They could be divided into

three groups. There were large males with the

obviously developed secondary sexual characters,

e.g., black head and erect nuchal crest. These

lizards ranged from about 90 to 114 mm SV and

in weight from 13 to 34 grams. There were also

smaller males (71 to 85 mm SV and 7 to 13 g) that

were sexually mature, but lacked the impressive

secondary sexual characters of their larger neighbors. In other words, they looked like females.

This is not particularly surprising, for in other

studies of Anolis' reproductive cycles (Licht and

Gorman, 1970) it has been noted that males are

physiologically sexually mature long before they

are fully grown and, in fact, before they could be

categorized as socially mature, i.e., assuming the

social role of an adult male.

But there was a remaining class of animals that

puzzled us in the field. Six male specimens (75, 85.

86, 90, 94, and 100 SV) lacked male secondary

sexual characters. Of interest is that three of these

were 90 to 100 mm long and were as large as some
NUMBER 176 35

black-headed males, yet all six had completely

regressed gonads, to the extent that we at first

labeled them "intersexes." Microscopic examination led to the conclusion that these were fully

regressed males. Nothing like this has been seen

in studies of Caribbean Anolis. Although most

tropical anole species show a moderate seasonal

cycle for gonad size, the testes are never completely

regressed; they are always obviously testes, and in

most species they produce sperm throughout the

year. Thus there is a testes weight cycle in Malpelo

Anolis. Furthermore, the population rarely shows

a large variation for testes condition, i.e., the males

are generally in phase with one another. In contrast, on Malpelo, we found some males that appeared fully reproductive and others that appeared

fully regressed.

The museum collections also had specimens

with fully regressed gonads. In the San Diego collection there were putative males of 69, 78, 79, 82,

and 83 mm SV that were fully regressed, yet males

as small as 74 that were reproductive. There is no

way to determine whether this is a seasonal phenomenon or whether it is a population phenomenon. The general lack of aggression that we

observed (p. 36) might be related to the large

number of nonreproductives in the population.

Male Anolis are often more conspicuous than

females and more likely to hold their ground

against an approaching collector. Thus they have

greater likelihood of ending up in museum jars.

Our collection consisted of 27 males and 14 females, a sex ratio of almost 2:1. A pool of our sample with the museum collections (and restricting

ourselves to animals 72 mm and greater) shows a

sex ratio of 62:27, favoring males. We find it hard

to believe that this is simply sampling artifact, but

serious demographic study would have to be undertaken to resolve the problem.

Reproduction in Captivity

Four females returned to Panama laid eggs, one

at a time. Time intervals (n = 18) between successive eggs varied from 4 to 27 days (x = 13.6).

Eggs were kept under ambient climatic conditions

in Panama (which must be close to those on Malpelo) and 13 hatched with an incubation time of

48 to 69 clays (x = 57.8). The hatchlings measured 25 to 30 mm SV (x

0.42 to 0.81 g (x = 0.62).

= 27.7) and weighed

Social Interactions

The anoles literally swarmed in our study area.

The most impressive observation was the relative

lack of aggressive encounters. There appears to be

a very strong tendency towards social facilitation,

i.e., when an anole took the initiative to approach

an object, others in the area were likely to follow

suit. They also appear to have a strong tendency

to examine any unusual object, making it very difficult to perform unbiased observation without a

blind. Thus the anoles frequently made active approaches to the observers, on several occasions

even climbing on the camera and tripod set up to

film their behavior. We could do little to avoid

them—backing off is not recommended on Malpelo's steep cliffs.

We interpret the social facilitation as an adaptation to patchy but superabundant food sources.

Thus it is likely that the carcass of a bird may

often serve as a focus for feeding of many anoles in

a given area. In the West Indies, a well-established

method for attracting large numbers of anoles is

to break open a termite nest. Under such conditions, large numbers will often gather and feed

actively with little aggressive interaction. The entire social organization of A. agassizi seems to be

based upon this type of situation.

Home ranges were large and activity was not

restricted to a few preferred perches. The following anecdotal observations support the conclusion

that home ranges overlap widely with residents

showing little aggression. Up to 8 anoles simultaneously would lick an orange, with up to 15 within

a meter of it. One morning, approximately a half

dozen lizards that had been active on a smooth

rock ledge moved laterally about 10 meters to a

cliff face. This occurred about a half hour after the

sun reached the ledge. Presumably, they were

abandoning a foraging area that had become too

hot. Many then gathered to try to catch a dangling

trout fly without showing any aggressive interaction. On one accasion, a group of about a dozen

anoles assembled at our portable radio and climbed

all over it, though it provided neither food nor

water. After some moments, all drifted away. This
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

is an example of the social facilitation referred to

above.

H. Wolda reported that as he turned over rocks

looking for invertebrates, he was followed by a

group of anoles that fed on the food that he exposed. The A. agassizi in a walk-in cage in Panama

gather around anyone who enters, as if expecting

food.

Occasionally we saw aggressive interactions between individuals, including chasing and some

display. The instances involved crested vs. crested

males, crested vs. noncrested males, and noncrested

vs. noncrested individuals of unknown sex. But we

emphasize that we saw remarkably little display in

comparison to our experiences with West Indian

anoles. The only display that we saw was a very

simple multiple bob. This varied somewhat in

amplitude and it was given sometimes with and

sometimes without the dewlap being spread. The

display was usually seen in seemingly aggressive

situations, but was also occasionally given in a

somewhat less vigouous form when a lizard moved

and stopped.

The captive anoles in Panama have shown little

in the way of additional kinds of display. We have

seen males jaw fencing, jaw locking and biting one

another. More elaborate displays have not been

seen, nor any other simple displays. Males in captivity that are courting approach the female with

the simple multiple bob, followed by pauses. This

is similar to courtship displays of other Anolis

species. Anolis agassizi has a display repertoire that

is more reduced than is typical for the genus and

appears more like a sceloporine lizard. This is probably a secondary simplification.

In a large outdoor cage in Panama, adult males

show evidence of having developed a rather complex dominance hierarchy.

On Malpelo, the anoles exhibited direct competition for food. Frequently, when an anole had

worried off a bit of orange pulp or picked up a

piece of "Chuckle" too big to swallow, immediately

it would run off with it. Sometimes it was pursued

by several other lizards, including much smaller

individuals that would try to seize the food from

its mouth. In these cases there was very little aggression shown.

No anoles attempted to monopolize the resource

by chasing or displaying at others. This was true

even of the first individual to arrive at a resource.

There was not an initial territoriality that subsequently broke down under repeated attacks as has

been described for gulls by Drury and Smith

(1968).

At the orange, when a new lizard arrived with a

rush, one already there might run off. In general,

large anoles supplanted smaller ones, but no organized dominance hierarchy was evident.

Introducing anoles tethered to a long stick at

various spots both close to and far from other

anoles produced a little interaction and in one

case an attack and bite. It also produced a little

display. The residents did not seem to be afraid of

the intruder, nor of the way it was presented but

in most cases they were just not very interested.

The social behavior in Anolis agassizi differs

from that described for any other species of anole.

This is most readily interpretable in terms of the

atypical structural niche, viz., in place of discrete

defensible perch sites (tree trunks, fence posts,

etc.) there are large areas of cliff face that are exposed to complete shade or complete sun for

many consecutive hours each day. In addition, food

resources are presumed to be much patchier than

in a typical tropical environment.

The basic difference in social organization of

this species lies in the reduced overt aggressive

interactions among individuals, and the high degree of social facilitation such that directional

movements of one individual are likely to prompt

similar investigatory movements in several other

individuals. In anthropomorphic terms, this species

is extremely curious.

Both males and females seem to have overlapping home ranges. They do not defend territories

or individual areas. There is a high degree of tolerance for the close approach of conspeciflcs at a

common resource, be it food, water, or perch site.

There are occasional overt aggressive interactions

when "individual distance" appears to be violated, but aggression is seemingly not associated with

violation of specific space (territory). Widely overlapping home ranges without aggression have been

observed in females of A. valencienni in Jamaica

(R. Trivers, Harvard University, pers. comm., and

A. biporcatus in Panama, A. S. Rand and pers. obs.).

But in both these species the males are much less

tolerant of members of the same sex.

Anolis agassizi not only has a reduced frequency

of display interactions but also seems to have a re-
NUMBER 176 37

duced display repertoire. This is in strong contrast

to the anoline species in the West Indies (Gorman,

1968) and to A. townsendi of Cocos Island (Carpenter, 1965). Our tentative explanation is: first,

that, with reduced aggression and territorial defense, there are fewer bits of information to be

communicated; and second, that the striking, permanently displayed color pattern and erect nuchal

crest of adult, reproductively active males assume

much of the communication function otherwise

transmitted by complex movements of head, dewlap, and tail (display action patterns). The only

West Indian anole known to have reduced display

is Anolis ferrens of Marie Galante which is like A.

agassizi in being unexpectedly large and in having

a very striking permanently erect crest (in this case

on the tail) in adult males.

We would suggest that the reduced aggression

is a direct adaptation to living under conditions

where resources are scattered so that a large home

range is necessary to include them. Furthermore,

the resources are often so abundant that there is

enough for all and consequently little advantage

gained by attempting to monopolize them. The

reduced amount of display permits efficient utilization of common resources. The social facilitation

("curiosity") aids in the discovery of these resources.

The social organization that we have described

for A. agassizi is very different from that described

in studies of A. sagrei (Evans, 1938), A. carolinensis

(Greenberg and Noble, 1944), A. lineatopus

(Rand, 1967), and A. nebulosus (Jenssen, 1970).

From these studies we have come to expect anoles

to show a more or less similar pattern. In these

species the adult male maintains a small home

range centering on one or more preferred perches.

The home range is defended against other adult

males as a territory and the resident gives frequent

assertion displays. An adult female maintains a

still smaller home range which she may defend

against other females and against similarly sized

subadult males. The territory of an adult male

typically overlaps those of one or more adult females. All of the species showing this general pattern of social behavior have similarities in their

ecologies. Evidence is accumulating from recent

studies of species with different ecologies (e.g., A.

aeneus, Stamps, 1973; A. valencienni, Trivers, pers.

comm.; A. biporcatus, A. S. Rand, pers. obs.) that

there is more variability between species of Anolis

in social organization than had been realized. We

have attempted to show in this paper that the

rather unusual social organization of A. agassizi

is an adaptation to its unusual enviroment. We

predict that A. agassizi will remain an extreme in

the continuum of social organization among anoles,

but that intermediates between it and the more

common types exist and, further, that the continuum will be expanded in several quite different

directions as species adapted to unusual ecologies

are described.

Literature Cited

Andrews, R. M.

1971. Structural Habitat and Time Budget of a Tropical

Anolis Lizard. Ecology, 52 (2):262-270.

Bond, J., and R. M. deSchauensee

1938. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part II: The Birds of

Malpelo Island, Colombia. Proceedings of the Academy of Natural Sciences of Philadelphia, 90:155-157,

plates 9-11.

Carpenter, C. C.

1965. The Display of the Cocos Island Anole. Herpetologica, 21 (4):256-260.

Drury, W. H., and W. J. Smith

1968. Defense of Feeding Areas by Adult Herring Gulls

and Intrusion by Young. Evolution. 22 (I): 193-201.

Dunn, E. R.

1939. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part III: The Lizards

of Malpelo Island, Colombia. Notulae Naturae of

the Academy of Natural Sciences of Philadelphia,

4:1-3.

Etheridge, R.

1960. The Relationships of the Anoles (Reptilia: Sauria:

Iguanidae): An Interpretation Based on Skeletal

Morphology. Unpublished Ph.D. Dissertation, University of Michigan. Ann Arbor: University Microfilms Inc.

Evans, T. L.

1938. Cuban Field Studies on Territoriality of the Lizard,

Anolis sagrei. Journal of Comparative Psychology,

25:97-125.

Fowler, H. W.

1938. The Fishes of the George Vanderbilt South Pacific

Expedition, 1937. Monographs of the Academy of

Natural Sciences of Philadelphia, 2: v + 349 pages,

10 plates. [Malpelo, pages 5-6.]

Gorman, G.

1968. The Relationships of Anolis of the roquet Species

Group (Sauria, Iguanidae), III: Comparative Study

of Display Behavior. Breviora, 284:1-31.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Gorman, G., and J. Boos

1972. Extinction of a Local Population of Anolis Lizards

through Competition with a Congener. Systematic

Zoology, 21:440-441.

Greenberg, B., and G. K. Noble

1944. Social Behavior of the American Cameleon (Anolis

carolinensis Voigt). Physiological Zoology, 17 (4):

392-439.

Heatwole, H., T. H. Lin, E. Villal6n, A. Muftiz, and A. Matta

1969. Some Aspects of the Thermal Ecology of Puerto

Rican Anoline Lizards. Journal of Herpetology,

3 (l-2):65-77.

Janssen, T. A.

1970. The Ethoecology of Anolis nebulosus (Sauria,

Iguanidae). Journal of Herpetology, 4 (1-2): 1-38.

Lazell, J. D., Jr.

1972. The Anoles (Sauria, Iguanidae) of the Lesser Antilles. Bulletin of the Museum of Comparative

Zoology, Harvard University, 143 (1): 1-115.

Licht, P., and G. C. Gorman

1970. Reproductive and Fat Cycles in Caribbean Anolis

Lizards. University of California Publications in

Zoology, 95:1-52.

Pianka, E. R.

1970. Comparative Autecology of the Lizard Cnemidophorus tigris in Different Parts of Its Geographic

Range. Ecology, 51 (4):702-720.

Philibosian, R.

1972. Territorial Behavior and Population Regulation in

Anoline Lizards. Unpublished Ph.D. dissertation,

University of California, Riverside.

Rand, A. S.

1964. Ecological Distribution in Anoline Lizards of

Puerto Rico. Ecology, 45 (4):745-752.

1967. Ecology and Social Organization in the Iguanid

Lizard Anolis lineatopus. Proceedings of U. S.

National Museum, 122 (3595): 1-79.

1969. Competitive Exclusion Among Anoles (Sauria:

Iguanidae) on Small Islands in the West Indies.

Breviora, 319:1-16.

Ruibal, R.

1961. Thermal Relations of Five Species of Tropical

Lizards. Evolution, 15:98-111.

Schoener, T. W.

1969. Size Patterns in West Indian Anolis Lizards, I: Size

and Species Diversity. Systemic Zoology, 18:386-401.

Schoener, T. W., and G. C. Gorman

1968. Some Niche Differences in Three Lesser Antillean

Lizards of the Genus Anolis. Ecology, 49 (5):819-830.

Schoener, T. W., and A. Schoener

1971. Structural Habitats of West Indian Anolis Lizards,

I: Lowland Jamaica. Breviora, 368:1-53.

Sexton, O. J., E. P. Ortleb, L. M. Hathaway, R. E. Ballinger,

and P. Licht

1971. Reproductive Cycles of Three Species of Anoline

Lizards from the Isthmus of Panama. Ecology,

52(2):2Ol-215.

Slevin, J. R.

1928. Description of a New Species of Lizard from

Malpelo Island. Proceedings of the California

Academy of Sciences, fourth scries, 16:681-684,

plates 25, 26.

Stamps, J. A.

1973. Displays and Social Organization in Female Anolis

aeneus. Copeia, 1973 (2):264-272.

Stejneger, L.

1900. Description of Two New Lizards of the Genus

Anolis from Cocos and Malpelo Islands. Bulletin

of the Museum of Comparative Zoology, 34 (6): 161-

164, 1 plate.

Williams, E. E.

1969. The Ecology of Colonization as Seen in the Zoogeography of the Anoline Lizards on Small Islands.

Quarterly Review of Biology, 44:345-389.

1972. The Origin of Faunas: Evolution of Lizard Congeners in a Complex Island Fauna: A Trial Analysis. Pages 47-89 in volume 6 in T. Dohzhansky, M.

Hecht, W. Steere, editors, Evolutionary Biology.

New York: Appleton, Century, Crofts.
Notes on the Natural History of

Diploglossus millepunctatus

(Sauria: Anguidae)

A. Ross Kiester

ABSTRACT

The natural history of the large anguid lizard

Diploglossus millepunclatus from Malpelo Island

is described. This species is characterized by its

abundance, flexible thermal regime, euryphagy,

and curiosity.

Introduction

The large anguid lizard Diploglossus millepunctatus is one of the most conspicuous elements of

the fauna of Malpelo Island. Because the island

is seldom visited little has been reported of its

natural history. Here we begin to fill in some of

the gaps in our knowledge of this species.

Diploglossus millepunctatus was first described

by O'Shaughnessy (1874) who listed the typelocality as "the north-west coast of America."

Slevin (1928) described Celestus hancocki from

Malpelo Island, noting that it was similar to the

unique type of O'Shaughnessy. Dunn (1939), who

had additional material, synonomized Slevin's

name on the basis of similarities of his series with

the type of millepunctatus and the fact that there

was some indirect historical evidence that the original specimen could indeed have been collected

on Malpelo. Dunn also placed the species in the

genus Diploglossus arguing that Celestus was not

recognizable. While Dunn's generic allocation of

the species millepunctatus has prevailed, the issue

A. Ross Kiester, Museum of Comparative. Zoology, Harvard

University, Cambridge, Massachusetts 02138.

of the validity of the genus Celestus requires further attention (Myers, 1973).

Diploglossus millepunctatus is among the largest

of the anguids, ranging up to 250 mm snout-vent

length with a tail length of 162 mm and a weight

of 268 grams. It is rather heavy-bodied with short

limbs. Older males possess the enlarged wide heads

that are characteristic of some other anguid genera

such as Gerrhonotus (Stebbins, 1954). The body

has a very shiny metallic appearance and like that

of other anguids is encased in a solid armor of

osteoderms. The tail tends to be rather blunt in

adults but is more typically tapered in the young.

The color pattern is dark black or brown-black

with small yellow or yellow-white flecks in great

profusion, as the specific name suggests (Figure

16).

Fir.tRF. 16.—Diploglossus niillepurictatus aggregated about a

young Blue-faced Booby (Sula dactylalra) which has just

been fed.

39
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ACKNOWLEDGMENTS.—I wish to thank T. L.

Chorba of Oxford University, G. C. Gorman of the

University of California, Los Angeles, J. A. Hoffman of the Smithsonian Astrophysical Observatory,

A. S. Rand of the Smithsonian Tropical Research

Institute; and W. M. Rand of the Massachusetts

Institute of Technology for assistance in collecting

data, and R. B. Huey of the Museum of Comparative Zoology for aid in the preparation of this

manuscript.

Habitat

Diploglossus millepunctatus is found virtually

everywhere on the island from next to the water's

edge to the top of the summit (376.4 m). Lizards

usually did not occur far from the crevices into

which they retreated. Otherwise, they scramble

over the bare rocks and cracks, sometimes moving

with agility in an almost serpentine fashion, but

also occasionally falling.

Although these lizards live next to the ocean

and are capable of swimming, we were unable to

get any individuals to enter the water voluntarily

as Slevin (1928) reported they would do.

Population Size and Structure

These animals are extremely common. No quantitative measurements of population density were

attempted, but rough counts indicated that a density of 1 per 10 square meters was perhaps a reasonable estimate. The population seems to consist

mostly of adults. Of the literally hundreds of animals that were observed only three young were

observed. Except for these individuals, the entire

population fell into a size range of approximately

180-260 mm snout-vent length. The juveniles were

approximately 75 mm snout-vent length and no

animals of an intermediate size were seen. This

striking size distribution might possibly be due to

the fact that younger animals tended to remain

hidden in crevices. However, the fact that some

young were seen while no intermediate sized animals were found seems to indicate that the observations reflect a real phenomenon, namely that

reproduction is seasonal or erratic and that our

sample was taken prior to a period of reproductive

activity.

Activity and Thermal Relations

Although no nights were spent on the island, it

seems likely that D. millepunctatus spends nights

inside crevices. Upon arriving on the island at 0800

or earlier we observed few lizards out in the open

but saw many in crevices. Later, the lizards moved

out to bask in the midmorning sun. By midday

most had retreated back into the crevices. During

the afternoon most remained in or adjacent to

crevices and only an occasional individual roamed

in the open.

Overall mean body temperature (cloacal temperature ± se) of 12 D. millepunctatus during

activity was 27.5 ± 0.64°C (range 24.8 to 32°C).

However, body temperatures shifted upward between morning and afternoon. The mean body

temperature of six lizards active between 0838 and

0910 was 25.8 ± 0.3°C, whereas the mean body

temperature of six lizards active between 1324 and

1342 was 29.3 ± 0.7°C. There was no overlap in

body temperature between these two samples, and

the difference between the samples is significant

(p = 0.01, Kolmogorov-Smirnov test, Siegel, 1956:

127).

Among anguids only Ophisaurus attenuatus is

reported to have a higher body temperature (Fitch,

1956:444). However, as with Anolis agassizi (p.

33), D. millepunctatus appears in general to have

no special physiological adaptations to the extreme thermal environment in which it lives. The

ready availability of the cooler crevices seems to

afford D. millepunctatus a means of cooling itself

when the temperatures on the exposed parts of the

island are extreme. On the other hand, much of

the island remains in the shade until the very late

morning, by virtue of the steep rock walls, and

this species can also be active at lower temperatures. Thus it appears to be flexible with regard

to the temperatures at which it is active.

Drinking and Water Relations

Diploglossus millepunctatus drinks readily in

captivity. On Malpelo standing fresh water was

observed in a few locations and water seepages

were seen in many of the crevices, although most

of the island was dry. It is impossible to know from

our observations what the seasonal pattern of water

availability is, or whether there is water available
NUMBER 176 41

in the deeper crevices. Rock surfaces on Malpelo

ranged from very porous to extremely smooth and

hard. In either case it seems that water would not

stand on the surface of the island for any appreciable time after a rain storm, either percolating

down into the rock, running off rapidly, or evaporating. Although it did not rain during our visit,

captive animals displayed an extremely rapid response to water. When kept in a large pen with no

available water, the animals would respond to the

first drops of water sprayed into the cage by running immediately to the nearest puddle that was

forming. This response seemed to be about as fast

a movement as the animals were capable of. They

responded by coming out of crevices or down off

the tops of large bricks that were in the cage. This

rapid response appears to be a specific behavioral

adaptation to a transiently available water supply.

Food and Feeding Behavior

Although this species will eat the usual kind of

prey items that are to be expected for a lizard of

this size, such as insects and invertebrates, the

bulk of the food taken by adults is quite out of the

ordinary for a lizard. There are small insects on

Malpelo, and in captivity D. millepunctatus will

readily take crickets and mealworms. Actual feeding on insects on the island was not observed, however, probably because the insects occur only inside

the crevices. Out in the open the lizards were observed to feed on a variety of bizarre foods. Both

Slevin (1928) and Dunn (1939) reported finding

FICURE 17.—Diploglossus millepunctatus feeding on a fish

regurgitated by the booby.

crabs in the stomachs of the animals they examined, and Slevin also reported finding feathers. We

observed this species feeding on carrion consisting

of both dead crabs (Gecarcinus malpilensis) and

a Blue-faced Booby (Sula dactylatra). In the case

of the booby, the lizards may have also been feeding on the insects attracted to the carcass.

Most of the feeding observed was connected with

the activities of living boobies. On two occasions

animals partially consumed fresh booby feces. The

most spectacular feeding behavior occurs when an

adult booby is feeding its young regurgitated fish.

When an adult bird returns and alights near its

young, the young bird begins to squawk loudly.

This noise attracts the attention of any nearby D.

millepunctatus which immediately run to the vicinity of the two birds. If any of the fish is dropped,

the waiting lizards immediately snatch the food

and drag it away. We observed this complete sequence of behavior on two occasions, and saw

lizards attracted to calling boobies on several others.

In one case, more than seven adult lizards that had

aggregated about two birds successfully carried off

and consumed an entire fish (Figures 16, 17). During such activity the birds appear oblivious to the

lizards. In captivity these lizards would readily eat

canned tuna or fish-flavored cat food.

It appears that the boobies provide a substantial

amount of food for the lizards, at least during the

nesting season. Obviously this source of food is

not available year round. The lizard's large stumpy

tail may possibly serve as a storage organ, and

several of the individuals which we examined had

large fat bodies in the abdominal region. Presumably the young feed more on the insects that are

available in the crevices and that may not be as

seasonal as the supply of fish.

Social Behavior

In general this species does not appear to have

a complex social behavior system. Individuals frequently ignore each other, although short chases

were occasionally observed. The only activity that

could be interpreted as a display is a particular

"head-up" posture in which a lizard sits with its

head held back so that it points almost directly

upwards. Although two lizards within eyesight of

each other could be observed to do this, no obvious

reaction was observed. Further observations will
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

be needed to confirm that this is indeed a display.

Social facilitation for feeding appears to be well

developed in this species. A running individual

seems to attract the attention of others who frequently give chase. This reaction appears to be in

contrast to the response elicited by a slowly moving

individual. When food or a source of food is present, it is not clear how much of the attraction is

due to the food itself and how much to the other

lizards; however, aggregation can be extremely

rapid. As mentioned above, seven or eight animals

aggregated about two boobies, and the aggregation

took place in a radius of over 5 meters in about 2

minutes. Rapid aggregation in itself is evidence for

social facilitation (Kiester and Slatkin, 1974), but

this point needs to be investigated experimentally.

One pair was observed copulating. They were

found in a crevice in the shade. The male held the

head of the female in his jaws, biting from the

side. His body was positioned laterally and intromission took place from the side. The pair was

found in copulation in the midmorning and remained in the same position for two and one-half

hours. The position is similar to that described

for Gerrhonotus multicarinatus (Fitch, 1935:18).

Reproduction

A series of 18 specimens collected on a single

day during the visit (early March) showed marked

reproductive activity. Females had many welldeveloped embryos, and males had large testes.

Females brought to Panama and maintained in

captivity gave birth to live young two months

later. Other females in this same group and some

females taken to Cambridge, Massachusetts, which

initially appeared gravid showed no embryos on

examination after their death some weeks later.

Thus it may be possible that resorption of embryos

can occur.

We may conjecture that D. millepunctatus will

have its reproductive cycle tied to that of the

booby. From our collections we know that the

lizards were in reproductive condition (and often

fat) near the end of the fledging period of the

boobies. If the amount of food available to the

adult lizards is markedly increased by the appearance of nestling boobies which need to be fed, then

breeding by the lizards might be expected to follow

this period.

Conclusions

This species appears to be very opportunistic,

ready to take advantage of a transiently available

resource on split second notice. Associated with

this opportunism is a high level of curiosity and a

tendency to respond to anything novel or disturbing in the environment. We found that individuals

of D. millepunctatus would approach human

beings or their artifacts with little regard for possible danger. This curiosity seems linked to the

unpredictable and meager food supply on Malpelo

Island where any disturbance may likely mean

food rather than danger for the lizards, since there

are no natural predators (with the possible exception of migrating raptors; Bond and deSchauensee,

1938). Thus the benefits of curiosity are great

while the disadvantages normally associated with

this behavior are reduced or absent. In this way

D. millepunctatus is typical of predators on remote

islands (Carlquist, 1965).

Literature Cited

Bond, J., and R. M. deSchauensee

1938. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part II: The Birds of

Malpelo Island, Colombia. Proceedings of the

Academy of Natural Sciences of Philadelphia, 90:

155-157, plates 9-11.

Carlquist, S.

1965. Island Life. Garden City, New York: The Natural

History Press.

Dunn, E. R.

1939. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part III: The Lizards

of Malpelo Island, Colombia. Notulae Naturae of

the Academy of Natural Sciences of Philadelphia,

4:1-3.

Fitch, H. S.

1935. Natural History of the Alligator Lizards. Transactions of St. Louis Academy of Sciences, 29:1-38.

1956. Temperature Responses in Free-living Amphibians

and Reptiles of North-eastern Kansas. University

of Kansas Publications of Museum of Natural History, 8:417-476.

Kiester, A. R., and M. Slatkin

1974. A Strategy of Movement and Resource Utilization.

Theoretical Population Biology. [In press.]

Myers, C. W.

1973. Anguid Lizards of the Genus Diploglossus in Panama with the Description of a New Species.

American Museum Novitates, 2523:1-20.
NUMBER 176 43

O'Shaughnessy, A. W. E. Slevin, J.

1874. Descriptions of New Species of Scincidae in the 1928. Description of a New Species of Lizard from MalCollection of the British Museum. Annals and pel o is

i an d. Proceedings of the California Academy

Magazine of Natural History, fourth series, 13: of Sciences fourt h series > 16:681-684, plates 25, 26.

298-301.

Siegel, S. Stebbins, R. C.

1956. Nonparamelric Statistics for the Behavioral Sci- 1954. Amphibians and Reptiles of Western North Amerences. New York: McGraw-Hill. ica. New York: McGraw-Hill.
A New Gecko from Malpelo Island

(Sauria: Gekkonidae: Phyllodactylus)

Raymond B. Huey

ABSTRACT

Phyllodactylus transversalis, a new gecko from

Malpelo Island, Colombia, is characterized by absence of tubercles on arms, legs, rear of head, and

tail; absence of ear denticulation; absence of abdominal plaque; slight enlargement of dorsal tubercles, which form only two distinct paravertebral

rows; terminal lamellae of digits distinctly widened

and truncate; 13 to 15 fourth toe lamellae; ground

color medium gray-brown with dark chocolate

bands. Source of this species is uncertain.

Introduction

The gekkonid genus Phyllodactylus is widely

distributed in the western hemisphere. Recent

reviews by Dixon (1962, 1964a, 1964b) and Dixon

and Huey (1970) have revised all New World

members of the genus except species from the

Galapagos Islands. Some 45 species are currently

recognized.

The collection of lizards from the SmithsonianU. S. Navy Expedition to Malpelo Island, Colombia, included two specimens of Phyllodactylus, a

genus not collected by the few previous expeditions to the island. Morphological characteristics

indicate distinctness of these specimens, which I

here describe following the methodology of Dixon

(1964a).

ACKNOWLEDGMENTS.—I thank J. R. Dixon of

Texas A & M University; G. C. Gorman of the

University of California at Los Angeles; J. B.

Graham and A. S. Rand of the Smithsonian Tropical Research Institute; A. E. Greer, A. R. Kiester,

Raymond B. Huey, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138.

and E. E. Williams of the Museum of Comparative Zoology for helpful comments on the manuscript. I acknowledge the collectors of the typeseries, A. S. Rand and H. Wolda of the Smithsonian Tropical Research Institute.

Phyllodactylus Iransversalis (new species)

FIGURE 18

HOLOTYPE.—Adult female, Museum of Comparative Zoology (MCZ 130042), collected by A. Stanley

Rand from under a rock near the first collecting

area on Malpelo Island, Colombia, February 1972.

PARATYPE.—Immature specimen (MCZ 130043)

collected from under a rock on Malpelo Island by

Hendrik Wolda, February 1972.

DIAGNOSIS.—Differs from the Central American

tuberculosus group, the Mexican delcampi group,

and most South American groups by having small

and scattered dorsal tubercles forming only two

paravertebral rows; differs from unctus group in

possessing dorsal tubercles; differs from gerrhopygus group by absence of abdominal plaque; from

the species microphyllus by having expanded terminal lamellae and a small nostril; and from inaequalis in having dark chocolate dorsal bars, more

fourth toe lamellae (13 to 15 vs. 10 to 12), smaller

head scales (third labial snout scales 32 vs. 20 to

24; scales between eye and nostril 14 to 15 vs. 11

to 12).

DESCRIPTION OF HOLOTYPE.—Rostral twice as wide

as high, its dorsal edge almost straight with a dorsal vertical groove one-half depth of rostral; 2

internasals, somewhat rounded, their median edges

in broad contact, bordered posteriorly by 7 small

granules and a postnasal on each side; nostril

44
NUMBER 176 45

surrounded by rostral, labial, internasal, and 2

postnasals; first supralabial in narrow contact

with ventral edge of nostril; shallow depression

between internasals, slight depression in frontal

region; 15 scales on line between nostril and eye;

scales in posterior loreal region slightly larger than

scales in mid-orbital region; 32 scales across snout

at level of third labial; 22 interorbital scales; eye

large, its diameter contained in snout length about

1.5 times; eyelid with 2 inner rows of granules, a

median row of larger scales, and 1 row of large

scales on the edge, the last 8 to 9 are pointed; ear

large, its diameter contained in the eye diameter

2.7 times; scales on anterior edge of ear opening

FIGURE 18.—Holotypc of Phyllodactylus transversalis, adult

female, MCZ 130042, dorsal view.

flattened, overlapping, those on posterior edge

smaller and granular; rear of head granular without intermixed tubercles; 5 supralabials and 4 infralabials to a point below center of eye; mental

bell-shaped with posterior median edge sharply

angular, as wide as long, bordered posteriorly by

2 postmentals; postmentals slightly longer than

wide, their median edges in broad contact, followed by transverse row of 5 flattened scales.

Dorsum with scattered, small, conical tubercles;

only 2 paravertebral rows evident, extending onto

neck; paravertebral rows separated by 3 to 5 irregular rows of small granules; 40 paravertebral tubercles between axilla and groin, separated by 0

to 2 granules; remaining dorsal tubercles not in

distinct rows; dorsal granules irregular; postanal

tubercles 3 on each side, not distinct; enlarged

abdominal plaque absent; venter with 27 scales

across belly, from throat to vent 69.

Dorsal surface of upper arm with rounded,

slightly elevated scales; forearm with slightly

smaller scales of similar shape; dorsal surface of

thigh and tibia with granular scales; claw long,

visible from above and below; terminal toepad

greatly enlarged, longer than wide, truncate;

fourth toe lamellae 13 to 14; tail partially regenerated, original tail stub with a few wide median

scales on ventral surface, devoid of tubercles on

dorsal surface, but with scattered, flattened tubercles at base.

Measurements (in mm): Snout-vent length 57,

axilla-groin length 26, length of leg 25, length of

arm 17, length of tail 22 + 18, length of head 15.5,

depth of head 5.7, width of head 11.2, length of

snout 6.4, diameter of eye 4.3, diameter of ear 1.6,

distance from eye to ear 5.7.

Color in Alcohol: Dorsum medium gray-brown

ground color with white speckles; 8 distinct broad

chocolate brown transverse bars from base of head

to base of tail, somewhat broken along midline;

width of bars slightly larger than ground interspaces; lateral area of trunk as dorsal ground; arms

and legs with dark chocolate ground suffused with

light scales and small light spots, without definite

pattern; top of head with suffusion of dark brown

and small light blotches, no definite pattern; dark

chocolate brown stripe from nostril to eye, blending with head color posterior to eye, bordered below by a pale whitish line from first infralabial

through ear; belly pale yellow.
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

VARIATION (based on paratype).—Similar to

holotype except in the following characters: 14

scales between nostril and eye; 24 interorbital

scales; 6 supralabial and 5 infralabial scales to a

point below center of eye; mental nearly triangular; no contact between postmentals, followed posteriorly by transverse row of 6 flattened scales;

venter with 30 scales across belly, from throat to

vent 67; 39 tubercles in paravertebral row between

axilla and groin; fourth toe lamellae 15-14; tail

regenerated 3 + 19; snout-vent length 36 mm.

NATURAL HISTORY.—Both geckos were collected

during the day under rocks. Because of the topography of Malpelo, these geckos are necessarily

scansorial. Dixon and Huey (1970:66) have shown

that scansorial Phyllodactylus have expanded toepads, a characteristic of P. transversalis.

Two large diurnal lizards are also present on

Malpelo (Diploglossus millepnnctatus and Anolis

agassizi) and eat crustaceans as well as booby regurgitations and feces. Ants and crickets were found

in the stomachs of P. transversalis (A. S. Rand,

pers. comm.).

Accompanying papers in this volume describe

Malpelo Island (Kiester and Hoffman, pp. 13-16)

and the biology of the diurnal lizards (D. millepunctatus, Kiester; A. agassizi, Rand, Gorman,

and Rand).

ZOOGEOGRAPHIC RELATIONSHIPS.—The dorsal crossbanding and the two rows of dorsal tubercles of

P. transversalis are distinctive. These unique characters suggest long isolation from source areas

and hinder analysis of affinities.

Phyllodactylus transversalis shares many external morphological characters with the inaequalis

group of Peru (Dixon and Huey, 1970:69). After

examining the type and paratype, however, J. R.

Dixon (pers. comm.) suggested that P. transversalis may be closer to certain Mexican species (P.

paucituberculatus, P. unctus, and P. delcampi).

There are no obvious affinities with Phyllodactylus

from the Galapagos. The source of this gecko is

thus equivocal.

ETYMOLOGY.—The specific name refers to the distinctive dorsal cross-bands (Figure 18).

Literature Cited

Dixon, J. R.

1962. The Leaf-toed Geckos, Genus Phyllodactylus, of

Northeastern South America. Southwestern Naturalist, 7:211-226.

1964a. Further Data on the Geckos (Phyllodactylus) of

Islands of the Extreme Southern Caribbean. Southwestern Naturalist, 9:203-206.

1964b. The Systematics and Distribution of the Lizard

Genus Phyllodactylus of North and Central America. New Mexico State University Science Bulletin,

64:1-139.

Dixon, J. R., and R. B. Huey

1970. Systematics of the Lizards of the Gekkonid Genus

Phyllodactylus of Mainland South America. Contributions in Science of the Los Angeles County

Museum, 192:1-78, 14 figures.
Electrophoretic Estimates of Genie

Variation in, and the Relationships of,

Anolis agassizi

T. Preston Webster

ABSTRACT

From data on proteins representing 30 genes, an

average individual of Anolis agassizi is estimated

to be heterozygous at 2.1 percent of its loci. This

low value is similar to that found in other solitary

anoles and in some anole populations of relatively recent origin, but it contrasts sharply with

results from some Greater Antillian species. Very

little genetic similarity was found in the comparison of A. agassizi to five congeners.

Introduction

An insular lizard population may have relatively

little genie variation as a result of random processes and natural selection acting during its establishment. The small size of the propagule, perhaps

only a single impregnated female, should limit

variation. This holds whether the alleles carried

by the founder or founders are a random or a selected sample of those in the parental population.

Subsequently, random allelic loss (genetic drift)

during the initial or secondary periods of small

population size and selection would further reduce

variation. A low level of heterozygosity could endure, but in a persistent population many new

polymorphisms might accumulate.

A considerable loss of genie variation may have

occurred during the arrival and life of Anolis

agassizi Stejneger on Malpelo Island. Malpelo may

have received anoline immigrants only once and,

as the source population is extinct, there certainly

T. Preston Webster, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138.

have been no recent immigrants bearing new alleles. On such a small island, secondary reductions

in population size are possible. The founders probably encountered more extreme conditions on

Malpelo than those to which they were adapted,

even if the source population, like those anoles

that have been successful colonists in the Caribbean (Williams, 1969), occurred in open, edge

situations. Anolis agassizi, however, is a long established population that has perhaps experienced

pressures and had time to reacquire heterozygosity.

In this study, genie variation in A. agassizi was

estimated by electrophoretic examination of enzymatic and non-enzymatic proteins. Since there is

no prediction to which this result can be compared,

it is considered in the context of similar estimates

from other anole populations. Secondarily, many

of the same proteins were used to estimate the

genetic difference between A. agassizi and five

congeners.

ACKNOWLEDGMENTS.—I thank the following for

their contributions to this study: G. C. Gorman of

the University of California at Los Angeles, T. A.

Jenssen of Virginia Polytechnic Institute and State

University, A. R. Kiester of the Museum of Comparative Zoology, and A. S. Rand of the Smithsonian Tropical Research Institute for collecting

specimens. J. B. Graham and A. S. Rand of the

Smithsonian Tropical Research Institute, and E. E.

Williams of the Museum of Comparative Zoology

commented on the manuscript. This study was

conducted in part while I was an NSF predoctoral

fellow and was supported by NSF grant B 019801X

to E. E. Williams, Museum of Comparative

Zoology.

47
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Materials and Methods

Plasma and hemolysate were obtained from 32

A. agassizi collected at the first landing site on the

west side of Malpelo. Six individuals were used for

chromosome studies and preserved, and the remaining 26 were homogenized in distilled water

(1:2, W:V). Homogenates were centrifuged and

stored at -80°C.

Electrostarch was used for horizontal starch gel

electrophoresis. Each protein was examined in

as many electrophoretic environments as convenient and feasible. In several cases this multiplicity

of observation increased the number of mobility

differences detected. Many of the buffer systems

and protein assays were those previously used in

studies of Anolis (Webster, Selander, and Yang,

1972; Webster and Burns, 1973). Additional techniques are those commonly used in studies of vertebrate proteins (Brewer, 1970; Shaw and Prasad,

1970; Selander et al., 1971).

Twenty-eight enzymatic and nonenzymatic proteins were examined: leucine aminopeptidase, two

peptidases, fumarase, creatine kinase, phosphohexose isomerase, eight dehydrogenases (lactate,

two malate, two isocitrate, alcohol, a-glycerophosphate, and 6-phosphogluconate), indophenol

oxidase, two aspartate aminotransferases, two phosphoglucomutases, an esterase, three "general proteins" (A,B,C), hemoglobin, transferrin, and three

plasma proteins. Like other anoles (Webster, Selander, and Yang, 1972), A. agassizi appears to have

four or five genes encoding the polypeptides that

are subunits of hemoglobin, but it is treated here

as if it were under two-gene control. Lactate dehydrogenase is encoded by two genes. Since each of

the remaining proteins appears to be encoded by

a single gene, the products of 30 genes have been

examined.

Many of the proteins studied in A. agassizi were

compared with their apparent homologues in these

congeners: A. garmani (Jamaica: Mandeville),

A. luciae (St. Lucia: Vigie Beach), A. squamulatus

(Venezuela: Parque Nacional Rancho Grande

"Henry Pihier"), A. marmoratus (Guadeloupe:

3.6 km SW Capesterre), and A. singularis (Haiti:

Savane Zombi). To obtain an estimate of differentiation between members of the same species

group, A. luciae was compared to A. extremus (St.

Lucia: Vigie Beach, a recently introduced population). Single individuals or pooled extracts were

used in these comparisons, but the variation in

each population has been studied and considered.

Genie Variation

In A. agassizi variation was detected at 4 of the

30 loci. One of 26 individuals was heterozygous

for Protein B, 2 of 26 were heterozygous for one of

the malate dehydrogenases, and 2 of 26 were heterozygous for one of the peptidases. Transferrin was

represented by 3 variants, with 14 of 32 individuals

heterozygous. The remaining 26 genes appear to

be monomorphic. Thus the average individual of

this population is estimated to be heterozygous at

2.1 percent of its loci (H = 0.021).

This result can be evaluated within the context

of a broad range of estimates of H from anole

populations (Table 3). Since a large proportion

of protein polymorphism is expected (Lewontin

and Hubby, 1966) and known (Boyer, 1972) to be

undetectable by electrophoresis, such values are

surely underestimates of the actual extent of genie

variation. Their comparison requires both that the

electrophoretically determined H be considered

as having some value as an index to actual genie

variation among structural loci or within the

genome as a whole and that some standard error

or arbitrary criterion for difference be assigned.

From a joint consideration of morphological

variation and electrophoretically determined H,

Soult et al. (1973) concluded that H has considerable power as an index to genie variation, even

when only 20 to 25 loci are examined. Two other

studies support this relationship. Kluge and Kerfoot (1973) examined several sets of vertebrate

populations, finding that within each the average

variability of a morphological character is correlated with the extent to which there has been divergence in its mean. Webster, Selander, and Yang,

(1972) provided the complementary relationship,

although their measure of morphological divergence was relatively crude. They found that among

Anolis species, those with higher values of H were

differentiated into more geographic races. The

genetic base for both morphological variation and

divergence probably involves many more genes

than the handful examined in the direct estimation of H. But whether these relationships are

simply evidence for the strength of H as an index
NUMBER 176 49

to heterozygosity at structural loci or if they extend treatment. I suggest that an H of 0.02 should be

it to other components of the genome is a matter considered indicative of less genie variation than

for further study. occurs in a population with an H of 0.08 or more.

The estimates of H for anoles are based on simi- The reader is encouraged to provide his own

lar sets of genes that received similar technical criterion.

TABLE 3.—Electrophoretic estimates of genie variation (H) in populations of Anolis lizards

Species Populations Loci H Source

CONTINENTAL SPECIES

A. carolinensis

Southeastern U.S.

GREATER ANTILLEAN SPECIES

A. stratulus

Puerto Rico

A. poncensis

Puerto Rico

A. evermanni

Puerto Rico

A. gundlachi

Puerto Rico

A. krugi

Puerto Rico

A. pulchellus

Puerto Rico

A. porcatus

Havana, Cuba

A. distichus dominicensis .

Bon Repos, Haiti

SOLITARY SPECIES

A. extremus

Barbados

A. roquet

Martinique

A. luciae

St. Lucia

A. marmoratus

Guadeloupe

A. agassizi

Malpelo

RELATIVELY RECENT POPULATIONS

A. carolinensis

South Bimini

A. angusticeps

South Bimini

A. distichus

South Bimini

A. sagrei

South Bimini

A. sagrei

Swan Island

29 0.036-0.057

22

22

22

22

22

22

24

21

0.024

0.035

0.070

0.082

0.104

0.129

0.079

0.090

22 0.036

21 0.041

27 0.012

21-23 0.014-0.033

31 0.021

29

25

27

26

28

0.064

0.000

0.043

0.009

0.052

Webster, Selander, and Yang,

1972:529

Soute, et al., 1973, fig. 1

Webster, in prep.

Soute, et al., 1973, fig. 1

Webster, in prep.

Webster, herein

Webster, Selander, and Yang,

1972:529

Lister and Webster, in prep.
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Unfortunately, H has not been estimated for

any of the South or Central American anoles.

Whether A. carolinensis, a secondarily continental

species, has more genie variation than A. agassizi

is questionable (Table 3).

Many of the Greater Antillean species represent

stocks long established on islands so large and biologically diverse that they probably have a continental influence on genie variation. Estimates of

H have a broad range, but many of the values are

higher than those obtained from anoles in any

other evolutionary or ecological context. Source

island populations of two good colonists in the

Caribbean, A. porcatus and A. distichus, appear

to be more variable than A. agassizi.

Like A. agassizi, many of the species of the Lesser

Antilles have been long established and occur in

the absence of congeners. Their islands, however,

tend to be larger than Malpelo and provide a more

typical anole environment. Estimates of H from

four species are nonetheless similar to that from

A. agassizi.

The South Bimini and Swan Island populations represent colonizations that are relatively recent (Williams, 1969). It is in such populations

that any loss of heterozygosity during and soon

after founding should be most visible. The populations of A. sagrei and A. angnsticeps on South

Bimini have little or no detectable genie variation,

although the A. angusticeps have some esterase

variation that was refractory to interpretation and

was omitted from the reported H of 0.00. Elsewhere in the Bahamas and the West Indies, however, these species have more variation (Webster,

in prep.; Lister and Webster, in prep.; see Table

3, A. sagrei, Swan Island). These results and those

for insular populations of Peromyscus (Selander

et al., 1971) and cave populations of Astyanax

(Avise and Selander, 1972) suggest that a major

loss of heterozygosity may frequently occur during

the colonization of islands; however, the South

Bimini populations of A. distichus and A. carolinensis seem to have been less affected.

Since several Greater Antillean species appear

to have as little genie variation as A. agassizi, there

is no compelling reason for concluding that it lost

much during and subsequent to its colonization of

Malpelo. If it did, it is likely that both random

and selective processes played a role, but their relative importance cannot be determined. It is

interesting to note that no variation was observed

for three esterases (only one of which was included in the estimate of H). These enzymes usually show complex patterns of variation in anole

populations, including some with little genie

variation. That the estimates of H for A. agassizi

and several other solitary anoles are fairly uniform

suggests that the net influence on genie variation

may be quite similar in these older populations.

If this consistency is confirmed by further studies

of solitary species, it will be appropriate to consider selective explanations.

Genetic Differentiation

On the basis of a protein phenotype including

the products of 24 genes, A. agassizi is quite distinct

from all five of the species with which it was compared (Table 4). Two of the species are also members of the latifrons series of the alpha section

(Etheridge, 1960): A. luciae (St. Lucia, Lesser

Antilles), like A. agassizi, has retained caudal autotomy, but A. squamulatus (Mainland, Venezuela)

has lost it. Since the latifrons series is defined by

character states that are ancestral for the alpha

section (Etheridge, 1960), it is not surprising that

the species within it should have diverged to such

an extent. Even two members of the apparently

monophyletic roquet species group (Gorman and

Atkins, 1969), A. luciae and A. extremus (native

to Barbados, Lesser Antilles), differ completely at

17 of the 29 genes examined. Anolis singularis is

as distinct from A. agassizi as are members of the

latifrons series; and the fourth alpha section species,

A. marmoratus, has no shared electrophoretic mobilities. The beta section A. garmani is no more

distinct than some of the alpha section species.

These findings shed no new light on the affinities

of A. agassizi. Except for very closely related populations, electrophoretically determined genetic difference is always an underestimate of genetic distance. When the species compared are as distantly

related as these, it even becomes unreliable as an

index to genetic distance. Thus the small differences in similarity that were observed in this study

should be considered meaningless. As concluded

previously (Webster, Selander, and Yang, 1972),

when only 20 to 30 proteins are examined, electrophoresis has little value in anoline taxonomy beyond the species group.
NUMBER 176 51

TABLE 4.—Similarity (j*,; Nei, 1972) of eleclrophoretic mobilities of proteins in

comparisons between Anolis agassizi and five congeners

Protein

aspartate aminotransferase-1

phosphohexose isomerase

leucine aminopeptidase

isocitrate dehydrogenase-2

lactatc dehydrogenase-1

lactate dehydrogenase-2

malate dehydrogenase-1

malate dehydrogenasc-2

ct-glycerophosphate

dehydrogenase

Sum of Similarities

A.

garmani

0.00

0.00

0.00

1.00

0.00

0.00

1.00

0.00

0.00

2.00

A.

marmoratus

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

A.

singular is

0.00

1.00

1.00

0.00

0.00

0.00

1.00

0.00

0.00

3.00

A.

squamulatus

1.00

0.00

1.00

0.00

1.00

1.00

0.00

0.00

0.50

4.50

A.

luciae

0.00

1.00

1.00

0.00

0.00

0.00

0.00

1.00

1.00

4.00

Values for the following are 0.00: 6-phosphogluconate dehydrogenase, aspartate aminotransferase-2, peptidase, isocitrate dehydrogenase-1, two phosphoglucomutases, fumarase, indophenol

oxidase, alcohol dehydrogenase, albumin, proteins A, B, and C.

Literature Cited

Avise, J. C, and R. K. Selander

1972. Evolutionary Genetics of Cave-Dwelling Fishes of

the Genus Astyanax. Evolution, 26:1-19.

Boyer, S. H.

1972. Extraordinary Incidence of Electrophoretically Silent Genetic Polymorphisms. Nature, 239:453-454.

Brewer, G. J.

1970. An Introduction io Isozyme Techniques. New York:

Academic Press.

Etheridge, R.

1960. The Relationships of the Anoles (Reptilia: Sauria:

Igiianidae): An Interpretation Based on Skeletal

Morphology. Unpublished Ph.D. dissertation, University of Michigan. Ann Arbor: University Microfilms Inc.

Gorman, G. C, and L. Atkins

1969. The Zoogeography of Lesser Antillean Anolis Lizards: An Analysis Based upon Chromosomes and

Lactic Dehydrogenases. Bulletin of the Museum of

Comparative Zoology, 138:53-80, 15 figures.

Kluge, A. G., and W. C. Kerfoot

1973. The Predictability and Regularity of Character

Divergence. The American Naturalist, 107:426-442,

10 figures.

Lewontin, R. C, and J. L. Hubby

1966. A Molecular Approach to the Study of Genie

Heterozygosity in Naural Populations, II: Amount

of Variation and Degree of Heterozygosity in Natural Populations of Drosophila pseudoobscura.

Genetics, 54:595-609.

Lister, B. C, and T. P. Webster

In prep. Genie Morphologic Variation in Island Populations of the Lizard Anolis sagrei.

Nei, M.

1972. Genetic Distance between Populations. The American Naturalist, 106:283-292.

Selander, R. K., M. H. Smith, S. Y. Yang, W. E. Johnson, and

J. B. Gentry

1971. Biochemical Polymorphism and Systematics in the

Genus Peromyscus, I: Variation in the Old-field

Mouse (Peromyscus polionotus). Studies in Genetics

(University of Texas Publication 7103), 6:49-90.

Shaw, C. R., and R. Prasad

1970. Starch Gel Electrophoresis of Enzymes: A Compilation of Recipes. Biochemical Genetics, 4:297-320.

Soule, M. E., S. Y. Yang, M. G. W. Weiler, and G. C. Gorman

1973. Island Lizards: The Genetic-Phenetic Variation

Correlation. Nature, 242:191-193, 2 figures.

Webster, T. P.

In prep. Electrophoretic Estimates of Genie Variation

in Populations of West Indian Anolis Lizards.

Webster, T. P., and J. M. Burns

1973. Dewlap Color Variation and Electrophoretically

Detected Sibling Species in a Haitian Lizard, Anolis

brevirostris. Evolution, 27:368-377.

Webster, T. P., R. K. Selander, and S. Y. Yang

1972. Genetic Variability and Similarity in the Anolis

Lizards of Bimini. Evolution, 26:523-535. 1 figure.

Williams, E. E.

1969. The Ecology of Colonization as Seen in the Zoogeography of Anoline Lizards on Small Islands.

Quarterly Rex'ieu' of Biology, 44:345-389, 17 figures.
Notes on the Chromosomes of

Anolis agassizi (Sauria: Iguanidae)

and Diploglossus millepunctatus

(Sauria: Anguidae)

Brad Stamm and George C. Gorman

ABSTRACT

The karyotypes of Anolis agassizi and Diploglossus millepunctatus are very similar. When compared to other species in their respective groups,

the karyotypes of the two Malpelo lizards are considered primitive.

During the past ten years description of reptilian karyotypes has moved from relative novelty to

virtually routine. Chromosome preparations of

Malpelo lizards were made directly from testes following methods described by Gorman, Atkins, and

Holzinger (1967).

Both Anolis agassizi and Diploglossus millepuncctatus have kartotypes consisting of six pairs of

metacentric macrochromosomes and 12 pairs of

microchromosomes, 2n = 36 (Figure \9a,b).

Among anguid lizards previously studied, this

karyotype was found in Diploglossus costatus (the

only other Diploglossus studied to date) but not

in the genera Gerrhonotus, Anguis, and Ophisaurus (Bury, Gorman, and Lynch, 1969). McDowell

and Bogert (1954) considered the diploglossines

the most primitive of the anguids.

Similarly, representatives of the most primitive

group of Anolis, the Lesser Antillean roquet group

(alpha group, latifrons series; Etheridge, 1960),

Brad Stamm, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138. George C. Gorman,

Department of Biology, University of California, Los Angeles,

California 90024.

have this same karyotype. Although many species

groups of Anolis show highly derived chromosomal morphology, a large section of the genus has

been conservative and retained the ancestral karyotype (for review, see Gorman, 1973).

Anolis agassizi is also a member of the latifrons

group. This species group has representatives on

the South American mainland as well as the Lesser

Antilles (the term roquet group follows Underwood, 1959). The Malpelo anole and the roquet

group all possess fracture planes in the caudal vertebrae, hence caudal autotomy is commonplace.

This character appears to have been secondarily

lost among living mainland forms. Thus the presumed ancestors of A. agassizi and the Lesser Antillean latifrons anoles are extinct, and the living

South American anoles are more derived (Etheridge, 1960).

Chromosomes tend to support this contention.

Many members of the roquet group have a karyotype virtually indistinguishable from A. agassizi.

Only a single mainland latifrons member has been

previously studied, A. jacare, and it has a derived

karyotype of 2n = 32, with a reduction in chromosome number (Williams et al., 1970). For comparison we add data on two other mainland latifrons

species, A. frenatus from Panama, and A. squamulatus from Rancho Grande, Venezuela. Webster,

Hall, and Williams, (1972) argued convincingly

that within Anolis, 12 metacentric macrochromosomes and 24 distinctly smaller microchromosomes

is the primitive condition. Anolis squamulatus (not

illustrated) has the typical primitive complement.

Anolis frenatus has 2n = 36, which is the typical

52
NUMBER 176

ft*

B S IIU « «"

IOJJ

•

FIGURE 19.—Lizard karyotypes: A, Anolis agassizi £, 2n = 36, 12 ca. metacentric macrochromosomes and 24 microchromosomes (pair 6 appears heteromorphic in this preparation, but we lack

sufficient numbers of metaphase plates, and have no data on females to confirm this); B,

Diploglossus millepunctatus $, 2n = 36, 12 ca. metacentric macrochromosomes and 24

microchromosomes; c, Anolis frenatus $, 2n = 36, size break between pairs 4 and 5, macrochromosomes do not break sharply from microchromosomes; pair 7 is intermediate in size.

saurian diploid number; but there is an enlarged

metacentric pair of "microchromosomes," and a

break in size between pairs four and five implying

a derived karyotype (Figure 19c).

The similarity between A. agassizi and the

roquet group of Lesser Antillean Anolis in both

osteology and karyotype substantiates the belief

that both retain primitive character states. We do

not argue for common origin between these two

groups. The similarity in karyotype between the

Diploglossus and Anolis of Malpelo is fortuitous.

This chromosomal formula is common to representatives of numerous lizard families, often

among the most primitive representatives, as determined osteologically (Gorman, 1973).

ACKNOWLEDGMENTS.—Travel to Malpelo by Gorman was supported by a grant from the National

Geographic Society. Laboratory work supported by

NSF Grant B 019801X to Dr. E. E. Williams.

Literature Cited

Bury, R. B., G. C. Gorman, and J. F. Lynch

1969. Karyotypic Data for Five Species of Anguid Lizards.

Experientia, 25:314-316.

Etheridge, R.

1960. The Relationships of the Anoles (Rcptilia: Sauria:
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Iguanidae): An Interpretation Based on Skeletal

Morphology. Unpublished Ph.D. dissertation, University of Michigan. Ann Arbor: University Microfilms Inc.

Gorman, G. C.

1973. The Chromosomes of the Reptilia: A Cytotaxonomic Interpretation. Chapter 12 in A. B. Chiarelli

and E. Capanna, editors, Cytotaxonomy and Vertebrate EvohUion. New York: Academic Press.

Gorman, G. C, L. Atkins, and T. Holzinger

1967. New Karyotypic Data on 15 Genera of Lizards in

the Family Iguanidae, with a Discussion of Taxonomic and Cytological Implications. Cytogenetics,

6:286-299.

McDowell, S. B., and C. M. Bogert

1954. The Systematic Position of Lanthanotus and the

Affinities of the Anguinomorphan Lizards. Bulletin

of the American Museum of Natural History, 105:

1-142.

Underwood, G.

1959. The Anoles of the Eastern Caribbean, Part III:

Revisionary Notes. Bulletin (Museum of Comparative Zoology), 121:191-226.

Webster, T. P., W. P. Hall, and E. E. Williams

1972. Fission in the Evolution of a Lizard Karyotype.

Science, 177:611-613.

Williams, E. E., O. A. Reig, P. Kiblisky, and C. Rivero-Blanco

1970. Anolis jacare Boulenger: A "Solitary" Anole from

the Andes of Venezuela. Brevoria (Museum of

Comparative Zoology), 353:1-15.
Sub tidal Communities of Malpelo Island

Charles Birkeland, David L. Meyer, James P. Stames,

and Caryl L. Buford

ABSTRACT

The subtidal communities of Malpelo are described

in terms of the pattern of spatial coverage of sessile

organisms. The Malpelo subtidal is characterized

by a prevalence of large barnacles (Balanus peninsularis) on steeply sloping rock walls and a highly

structured community of hermatypic corals on

gradually sloping substrata, both communities extending to a depth of about 30 meters. The rock

wall communities of Malpelo have significantly

more free space available for recruitment (38.6%)

than do comparable communities in continental

regions (7.3%). Size distribution suggests that recruitment is a greater risk, but conditions for adults

are better on Malpelo than in continental regions.

Observations suggesting the importance of fish

grazing are discussed. An expanded depth range

at Malpelo provides clearer zonation of the coral

community than along the Panamanian coast. Although coral growth is rich in terms of surface

coverage and size of colonies, a true reef is not

formed.

Introduction

The subtidal communities at Malpelo had never

been observed before our brief visit, and thus our

objective was to describe these communities in

quantative terms that would be useful for comparisons to other areas and for any return visits to

the island.

We have utilized the relative surface area coverage of macroinvertebrate species as the quantitative expression of community composition. Since

Charles Birkeland, David L. Meyer, James P. Stames, and

Caryl L. Buford, Smithsonian Tropical Research Institute,

P. O. Box 2072, Balboa, Canal Zone.

the Malpelo subtidal consists almost entirely of

vertical rock walls, surface area data were easily

acquired from underwater photographic transects,

using color and black-and-white film. Relative surface area coverage was then estimated by tabulation of randomly located points on the projected

transparencies and by planimetry (Tables 5,6,8).

Photographs used for the quantitative survey are

on file with C. Birkeland. We found that such a

method was most efficient for acquiring basic information from a large, unfamiliar, area within a

limited time. Photographic transects have the following additional advantages for primary survey of

an unexplored underwater area: First, size distribution of the predominant sessile organisms can

sometimes be used to deduce aspects of the age or

recent history of a community, such as the frequency or heterogeneity of disruptive factors.

Second, the extent of available unoccupied space

in a habitat is often an important limiting factor

for sessile organisms. Finally, the habitat or nursery of certain motile animals is often defined by

specific sessile organisms and, therefore, the sessile

organisms may provide the best definition of the

community as a whole (e.g., coral reef, oyster bed,

kelp bed, sea grass bed, etc.).

Dives were made at each of eight stations (Figure 20). At all stations except Station 6, the basaltic rock wall sloped steeply, sometimes vertically,

to a depth of about 37 meters on Malpelo itself or

about 49 meters on the offshore pinnacles (Stations

4 and 5). Sometimes a talus slope of large boulders

provided a more gentle slope in deeper water (below about 20 m; Stations 3, 7, and 8). Station 6 was

the only area found with a gradual slope and it

was the only location of a rich community of hermatypic corals.

55
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 5.—Percent coverage of primary substrata on rock walls of Malpelo Island,

1 to 3 March 1972

Depth range (meters)

0-3 3-6 6-27 27-37 37-50

No. of points (sample size x 50)

"OPEN SPACE"

450 1350 2800

bare rock or sediment ..

•crustose coralline algae

"OCCUPIED SPACE"

"Balanus peninsularis (live)

*Balanus peninsularis (dead, test)

Balanus peninsularis (total)

*Lobophora variegata

•"algal scuz"

"red algal crust"

branching coralline algae

(mostly Amphiroa spp.)

*Polyfibrospongia sp

•orange flat sponge

yellow flat sponge

red flat sponge

•white flat sponge

white lumpy sponge

blue sponge

greenish gray sponge

gray-brown sponge

hydroid

bryoz'oan

Lichenopora sp

Pacifigorgia spp

Lophogorgia alba

gorgonacean

Errinopora pourtalesii

Tubastraea aurea

Tubastraea aurea (dead)

Pocillopora robusta

Porites spp

Astrangia sp

Pavona varians

Ostrea irridescens

Ostrea irridescens (dead)

Ostrea fisheri

Didemnum candidum

Didemnum sp

Polyandrocarpa tincta

2.9

3.3

500 350

1.1

17.8

5.3

16.8

22.1

11.6

4.9

-

5.1

30.7

_

-

0.4

_

_

_

_

_

_

_

1.6

30.1

14.0

23.6

37.6

3.8

6.0

0.7

0.1

6.7

0.1

0.1

0.1

_

_

0.1

_

_

_

_

4.6

34.0

5.8

8.7

14.5

1.0

5.2

1.2

0.8

1.1

1.6

0.5

1.6

0.3

0.5

-

_

-

-

_

5.2

24.6

1.8

6.6

8.4

18.8

2.2

_

6.0

0.2

-

6.2

-

-

05

-

-

_

5.4

20.3

1.1

2.8

3.9

12.0

1.4

-

13.1

0.6

0.6

11.1

0.6

-

-

0.3

1.7

7.7

0.1

4.0

_

_

_

0.3

0.1

0.4

_

_

-

-

0.1

-

7.2

0.4

_

+

10.0

0.6

0.3

2.4

0.5

0.9

9.0

-

0.5

3.0

-

-

1.7

3.3

_

-

2.6

-

-

7.6

0.6

0.2

-

-

1.4

-

6.4

-

0.8

5.4

1.4

0.8

—

0.0

-

6.6

2.3

1.1

-

-

-

-

-

8.0

-

-

2.6

+

_

• Predominant categories. An organism that occupies over 10 percent of the primary substrata

is arbitrarily denned as predominant.

+ indicates that the organism was present in photographs of the area, but it did not appear

as data in surface-coverage counts from contact of random points.
NUMBER 176 57

TABLE 6.—Proportion of space open for invasion, as opposed to space occupied

by macroscopic sessile organisms other than crustose coralline algae, comparing

Malpelo Island and the continental coast of the eastern tropical Pacific (coverage

estimated by number of rectangular coordinate points—taken from a random

number table—falling on the image of each category; 50 points tallied from each

quadrat photograph; several points in the continental photographs fell into

shadow so were not determined)

Region

Oceanic (Malpelo)

Continental (Taboguilla Is.,

Panama and Playas del

Coco, Costa Rica)

No. of

quadrats

56

Open

space

1081

213

Occupied

space

1719

2705

Total no.

of points

2800

2918

Mean percent

of open space

38.6

x* = 800, 1 d.f.

Reject Ho: conclude significantly more "open space" exists in subtidal communities of Malpelo

than in comparable communities ©t mainland.

ACKNOWLEDGMENTS.—We were assisted during

diving work at Malpelo by D. B. Macurda, Jr., of

the University of Michigan and W. L. Smith of

the State University of New York at Stony Brook.

We thank D. Dexter of San Diego State University,

and J. B. Graham of the Smithsonian Tropical

Research Institute for constructive review of the

manuscript. The list of invertebrates was compiled

with the generous assistance of ten authorities.

Maureen Downey of the National Museum of Natural History, Smithsonian Institution, contributed

her list of asteroids to the Appendix of this paper.

John S. Garth of the Allan Hancock Foundation

identified the xanthid, dynomenid, and majid

crabs; and Janet Haig, also with the Allan Hancock Foundation, identified the porcellanid and

diogenid crabs. L. G. Abele of Florida State University completed the list of crabs and provided a

complete list for the decapods. Joseph Rosewater

of the National Museum of Natural History,

Smithsonian Institution, confirmed and corrected

the lists of mollusks. Alan J. Kohn of the University of Washington verified the species of Conus

and Dora P. Henry, also from the University of

Washington, identified the barnacles. Peter W.

Glynn of the Smithsonian Tropical Research Institute confirmed and corrected our identification of

scleractinian corals. J. W. Wells of Cornell UniTABLE 7.—Colony-polyp-number distributions in populations of Tubastraea

aurea at Malpelo compared with colony polyp number distributions in two

areas near the continental coast (counts were made from quadrat photographs)

Region

A

Total no.

colonies

examined

B

No. solitary

polyps

C

No. colonies

with 10 or

more polyps

Mean no.

polyps per

colony

(±SE)»

Oceanic (Malpelo)

Continental (Taboguilla Is.,

Panama and Playas del

Coco, Costa Rica)

372

408

88

135

102

92

11.6±0.8

10.0±0.8

• Solitary polyps not included in analysis (N = A —B).

X* = 6.78, 2 d.f. (categories compared were B, C, and A-B-C).

Reject Ho: conclude significantly fewer solitary polyps in proportion to number of larger

colonies (> 10 polyps) in a single sample of Malpelo populations when compared with a 3

year sample of several mainland populations.
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 8.—Zonation of the coral community of Malpelo Station 6: percent coverage by each species, bare rock, or dead coral substrata (data acquired by planimeter measurements on tracings of projected photographs)

92 & 10.7

Depth range (meters)

123 13.8 & 15.4 16.9 18.4 26.1 & 27.6

No. of planimeter samples

PERCENT OF SURFACE

Live hermatypic corals

Dead coral substrata

Rock or sediment substrata

PERCENT OF LIVE CORAL SURFACE

Pocillopora robusta

Porites spp

Pavona clivosa

Agariciella planulata

8 10 13

57.2

24.5

18.3

94.3

5.7

_

69.3

27.6

3.1

80.5

19.5

_

74.8

22.7

2.5

14.4

77.3

8.3

89.2

10.8

—

2.4

43.2

54.4

58.6

29.8

11.6

80.5

19.5

42.7

18.0

38.4

47.4

_

52.6

versity identified the hydrocoral and George Hechtel of the State University of New York at Stony

Brook identified the sponge.

500 Meters

o

4

FIGURE 20.—Subtidal collection stations at Malpelo Island.

Rock Wall Communities

The steep rock walls (Figure 21) were predominantly occupied by the large barnacle Balanus

peninsularis (Table 5). Sixty to 80 percent of the

Balanus space was occupied by empty tests (Figure

22) which remained on the walls after the death of

the barnacles because their predators were snails

(probably Thais planospira and possibly Purpura

pansa) rather than sea stars. These tests provide a

major source of shelter or habitat for many polychaetes, mollusks, crustaceans, and fishes (Entomacrodus chiostictus) and a substratum for hydroids, sponges, and tunicates. The prevalence of

barnacles is reflected in the white sand sediment

which encircles the island at the base of the rock

wall. This sand is composed almost entirely of fragments of Balanus tests.

A clear zonation was observed on the vertical

walls (Table 5). To a depth of 3 meters, the large

black sponge Polyfibrospongia sp. occupied over

30 percent of the space. The brown alga Lobophora variegata was also prevalent above 3 meters.

Both Polyfibrospongia and Lobophora decreased

rapidly in abundance between 3 and 6 meters

where Balanus tests occupied nearly 37 percent of

the space. From 6 to 27 meters the substratum was
NUMBER 176 59

FICURE 21.—Vertical rock wall community; Station 3, 15 m depth. Prevalent animals are Pacifigorgia media (left), Porites sp. (right), and Balanus peninsularis.
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

very clear of "scuz" (filamentous algae, hydroids,

etc.) and was covered mostly with crustose coralline algae and scattered patches of anthozoans, i.e.,

hermatypic corals (mostly Poriles spp.) an ahermatype (Tubastraea aurea), and gorgonaceans

(Pacifigorgia spp.). Below 27 meters, filamentous

algae, small flat encrusting sponges, oysters (Ostrea

irridescens), and the violet branching hydrocorals

(Errinopora pourtalesii) became prevalent. Although occupying only 7.6 percent of the substratum, the vivid violet hydrocorals appeared as a

"field of Errinopora" below 27 meters. Certain

motile animals were also found only at depths of

27 meters or below: the basket star, Astrodictyum

panamense, and the sea stars, Tamaria sp. and

Narcissia gracilis.

The most striking aspect of the occupation of

the subtidal substrata at Malpelo was the amount

of "open" or "free" space. Table 6 compares the

amount of open space in 59 permanent quadrat

photographs from small Pacific islands less than

14 km from the mainland in Costa Rica and Panama with the amount in 56 survey photographs

from Malpelo. Bare rock or space occupied by

crustose coralline algae is available as attachment

space for other sessile or sedentary organisms. Between the depths of 6 and 27 meters near the mainland, an average of 7.S percent of the space is

FICURE 22.—A typical example of the rock wall community; Station 7, 10 m depth (1 = Balanus

peninsularis, 2 = Porites sp., 3 = Tubastraea aurea, 4 = Pacifigorgia media).
NUMBER 176 61

available as open space in rock wall communities.

In comparable rock wall communities at Malpelo,

38.6 percent of the space is available.

Many of the prevalent sessile organisms were

common to continental and oceanic rock wall communities. The oceanic Malpelo populations for

both Tubastraea aurea and Lophogorgia alba appeared to be characterized by size distributions in

which larger individual colonies were more prevalent and less recruitment was observed in comparison with the continental populations. The number

of polyps in each Tubastraea aurea colony in Malpelo photographs were counted and the polypnumber distribution (presumed to be a form of

age distribution) was compared with polyp-number

distribution in Panama mainland quadrats (Table

7). The distributions differ significantly by chi

square when the proportions of recruitment for

solitary polyps, colonies of 2 to 9 polyps, and colonies of 10 or more polyps are compared. The mainland data are from several areas through all seasons over a 3-year period while the Malpelo data

are from a single survey. The significant difference

in age distribution could be the result of a particularly poor year, or time of year, in which the Malpelo survey was made, rather than to differences

between oceanic and continental populations. If

the solitary polyps are excluded from the analysis, the polyp-number distributions do not differ

significantly by Mest. Nevertheless, we predict that

more data from further surveys would confirm a

general trend towards proportionately less recruitment and longer adult life in oceanic populations.

A similar trend is evident for closely related

species living on the island and mainland, respectively. For instance, Balanus trigonus (adults ca.

0.8 cm basal diameter) is the prevalent species in

continental areas examined while Balanus peninsularis (adults ca. 6 cm basal diameter) is the prevalent species at Malpelo. The same generalization

could be drawn comparing colonial tunicates of

the genus Didemnum.

In summary, the oceanic rock wall communities

at Malpelo had over five times as much open space

as the continental rock wall communities. Since

space is a limiting resource for sessile benthic organisms on solid substrata, the abundant availability of open space at Malpelo implies that recruitment is a more difficult problem for these populations. Once established, however, an individual

lives under less crowded conditions. In other words,

an individual may be subjected to greater risk initially, but may attain greater fecundity if it survives until adulthood.

The reason for reduced recruitment success is

unknown but two possible causes can be hypothesized: isolation from other populations and heavy

grazing by fishes. Isolation may reduce recruitment

of shallow benthic organisms because most, perhaps 85 percent of the species (Thorson, 1961:456),

have pelagic larval stages for which the mean duration is about 2 weeks. Only about 30 percent have

planktonic stages longer than 5 weeks (Thorson,

1961:459). Malpelo is small, isolated, and surrounded by deep water. It seems likely that a large

portion of the larvae may be carried away before

settlement can occur. The water moving past Malpelo comes predominately from the northeast

where the nearest shallow water is around the Azuero Peninsula, 367 km away. The fauna of Cobb

Seamount, 430 km west of Washington State in the

northern Pacific, has large populations of brooding species usually found only in the intertidal or

shallow subtidal (Birkeland, 1971). Once a population of a brooding species is established on an

isolated pinnacle, it may have a generally better

chance of maintaining a population than species

with planktonic larvae. The emplacement of settling plates at Malpelo would provide particularly

interesting quantitative data for comparison with

data from plates set nearer the mainland.

The sessile ciliate protozoan Folliculina was

found on some of the Malpelo algae and it may

have arrived by rafting on drifting algae. However,

Scheltema (1973) has recently presented data showing that dispersal of this organism can also occur

by attachment to the shells of veliger larvae. Folliculina is often found on the larger species of

veligers which are "long-distance larvae from shoalwater bottom-dwelling organisms of the shelf"

(Scheltema 1973:123).

The effects of grazing fish in producing open

space is implied by some general observations concerning algae and is based on the assumption that

grazers of algae also remove recently settled benthic animals. Algae and juvenile sessile organisms

were very sparse on Malpelo (Tables 5, 6), yet the

exceptionally clear water should provide favorable

growth conditions for algae. As an example, one

small sprig of Sargassum was found at 46 m. The
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

I

FIGURE 23.—"Shingle" formation of Porites sp. on the rock wall community.
NUMBER 176 63

substrata in shallower water presumably had

enough available light and other conditions for

growth of Sargassum. The large schools of acanthurids may have kept the algae (and settling larvae)

under heavy grazing pressure. This hypothesis

could be simply tested by the emplacement of fish

exclusion cages for 4 to 12 weeks.

A species list from the collections and observations made during the survey is given in the Appendix. The most prevalent of all animals, Balanus

peninsularis (Table 5), is known from the Galapagos but not from the mainland. Since its test

provides a major source of shelter or habitat for

motile animals and a substratum for sessile organisms, the prevalence of B. peninsularis may be

the major factor contributing to the strong faunal

similarities between Malpelo and the Galapagos

Islands. Many familiar tropical eastern Pacific

mainland species were conspicuous by their absence: Muricea and antipatharians (the usually

present arborescent coelenterates), and the sea stars

Pharia and Phataria. Scarid fishes, hermit crabs,

and gastropods were remarkably scarce.

Coral Communities

On the steeply sloping rock walk typical of the

Malpelo subtidal, hermatypic corals occur only

as a veneer interspersed with other growth (Figure

22) or in some cases as shingled, overhanging

FIGURE 24.—Porites heads in the coral community; .Station 6, 18 m depth.
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 25.—Agariciella planulata; Station 6, SO m depth.

masses (Figure 23). Table 5 shows that the grand

average surface area coverage by hermatypic corals

on steep rock walls is only about 10 percent. In

contrast, a more gradual slope at Station 6 supports

an extensive coral community (Figure 24) extending to 30 m depth. Table 8 shows that hermatypic

corals dominate the surface area coverage of this

slope. At 15 meters depth, a continuous growth of

Porites sp. measured 6.7 meters across by 2 meters

high. At 28 meters, a continuous growth of Agariciella planulata covered an area of 3 by 1.8 meters.

Figure 25 illustrates that coral coverage at about

30 meters is extensive and not merely confined to

isolated colonies.

The rich development of hermatypic corals at

Station 6 suggests that conditions for coral growth

are highly favorable at Malpelo. The lack of coral

development elsewhere around Malpelo may be

principally due to two factors, both related to the

prevalence of vertical rock walls around the island.

Vertical surfaces are frequently shaded due to sun

angle and substratal irregularities; thus, much of

the light falling on them is reduced in intensity

depth

in feet

water level

100

FIGURE 26.—Profile diagram of the coral community at Station 6. As the diver faces shore,

transect A is to the left and transect B is to the right with a sand and cobble channel separating

them (r = Pocillopora robusta, s = Porites spp., c = Pavona clivosa, and v = Agariciella

planulata).
NUMBER 176 65

FH.URE 27.—Nicks from fish grazing on Porites sp.

and quality by scattering or by passing obliquely

through the water mass. By contrast, intensity on

horizontal surfaces at a similar depth would be

greater. Calcification in corals is enhanced by the

photosynthesis of their symbiotic zooxanthellae

(Goreau, 1959) and it is likely that the small

amount of coral noted along vertical walls at Malpelo is partly due to lower light intensities.

The second factor possibly contributing to the

lack of extensive coral development at Malpelo is

simply the lack of horizontal support on which

corals can build a reef. Figure 23 shows that corals

growing on the vertical faces tend to extend out

into the water mass to form a more horizontal surface which would better intercept light. This process can proceed only so far until erosion by burrowing animals causes collapse. Furthermore, subtidal talus piles and the eroding aspect of the cliffs

above suggest that rockfalls are a frequent occurrence at Malpelo. Some talus appeared to be of

recent origin, judging from the low numbers of

large encrusting organisms. These rockfalls prob-
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ably constitute a major disruptive phenomenon

which prevents dense coral buildup.

Even at Station 6 the prolific coral growth was

not developed above 10 meters depth, and it was

not clear that the living corals were growing on a

pre-existing coral framework. We thus hesitate to

refer to Station 6 as a coral reef. It is possible that

the development of a true reef is prevented at

Malpelo by other factors, one of which may be

temperature. Surface temperatures were usually

about 26.5°C, but an abrupt thermocline was

formed at various depths. Temperatures below the

thermocline were as low as 19.5°C and it has been

suggested that this is cool enough to limit coral

growth (Glynn and Stewart, 1973).

To the casual observer, the pattern of coral zonation is more clearly expressed at Malpelo than in

continental eastern Pacific coral communities because the greater depth range at Malpelo allows

the lower zones to spread out over a larger area

(Figure 26). For instance, Pocillopora spp., Pavona gigantea, and Pavona varians are all intermixed

between 6 and 7 meters at Uva Island, near the

coast of Panama (cf. Figure 8 in Glynn, Stewart,

and McCosker, 1972) while Pocillopora robusta,

Pavona clivosa, and Agariciella planulata are separated by depth at Malpelo (Figure 26).

As on coral reefs in continental regions of the

tropical eastern Pacific (Glynn, Stewart, and McCosker, 1972), signs of the effects of grazing by

fishes are prevalent. The edges of the ridged extensions of the large Pavona clivosa colonies were

lined with nicks scraped by browsing fishes. Figure 11 in Glynn, Stewart, and McCosker (1972)

is a typical illustration of this phenomenon. Occasionally, concentrations of nicks also occurred

on sheets of Porites (Figure 27). Only three individuals of the scarid fish, Scarus rubroviolaceus,

were seen at Station 6 and none were observed at

other stations. The tetraodontid, Arothron meleagris, and balistids were common at all stations and

probably these fishes, rather than scarids, were responsible for most of the damage to corals (Figure

27).

Literature Cited

Birkeland, C.

1971. Biological Observations on Cobb Seamount. Northwest Science, 45:193-199, 2 figures.

Glynn, P. W., and R. H. Stewart

1973. Distribution of Coral Reefs in the Pearl Islands

(Gulf of Panama) in Relation to Thermal Conditions. Limnology and Oceanography, 18:367-379, 8

figures.

Glynn, P. W., R. H. Stewart, and J. E. McCosker

1972. Pacific Coral Reefs of Panama: Structure, Distribution and Predators. Sonderdruck aus der Geologischen Rundschau, 61:483-519, 15 figures.

Goreau, T. F.

1959. The Physiology of Skeleton Formation in Corals,

I: A Method for Measuring the Rate of Calcium

Deposition by Corals under Different Conditions.

Biological Bulletin, 116:59-75, 5 figures.

Scheltema, R. S.

1973. On an Unusual Means by which the Sessile Marine

Ciliate Folliculina simplex Maintains Its Widespread Geographical Distribution. Netherlands

Journal of Sea Research, 7:122-125, 2 plates.

Thorson, G.

1961. Length of Pelagic Larval Life in Marine Bottom

Invertebrates as Related to Larval Transport by

Ocean Currents. In M. Sears, editor, Oceanography.

American Association for Advancement of Science

Publication, 67:455-474, 3 figures.

Appendix

A List of Marine Invertebrates Identified from

Malpelo Island

PROTOZOA

Folliculina sp.

PORIFERA

Polyfibrospongia sp.

COELENTERATA

Hydrozoa

Errinopora pourtalesii (Dall)

Anthozoa

Pacifigorgia media (Verrill)

Pacifigorgia sp.

Lophogorgia alba (Duchassaing and Michelotti)

Pocillopora robusta Verrill

Pavona clivosa, Verrill

Pavona varians Verrill

Agariciella planulata (Dana)

Porites spp.

Astrangia sp.

Tubastraea aurea (Quoy and Gaimard)

MOLLUSCA

Gastropoda

Diodora saturualis (Carpenter)

Seila assimilata (C. B. Adams)
NUMBER 176 67

Epitonium billeeanum (DuShane and Bratcher)

Hipponix grayanus Menke

Hipponix panamensis C. B. Adams

Crepidula aculeata (Gmelin)

Capulus sp.

Erato (?columbella Menke)

Cypraea albuginosa Gray

Cypraea isabellamexicana Stearns

Cypraea teres pellucens Melvill

Simnia aequalis (Sowerby)

Cymatium pileare (Linnaeus)

Colubraria ochsneri Hertlein & Allison

Muricanthns princeps (Broderip)

Coralliophila sp.

Coralliophila Qneriloides [Lamarck])

Quoyula madreporarum (Sowerby)

Thais planospira (Lamarck)

Acanthina brevidentata (Wood)

Purpura pansa Gould

Morula lugubris (C. B. Adams)

Anachis (?coronata [Sowerby])

Mitrella sp. 1

Mitrella sp. 2

Fasciolaria princeps Sowerby

Conus diadema Sowerby

Conus dalli Stearns

Tylodina fungina Gabb

Chromodoris sedna (Marcus)

Bivalvia

Area mutabilis (Sowerby)

Barbatia reeveana (Orbigny)

Septifer zeteki Hertlein and Strong

Lithophaga aristata (Dillwyn)

Lithophaga plumula (Hanley)

Isognomon recognitus (Mabille)

Ostrea fisheri Dall

Ostrea Qiridescens Hanley)

ARTHROPODA

Cirripedia

Balanus peninsularis Pilsbry

Tetraclita stalactifer (Lamarck)

Heteralepas quadrata (Aurivillius)

Lepas anserifer Linnaeus

Decapoda

PALAEMONIDAE

Brachycarpus biunguiculatus (Lucas, 1849), circumtropical

Harpiliopsis depressus (Stimpson, 1860), Indo-Pacific-western

America

Pseudocoutierea elegans Holthuis, 1951, western America

Veleronia laevifrons Holthuis, 1951, western America

Peridimenaeus hancochi Holthuis, 1951, western America

GNATHOPHYLLIDAE

Gnathophylloides mineri Schmitt, 1933, circumtropical (?)

ALPHEIDAE

Alpheus lottini Gue"rin, 1830, Indo-Pacific-western America

A. malleator Dana, 1852, western America, eastern & western

Atlantic

A. grahami Abele, new species, endemic to Malpelo (see

paper by Abele in this volume)

Synalpheus charon (Heller, 1861), Indo-Pacific-western

America

S. nobilii Coutiere, 1909, western America

S. digueli Coutiere, 1909, western America

S. biunguiculatus (Stimpson, 1860), Hawaii, Clipperton,

Malpelo

5. bannerorum Abele, new species, endemic to Malpelo (see

paper by Abele in this volume)

Pomagnathus corallinus Chace, 1937, western America

HlPPOLYTIDAE

Lysmata galapagensis Schmitt, 1924, Galapagos Island, Malpelo

L. trisetacea (Heller, 1861), Indo-Pacific-offshore islands

Palinuridae

Panulirus penicillatus (Oliver, 1791), Indo-Pacific-offshore islands

PORCELLANIDAE

Petrolisthes haigae Chace, 1962, western America

P. glasselli Haig, 1957, western America

Pachycheles biocellatus (Lockington, 1878), western America

Clastotoechus diffractus (Haig, 1957), western America

PAGURIDAE

Pagurus sp., western America

DlOCENIDAE

Aniculus elegans Stimpson, 1858, western America

Trizopagurus magnificus (Bouvier, 1898), western America

DYNOMENIDAE

Dynomene ursula Stimpson, 1859, western America

PORTUNIDAE

Euphylax dovii Stimpson, 1860, western America

XANTHIDAE

Qitadrella nitida Smith, 1869, western America

Trapezia digitalis Latreille, 1825, Indo-Pacific-western

America

T. ferruginea Latreille, 1825, Indo-Pacific-western America

Menippe obtusa Stimpson, 1859, western America

Domecia hispida Eydoux and Souleyet, 1842, Indo-Pacificwestern America

Medaeus spinulifer Rathbun, 1898, western America

Pilumnus pyginaeus Boone, 1927, western America

Globopilutnnus xantusii (Stimpson, 1859), western America

Ozius perlatus Stimpson, 1859, western America

Carpilodes cinctimanus (White, 1847), Indo-Pacific-western

America
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

GRAPSIDAE

Grapsus grapsus (Linnaeus, 1758), western America, eastern

and western Atlantic

GECARCINIDAE

Gecarcinus mapilensis Faxon, 1893, endemic to Malpelo

MAJIDAE

Teleophrys cristulipes Stimpson, 1959, western America

Mithrax pygmaeus Bell, 1835, western America

Microphrys platysoma (Stimpson, 1859), western America

Lissa tuberosa Rathbun, 1898, western America

ECHINODERMATA

ASTEROIDEA

Leiaster callipeplus Fisher

Narcissia gracilis malpeloensis, new subspecies

Tamaria stria Downey, new species, endemic to Malpelo (see

paper by Downey in this volume)

Mithrodia bradleyi Verrill

Nidorellia armata (Gray)

Asterope carinifera Miiller and Troschel

OPHIUROIDEA

Astrodictyum panamense (Verrill)

Ophiocoma aethiops Liitken

Ophiocoma alexandri Lyman

Ophiactis savignyi (Miiller and Troschel)

Ophiotheta mirablis Verrill

ECHINOIDEA

Diadetna mexicanum A. Agassiz

Eucidaris thouarsii (L. Agassiz and Desor)

Tripneustes depressus A. Agassiz

Echinometra sp.

CHORDATA

ASCIDIACEA

Polyandrocarpa (Eusynstyela) sp. aff. tincta (Van Name)

Didemnum (Didemnum) sp. aff. candidum Savigny

Didemnum (Didemnum) sp. aff. moseleyi (Herdman)
The Macruran Decapod Crustacea

of Malpelo Island

Lawrence G. Abele

ABSTRACT

Eighteen species of Macrura are reported from

Malpelo Island. Two new species of Alpheidae,

Alpnens grahami and Synalpheus bannerorum, are

described and S. nobilii and S. digueti are redescribed. Notes are given on Periclimenaeus hancocki, and Gnathophylloides mineri is recorded

from the eastern Pacific.

Ecologically about half of the 43 decapods known

from Malpelo are commensals with other invertebrates, mostly Pocillopora coral. Two species are

semiterrestrial or terrestrial and the remaining

species occur in a variety of subtidal habitats.

Zoogeographically decapod fauna of Malpelo (43

spp.) is most closely related to that of the western

American mainland with 62 percent or 27 species

occurring there. Nine species (20%) occur in the

Indo-Pacific region and seven of these reach the

western American mainland. There are three endemic species, two circumtropical species, and two

species known only from offshore islands.

The very limited data available at present suggest a species-area effect on the shallow water

decapods associated with oceanic islands.

Introduction

The first report of a decapod crustacean from

Malpelo Island was that of Faxon (1893) who described a land crab, Gecarcinus malpilensis, based

on material collected by the Albatross. This species

was considered to be a synonym of G. planatus by

Rathbun (1918), but Turkay (1970) recognized

Lawrence G. Abele, Smithsonian Tropical Research Institute,

P. O. Box 2072, Balboa, Canal Zone. Present address: Department of Biological Science, Florida State University, Tallahassee, Florida 32306.

G. malpilensis as a species endemic to Malpelo. In

1948, Garth reported on the Askoy collections,

which included three species taken at Malpelo: an

intertidal crab, Grapsus grapsus, the land crab,

Gecarcinus malpilensis, and a pelagic portunid,

Euphylax dovii, which was abundant in the waters

around the island. No additional material of decapod crustaceans appears to have been collected at

Malpelo until the Smithsonian Tropical Research

Institute and U. S. Navy expedition visited the

island in 1972.

Several members of the expedition independently made collections of decapods and sent them

to various individuals for study. I was unaware that

both Janet Haig and John S. Garth were studying

material from Malpelo when I began studying the

present collections. My study was then restricted

to the macrurans, the subject of the present report.

A complete list of the decapods reported from Malpelo is included in the Birkeland et al. appendix

(pp. 66-68, herein), and, therefore, it is unnecessary to repeat here.

The macrurans collected at Malpelo Island form

a small but interesting collection of 18 species. Two

species were previously undescribed, three species

are recorded for the first time since the original

description, the genus Gnathophylloides is recorded from the eastern Pacific for the first time

and many of the remaining species were previously

known from only a few records.

Collections of macrurans came primarily from

two localities on Malpelo Island referred to in the

descriptions as: "Malpelo coll. no. 3," which was

made on the southeastern section of the island

among coral in 10 m depth on 2 March 1972, and

"Malpelo coll. no. 4," which was made on the west

side of the island in 15 m depth on 3 March 1972.

69
70

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

"Malpelo coll." refers to general collecting around

the island from 29 February to 3 March 1972. The

abbreviation cl refers to carapace length excluding

the rostrum, tl to total length excluding the rostrum unless otherwise specified and USNM to the

National Museum of Natural History, Smithsonian

Institution, Washington, D.C. All of the material

will eventually be deposited in that institution.

ACKNOWLEDGMENTS.—I thank Jeffrey B. Graham

for entrusting me with the study of this interesting

collection. Charles Birkeland supplied me with a

collection of shrimps and the color slide of Panulirus penicillatus. The following three individuals

contributed so much to this study that completion

would not have been possible without their aid:

Dora M. Banner and Albert H. Banner of the University of Hawaii, examined alpheid material and

loaned material of Alpheus crockeri. Fenner A.

Chace, Jr., of the Smithsonian Institution, has

once again given his time and energy to the completion of a study. He loaned material of Periclimenaeus pearsei and compared material of Pseudocontierea, Veleronia, and Gnathophylloides with

type and other material in the Smithsonian Institution collections.

PALAEMONIDAE

Brachycarpus biunguiculatus (Lucas, 1849)

Palaemon biunguiculatus Lucas, 1849:45, pi. 4: fig. 4.

Brachycarpus biunguiculatus (Lucas).—Holthuis, 1952:3, pi.

1—Chace, 1962:606; 1966:625; 1972:18.

MATERIAL.—3 males, 6 non-ovigerous females, 5

ovigerous females, 1 juvenile, Malpelo coll. no. 3;

2 males, 2 non-ovigerous females, Malpelo coll.

no. 4.

MEASUREMENTS.—Males tl (including rostrum)

36.1-40.7 mm; non-ovigerous females tl 22.7-28.7

mm; ovigerous females tl 31.5-41.3 mm.

DISTRIBUTION.—This species is circumtropical in

distribution occurring on most oceanic islands.

REMARKS.—All of the Malpelo material agrees

with the description and illustrations of B. biunguiculatus, which has an extensive synonomy discussed by Holthuis (1952). A second recognized

species of Brachycarpus, B. holthuisi, has been

described from deeper waters (30-60 m) off Brazil

by Fausto Filho (1966). The present species occurs

in shallow waters in a wide variety of habitats: on

coral reefs, in tide pools, on grass flats, and among

subtidal rocks (Chace, 1972).

Periclimenaeus hancocki Holthuis, 1951

FIGURE 28B-D

Periclimenaeus hancocki Holthuis, 1951:97, pi. 29a-k.

MATERIAL.—1 male, 1 ovigerous female, Malpelo

coll.

MEASUREMENTS.—Male cl 6.0 mm, ovigerous female cl 8.4 mm.

DISTRIBUTION.—The unique holotype was collected from Pinas Bay on the Pacific coast of Panama. It is now reported from Malpelo Island.

REMARKS.—The male specimen agrees with the

description of the unique male holotype given by

Holthuis (1951). The lamella of the scaphocerite,

however, does not extend as far beyond the lateral

tooth as figured by Holthuis. The female specimen

differs in several respects. The rostrum is relatively

shorter than that of the male and it is armed with

four rather than five dorsal teeth. Holthuis had

only the male holotype of P. hancocki available

when he noted certain differences between this

FICURE 28.—Periclimenaeus pearsei Schmitt: A, chela of first

pereiopod. Periclimenaeus hancocki Holthuis: B, chela of first

pereiopod; c, telson and left uropod of female; D, anterior

portion of carapace of ovigerous female. [Scale = 1 mm for

A, B; 2 mm for c, D.]
NUMBER 176 71

species and the closely related species, P. pearsei

Schmitt, 1932 from the western Atlantic and these

are listed in his key (1951:78). The present female,

and to a lesser extent the male, bridge these differences with one important exception: the form of

the dactylus of the chela of the first pereiopods. In

P. pearsei (Figure 28A), of which two males and

two females (USNM 65113, 65114) were available

for comparison, the dactylus is high and convex

while in P. hancocki (Figure 28B) it is low and not

nearly as wide. This distinct and so far constant

difference seems to warrant continued specific recognition of these two closely related species.

The eggs of both species are small (about 0.45

X 0.3 mm) and numerous.

Harpiliopsis depressus (Stimpson, 1860)

Harpilius depressus Stimpson, 1860:38.

Periclimenes pusillus Rathbun, 1906:921, fig. 71, pi. 24: fig.

7 Bruce, 1970:306, fig. 1.

Harpiliopsis depressus (Stimpson) Holthuis, 1951:70, pi.

21, pi. 22a-f.

MATERIAL.—1 male, Malpelo coll. no. 3.

MEASUREMENTS.—Male cl 2.5 mm.

DISTRIBUTION.—Red Sea and Seychelles Islands,

islands of the central Pacific and the west coast of

America from the Gulf of California to Colombia.

REMARKS.—Bruce (1970) examined the status of

Periclimenes pusillus Rathbun from Hawaii and

concluded that it represents a juvenile of Harpiliopsis depressus. Two color forms of this species

occur on Pocillopora coral heads in Panama and

Australia. Their status is being investigated by A.

J. Bruce.

The translucent form of this species is one of the

most abundant Pocillopora commensals in Panama (Abele, 1972).

Pseudocoutierea elegans Holthuis, 1951

Pseudocoutierea elegans Holthuis, 1951:182, pi. 55a-r [pis.

55 and 57 reversed, legends are not].—Chace, 1972:45.

MATERIAL.—3 males, 6 non-ovigerous females, 7

ovigerous females, Malpelo coll.

MEASUREMENTS.—Males tl 7.1-7.3 mm, nonovigerous females tl 8.1-8.4 mm, ovigerous females

// 7.4-8.6 mm.

DISTRIBUTION.—California, Santa Catalina Island.

Mexico, Baja California off Ildefonoso Island,

Guerrero south of White Friars. Ecuador, Galapagos Islands, Bindloe Island. It is now reported from

Malpelo Island.

REMARKS.—The present material differs in several respects from the description of P. elegans

given by Holthuis (1951). All of the Malpelo specimens are smaller (tl 7.1-8.6 mm) than those listed

by Holthuis (tl 9-16 mm). The lamella of the scaphocerite is less produced in the Malpelo specimens than those figured by Holthuis. In the original description he notes that larger specimens

tend to have the dactylus of the larger second pereiopod robust with a large proximal tooth. Even the

smallest of the Malpelo specimens have this characteristic. The rostrum of the Malpelo material

seems to be shorter than Holthuis' material. The

Malpelo males lack a spine on the posterior margin of the third pleuron, which is present in Holthuis' figure (see Chace, 1972:45). Dr. Fenner A.

Chace, Jr., compared Malpelo material with paratypes of P. elegans and noted that the Malpelo

specimens seem to have the cornea set less obliquely on the eyestalk and the general body form

more depressed.

Given, however, the relatively few specimens

available from such widely scattered localities and

the variation known in other pontoniids, it seems

best to refer the present material to P. elegans

until more material is available.

Veleronia laevifrons Holthuis, 1951

Veleronia laevifrons Holthuis, 1951:199, pi. 63f-m.

MATERIAL.—3 males, 3 non-ovigerous females, 4

ovigerous females, Malpelo coll.

MEASUREMENTS.—Males tl 4.2-4.9 mm, nonovigerous females tl 6.4 mm, ovigerous females tl

5.4-6.6 mm.

DISTRIBUTION.—Ecuador, off Cape San Francisco

and Santa Elena Bay, Galapagos Islands (Hood

Island). The species is now reported from Malpelo

Island.

REMARKS.—Dr. Fenner A. Chace, Jr., compared

the present material with paratypes of V. laevifrons from Hood Island and concluded they were

conspecific.

Schmitt (in Holthuis, 1951:200) gives color notes

on specimens of this species that were found on a

gorgonian.
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

GNATHOPHYLLIDAE

Gnathophylloides mineri Schmitt, 1933

Gnathophylloides mineri Schmitt, 1933:7, fig. 3; 1935:167, fig.

31.—Lewis, 1956:288, figs. 1, 2.—Chace, 1972:52 Bruce,

1973:27.

MATERIAL.—1 ovigerous female.

MEASUREMENTS.—Ovigerous female cl 1.9 mm,

tl 5.5 mm.

DISTRIBUTION.—Southeastern Florida, Yucatan,

and Caribbean Sea (Chace, 1972). The species is

now reported from the eastern Pacific at Malpelo

Island.

REMARKS.—Dr. Fenner A. Chace, Jr., compared

the Malpelo specimen with Atlantic material of

G. mineri and was unable to find any obvious differences such as those noted by Bruce (1973) in

his comparison of G. mineri with G. robustus.

In the Atlantic the species occurs under stones

or in coral, but it is usually found on the spines of

sea urchins (Tripneustes and Lytechinus). A

species of Tripneustes, T. depressus, is known to

occur at Malpelo Island (Birkeland et al., Appendix, this volume).

ALPHEIDAE

Alpheus lottini Guerin-Meneville, 1830

Alpheus lottini Guerin-Meneville, 1830, pi. 3 Holthuis,

1958:22.

Alpheus ventrosus H. Milne Edwards, 1937:352.—Banner,

1958:164, fig. 4.

Alpheus laevis Randall, 1839:141.

Crangon ventrosa (H. Milne Edwards).—Banner, 1953:84, fig.

28.

Crangon latipes Banner, 1953:82, fig. 27.

MATERIAL.—2 ovigerous females, 1 juvenile,

Malpelo coll. no. 3.

MEASUREMENTS.—Ovigerous females cl 5.2, 8.2

mm, juvenile cl 2.9 mm.

DISTRIBUTION.—Red Sea and South Africa, across

the Central Pacific to western America where it

occurs from Baja California to at least Panama.

It is now reported from Malpelo.

REMARKS.—This species is a characteristic member of the Pocillopora coral community where it

usually occurs in male-female pairs. Patton (1973)

has dealt with aspects of the biology of this colorful species.

Alpheus malleator Dana, 1852

Alpheus malleator Dana, 1852:557; 1855, pi. 31: fig. 9a-h—

Crosnier and Forest, 1966:240, fig. 10.

MATERIAL.—11 males, 2 non-ovigerous females,

2 ovigerous females, 7 juveniles.

MEASUREMENTS.—Males cl 7.8-11.9 mm, nonovigerous females cl 5.2-5.4 mm, ovigerous females

cl 7.8-11.4 mm.

DISTRIBUTION.—Western Atlantic: southern Florida to Sao Paulo, Brazil. Eastern Atlantic: Senegal

to the Congo. Eastern Pacific: Gulf of California

to Ecuador, the Galapagos Islands, and now Malpelo Island.

REMARKS.—None.

Alpheus grahami new species

FIGURE 29A-C, F.-J

MATERIAL.—2 ovigerous females (smaller specimen is holotype), 2 juveniles, Malpelo coll. no. 3.

MEASUREMENTS.—Ovigerous female holotype cl

11.1 mm, paratypes, ovigerous female cl 12.3 mm,

juveniles cl 3.7, 4.2 mm.

TYPE-LOCALITY.—Southeastern side of Malpelo

Island, Colombia in 10 m depth among coral.

DISTRIBUTION.—Known only from the typelocality.

DESCRIPTION.—The rostrum is short and triangular with an acute apex; it extends to about the middle of the basal antennular segment. The orbital

teeth are triangular, subacute, and extend almost

to the apex of the rostrum.

The first three abdominal pleura are rounded;

the fourth and fifth are bluntly angled and the

sixth is subacute. The sixth segment is slightly

longer than the fifth and is about 0.7 of the length

of the telson. The telson has the length about 1.8

times the width. The distal lateral angles are

armed with a pair of spines, the inner one being

longer and stronger. Two pairs of dorsal spines are

present; the anterior pair is placed slightly less

than half the distance from the anterior margin

and the posterior pair is located about threefourths the distance from the anterior margin. The

uropods extend well beyond the distal margin of

the telson. The lateral uropod is armed with a

strong movable spine at the distal lateral angle

that is adjacent to a smaller immovable spine.
NUMBER 176 73

The stylocerite extends distinctly beyond the

distal margin of the basal antennular segment. The

visible portion of the basal segment is about threefourths the length of the second segment. The

third segment is about three-fourths the length of

the second segment.

The basicerite is unarmed dorsally, the angle

being blunt. The lateral spine is short and acute,

extending to about half the length of the stylocerite. The scaphocerite is widest proximally and

does not extend beyond the antennular peduncle.

The third maxilliped extends beyond the carpocerite by the distal third of the ultimate segment.

The distal portion of the ultimate segment is

armed with strong setae but no spines.

The first pereiopods are strong and unequal.

The ischium of the major cheliped is armed with

about four spinules on the inferior margin and a

single spine on a lobe on the distal superior margin. There are about 10 movable spinules on the

FIGURE 29.—Alpheus grahami, new species: A, anterior portion of carapace; B, third pereiopod;

c, small chela of first pereiopod; E, telson; F, third through fifth abdominal pleura; c, anterior

portion of major chela of first pereiopod; H, dactylus of third pereiopod; i, second pereiopod;

j , major chela of first pereiopod. Alpheus crockeri (Armstrong): D, anterior portion of major

chela of first pereiopod. [Scale -• 2 mm for A, B, D, E, I; 4 mm for c, F-H, J.]
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

inferior margin of the merus; the distal margin is

unarmed. The major chela is about three times as

long as high. There are no superior notches but a

sulcus is present on the medial surface below the

superior margin. It extends from the base of the

movable finger to about the middle of the chela

where it bifurcates; the upper portion extends

only a short distance but the lower portion extends

downward almost to the proximal margin of the

chela. The lower margin of the chela is compressed

proximal to the base of the immovable finger resulting in the formation of a weak inferior notch

adjacent to two weak tubercles. The distal margin

of the chela is armed with a pair of large, acute

teeth, one on either side of the movable finger. Below the lateral (outside) tooth, the margin of the

chela leading to the immovable finger is high and

crenulate. The movable finger is high, slightly compressed, and blunt, although in the single chela

available the apex may be damaged. The ischium

of the minor chela is armed with about six spinules

on the inferior margin and a spinule on a lobe on

the distal superior margin. The merus is armed

with about 11 spinules on the inferior margin; the

distal margin is unarmed. The chela is about 1.7

times as long as high. The movable finger is almost 0.6 the length of the chela and is somewhat

flattened dorsally. The distal margin of the palm

is armed with two strong teeth, one on either side

of the movable finger. The tips of both fingers are

acute; the hand is not balaeniceps in form. The

second pereiopods are subequal and extend beyond

the carpocerite by the distal three segments of the

carpus. The ischium is slightly longer than the

merus and both are shorter than the carpus. The

carpus is subdivided into five segments; the first

is the longest being slightly longer than the second and the fifth, which are subequal in length.

The third and fourth segments are subequal in

length and are slightly less than half the length of

the first. The movable finger is slightly more than

half the length of the chela. The fingers are unarmed. The third through the fifth pereiopods are

similar in form. Three of the four available specimens have a small movable spinule at the base of

the ischium of pereiopods three and four. This

spinule is relatively large on the two small specimens (cl 3.7, 4.2 mm), but it is small and difficult

to see on one of the larger females (cl 12.3 mm)

and it is absent on the other female (cl 11.1 mm).

The third pereiopod has the ischium less than

half the length of the merus. The merus is slightly

less than five times as long as wide; it is about 1.5

times the length of the carpus, about 1.3 times the

length of the propodus and about 5 times the

length of the dactylus. The propodus is armed with

about nine movable spines on the inferior margin.

The dactylus has a minute accessory tooth, surrounded by setae, on the superior margin. The

eggs are small and numerous.

REMARKS.—This species seems to be most closely

related to Alphens crockeri (Armstrong, 1941)

(= Crangon tuthilli Banner, 1953) known from

the Central Pacific Ocean, Thailand (Banner,

1953; Banner and Banner, 1966) and the eastern

Atlantic (Crosnier and Forest, 1966). Three specimens, through the courtesy of D. M. and A. H.

Banner, of A. crockeri were available for comparison: 1 male cl 8.0 mm, 2 non-ovigerous females cl

5.0, 6.9 mm from Oahu, Hawaii. The two species

are very similar, and were it not for the extreme

differences in the form of the major chela, other

differences would have been attributed to variation. In A. crockeri (Figure 29u), the movable finger is greatly compressed, with the superior margin

being defined by a knife-like ridge; it is twisted

in a horizontal plane and the apex is bulbous. The

immovable finger has a large, high proximal tooth

present and is twisted in a horizontal plane. In A.

grahami (Figure 29c, F), the movable finger is

robust and only slightly compressed; it is not

twisted and the width is the same throughout. The

immovable finger lacks a proximal tooth and is

not twisted.

Other differences between the two species are

slight but are constant in the material examined.

In A. crockeri the stylocerite is narrow and extends

almost to the distal margin of the basal antennular segment, while in A. grahami the stylocerite is

more robust and extends distinctly beyond the

basal segment. The antennular peduncle of A.

crockeri is slender with the second segment having

the length about twice the width, while in A. grahami the length is distinctly less than twice the

width. The scaphocerite of A. crockeri is slender

and extends to or beyond the distal margin of the

antennular peduncle, while in A. grahami it is

shorter than the antennular peduncle. The pereiopods of A. crockeri are slender with the length of

the merus being about six times the width while in
NUMBER 176 75

A. grahami the length is about five times the width.

From illustrations in the literature (Banner and

Banner, 1966; Crosnier and Forest, 1966) it

would appear that the rostrum of A. crockeri is

longer and more acute than that of A. grahami,

but the specimens of A. crockeri from Oahu,

Hawaii have a rostrum similar to that of A. grahami.

ETYMOLOGY.—The species is named for Jeffrey

B. Graham, who collected the specimens and entrusted them to me for study.

Synalpheus biunguiculatus (Stimpson, 1860)

Alpheus biunguiculatus Stimpson, 1860:31.

Synalpheus biunguiculatus (Stimpson) Banner, 1953:32,

fig. 9.

MATERIAL.—1 ovigerous female, 2 specimens,

Malpelo coll. no. 3; 1 specimen, Malpelo coll. no.

4; 1 specimen, Malpelo coll.

MEASUREMENTS.—Ovigerous female cl 4.2 mm;

other specimens cl 2.7-5.7 mm.

DISTRIBUTION.—Hawaiian Islands (Banner, 1953),

Clipperton Island (Chace, 1962), and now Malpelo

Island.

REMARKS.—The Malpelo material all lack the

major chela making the identification somewhat

uncertain. They agree in most respects with the

description and illustrations of Banner (1953).

The carpocerite in the Malpelo material is somewhat longer than the spine of the scaphocerite, but

the species is variable in this and other characters

(Chace, 1962).

Synalpheus nobilii Coutiere, 1909

FIGURE 30

Synalpheus nobilii Coutiere, 1909:40, fig. 22.—Schmitt, 1924:

162, fig. 39.—Sivertsen, 1934:22 Schmitt, 1939:12, 24.—

Chace, 1962:613.

MATERIAL.—23 specimens (6 ovigerous), Malpelo

coll. no. 3; 4 specimens, Malpelo coll. no. 4; 10

specimens, Malpelo coll.

MEASUREMENTS.—Ovigerous females cl 4.1-4.9

mm, other specimens cl 1.8-6.2 mm.

DISTRIBUTION.—St. Helena, Ecuador (Coutiere,

1909), Galapagos Islands (Schmitt, 1924), Clipperton Island (Chace, 1962) and now Malpelo Island.

Specimens were also examined from Punta Paitilla,

Panama.

DESCRIPTION.—The rostrum is narrow and acute,

not extending to the distal margin of the basal antennular segment. It is on a higher level and extends beyond the orbital teeth, which are small

acute extensions on the orbital hoods.

The third abdominal pleuron is obtusely

rounded, the fourth and fifth are bluntly angled

and the sixth is acute. The fifth segment is slightly

shorter than the sixth, which is slightly more than

half the length of the telson. The telson is slightly

longer than its anterior width. It narrows distally

and has a distinct constriction proximally. The

telson is armed with two pairs of spines at the distal lateral angles; the lateral spine is about onethird the length of the medial. There are two pairs

of dorsal spines present; the first pair is located

slightly anterior of the middle and the second is

located about four-fifths of the distance from the

anterior margin. The uropods extend well beyond

the distal margin of the telson. The lateral branch

is armed with a strong movable spine situated between two immovable spines at the level of the

diaeresis.

The stylocerite is long and slender extending to

about the middle of the second antennular segment. The visible portion of the basal segment is

slightly longer than the second segment, which is

distinctly longer than the third segment.

The basicerite is armed with strong dorsal and

lateral spines; the dorsal spine is about half the

length of the lateral. The lateral tooth of the scaphocerite extends beyond both the antennular peduncle and the carpocerite.

The third maxilliped has the apex armed with

nine strong, dark-colored spines arranged in a circle. The ultimate segment extends slightly beyond

the scaphocerite.

The first pereiopods are strong and unequal.

The superior distal margin of the merus is armed

with a tooth. The major chela has the length

slightly less than three times the height. The distal

margin of the palm is unarmed. A tubercle is present on the medial margin of the palm just below

the distal margin. The immovable finger is armed

with two teeth proximally. The movable finger is

slightly less than one-third the length of the chela.

The superior distal margin of the merus of the

smaller chela is armed with a tooth. The smaller
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

chela has the length slightly more than three times

the height. The distal margin of the palm is unarmed. The fingers are unarmed; the movable one

is about one-third the length of the chela. The second pereiopods are equal and extend beyond the

scaphocerite by the distal segment of the carpus.

The ischium is about three-fourths the length of

the merus, which is slightly shorter than the carpus. The carpus is subdivided into five segments;

the first is slightly less than five times the length

of the second, third, and fourth, which are subequal, and about twice the length of the fifth. The

movable finger is about half the length of the chela. The third through the fifth pereiopods are

similar. The merus of the third is about three

times as long as wide and is unarmed. It is about

twice the length of the carpus and about 1.3 times

the length of the propodus. The propodus is armed

with six movable spines, the sixth being paired,

on the inferior surface. The carpus has a distal

extension on the surperior margin and a pair of

spines on the distal inferior margin. The dactylus

is bifid with a small ventral protuberance; the ventral tooth is wider than the dorsal and slightly

shorter. The eggs are small and numerous.

REMARKS.—Specimens of this species have been

collected from the rocky intertidal zone in the Bay

of Panama. The material from Malpelo Island

came from among coral and coral rubble in about

10 m depth.

FIGURE SO.—Synalpheus nobilii Coutiere: A, anterior portion of carapace; B, second pereiopod;

c, third pereiopod; D, small chela of first pereiopod; t, anterior portion of major chela of first

pereiopod; F, distal portion of dactylus of third pereiopod, c, major chela of first pereiopod;

H, telson and left uropod. [Scale = 4 mm for E, C; 2 mm for A-D, H; 1 mm for F.]
NUMBER 176 77

Synalpheus digueti Coutiere, 1909

FIGURE 31

Synalpheus digueti Coutiere, 1909:48, fig. 28 Chace, 1937:

123.

MATERIAL.—4 ovigerous females, 36 specimens,

Malpelo coll. no. 3; 3 specimens, Malpelo coll. no.

4; 16 ovigerous females, 40 specimens, Malpelo

coll.

MEASUREMENTS.—Ovigerous females cl 5.2-8.4

mm, other specimens cl 3.0-7.6 mm.

DISTRIBUTION.—Baja California (Coutiere, 1909;

Chace, 1937). The species is now reported from

Malpelo Island. Material from Taboguilla Island

and the Pearl Islands, Panama, was also examined

during the present study.,

DESCRIPTION.—The rostrum is narrow, subacute,

and short; extending to about one-fourth of the

length of the basal antennular segment. The orbital teeth arise from the anterior margin of the carapace; they are triangular, subacute, almost twice

as broad as the rostrum, and are subequal in

length to it.

The third through the fifth abdominal pleura

are bluntly angled; the sixth is acute. The fifth

segment is almost equal in length to the sixth

which is slightly more than half the length of the

telson. The telson is longer than wide with two

pairs of spines on the distal lateral angles; the

FIGURE 31.—Synalpheus digueti Coutiere: A, anterior portion of carapace; B, anterior portion of

carapace of aberrant specimen; c, second pereiopod; D, minor chela of first pereiopod; E, third

pereiopod; F, telson and left uropod; c, anterior portion of dactylus of third pereiopod; H, major

chela of first pereiopod; i, anterior portion of major chela of first pereiopod. [Scale = 2 mm

for A-F; 1 mm for c; 4 mm for H, I.]
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lateral spine is slightly more than half the length

of the medial. The distal margin of the telson is

convex and extends to about two-thirds the length

of the medial spine. There are two pairs of dorsal

spines; the first pair is located at about the middle

of the telson and the posterior pair is located

slightly more than two-thirds of the distance from

the anterior margin. The uropods extend beyond

the distal margin of the telson. The lateral branch

is armed with a strong movable spine situated between two immovable spines at the level of the

diaeresis.

The stylocerite is long and acute, extending

slightly beyond the middle of the second antennular segment. The first antennular segment is about

1.4 times the length of the second and about twice

the length of the third.

The basicerite is armed with dorsal and lateral

spines; the dorsal spine extends to about one-fifth

of the length of the lateral. The lateral tooth of

the scaphocerite is long and acute and extends

beyond the antennular peduncle. The carpocerite

is long and broad, extending well beyond the lateral

tooth of the scaphocerite.

The first pereiopods are robust and unequal.

The merus of the major cheliped is unarmed. The

chela is about 2.6 times as long as high. The distal

margin of the palm is armed with a downwardly

directed spine. There are two tubercles on the

lateral margin of the palm at the insertion of the

movable finger. The distal portion of the immovable finger is compressed. The movable finger is

slightly more than one-fourth the length of the

chela. The merus of the smaller cheliped is unarmed, as is the chela. The length of the chela

is about 2.5 times the height. The movable finger is

about half the length of the chela. The second

pereiopods are subequal. The ischium is unarmed

and is about three-fourths the length of the merus.

The merus is unarmed and is about four-fifths the

length of the carpus. The carpus is subdivided into

five segments; the first is over five times the length

of the second, third, and fourth which are subequal

and which are half the length of the fifth. The

movable finger is slightly more than half the

length of the chela. The third through the fifth

pereiopods are similar. The ischium and merus

are unarmed. The ischium is slightly less than onethird the length of the merus. The carpus has an

extension on the distal superior margin and a pair

of spines on the distal inferior margin. The carpus

is about half the length of the merus. The propodus is about four-fifths the length of the merus

and is armed with seven movable spinules on the

inferior margin; the last spinule is paired. The

dactylus is bifid; the superior tooth is slightly

longer than the inferior tooth and they are subequal in width at the base.

The eggs are small and numerous.

In fresh material the body is translucent, the

appendages are blue, and the tips of the fingers of

the first pereiopods are red.

REMARKS.—The rostrum was somewhat variable

in form often being wider than figured here. One

specimen (Figure 31B) has a highly aberrant rostrum but was normal in other features. Three of

the 99 specimens examined lack a tooth on the

distal margin of the major chela. In other features

the species was not variable.

In the Gulf of Panama Synalpheus digueti

occurs as male-female pairs (the female is usually

ovigerous) in Pocillopora coral heads.

Synalpheus charon (Heller, 1861)

Alpheus charon Heller, 1861:27.

Synalpheus helleri de Man, 1911:246, pi. 8: fig. 37.

Synalpheus charon (Heller).—Banner, 1953:37, fig. 11.—

Banner and Banner, 1964:88.

Synalpheus charon obscurus Banner, 1956:329, fig. 5.

Synalpheus charon charon (Heller) Banner, 1956:331.

MATERIAL.—3 ovigerous females, 11 specimens,

Malpelo coll. no. 3.

MEASUREMENTS.—Ovigerous females cl 4.6-5.3

mm, other specimens cl 2.6-4.9 mm.

DISTRIBUTION.—Red Sea across the Pacific to

Baja California (Banner and Banner, 1966; Chace,

1937). During the present study, material was also

examined from the Pearl Islands and Taboguilla

Island in the Gulf of Panama.

REMARKS.—This species is a characteristic member of the Pocillopora coral community, occurring

deep inside the coral at the base of the branches

(Banner and Banner, 1964). It is common in this

habitat in the Gulf of Panama where it usually

occurs in male-ovigerous female pairs. The body

and appendages of fresh material is an even red

color.
NUMBER 176 79

Synalpheus bannerorum, new species

FIGURE 32

MATERIAL.—5 ovigerous females (holotype tl 11.1

mm, cl 4.4 mm), 23 specimens, Malpelo coll. no. 3

(paratypes); 2 specimens, Malpelo coll. no. 4.

MEASUREMENTS.—Ovigerous females cl 4.0-4.7

mm, tl 9.9-11.1 mm, other specimens cl 3.3-5.0 mm,

// 8.5-11.4 mm.

TYPE-LOCALITY.—Southeastern side of Malpelo

Island, Colombia, in 10 meters depth among coral.

DISTRIBUTION.—Known only from the typelocality.

DESCRIPTION.—The rostrum is narrow and acute;

it does not extend to the distal margin of the basal

antennular segment. It is on a higher level than

the orbital teeth, which are small, acute, directed

slightly medially, and do not extend to the apex

of the rostrum.

The first five abdominal pleura are rounded; the

sixth is subacute. The fifth segment is subequal in

length to the sixth, which is three-fourths the

length of the telson. The anterior width of the telson is greater than the length; it narrows distinctly

in the posterior half. The posterior margin is less

than one-half the width of the anterior margin.

The distal lateral angles of the posterior margin

are acute and armed with a pair of spines; the

lateral pair being shorter and more robust than

the medial. Both extend well beyond the lateral

angles of the telson. The dorsal surface of the telson is armed with a pair of strong spines; the first

pair is located about two-thirds of the distance

from the posterior margin and the second pair is

located about one-third of the distance from the

posterior margin. Both pairs are set in well from

the lateral margins. The lateral angle of the outer

uropod is armed with a long movable spine set

FIGURE 32.—Synalpheus bannerorum, new species: \, anterior portion of carapace; B, major chela

of first pereiopod; c, anterior portion of major chela; D, second pereiopod; E, anterior portion of

dactylus of third pereiopod; F, third pereiopod; c., telson and left uropod; H, minor first pereiopod. [Scale = 2 mm for A-D, F, H; I mm for E, G.]
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

between a pair of smaller acute immovable spines

at the level of the diaeresis. The medial uropod is

longer than the lateral, which is subequal in length

to the telson.

The stylocerite is acute and extends to about the

middle of the second antennular segment. The

basal antennular segment is longer than the second

and third, which are subequal in length.

The basicerite is armed with strong, acute dorsal

and lateral spines; the dorsal spine extending to

slightly over one-third of the length of the lateral.

The scaphocerite is reduced in size but the lateral

spine is strong, extending beyond both the antennular peduncle and the carpocerite. The carpocerite extends slightly beyond the antennular peducle.

The third maxilliped has the tip of the apex

armed with six dark-colored spines. It extends beyond the carpocerite by the distal third of the

ultimate segment.

The first pereiopods are unequal and robust.

The merus of the major cheliped is armed with a

tooth on the distal superior margin. The carpus

is armed with two acute extensions. The major

chela is about 2.3 times as long as high. The distal

margin of the palm is armed with a small downwardly directed spine. There is a large tubercle at

the base of the immovable finger which is followed

by a deep depression and then another tubercle.

The other side of the finger is also depressed giving

a pinched appearance to the base of the finger.

The immovable finger is armed with two strong,

acute teeth in the proximal portion. The movable

finger is more than one-third the length of the

chela and has two distinct tubercles on the proximal dorsal surface. There is a series of about five

unequal tubercles on the medial surface of the

palm between the acute distal spine and the base

of the immovable finger. The merus of the minor

cheliped is armed with an acute spine on the superior distal margin. The carpus is cup-shaped. The

chela is unarmed; the movable finger is slightly

less than half the length of the chela. The second

pereiopods are subequal. The fingers are unarmed

and the movable finger is slightly more than half

the length of the chela. The carpus is subdivided

into five segments; the first is about six times the

length of the second, which is subequal in length

to the third and fourth, which are about half the

length of the fifth. The ischium is about half the

length of the carpus and about three-fifths the

length of the merus. The third through the fifth

pereiopods are similar and robust. The ischium is

unarmed. The merus of the third is armed with

four to six movable spines. The length is slightly

more than three times the width. The carpus is

armed with two distal spines. The propodus is

armed with about nine movable spinules on the

inferior margin. The dactylus has the superior

tooth longer and slightly more than half the width

of the inferior. The merus of the fourth pereiopod

is armed with three or four movable spines. The

merus of the fifth pereiopod is unarmed. The eggs

are small and numerous.

REMARKS.—This species differs from all known

eastern Pacific species in the form of the meri of

the third and fourth pereiopods and in the form

of the major chela. It is the only western American

species in which the meri of the third and fourth

pereiopods are armed and the constriction of the

immovable finger of the major chela appears also

to be unique. There is one east American species

of Synalpheus (S. dominicensis Armstrong, 1949)

from the Dominican Republic, which has the meri

of pereiopods three and four armed. Synalpheus

bannerorum can be distinguished from S. dominicensis by the form of the anterior portion of the

carapace, by the armature of the telson, by the

form of the major chela and by the form of the

dactyi of pereiopods three through five. In S. dominicensis the medial margins of the orbits meet the

rostrum at a sharp angle while they blend into the

carapace before they meet the rostrum in S. bannerorum. The dorsal spines of the telson are both

located in the posterior half of the segment in S.

dominicensis, while in S. bannerorum only the posterior pair is located in the posterior half. The

major chela of S. dominicensis is not compressed

at the base of the immovable finger as it is in S.

bannerorum. The dactylus of the third pereiopod

in S. dominicensis has the lower tooth less divergent than that of S. bannerorum. Synalpheus bannerorum is also related to Indo-Pacific members

of the genus, but the form of the chela appears to

distinguish it from all other species of the genus.

ETYMOLOGY.—This species is named for Dora M.

Banner and Albert H. Banner who have contributed so much to the knowledge of the Alpheidae

and who have shown interest in the present study.
NUMBER 176 81

Pomagnathus corallinus Chace, 1937

Pomagnathus corrallinus Chace, 1937:124, fig. 5.—Schmitt,

1939:12.—Chace, 1962:612.

MATERIAL.—3 ovigerous females, Malpelo coll.

no. 3.

MEASUREMENTS.—Ovigerous females cl 4.6, 4.7,

5.4 mm.

DISTRIBUTION.—Arena Bank, Baja California

(Chace, 1937), Clipperton Island (Schmitt, 1939;

Chace, 1962). Specimens from Taboguilla Island

in the Gulf of Panama were also examined during

the present study.

REMARKS.—The present material agrees with the

description and illustrations of Chace (1937). The

species commonly occurs on Pocillopora coral in

the Gulf of Panama.

HIPPOLYTIDAE

Lysmata galapagensis Schmitt, 1924

Lysmata galapagensis Schmitt, 1924:165, fig. 41 Sivertsen,

1934:22 Hult, 1939:6.—Schmitt, 1939:12—Chace, 1962:

616.

MATERIAL.—2 males, Malpelo coll. no. 3.

MEASUREMENTS.—Males cl 2.6, 3.5 mm.

DISTRIBUTION.—This species was previously

known from two records off Santa Cruz Island

in the Galapagos Islands, and is now reported from

Malpelo Island.

REMARKS.—Both specimens agree in all respects

with the description and illustration of Schmitt

(1924). The rostral formula of both specimens

was one postorbital, five preorbital and one subapical ventral teeth. The occurrence of this species

with L. trisetacea offers additional support (Schmitt, 1939; Chace, 1962) for the specific distinctness of these two species.

Lysmata trisetacea (Heller, 1861)

Hippolyte trisetacea Heller, 1861:29.

Hippolysmata paucidens Rathbun, 1906:913, pi. 24: fig. 4.

Lysmata paucidens (Rathbun).—Schmitt, 1939:12.

Lysmata trisetacea (Heller) Holthuis, 1947:19, 65— Chace,

1962:614.

MATERIAL.—6 males, 1 immature male, 5 nonovigerous females, 9 ovigerous females, Malpelo

coll. no. 3.

MEASUREMENTS.—Immature male cl 1.8 mm,

males cl 2.9-3.8 mm, non-ovigerous females cl 2.9-

6.0 mm, ovigerous females cl 4.0-5.1 mm.

DISTRIBUTION.—The species has previously been

reported from the Red Sea, western Indian Ocean,

the Malay Archipelago, Micronesia, New Zealand,

Hawaiian Islands, Clipperton Island (Holthuis,

1958; Chace, 1962), and now Malpelo Island.

Neither this species nor L. galapagensis appear to

occur on the western American mainland. Lysmata

californica (Stimpson, 1866) occurs in California

and is common in the Gulf of Panama while L.

porteri (Rathbun, 1907) occurs in Chile. The record of Sivertsen (1934) for L. intermedia (Kingsley, 1878), a western Atlantic species, is based on

a single specimen from the Galapagos Islands,

which appears to be within the range of variation

of L. californica (see Sivertsen, 1934, pi. 2: figs.

9-15; and Limbaugh, Pederson, and Chace, 1961,

fig. 7).

REMARKS.—Chace (1962) discussed variation in

this species and noted three differences between

material from Clipperton Island and material from

the Hawaiian Islands. In some Hawaiian specimens

the posterior ventral rostral tooth is behind or

opposite the anterior dorsal tooth while in most

of the Clipperton specimens both ventral teeth,

when present, are well in advance of the anterior

dorsal tooth. In larger adults from Hawaii the

fused part of the upper antennular flagellum is

not much shorter than the free portion of the

shorter branch while in large Clipperton specimens the fused portion is distinctly less than half

the length of the free portion. The average number of carpal segments in Hawaiian material (23)

tended to be higher than that of Clipperton material (21).

These three characters were examined in the

Malpelo material with the following results. The

rostral formula was (number of specimens in parentheses): 2+2/2 (16), 2+3/2 (1), 2+2/3 (1),

2+2/1 (1), 1+2/2 (1), 1+2/1 (1). All specimens

with two ventral teeth had them both placed well

in advance of the anterior dorsal tooth; those with

one ventral tooth had it placed well in advance of

the anterior dorsal tooth. The Malpelo material

agreed also with the Clipperton material regarding

the relative lengths of the fused and free portions

of the upper antennular flagellum. The number

of carpal segments in Malpelo material varied
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

from 21-26 distributed as follows (number of specimens in parentheses): 24 (5), 22 (3), 21 (2), 23

(2), 25 (1), 26 (1). The average was slightly more

than 24 agreeing more with the Hawaiian material

than with the Clipperton material. These differences warrant further consideration when comparing material from throughout the range of L. trisetacea as suggested by Chace (1962) and Holthuis

(1947).

PALINURIDAE

Panulirus penicillatus (Olivier, 1791)

Astacus penicillatus Olivier, 1791:343.

Panulirus penicillatus (Olivier).—Holthuis, 1946:125.—Holthuis and Loesch, 1967:217. pi. 8.

MATERIAL.—None, the species was identified from

color slides.

MEASUREMENTS.—Cl 91-137 mm (Holthuis and

Loesch, 1967).

DISTRIBUTION.—This species occurs in the IndoWest Pacific region: Red Sea, Southeast Africa to

Korea, Formosa, and Polynesia. In the eastern Pacific it occurs at Revillagigedo Islands, Clipperton

Island, Cocos Island, Galapagos Island (Holthuis

and Loesch, 1967), and now is reported from Malpelo Island. It has not been found on the western

American mainland.

REMARKS.—Although no material of this species

was collected, comparison of color slides taken by

Dr. Charles Birkeland with the color plate of this

species given by Holthuis and Loesch (1967) permitted identification.

Ecology

The short time available to members of the expedition did not permit detailed collecting notes on

so many different groups of organisms (see Appendix in Birkeland et al., this volume). However,

some observations are possible based on published

information and my own experiences in Panama

(Abele, 1972).

Perhaps the most interesting observation is the

large percentage of decapods from Malpelo Island

which are commensals. Almost half of the 43

species occur in close association with some other

invertebrates. Most occur with Pocillopora coral,

including the following: Harpiliopsis depressus,

Alpheus lottini, Synalphens charon, S. digueti,

Pomagnathus corallinus, Petrolisthes haigae, P.

glasselli, Pachycheles biocellatns, Pagurus sp., Trapezia digitalis, T. ferruginea, Domecia hispida,

Medaeus spimilifer, Carpilodes cinclimanus, Teleophrys cristulipes, and Mithrax pygmaeus. Both

Veleronia laevifrons and Pseudocoutierea elegans

are associated with gorgonians. Gnathophylloides

mineri occurs on the spines of sea urchins and

Periclimenaeus hancocki probably occurs with

sponges.

Two species, Grapsus grapsus and Gecarcinus

malpilensis, occur intertidally and supratidally,

respectively.

The remaining species occur in a variety of subtidal habitats, mostly with rocks and coral rubble.

Distribution

The decapod fauna of Malpelo Island has affinities with both the western American mainland and

Indo-Pacific decapod faunas (p. 67). Twenty-seven

species, 63 percent of the Malpelo fauna, occur on

the western American mainland. Nine species, 21

percent of the fauna, occur in the Indo-Pacific region and seven of these reach the western American mainland. The other two, Lysmata trisetacea

and Panulirus penicillatus occur from the IndoPacific only to the offshore islands of the eastern

Pacific. Three species, Gecarcinus malpilensis, Alpheus grahami, new species, and Synalpheus bannerorum, new species, appear to be endemic to

Malpelo Island. Two species, Brachycarpus biunguiculatus and probably Gnathophylloides mineri

are circumtropical in distribution. Lysmata galapagensis is only known from the Galapagos Islands

and Malpelo Island, while Synalpheus biunguiculatus is known from Hawaii, Clipperton Island,

and Malpelo Island.

The decapod fauna of Malpelo Island then has

its greatest affinities (63%) with that of the western American mainland, next is the Indo-Pacific

region (21%), then endemic (7%), circumtropical

(4%), and island species (4%).

Comparison with Other Islands

A detailed comparison, at the species level, of

the decapod faunas of the offshore islands of the
NUMBER 176 83

fe 1.8.

I

8 1.7

I.OC PERIMETER OF ISLAND

FIGURE 33.—The relationship between island perimeter and

the number of decapod crustacean species present at (1),

Malpelo, (2) Clipperton, and (3) Cocos islands. Refer to text

for discussion.

eastern Pacific will have to await more data on

other islands.

Some comparisons can be made, however, between numbers of species of decapods associated

with oceanic islands (Figure 33). There are 25

species of decapods known from Saint Helena

(Chace, 1966; 1968), an isolated island in the

south Atlantic. Twenty percent, five species, are

endemic. There are 58 species of decapods known

from Clipperton Island (Chace, 1962; Garth, 1965)

in the eastern Pacific. None of these species is

endemic. I have found about 100 species

known from Cocos Island in the eastern Pacific of

which at least six (6%) are endemic. Malpelo Island has a decapod fauna of 43 species with three

(7%) of them being endemic.

Within a given region there generally exists an

orderly relation between numbers of species and

the size of the area sampled. This relationship has

been worked out for a number of terrestrial organisms occurring on islands (MacArthur and Wilson,

1967). In comparing shallow water organisms associated with islands, it seems more appropriate

to use island perimeter rather than island area. A

comparison between island perimeter and numbers

of decapod species of three eastern Pacific islands

(Saint Helena, having a different faunal source,

is not included), Malpelo, Clipperton, and Cocos,

is shown in Figure 33. The relationship is striking, although the slope (z=0.56) is higher than

that reported for other area-species curves (z =

0.20-0.35; MacArthur and Wilson, 1967). The

present curve may not be representative being

based on only three islands and is presented here

primarily to stimulate interest in examining this

relationship in other marine organisms.

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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1966. Brachycarpus holthuisi, Nova Especie de Crustaceo

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Galapagos, to the West Coast of Mexico, and in

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Garth, J. S.

1948. The Brachyura of the "Askoy" Expedition with

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1946. The Stenopodidae, Nephropsidae, Scyllaridae and

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the Snellius Expedition. Part XIV in Biological

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1947. The Decapoda of the "Siboga Expedition, IX: The

Hippolytidae and Rhynchocinetidae Collected by

the Siboga and Snellius Expeditions with Remarks

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1951. The Subfamilies Euryrhynchinae and Pontoninae.

Part I in A General Revision of the Palaemonidae

(Crustacea Decapoda Natantia) of the Americas.

Allan Hancock Foundation Occasional Papers, 11:

1-332, 63 plates.

1952. The Subfamily Palaemoninae. Part II in A General

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1837. Histoire Naturelle des Crustaces, Comprenant

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1918. The Grapsoid Crabs of America. Bulletin of the

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1924. The Macrura and Anomura Collected by the Williams Galapagos Expedition, 1923. Zoologica (New

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51 (5/6):333-354, 11 figures.
Asteroidea from Malpelo Island

with a Description of a New Species

of the Genus Tamaria

Maureen E. Downey

ABSTRACT

One new species, Tamaria stria, and a new subspecies, Narcissia gracilis malpeloensis, are described from shallow waters of Malpelo Island. A

collection totaling six species of starfish is described,

with notes on distribution.

Introduction

In the course of the faunal survey of the shallow

waters (0-50 m) of Malpelo conducted by C. Birkeland and his colleagues, only six species of starfish

were collected; however, all are species with interesting or unusual distribution patterns, and included in the collection are a new species of the

genus Tamaria and a new subspecies of Narcissis

gracilis. The abbreviation R refers to the major

radius from center of disc to tip of arm; and r refers to the minor radius, from center of disc to

interradial margin.

ACKNOWLEDGMENTS.—I wish to thank Charles

Birkeland and David Meyer of the Smithsonian

Tropical Research Institute for the opportunity to

examine this collection, and Dr. David L. Pawson,

Chairman, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, for reading the manuscript and

offering helpful comments and suggestions.

Maureen E. Downey, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

OREASTERIDAE

Nidorellia armata (Cray)

Pentaceros (Nidorellia) armata Gray, 1840:277; 1866:7, pi. 14:

fig. 1-3.

Oreaster armatus (Gray).—Mueller and Troschcl, 1842:52 —

Lutkcn, 1864:148—Bell, 1884:79.—von Martens, 1865:433.

Goniodiscus armatus (Gray).—Lutken, 1859:75.

Goniodiscus conifer Mobius, 1859:10, pi. 3: fig. 5-6.

Nidorellia armata (Gray).—Verrill, 1867:280—Perrier, 1876:

67—Viguier, 1878:193.—Leipoldt, 1895:634—H. L. Clark,

1910:332, pi. 4: fig. 2.—Doderlein, 1916:418; 1936:316, pi.

21: fig. l-6a.—Boone, 1926:3, pi. 1; 1933:73, fig. 2.—

Ziesenhenne, 1937:216.—H. L. Clark, 1940:333.—Steinbeck

and Ricketts, 1941:381, pi. 10: fig. 1, pi. 11: fig. 2—Ely,

1945:15.—Caso, 1943:9, 66, pi. 22: fig. 1-2, pi. 23: figs. 1-2;

1953:221; 1961:63, figs. 22-23.—H. L. Clark, 1958:95.—Caso,

1962:63, figs. 22-24.

Nirodella armata (Gray).—Stanek, 1955:48.

Nidorellia armata is represented in this collection by one specimen, quite large and heavy (R

10 cm, r 6.5 cm, dry weight 177 grams). The marginals, particularly the distal ones, are tremendously swollen and most bear 1 or 2 stout spines or

low tubercles. The much smaller principal plates

of the abactinal surface usually bear 1 or 2 low

tubercles. The papular areas of the abactinal surface are covered with pedicellariae of the split

granule type. This is a common shallow water

species of the eastern tropical Pacific, known from

lower California to Peru and the Hawaiian Islands.

OPHTOIASTERTOAE

Leiastef callipeplus Fisher

Leiaster callipeplus Fisher, 1906:1083, pi. 30: figs. 1, la, pi.

31: fig. 3.

86
NUMBER 176 87

Leiaster callipeplus is a Hawaiian species, reported by Fisher from Maui, Lanai, Kauai, and

Bird Islands, in 58-124 meters. One fine specimen,

collected at Malpelo in 49 meters at the base of a

rock wall (Station 5, Birkeland et al., Appendix,

this volume) measures R 20 cm, r 2.9 cm. Leiaster

callipeplus has also been collected near the central

American mainland in the Gulf of Chiriqui, Panama (Birkeland, pers. comra.).

Narcissia gracilis malpeloensis, new subspecies

Narcissia gracilis malpeloensis is represented in

this collection by six specimens. The species, described by A. H. Clark (1916:58) has previously

been collected from Baja California to the Galapagos Islands. The average size of these specimens

is R 12 cm, r 2.5 cm. The Malpelo specimens represent a new subspecies of N. gracilis, differing from

the type-specimen and Clark's description in the

following respects:

Narcissia gracilis gracilis: Pedicellariae abundant

on all surfaces, of 2 slender valves with expanded

toothed tips, lying in alveoli; granules hemispherical; subambulacral spines in 3 rows; madreporite

round; papulae single; and proximal marginals

longer than broad.

Narcissia gracilis malpeloensis: Few or no pedicellariae—when present, of 2 stout curved untoothed valves of uniform thickness, not in alveoli;

granules flat-topped, polygonal; subambulacral

spines in one row, plus other spines not in rows;

madreporite triangular; papulae usually double;

and proximal marginals broader than long.

Although the above differences are minor, and

abundant pedicellariae versus few or no pedicellariae might be accounted for by the difference in

size (holotype: R 54 mm, r 8.5 mm) between these

specimens and the type (starfishes frequently have

many pedicellariae when young and few or none

when fully grown), the very dissimilar character of

the pedicellariae, plus the other variations listed

above seem, in view of Malpelo's isolated position,

to warrant separation at the subspecific level.

Tamaria stria, new species

FIGURE 34

DESCRIPTION.—R 44 mm, r 8 mm. Disc small,

arms five, long, blunt, cylindrical. Carinal, adradial, superomarginal, and inferomarginal plates

diamond-shaped, slightly overlapping distal corner

of preceding plate, connected transversely to adjacent row of plates by small, irregular, secondary

plates. Six rows of papular areas, 6-10 pores per

area. Three rows of rounded actinal intermediate

plates, becoming two distally. Entire body, up to

adambulacral armature, covered with close uniform coating of granules, those in papular areas

slightly smaller than those on plates. Adambulacral furrow spines two, one small, acicular, the other

heavy, flared at tip, broad in plane of furrow. Subambulacral spines single, heavy, thick, longer than

broad. No pedicellariae noted. Madreporite small,

round, covered with fine radiating gyri. Anus

subcentral, inconspicuous. Ocular plates large,

rounded, raised, mostly bare, with few granules.

Color in life is reddish orange; dried, bright

orange with blue papular areas.

MATERIAL EXAMINED.—Holotype, USNM El 1838,

Station 4, Malpelo Island, 49 m, on rubble near

base of vertical rock wall. Paratypes, USNM

El 1839, 4 specimens, Stations 4 and 5, Malpelo

Island, 36-49 m.

ETYMOLOGY.—The species name is the latinized

acronym for the Smithsonian Tropical Research

Institute.

DISCUSSION.—Only one other species of Tamaria,

T. obstipa Ziesenhenne, has been described from

the eastern tropical Pacific, Tamaria obstipa is

known from the type-locality, Cocos Island, Costa

Rica, and from James Island, Galapagos Islands.

The present species agrees closely with T. obstipa,

but differs in having secondary connecting plates

not only intermarginally but between all abactinal plate rows; in having a carinal series, lacking

in T. obstipa; in the nature of the adambulacral

armature; and in the lack of pedicellariae. The

number of pores per papular area is smaller in T.

stria (6-10) but is within the range of T. obstipa

(6-24).

MITHRODIIDAE

Mithrodia bradleyi Verrill

Mithrodia bradleyi Verrill, 1867:288— Perrier, 1878:77.—

Sladen, 1889:539.—Fisher, 1906:1094, pi. 36: figs. 1, 2, pi.

37: figs. 1-3; 1925:68; 1928:491.—H. L. Clark, 1910:327, pi.

6: fig. 1.—A. H. Clark, 1946:9.—Steinbeck and Ricketts,

1941:380, pi. 23: fig. 3.—Ely, 1945:27, pi. 8A, B.—Caso, 1944:
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 34.—Tamaria stria, new species, holotype, USNM El 1838.
NUMBER 176 89

253; 1953:214; 1961:92, figs. 34-36; 1962:92, figs. 34-36;

1963:300.—Engel, John, and Cherbonnier, 1949:1.

Mithrodia clavigera Perrier, 1875:360.—Ives, 1889:171.

Three specimens of Mithrodia bradleyi were collected at Malpelo at depths from 12 to 14 meters;

the known distribution for this species is from Baja

California to the Galapagos Islands. The Malpelo

specimens have many more pedicellariae than

specimens from Baja California in the collections

of the National Museum of Natural History,

Smithsonian Institution, but correspond well with

material from the Colombian coast.

PORANIIDAE

Asteropsis carinifera (Lamarck)

Asterias carinifera Lamarck, 1816:556.

Aster ope carinifera (Lamarck).—Mueller and Troschel, 1840:

104.—H. L. Clark, 1920:33, pi. v: fig. 2.

Asteropsis carinifera (Lamarck): Mueller and Troschel, 1840:

322; 1842:63.—A. M. Clark, 1967:37.—McKnight, 1968:713.

Gymnasterias inermis Gray, 1840:278.

Gymnasterias spinosa Gray, 1840:278.

Gymnasterias carinifera (Lamarck).—deLoriol, 1885:67, pi.

20: figs. 7-10.—Leipoldt, 1895:649, pi. 32: fig. 13.

One specimen of Asteropsis carinifera was collected at Malpelo on a vertical rock wall at 36

meters. It is distributed throughout the IndoPacific and the tropical eastern Pacific. This species

is apparently attacked by the shrimp Hymenocera

(Birkeland, pers. comra.), well known as a predator on Acanthaster plancii, the crown-of-thorns

starfish.

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1949. The Genus Mithrodia Gray, 1940. Zoologisch Verhandelungen (Leiden), 2 (1949): 1-39, 12 figures.
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Fisher, W. K.

1906. The Starfishes of the Hawaiian Islands. Bulletin of

the United States Fish Commission, 23 (3,1903):987-

1130, 49 plates.

1925. Sea Stars of the Tropical Central Pacific. Bulletin

of the Bernice P. Bishop Museum, 27:63-87, plates

5-8.

1928. Sea Stars from the Arcturus Oceanographic Expedition. Zoologica, 8:487:493.

Gray, J. E.

1840. A Synopsis of the Genera and Species of the Class

Hypostomata (Asterias, Linnaeus). Annals and

Magazine of Natural History, series 1, 6:175-184,

275-290.

1866. Synopsis of the Species of Starfish in the British

Museum. 17 pages, 16 plates. London: John Van

Voorst.

Ives, J. E.

1889. Catalogue of the Asteroidea and Ophiuroidea in

the Collection of the Academy of Natural Science

of Philadelphia.

Lamarck, J. B.

1816. Histoire naturelle des animaux san vertebres. Edition 1, volume 2, 568 pages. Paris.

Leipoldt, F.

1895. Asteroidea der "Vettor-Pisani" Expedition (1882-

1885). Mit Anhang: Die von F. Orsini im rothen

Meere gesammelten Asteroideen. Zeitschrift wissenschaften Zoologische, 59:545:654, plates 31, 32.

de Loriol, P.

1885. Catalogue Raisonne des Echinodermes recueillis

par M. V. de Robillard a Tile Maurice, II: Stellerides. Memoire Societe physique Histoire naturelle

Geneve, 29 (4): 1-84, plates 7-22.

Lutken, C.

1859. Bidrag til Kundskab om de ved Kysterne af Mellemog Syd-Amerika levende Arter af Sdstjerner.

Videnskabelige Meddelelser fra den naturhistoriske

Forening i Kjovenhavn, pages 25-96.

1864. Kritiske Bemaerkninger om forskjellige Sostjerner

(Asterider), med Beskrivelse af nogle nye Arter.

Videnskabelige Meddelelser fra den naturhistoriske

Forening i Kjovenhavn, 8-12:135.

von Martens, E.

1865. Ueber zwei Seesterne von Costa Rica. Annals and

Magazine of Natural History, 1865:433.

McKnight, D. G.

1968. Some Echinoderms from Tongatabu Island and the

South Minerva Reef. New Zealand Journal of Marine and Freshwater Research, 2:712-715.

Mobius, K.

1859. Neue Seesterne des Hamburger und Kieler Museums. 14 pages, 4 plates.

Mueller, J., and F. H. Troschel

1840. [Not seen, title unknown.] Akademie Wissenshaften. Monatsberichtc der Koniglichen Preuss.

1842. System der Asteriden. 134 pages, 12 plates. Braunschweig: Friedrich Viewig und Sohn.

Perrier, E.

1875. Revision de la collection de stellerides du Museum

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1876. Revision dc la collection de Stellerides du Museum

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Sladen, P.

1889. Report on the Asteroidea collected by H.M.S. Challenger. Pages 1-893 of volume 30 in Report on the

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Stanek, V. J.

1955. La Beaute de la nature. 373 pages, illustrated.

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Steinbeck, J., and E. F. Ricketts

1941. Sea of Cortez. Pages 10 + 598, 40 plates. New York.

Verrill, A. E.

1867. Notes on Radiata. Transactions of the Connecticut

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4-10.

Viguier, C.

1878. Anatomie comparee du squelette des Stellerides.

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Ziesenhenne, F.

1937. The Templeton Crocker Expedition: Echinoderms

from the West Coast of Lower California, the Gulf

of California and Clarion Island. Zoologica, 22 (15):

209-239, figure 2.
Fishes Collected at Malpelo Island

John E. McCosker

and Richard H. Rosenblatt

ABSTRACT

Seventy species of shore fishes are reported from

Malpelo Island, the majority of them previously

unrecorded from that locality. Four species are

undescribed and may be endemic to Malpelo. No

new taxa are described. On the basis of these collections, it appears that the fauna is primarily

eastern Pacific mainland in composition, although

five species are shared only with the Galapagos and

Cocos islands.

The fish collections made by the expedition contain the first shore fishes taken by rotenone ichthyocides and diving at Malpelo. Previous collections

of Malpelo fishes were limited to those caught by

hook and line (Fowler, 1938, 1944) or dipnetted

beneath surface night lights (Clemens, 1957; Clemens and Nowell, 1963). Important material representing several new species and many new locality

records was obtained by the STRI expedition. The

collections, however, were limited in scope and are

not sufficiently representative of the probable

total ichthyofauna to allow any definitive statements concerning the faunal composition.

Malpelo occupies an important zoogeographic

location in that it arises from deep water beyond

the continental shelf, and could provide a

"stepping-stone" to the Galapagos Archipelago for

shore fishes. The known fauna is mainly eastern

Pacific mainland in origin. However, there are

certain similarities to both the Galapagos and

Cocos islands. This is evidenced by the presence

John E. McCosker, Steinhart Aquarium, California Academy

of Sciences, Golden Gate Park, San Francisco, California

94118. Richard H. Rosenblatt, Scripps Institution of Oceanography, La Jolla, California 92037.

of Mycteroperca olfax, Apogon atradorsatus,

Eupomacentrus arcifrons, and E. beebei, species

otherwise known only from those localities. Other

specimens collected included fishes previously

though to be Galapagos endemics, e.g., Labrisomus

dendriticus and possibly a deep water species of

Lythrypnus. The presence of certain reef-associated

Indo-Pacific species, e.g., Gymnothorax flavimarginatus, Zanclus canescens, and Acanthurus glaucopareius, is relatable to hydrographic and ecological conditions found on other eastern tropical Pacific offshore islands (Rosenblatt, McCosker, and

RubinofF, 1972). Included in the Malpelo collections were undescribed, and apparently endemic,

species of Chriolepis, Acanthemblemaria and Axoclinus, and an undescribed genus and species of

tripterygiid.

The following annotated list of fishes from Malpelo Island is based primarily on those collected

by J. B. Graham and party at two localities, and

underwater photographs and sight records made

by C. Birkeland (pers. comm.). Fishes from the

first locality (SIO 72-96) were collected along a

sloping basalt and sparsely covered Pocillopora

bottom between the surface and 6 meters along the

western edge of the island. The second collection

(SIO 72-97) is a composite of small east and west

shore stations, from the surface to depths of 20

meters. Fishes and photographs are deposited in

the Marine Vertebrates Collection of the Scripps

Institution of Oceanography (SIO), University of

California at San Diego. Verified records from

hook and line and night-light stations are also included. The basis of identification is indicated by

the following abbreviations: * specimens collected

and deposited at sio,

 s sight record by STRI diving

party,

 p photographed underwater by C. Birkeland,

 1 recorded by Fowler (1938), 2 recorded by

91
92

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Fowler (1944),

 3 recorded by Clemens (1957),

4

 recorded by Clemens and Nowell (1963).

CARCHARHINIDAE

Carcharhinus galapagensis (Snodgrass and Heller)l

C. falciformis (Muller and Henle)2 as C. malpeloensis

C. leucas (Muller and Henle) S

Triaenodon obesus Ruppell S

SPHYRNIDAE

Sphyrna sp. S

MURAENIDAE

Gymnothorax flavimarginatus (Riippell).P Abundant in the

West Indo-Pacific and Hawaii, and several eastern Pacific

offshore and nearshore localities (Rosenblatt, et al., 1972).

G. castaneus Jordan and Gilbert P

G. dovii (Gunther)P

Muraena lentiginosa Jenyns P

HOLOCENTRIDAE

Myripristis leiognathus Valenciennes • 3

Holocentrus suborbitalis (Gill) S

KUHLIIDAE

Kuhlia taeniura (Cuvier) S.1,2

APOGONIDAE

Apogon atradorsatus Heller and Snodgrass.* Previously

known only from the Galapagos and Cocos islands.

SERRANIDAE

Epinephelus dermatolepis Boulenger •

E. labriformis (Jenyns) •

Paranthias furcifer (Valenciennes) *1

Mycteroperca olfax (Jenyns).l Previously known from the

Galapagos, Malpelo, and Cocos islands. Phillip C. Heemstra

had kindly reexamined Fowler's specimens (ANSP 89121)

and found their gill rakers and dorsal spine profile to

agree with Galapagos specimens of M. olfax.

MULLIDAE

Pseudupeneus grandisquamis (Gill).4 Based on surface dipnetted prejuvenile specimens.

Mulloidichthys dentatus (Gill) S

CARANGIDAE

Selar crumenophthalmus (Bloch) 1

Caranx caballus GiintherS

C. vinctus Jordan and Gilbert 3

C. tnelampygus Cuvier and Valenciennes P

Elagatis bipinnulatis (Quoy and Gaimard) 2

LUTJANIDAE

Lutjanus Jordan; (Gilbert) l.P

Lutjanus viridis (Valenciennes) 1.2.P

POMACENTRIDAE

Chromis atrilobata (Gill) •*,•

Eupomacentrus arcifrons (Heller and Snodgrass).'.• Known

only from Malpelo, the Galapagos, and Cocos islands.

E. beebei (Nichols).* Previously known only from the Galapagos and Cocos islands.

Microspathodon dorsalis (Gill) *

LABRIDAE

Thalassoma lucasanum (Gill) •

Bodianus diplotaenia (Gill) !»•

SCARIDAE

Scar us rubroviolaceus BleekerS

KYPHOSIDAE

Sectator ocyurus (Jordan and Gilbert) S

Kyphosus sp. S

CHAETODONTIDAE

Holacanthus passer Valenciennes I.*

Heniochus nigrirostris (Gill) P

Pomacanthus zonipectus (Gill) S

CIRRHITIDAE

Cirrhitichthys oxycephalus (Bleeker) •

Cirrhilus rivulatus Valenciennes 1,2,*

SCORPAENIDAE

Scorpaenodes xyris (Jordan and Gilbert) •

Scorpaena plumieri Bloch •

AULOSTOMATIDAE

Aulostomus chinensis L.S

FISTULARIIDAE

Fistularia petimba Lac^pede S

GOBIIDAE

Lythrypnus sp. A *

Lythrypnus sp. B •

Chriolepis lepidotus Findley (see paper by Findley in this

volume)
NUMBER 176

GOBIESOCIDAE

Tomicodon cf. petersi (Garman)#

Arcos decor is Briggs •

Labrisomus cf. dendriticus (Reid).* Previously known only

from the Galapagos Islands.

Parastathmonotus culebrai (Seale) •

TRIPTERYGIIDAE

Axoclinus new species •

Genus and species.* An undescribed species, closely related

to Enneapterygius corallicola Kendall and Radcliffe from

the Galapagos Islands, and an undescribed species from

Panama and Costa Rica.

CHAENOPSIDAE

Acanthemblemaria new species •

BLENNIIDAE

Ophioblennius steindachneri Jordan and Evermann 3,4,*

Entomacrodus chiostictus (Jordan and Gilbert) *

HypsobUnnius brevipinnis (Giinther) •

Plagiotremus azaleus (Jordan and Bollman) *

ANTENNARIIDAE

Antennarius sanguineus Gill *

ACANTHURIDAE

Acanthurus glaucopareius Cuvier •

A. xanthopterus Valenciennes S

Prionurus laticlavius (Valenciennes) S

Zanclus canescens (Linnaeus) S

CANTHIGASTERIDAE

Canthigaster punctatissima (Giinther) •

TETRAODONTIDAE

Arothron meleagris (Bloch and Schneider) •

BALISTIDAE

Sufflamen verves (Gilbert and Starks) 1

Melichthys niger (Bloch).l.P As M. buiva in Fowler, 1938.

Canthidermis maculatus (Bloch). As C. sp. in Clemens, 1957.

Cantherhines dumerilii (Hollard) S

Alutera scripta (Osbeck) S

Literature Cited

Clemens, H. B.

1957. Fishes Collected in the Tropical Eastern Pacific,

1954. California Fish and Game, 43:299-307.

Clemens, H. B., and J. C. Nowell

1963. Fishes Collected in the Eastern Pacific during Tuna

Cruises, 1952 through 1959. California Fish and

Game, 49:240-264.

Fowler, H. W.

1938. The Fishes of the George Vanderbilt South Pacific

Expedition, 1937. Monographs of the Academy of

Natural Sciences of Philadelphia, 2: v + 349 pages,

10 plates. [Malpelo, pages 5-6.]

1944. Results of the Fifth George Vanderbilt Expedition

(1941). Monographs of the Academy of Natural

Sciences of Philadelphia, 6:57-529.

Rosenblatt, R. H., J. E. McCosker, and I. Rubinoff

1972. Indo-West Pacific Fishes from the Gulf of Chiriqui,

Panama. Contributions in Science of the Los Angeles County Museum, 234:1-18, 3 figures.
A New Species of Goby from Malpelo Island

(Teleostei: Gobiidae: Chriolepis)

Lloyd Talbott Findley

ABSTRACT

Two specimens representing a new species of the

gobiid fish genus Chriolepis were collected during

the 1972 Smithsonian Institution-U.S. Navy Expedition to Malpelo Island. The new species is herein

described, and its taxonomic relationships are

briefly discussed.

Introduction

METHODS.—Measurements were made with dial

calipers to the nearest 0.1 mm, and are expressed

as percent of standard length (SL) in Table 9.

Counts and measurements requiring explanation

are as follows: Head length is distance from anterior margin of upper lip to upper attachment of

opercular membrane. Head depth is depth of head

at the vertical, posterior margin of preoperculum.

Head width is the maximum width between cheeks.

Depth at anal fin origin is shortest distance between base of anal spine and dorsal fin base. Postorbital distance is the distance from posterior rim

of eye to upper attachment of opercular membrane. Transverse scale rows is the number of scale

rows, counted upward and forward, between the

last anal ray and dorsal fin base. The last dorsal

and anal rays, branched to their bases, were

counted as one.

The holotype and paratype (the only known

specimens) are deposited in the National Museum

of Natural History (USNM), Smithsonian Institution, Washington, D. C.

ACKNOWLEDGMENTS.—Thanks are due the followLloyd Talbott Findley, Department of Biological Sciences,

University of Arizona, Tucson, Arizona 85721.

ing persons: C. E. Dawson of the Gulf Coast Research Laboratory, Mississippi, J. E. McCosker of

the California Academy of Sciences, and D. A.

Thomson of the University of Arizona for reading

and improving the manuscript; J. B. Graham of

the Smithsonian Tropical Research Institute for

providing collection data and editorial advice; and

Jenean Thomson of the University of Arizona

for the illustration. I thank C. E. Dawson and D. F.

Hoese of the Australian Museum for encouragement, and R. J. Lavenberg of the Los Angeles

County Museum of Natural History for examination of the type-species. This paper is a contribution from the University of Arizona Marine Biology Program.

Chriolepis lepidotus, new species

FIGURE 35

HOLOTYPE.—USNM 211456, adult male, 30.0 mm

SL, Malpelo Island, Colombia, east side of island,

depth ca. 10 m, above the coral zone, cobble bottom with some calcareous sand and boulders, very

sparse algal growth; collected by J. B. Graham

using rotenone ichthyocide and SCUBA, 2-3 March

1972.

PARATYPE.—USNM 211457, adult male, 29.1 mm

SL; collected with holotype.

DESCRIPTION.— (See Table 9 for meristic and morphometric data). A robust species. Head plump,

with wide cheeks, more or less rounded; depth at

preoperculum 0.71 (holotype) and 0.68 (paratype)

of head width. Mouth oblique, forming an angle

about 40°-45° with body axis, the posterior angle

reaching a point approximately under rear margin

of pupil. Interorbital narrow, about one-third or

94
NUMBER 176 95

less of eye length. Pectoral base broad. Gill slit

narrow; opercular membrane attachments at upper

and lower edges of pectoral base in advance of insertion of fin rays.

Gill rakers short on first arch, those on second

and following arches shorter; 3+8=11 on outer

face of first arch; 9 on lower limb of second arch.

Pseudobranchiae in four small tufts.

Head without pores or barbels. Sensory papillae

on head relatively large; in rows below eye, on

cheek, at edge of preoperculum, on operculum, on

snout, and on dorsal and ventral surfaces of head

TABLE 9.—Counts and measurements of the holotype and paratype of Ghriolepis lepidotus

Characters

Sex

Standard Length (mm)

COUNTS

Dorsal fin rays

Anal fin rays

Pectoral fin rays (left/right)

Pelvic fin rays

Segmented caudal fin rays

Branched caudal fin rays

Procurrent caudal fin rays

(upper/lower)

Gill rakers (outer face of first arch)

Longitudinal scale rows

Transverse scale rows

PROPORTIONS (% of SL)

Head length

Head depth

Head width

Base of second dorsal fin

Depth at anal fin origin

Pectoral fin length

Pelvic fin length

Caudal fin length

Caudal peduncle length

Caudal peduncle depth

Eye length

Snout length

Mouth length

Width between mandible tips

Interorbital width

Width between posterior nostrils

Length between anterior &

posterior nostrils

Postorbital distance

Holotype

(USNM

211456)

30.0

VII-1,10

1,9

20/20

1,5

17

15

5/5

11

ca.34

13i/2

33.3

15.5

21.8

28.5

16.8

27.3

205

24.7

16.0

13.7

6.7

7.6

13.4

10.2

2.4

4.1

2.9

19.6

Paratype

(USNM

211457)

$

29.1

VII-1,10

1,9

20/20

13

17

14

6/5

11

ca.35

13i/2

32.2

15.6

23.5

275

175

24.7

21.0

23.5

13.6

13.8

6.2

6.9

125

10.1

1.9

5.1

2.3

19.1

including mandibular area, and similar to other

species in the genus (Figure 35). A low, shallowly

V-shaped, transverse, fleshy crest on head behind

eyes. A low, short, fleshy "keel" extending medially

between nostrils. Nostrils tubular; anterior tube

narrow, its length about equal to pupil diameter;

posterior tube wider, its length about one-half that

of anterior tube. Tongue tip truncate.

Teeth in rows in both jaws, relatively large,

sharp, mostly caniniform, slightly to distinctly recurved, mostly depressible and with faint yellowish

apical tips; outer row teeth in upper jaw enlarged;

outer and inner row teeth in lower jaw enlarged.

Lower jaw with (1) abbreviate outer row of large,

slender, recurved, well-separated teeth confined to

anterior portion of jaw; (2) behind outer row anteriorly is a patch of many small (about % to 14

length of outer row teeth) closely set, less recurved

teeth in irregular rows, becoming fewer in number,

more widely spaced, and confined to one row posteriorly at side of jaw; (3) inner row of large, slender,

recurved, well-separated teeth anteriorly, becoming more widely spaced and stouter posteriorly,

until 2 to 3 much enlarged and recurved "fanglike"

teeth at side of jaw, followed by a few, much

smaller, closely set teeth. Upper jaw with (1) outer

row of large, slender, recurved, well-separated

teeth anteriorly and at side of jaw (not absent

posteriorly as in outer row of lower jaw); (2) behind outer row anteriorly is a patch of many small,

closely set, less recurved teeth in irregular rows,

becoming fewer in number posteriorly; (3) inner

row of many small, closely set teeth, becoming more

widely spaced posteriorly. No teeth on vomer or

palatines.

First dorsal fin with seven flexible spines; bases

of first five spines equally spaced; the last two

spines widely separated from the first five, their

bases widely spaced. Third and fourth spines slightly the longest, but no spine notably longer than

any other, and none filamentous.

Second dorsal fin separated from first dorsal fin

by distance slightly more than eye length. First

element of both second dorsal and anal fins a flexible spine; remaining elements are subequal, segmented, branched rays, with the last branched to

their bases.

Pectoral fins when adpressed reach past tips of

pelvic fins to level of end of spinous dorsal; all

pectoral rays branched.
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Origin of pelvic fins in advance of pectoral base.

Pelvic fins short, not reaching anus; separated and

diverging, not united by a membrane into an adhesive disc as in most gobiids; interspinal membrane absent, but pelvic bases slightly connected

by a very low, inconspicuous frenum. Pelvic spine

flexible; segmented pelvic rays well developed and

branched, most with three main branches, the

innermost ray branching less than others, but not

reduced or splinted to the fourth. Fourth ray

slightly the longest, then the third, then the fifth

(which is almost equal to the third). Caudal fin

short, rounded.

Urogenital papilla of male long, slender, attenuate; with a few scattered melanophores.

SQUAMATION.—Four enlarged strongly ctenoid

basicaudal scales (some lost from both specimens);

the outer two (at upper and lower angles of caudal

fin base) slightly narrower and longer than the

inner two. Body extensively scaled. Scales along

midline extending forward from hypural to near

axil, beneath origin of spinous dorsal fin (Figure

35), in about 34-35 more or less irregular longitudinal rows; difficult to enumerate (some scales lost

from both specimens, but scale pockets usually

discernible). Fewer scales in anterior rows; in a

wedge-shaped pattern as approaching axil. Transverse scales difficult to enumerate, about 13i/£

in an irregular row (first scale at base of last anal

ray very narrow in comparison to scales above it,

counted as i/2 scale).

Posterior scales large, with prominent ctenii; becoming progressively smaller, thinner, more embedded, and with fewer and shorter ctenii, until

cycloid anteriorly. Ctenii becoming progressively

restricted to central portion of posterior edge of

each scale proceeding anteriorly, until completely

lost at about twelfth scale along midline in advance of hypural. Anteriormost scales cycloid,

minute, embedded, and difficult to see.

A naked area beginning near base of sixth dorsal spine, equivalent to one scale width from base

of sixth spine to below base of fifth spine, to two

scale widths below fourth spine, becoming wider

and sloping downward and forward to near upper

margin of pectoral base. A lower naked area from

under pectoral base sloping downward and backward to near midline of venter, continuing as a

narrow strip around anus to about base of fourth

anal ray. Head, nape, chest, and venter scaleless.

COLORATION IN ETHYL ALCOHOL.—Background

color of head and body brownish yellow. Head

with a number of prominent black spots, especially

laterally and ventrally. An elongate spot at midposterior edge of eye (diffuse in paratype); one

behind eye at dorsal edge of cheek near end of low

transverse dorsal crest; one at dorsal margin of

preopercular-opercular junction; one at midanterior edge of operculum which begins a row of

about 9 or 10 widely spaced spots lying in preopercular-opercular groove, following curvature of

groove downward and forward and onto the immediately adjacent branchiostegal membrane, then

continuing forward ventrally below lower jaw to

FIGURE 35.—Chriolepis lepidotus, holotype, USNM 211456, male, 30.0 mm SL, Malpelo Island,

Colombia. Most scales not shown; the dashed line indicates anterior limit of squamation. The

row of specialized basicaudal scales is a composite from paratype and right side of holotype.

Sensory papillae on head shown as small circles and are a composite from both specimens.
NUMBER 176 97

near symphysis. This row is partially duplicated by

a shorter row of about 5 spots beginning at lower

posterior edge of preoperculum, running forward

below lower jaw to near symphysis, the spots lying

close to and opposite the spots of row described

above. About 5-8 widely spaced spots scattered over

cheek (more prominent on paratype). A spot (on

paratype) at posterior end of interorbital, followed by profuse minute melanophores on the low

transverse dorsal crest. A spot at posterior dorsal

corner of operculum. A prominent spot on upper

and lower angles of outer pectoral fin base; a more

distal irregular spot at bases of fifth to eighth pectoral fin rays (slightly larger on paratype). A small

spot on inner pectoral fin base near upper margin.

A black bar below anteroventral edge of eye,

passing obliquely downward across suborbital and

jaws, terminating in anteriormost ventral spot of

the series described above. A fainter dark bar extends from midventral edge of eye downward across

suborbital region, and curves around end of lower

jaw. Snout, nostrils, upper jaw, and anterior portion of lower jaw densely peppered with minute

melanophores.

Two wide, transverse, light brown bars across

dorsal surface of head, the posterior is across the

nape and bisected at either side by pectoral fin.

Five wide, vertical, light brown bands on body,

most noticeable dorsally as dark saddles, becoming

more diffuse ventrally, and terminating near midventral line. Brownish yellow interspaces between

bands with a few scattered melanophores. Each

band with one or two adjacent, prominent, black

spots at midline. Anteriormost body band crosses

dorsum between first and fifth dorsal spines, and

passes downward behind pectoral fin; second band

between sixth spine and end of spinous dorsal fin;

third band between second and fifth dorsal rays;

fourth band between seventh and ninth rays; fifth

band behind end of second dorsal fin on anterior

portion of caudal peduncle. Dorsal segments of

bands 3 and 4 appearing as darker blocks offset

posteriorly from their ventral counterparts below

midline, with dorsal block of band 3 most noticeable and lying over the interspace between third

and fourth ventral blocks. An irregular dark blotch

behind fifth band at dorsal edge of caudal peduncle, adjacent to first procurrent ray. A narrow

vertical bar of darker pigment underlying basicaudal row of specialized ctenoid scales.

Dorsal fins with about four or five oblique rows

of dark lines, noticeable as prominent black spots

on spines and rays (membranes torn in both specimens). A dark horizontal bar on anal fin membrane, running the length of fin near its edge. A

few faint diffuse spots on caudal rays suggest three

or four vertical lines. Pectoral fins slightly dusky

on interadial membranes. Pelvic fins dark.

Chest and venter whitish, with numerous minute

melanophores, appearing "peppered" on microscopic inspection. Inside of gill chamber with a

dark horizontal stripe ventrally, in advance of

lower pectoral base; a few melanophores dorsally

in area surrounding pseudobranchial tufts.

RELATIONSHIP.—The general head shape, relatively high number of fin rays, and general color

pattern suggest a relationship with a species complex of Chriolepis, most of whose members in the

eastern tropical Pacific are undescribed. Among

the described species, C. lepidotus is perhaps closest to C. minutillus Gilbert, 1891, and C. tagus

Ginsburg, 1953 (judging by original description

only). Complete discussion of relationships within

the genus Chriolepis and allied genera of sevenspined gobiids with separate pelvic fins is deferred

pending completion of a revision in progress by

the author of all known species in the eastern tropical Pacific.

COMPARISONS.—Chriolepis lepidotus differs from

other described congeners in the eastern tropical

Pacific in the extent of its squamation; its scales

extend forward to beneath the origin of the spinous dorsal fin. Among the other described species,

C. minutillus Gilbert, 1891 (Gulf of California),

most closely approximates this condition; its scales

extend forward to beneath or slightly in advance

of midspinous dorsal. From the description of C.

tagus Ginsburg, 1953 (Galapagos Islands), and in

C. zebra Ginsburg, 1938 (Gulf of California),

scales only extend forward to beneath the end of

the spinous dorsal, or slightly posterior to this position in the latter species.

The new species also differs from other described

eastern tropical Pacific congeners in having the

highest number of pectoral fin rays (20 vs. 19 in

C. tagus; 17 or 18 in C. zebra; 15 or 16 in C.

minutillus.) The prominent black spots on the

head further serve to distinguish C. lepidotus from

the other species described from these waters.

Chriolepis lepidotus further differs from C. min-
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

utillus in having a larger mouth (posterior angle

extends to below posterior margin of pupil vs. to

below midpupil in C. minutillus), shorter pelvic

fins (when adpressed, not reaching anus vs. reaching or almost reaching anus in C. minutillus), and

more gill rakers on the outer face of the first arch

(11 vs. 8 or 9 in C minutillus).

The new species further differs from C. zebra in

having a rounded rather than a depressed head, a

larger mouth (posterior angle extends only to below midpupil in C. zebra), and more second dorsal

fin rays (11 vs. 10 in C. zebra).

Chriolepis lepidotus differs from the description

of C. tagus in having fewer second dorsal fin rays

(12 in C. tagus), and fewer anal fin rays (10 vs. 11

in C. tagus).

DISTRIBUTION.—Chriolepis lepidotus is presently

known only from Malpelo Island, Colombia.

ETYMOLOGY.—From the Greek lepidotos (scaly),

referring to its squamation, the most extensively

known in the genus.

REMARKS.—The genus Chriolepis was erected by

Gilbert (1891:557-558) for the species C. minutillus, which he described as being totally scaleless on

the basis of a single specimen trawled by the Albatross in the Gulf of California. His description was

in error, since the holotype retains a single basicaudal scale, as mentioned by Ginsburg (1938:111;

1953:21), and confirmed by examination of the

type by D. F. Hoese and R. J. Lavenberg (pers.

comm.). Recently collected material of C. minutillus under study by the author shows that, indeed,

squamation is extensive in this species, less so than

in C. lepidotus, but more so than in C. zebra, for

example.

Species of Chriolepis are sublittoral in habitat

and secretive, hiding under rocks and in crevices.

Until recently, few specimens were known, but

with the introduction of SCUBA and rotenone ichthyocide collecting techniques by diving ichthyologists, specimens have begun to accumulate. These

will allow for a comprehensive systematic, zoogeographic, and evolutionary treatment of the genus

and its allies (briefly treated by Bohlke and Robins, 1968:129-131) in the eastern tropical Pacific.

There remain about five undescribed species of

Chriolepis in these waters.

Literature Cited

Bohlke, J. E., and C. R. Robins

1968. Western Atlantic Sevcn-spined Gobies, with Descriptions of Ten New Species and a New Genus,

and Comments on Pacific Relatives. Proceedings of

the Academy of Natural Sciences of Philadelphia,

12O(3):45-174.

Gilbert, C. H.

1891. Scientific Results of Explorations by the United

States Fish Commission Steamer Albatross, XXII:

Descriptions of Thirty-four New Species of Fishes

Collected in 1888 and 1889, Principally among the

Santa Barbara Islands and in the Gulf of California. Proceedings of the United States National

Museum, 14 (880):539-566.

Ginsburg, I.

1938. Eight New Species of Gobioid Fishes from the

American Pacific Coast. Allan Hancock Pacific

Expeditions, 2(7):109-121.

1953. Ten New American Gobioid Fishes in the United

States National Museum, Including Additions to a

Revision of Gobionellus. Journal of the Washington

Academy of Sciences, 43(1): 18-26.

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