The Biological Investigation of Malpelo Island, Colombia

The Biological Investigationof Malpelo Island, Colombia

JEFFREY B. GRAHAMEDITOR

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 176

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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y • N U M B E R 1 7 6


Jeffrey B. GrahamEDITOR

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, 35figures, 1975.—The results of joint Smithsonian Institution and U. S. Navy ter-restrial and marine investigations of Malpelo Island, Republic of Colombia arereported 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 ofMalpelo and the behavior and natural history of the lizards Anolis agassizi andDiploglossus millepunctatus are described and discussed. Genie variability inA. agassizi has been investigated and karyotypes of A. agassizi and D. millepunc-tatus are reported. The ecology of the island's benthic marine communities isdetailed and papers listing and discussing zoogeographically interesting featuresof the island's crustacean, starfish, and fish species are included. The geology ofMalpelo is briefly described and an improved map of the island is presented.The importance of Malpelo Island in the understanding of biogeographic prob-lems in the eastern tropical Pacific Ocean is reviewed.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recordedin the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5095. SERIES COVERDESIGN: The coral Montastrea cavemosa (Linnaeus).

Library of Congress Cataloging in Publication DataMain 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. Marinebiology—Malpelo Island. 3. Malpelo Island. I. Graham, Jeffrey B., ed. II. SmithsonianInstitution. III. United States. Navy. IV. Series: Smithsonian Institution. Smithsonian con-tributions to zoology, no. 176. [DNLM: 1. Ecology. 2. Lizards. 3. Marine biology. W1SM454Nno. 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 OfficeWashington, D.C. 20402 - Price $2.20 (paper cover)

Contents

Page

INTRODUCTION, by Jeffrey B. Graham 1TERRESTRIAL BIOLOGY OF MALPELO ISLAND: A HISTORICAL REVIEW,

by George C. Gorman and Terence L. Chorba 9RECONNAISSANCE AND MAPPING OF MALPELO ISLAND, by A. Ross Kiester

and Jeffrey A. Hoffman 13FIELD OBSERVATIONS ON THE GEOLOGY OF MALPELO ISLAND,

by Jeffrey A. Stead 17THE ECOSYSTEM ON MALPELO ISLAND, by Henk Wolda 21NATURAL HISTORY, BEHAVIOR, AND ECOLOGY OF Anolis agassizi,

by A. Stanley Rand, George C. Gorman, and William M. Rand 27NOTES ON THE NATURAL HISTORY OF Diploglossus millepunctatus

(Sauria: Anguidae), by A. Ross Kiester 39A NEW GECKO FROM MALPELO ISLAND (SAURIA: GEKKONIDAE:

Phyllodactylus), by Raymond B. Huey 44ELECTROPHORETIC ESTIMATES OF GENIC VARIATION IN, AND THE

RELATIONSHIPS OF, Anolis agassizi, by T. Preston Webster 47NOTES ON THE CHROMOSOMES OF Anolis agassizi (SAURIA: IGUANIDAE)

AND Diploglossus millepunctatus (Sauria: Anguidae), by Brad Stammand 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 ; 69ASTEROIDEA FROM MALPELO ISLAND WITH A DESCRIPTION OF A NEW

SPECIES OF THE GENUS Tamaria, by Maureen E. Downey 86FISHES COLLECTED AT MALPELO ISLAND, by John E. McCosker and

Richard H. Rosenblatt 91A NEW SPECIES OF GOBY FROM MALPELO ISLAND (TELEOSTEI:

GOBIIDAE: Chriolepis), by Lloyd Talbott Findley 94

i n


Introduction

Jeffrey B. Graham

On 28 February 1972 a group of 17 scientistsrepresenting the Smithsonian Tropical ResearchInstitute, the Republics of Panama and Colombia,and several U.S. universities embarked on a sixday expedition to investigate the marine and ter-restrial biota of Malpelo Island, Colombia, a smallisolated Pacific island 270 miles to the west of Co-lombia and south of Panama. The expedition, ajoint project of the U.S Navy and the SmithsonianTropical Research Institute, was undertaken togain a more comprehensive understanding ofecological processes and the natural history ofplants and animals on Malpelo. The scientificparty consisted mostly of biologists who made col-lections, carried out field observations, and con-ducted experiments on the island and in the watersaround it. (The Malpelo scientific party consistedof 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 TropicalResearch Institute (STRI); O. Arroyo, ColombianInstitute of Natural Resources (INDERENA); J. Bar-reto S., Universidad George Tadeo Lozana, Colom-bia; G. C. Gorman, University of California LosAngeles (UCLA); J. A. Hoffman, Smithsonian As-trophysical Laboratory; A. R. Kiester, Museumof Comparative Zoology; D. B. Macurda, Univer-sity of Michigan; W. M. Rand, MassachusettsInstitute 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 Hydro-graphic Center. R. J. Kinney, USN, served as ex-pedition physician. The geology of Malpelo wasstudied and a more accurate map of the island wasmade by an exploration team.

The scientists were transported from Panama toMalpelo by the U.S.S York County (LST 1175)commanded by LCDR Lauren Seeber. At Malpelo,the scientists were supported by York County crew-men who did everything from prepare box lunchesto operate small boats and assist in the collectionand sort of specimens. While on station the YorkCounty conducted sea trials and made numerousdepth soundings and positional sightings of theisland. This work, while helping to establish thatthe peaks on Malpelo are some 125 meters higherthan previous maps indicated, will also improvethe hydrographic charts of this area.

The Smithsonian-U.S. Navy Expedition to Mal-pelo marked the fourth time that Smithsonianscientists and their colleagues have been able touse a U.S. Navy vessel for research purposes. InMarch 1970 biological investigations of the SecasIslands in the Gulf of Chiriqui, Panama, were con-ducted by scientists aboard the U.S.S. TraverseCounty. Later in the same year the U.S.S. Wal-worth County supported marine studies at CoibaIsland, Panama. In 1971 the U.S.S. Terre BonneParish carried a research team to Bocas del Toro,Panama, for three days of field work. In the sum-mer of 1973, while the volume on Malpelo was

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

being compiled, the fifth and most ambitious ex-pedition was undertaken when the U.S.S. Spartan-burg County took 25 scientists to Cocos Island,Costa Rica, 550 miles southwest of Panama.

These expeditions have been fruitful. The larg-est and best developed coral reef formations to befound thus far in the eastern tropical Pacific werediscovered and subsequently studied by PeterGlynn and colleagues during the Gulf of Chiriquitrip (Glynn, Stewart, and McCosker, 1972). Previ-ous to these investigations it was thought that nowell-developed coral reefs occurred in the easterntropical Pacific. The coral-eating starfish Acanth-aster planci was found in great abundance andseveral species of Indo-Pacific corals were also dis-covered by Glynn and his co-workers during thesame expedition. Our understanding of the faunalcomposition and zoogeography of eastern tropicalPacific fishes has benefited significantly from Navy-supported research in this region (Rosenblatt,McCosker and RubinofF, 1972).

The use of Navy ships is a valuable addition tothe research capacity of the Smithsonian TropicalResearch Institute. Navy vessels allow many scien-tists to participate in the expeditions and it ispossible to visit localities beyond the range of smallresearch vessels. Needless to say, costs and thelogistical support problems are also vastly reduced.

ACKNOWLEDGMENTS.—It is a pleasure to acknow-ledge Admiral C. H. Griffiths, Commander of theUnited States Navy, Fifteenth Naval District, FortAmador, Canal Zone, and his staff who madearrangements for the Malpelo Expedition. We alsothank Captain Lauren Seeber and the officers andmen of the U.S.S. York County for supporting ourinvestigations. We express our appreciation to theForeign Ministry of the Republic of Colombia forpermitting us to visit Malpelo Island and to theU.S. Army Southern Command, 352nd AviationDetachment, for providing an aerial reconnaissanceof the island. I would like to thank my colleagues:L. G. Abele of Florida State University, D. Dexterof San Diego State University, J. McCosker of theCalifornia Academy of Sciences, and A. S. Rand,I. RubinofF, and R. R. Warner, all of the Smith-sonian Tropical Research Institute, for their ad-vice and assistance in the preparation of this vol-ume; I acknowledge the technical assistance ofI. Downs, A. Kourany, and V. Vergel of the Smith-sonian Tropical Research Institute.

Island Biology and Its BiogeographicalSignificance

The biological investigation of islands hasmade a valuable contribution to the studies ofevolution, biogeography, and ecology. Islands, byvirtue of their differences in size, shape, climaticconditions, and distances from continents, presenta stepped progression of natural experimentalareas (MacArthur, 1972). Darwin's ideas on thenatural variability and selection of species devel-oped in part through his observations at the Gala-pagos Islands. Some biologists investigate islandsto test different aspects of modern evolutionarytheory. Islands, however, are intrinsically inter-esting because each is unique, and they afford bio-logists the chance to study the effects of isolationon the resident species. Island populations mayhave greatly expanded or otherwise modifiedniches and may have decreased levels of geneticvariability or, if isolated for a long period, endemicspecies may be formed (MacArthur and Wilson,1967).

Malpelo Island (3°51'07"N, 81°35'40"W) is oneof several oceanic islands in the eastern tropicalPacific. It is the only island on the Malpelo Ridge(Figure 1) a solitary volcanic submarine ridge that

extends in a northeast-southwest direction with alength of 150 miles and a width of 50 miles (Chase,1968). An oceanic island, Malpelo occurs outsidethe continental shelf and it is separated from themainland of Central America by depths greaterthan 1800 fathoms. Malpelo has never been con-nected, even by shallow water, with any otherislands or the mainland.

Malpelo's distance from the mainland and thedepths of water between the island and the main-land are significant barriers to the colonization ofboth terrestrial and shallow-water marine organ-isms. Colonization therefore must occur throughsea surface currents which would transport raftingterrestrial organisms and the planktonic larvae andsubadults of marine forms to the island. Malpelois closer to the mainland than Cocos and the Gala-pagos islands (Figure 2) and it is reasonable toassume it serves as a stepping stone for marineforms enroute, via oceanic currents, to theseislands.

Consideration of the complex pattern of surfacecirculation in the eastern tropical Pacific Ocean

NUMBER 176

(Wyrtki, 1965) provides insight as to how surfacecurrents may serve for transport from the mainlandto Malpelo and beyond to Cocos and the Gala-pagos islands. Only general features of surfacecirculation in the area of these islands for themonths of January and July are presented (Figure2) and the reader is referred to Wyrtki's paper fordetails on monthly current vectors, velocities, andthe persistence of flow for the different areas in theeastern tropical Pacific.

In the months from January to March the trade-winds cause a strong current flow (0.5 knots) outof the Gulf of Panama towards Malpelo and theGalapagos islands (Figure 2). A small anticyclonic

gyre that borders the mainland and encompassesCocos and Malpelo islands is also formed at thistime. If winds blow strongly for long periods thecondition known as "El Nino" occurs. This resultsin the transport of a larger than normal amountof warm surface water, which displaces the Perucurrent to the southwest away from South Americaand the Galapagos Islands. During £1 Nino theGalapagos Islands may be completely bathed inwarm tropical waters.

The Equatorial Countercurrent (Figure 2) doesnot penetrate very far into the eastern tropicalPacific from January to April, but from May toDecember this current is strong and reaches the

FIGURE 1.—Submarine topography of the central eastern tropical Pacific Ocean in the vicinityof Malpelo, Cocos, and the Galapagos islands. The lines are 600, 1000, and 1200 fathoms. Dataare from Chase (1968).


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 PacificOcean. 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. Themajor part of the Equatorial Countercurrentflows northwest along Central America, while theremainder flows south and then turns west as partof the South Equatorial Current. From May toDecember Clipperton, Cocos, and Malpelo islandslie in the swath of the Equatorial Countercurrent,and the Galapagos Islands occasionally receivewater from the southern part of this current fromAugust to October. The combination of strongsouthern surface flow in the months from Januaryto May and the circulation pattern of the Equato-rial Countercurrent probably account for the dis-persal of mainland marine organisms to theseislands.

Another phenomenon that is of special interestto marine zoogeographers is the presence and dis-persal patterns of Indo-Pacific organisms (corals,mollusks, crustaceans, and fishes) at the easterntropical Pacific islands and on the Central Ameri-can mainland (Costa Rica and western Panama).This has been recently reviewed for fishes byRosenblatt, McCosker, and Rubinoff (1972). Lar-vae of these animals are apparently transportedacross 1000 miles of open sea by the EquatorialCountercurrent, which, as was shown above, ex-tends to the mainland of Central America andsweeps across some oceanic islands for long periodsof the year. It is not presently known how manyspecies have successfully completed this transit orwhether they more frequently colonize oceanic is-lands or the mainland.

Summary of the Malpelo Expedition

The amount of information that was accumu-lated in the four full days of field work on Malpelois gratifying. Part of the expedition's success lay inwhat was learned about the island by its earlierexplorers. Gorman and Chorba have traced thehistory of Malpelo's exploration, which goes backto 1542. Early visitors to Malpelo had the impres-sion that the flow of energy to the terrestrial eco-system had to be sustained by the sea and that thiswas done mainly by birds. While acknowledgingthat the success of the land crab (Gecarcinus) andthe large lizard (Diploglossus) definitely dependson the activity of birds, Wolda has found that theterrestrial community is much too large and com-plex for this to be the only pathway. He concludes

that lichens and mosses constitute the importantlower trophic levels for the land community.

Malpelo's endemic lizards were well studied onthis expedition. A new species of gecko (Phyllo-dactylns transversalis) was discovered and this isdescribed by Huey. This species, while similar inmany ways to Peruvian species has its closest affinityto Mexican Phyllodactylus. The depth of biologi-cal knowledge about mainland and island Anolis,together with their extensive research experiencewith anoles enabled Rand, Gorman, and Rand toask specific questions about the ecology and be-havior of Malpelo's A. agassizi. The Malpelo anoleis larger than most other anoles isolated on islandsand it reproduces seasonally. Its social behavior isstrikingly different from that of other Anolis; itis not territorial, there is little aggression, and itsdisplays are few and simple. Anolis agassizi is anactive, curious lizard that is readily attracted tothings that appear in its habitat: scientists, theirequipment, and their fruit and candy as well.Their curiosity and the remarkable tolerance ofindividuals for each other, these authors believe,enables A. agassizi to take advantage of resourcessuch as water and food that have variable distribu-tions in time and space on Malpelo. Kiester de-scribes Diploglossus millepunctatus as curiousand opportunistic and draws basically the sameconclusions about the ecological significance ofthis behavior as were made for A. agassizi. Interest-ingly, Kiester postulates that D. millepunctatus,because of its dependence on birds, may have itsreproductive seasonality synchronized with thatof the nesting seabirds. This awaits further study.Webster measured a low degree of genie variabilityin A. agassizi. From what is presently known thisseems to be a characteristic of old, solitary anolepopulations. Stamm and Gorman found that thekaryotypes of A. agassizi and D. millepunctatusare similar to the more primitive groups in thesetwo genera. Taken together these observationsmight suggest that Malpelo's Anolis and Diplo-glossus 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 thiswith certainty.

Kiester and Hoffman surveyed the island and,with special climbing equipment, scaled its sum-mits. Sightings from the peaks and from the bridgeof the York County enabled them to determine


FIGURE 3.—Left: Aerial view (4000^ of Malpelo Island. The northern end of the island is inthe foreground. Right: A York County landing craft approaching the west side of MalpeloIsland.

that the highest elevations of Malpelo are almost125 meters higher than early maps indicated. Steadconducted a geological survey of Malpelo. Hisdescription of the island's weathering, togetherwith a compilation of the York County's depthsoundings around Malpelo enables us to appre-ciate the original size of this rock. Once 8 to 10times its present size, Malpelo has been steadilyworn by the persistent pounding of the sea. Whenwe brought our small boats close to shore andtried to land on the island we keenly sensed theintensity of the struggle between Malpelo and thesea (Figure 3).

Rough water and the island's steep slope madegetting from the boat to shore and returning amajor undertaking for the terrestrial party. Ex-ploration ended prematurely for one scientist inthis group who severly injured his arm while tryingto get from the island into the boat. In spite ofthese obstacles none in the terrestrial group electedto spend their nights on Malpelo and, at days end,all were glad to return to the haven of the YorkCounty. The warship took on a curious appearanceat this time of day. Its main deck was littered withbiological specimens and collecting equipment andwas crowded with seamen who were rapidly trans-formed into eager assistants, audiences, or both.The crewmen relished this and took great pleasurein hearing about the discoveries and adventuresof the day. Accounts were passed rapidly to thesailors on watch and expedition progress reportswere included in the ships daily bulletin. News of

the discovery of the new Phyllodactylus species atMalpelo may have traveled faster than any othersjiecies discovery in history as it was flashed thesame day to the ship's home port at Little Creek,Virginia. Sorting, preservation, and labeling con-tinued past dark while at the same time, in otherparts of the ship, other men worked, and scubatanks, thermos bottles, and lunch containers wererefilled for use the next day. When work wasfinished for the day, showers, warm food, a recount-ing of the day's adventures, a soft bed, and last,but by no means least, the evening movie fullyrestored us for the next day.

Investigation of Malpelo's marine habitat wasalso rewarding. The water was so rough, however,that the small boats we had brought for the divingteams were unsafe and diving operations had tobe staged from one of the York County's boats. Weinitially tried to use the ship's landing crafts butthese had so much freeboard that they were notpractical. After struggling through a day's divingfrom the landing craft, I related this difficulty toCaptain Seeber, who immediately offered us theuse of his launch. Thus it came to be that theelegantly furnished Captain's Gig became ourdiving boat. No doubt the launch has seen betterdays both before and since we loaded it with divers,their gear, and specimens. But it proved an invalu-able asset at Malpelo. There were many sharksaround the island; they were seen at the surfaceand on every dive. No particularly dangerous en-counters occurred with sharks during our dives,

NUMBER 176

but their omnipresence was sobering and at timestheir proximity was slightly unnerving.

Birkeland and colleagues completed their ambi-tious project of studying and characterizing thesubtidal benthic communities at Malpelo. Thesmooth vertical subtidal walls are dominated bylarge, sparsely distributed barnacles. Recruitmentsuccess for transforming barnacle larvae is low butonce they are established, adults may live for along time. On the gently sloping southeastern edgeof Malpelo, Birkeland's group found extensivecoral development. Although they do not consti-tute a true coral reef, these corals are luxuriantand depth zonation is marked for different species.The occurrence of corals to as deep as 30 meters,about the deepest known in the eastern tropicalPacific, is attributable to the clear water aroundMalpelo. A list of all the marine invertebratesknown from Malpelo is appended to the Birkelandet al. paper. Abele records 43 species of decapodCrustacea (18 species of macrurans) at Malpeloand describes two new species of snapping shrimp.Abele lists the distribution of each species and, bydrawing upon his other recently completed studies,he makes an interesting observation about thespecies diversity of decapod crustaceans at Malpelo,Cocos Island, and several other eastern tropicalPacific localities. The diversity of starfish at Mal-pelo is low and Downey lists only six species inher paper. Three species occur throughout theeastern tropical Pacific and a new species of Tam-aria and a new subspecies of Narcissia are de-scribed for Malpelo. Until Downey's work, oneMalpelo starfish (Leiaster) was previously knownto occur only at the Hawaiian Islands.

Our knowledge of the Malpelo fish fauna is stillincomplete but McCosker and Rosenblatt reportthat 70 species of fish are now known for the island.In the specimens collected in the Smithsonian-Navy Expedition, McCosker and Rosenblatt foundseveral new species, as well as a new genus of fish;all awaiting description except the new speciesof goby (Chriolepis) described by Findley. TheAbele, and McCosker and Rosenblatt papers dem-onstrate several interesting features that the deca-pod crustaceans and fishes of Malpelo have incommon. The majority of species in both groupsare Central American mainland forms; there arealso Indo-Pacific species and some endemics. Bothgroups contain species that were previously

thought to be endemic at Cocos or the Galapagosislands or were known only from these two locali-ties. These findings strongly suggest that Melpe-lo is a stepping-stone for marine mainlandspecies enroute to more distant islands and, as thesurface current patterns indicate, for Indo-Pacificspecies as well. Some of the terrestrial invertebrateslisted by Wolda appear to be most closely relatedto Indo-Pacific groups, which prompts the con-clusion that these were rafted from the old worldto Malpelo via the Equatorial Countercurrent.The exchange of species between Malpelo, Cocos,and the Galapagos Islands is implied by their simi-lar fish and decapod faunas and by the markedfaunal similarity observed by Birkeland et al. be-tween Malpelo and the Galapago Islands. Furtherwork with these faunas will clarify these relation-ships.

Although much was learned at Malpelo theshort time for study has left questions and the biotaof the island would doubtlessly be better under-stood if more sampling, observation, and experi-mentation could be conducted. It was agreed thatadditional visits should be made, ideally at dif-ferent times of the year so that the possible influ-ence of seasons could be assesed. That there ismuch to learn is evident when it is pointed outthat we still do not know how much rain annuallyfalls on Malpelo. Murphy (1945) suggested thatthe island never receives rain, but Mapelo doeslie within the range of the convergence of the tradewinds (Intertropical Convergence Zone, Wyrtki,1965), which suggests a seasonal rainfall. The is-land does have weathered, phosphatized rock(McConnell, 1943), and seepage marks, drystream beds, and pools of water were found by theSmithsonian-Navy expedition. All of these suggestthat rain is not uncommon but expeditions to theisland have practically all been during the monthsof the dry season (December to March) and wecan only speculate about amounts of rain in therest of the year.

This volume, by reviewing and compiling Mal-pelo's literature and by reporting results to dateof the most thorough investigation of the islandsets the stage for future exploration. It describesMalpelo but it cannot convey the impressivescenario of that island. When viewed objectivelyit is an isolated, inhospitable island, which by these

8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

virtues has remained a natural laboratory for thestudy of evolution.

When seen first hand, however, Malpelo is alarge grotesque rock that rises rudely, ominouslyout of a monotonous sea. As we contemplated thisrock we reminded ourselves, and took pleasure inthe fact, that it was a sight that few people haveever 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. McCosker1972. Pacific Coral Reefs of Panama: Structure, Distribu-

tion, and Predators. Sonderdruck aus der Geolo-gischen 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. Bul-

letin 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. Rubinoff1972. Indo-West Pacific Fishes from the Gulf of Chiriqui,

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

Wyrtki, K.1965. Surface Currents of the Eastern Tropical Pacific

Ocean. Bulletin of the Inter-American TropicalTuna Commission, 9:271-304.

Terrestrial Biology of Malpelo Island:A Historical Review

George C. Gormanand Terence L. Chorba

ABSTRACT

Records of the exploration of Malpelo Island dateback to 1542. Early biological investigations mainlyconcerned the terrestrial fauna, particularly theisland's lizards. The complete list of Malpelo'savifauna is presented. The etymology of Malpelois discussed.

That the biology of Malpelo would be poorlyknown should surprise few people. The islandcouples geographic isolation with hazardous acces-sibility. It also lacks spectacularly unique denizenssuch as Komodo dragons or giant tortoises. Thusinterest in Malpelo is primarily restricted to thededicated naturalist who can appreciate that thevery isolation makes it a laboratory of evolution,and who realizes that a species need not be thelargest or the strangest to be of genuine biologicalinterest. The joint Smithsonian-U.S. Navy Expedi-tion to Malpelo was not the first to visit this uniquecorner of the world; others have stopped on theisland, perhaps while enroute to the Galapagos, orwhile collecting marine life off the coast of SouthAmerica. Our expedition probably was the first tobe dedicated to the Malpelo ecosystem per se, toattempt to collect and identify as many of theliving forms as was feasible, to examine the be-havior and ecology of at least some of the species,

George C. Gorman, Department of Biology, University ofCalifornia, Los Angeles, California 90024. Terence L. Chorba,student in Final Honour School of Psychology, Philosophy,and Physiology, Oxford University, and member of BrasenoseCollege, Oxford, England.

and to attempt to understand the flow of energyin the system. The expedition will have been suc-cessful if, for no other reason, others are inspiredto visit this unique massive rock to pursue furthersome of the interesting problems raised in thesepages.

The first record of a landing on Malpelo thatwe know of took place in the late 18th century.The Spanish Commodore, Alejandro Malaspina(in Malaspina and Bustamente, 1885:551) quot-ing one of his officers wrote:

One of the shipmasters in the local trade, favoured by fineweather and a smooth sea, once landed on a shelf of rock onthe north face of the island, and after climbing thirty stepsdown hewn out by hand, came upon a large pool of rain-water, which was not overclean and had some bird feathersin 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 beensaid, but Malapsina either was not familiar withany previous mention of the island by the chroni-clers, or did not consider what they said to be ofsufficient importance.

Pedro de Cieza de L£on (1881:92) wrote thatthe Spanish colonial administrator in Peru, Cristo-bal Vaca de Castro, en route from Panama toBuenaventura in 1542 "came to a rocky islet whichthe sailors call Mai Pelo."

Literally, Malpelo means "bad hair." The originof this name is unknown but the island appears ona parchment map of the world of 1550 (Desceliers,1550) that is in the British Museum. The name onthe map is "ye mallabry." The term "malabrigo"was frequently used by cartographers to designateislands and bays. Literally, "malabrigo" means


shelterless. Such a name would be well deserved.Another possibility, however, is that the word"Malpelo" has roots in the Latin malveolns (liter-ally "inhospitable" or "spiteful"), which in thevernacular could have degenerated to "malbolo"and at some point been interpreted as "malpelo."

At the time that Vaca de Castro sailed pastMalpelo, though perhaps unknown to the Crown,the island was a territory of Spain by virtue of thePapal Bull issued in 1493 by Pope Alexander VI,and the Treaty of Tordesillas (1494) under whichSpain and Portugal divided the non-Christianworld in half. Subsequently, Malpelo passed auto-matically to Peru and later to Colombia withoutdispute. Inaccessible, uninhabitable, lacking arableland and suitable anchorage, it is not the kind ofreal estate for which one battles furiously.

One of the first scientists to discuss life on Mal-pelo was C. H. Townsend (1895), who visitedthe island in March 1891 on board the U.S. FishCommission Steamer Albatross. During the briefvisit, Townsend collected a Swallow-tailed Gull(Creagrus furcattis) and noted that seabirds ofseveral species swarmed about the inaccessiblesummit.

Townsend also collected the first reptiles withthe locality data "Malpelo Island." These lizardswere described as Anolis agassizi (Iguanidae) byStejneger (1900:163), who reported:

Mr. Charles H. Townsend, who collected these specimens inMalpelo, informs me that they were running over the rocksnear the water. The island was too steep to afford a landing,but the lizards were shot off or whisked off the face of thecliffs, thus falling into the water, whence they were securedby the collector.

Two years prior to the publication of Town-send's report, Faxon (1893) described the commonland crab of Malpelo as Gecarcinus malpilensis(Gecarcinidae) from a single male specimen col-lected by the same expedition in which Townsendcollected the lizards. No notes other than a descrip-tion were provided.

In November and December of 1927, CaptainA. G. Hancock made a trip to the Galapagos. J. R.Slevin of the California Academy of Sciences wasalong, and when the boat passed near Malpelo,Slevin and a colleague were able to effect a landingfrom a small skiff on the northeast corner of theisland. They collected 27 Anolis (a medium-sizedlizard) and 10 specimens of a large anguid lizard

which Slevin (1928) named Celestus hancocki. Heclaimed that this lizard was most closely related toCelestus millepunctatus O'Shaughnessy knownfrom a single specimen of unknown provenance,but that it differed from C. millepunctatus inseveral features of scutellation and limb develop-ment.

Slevin did not explore the island. All his speci-mens of Celestus were taken within a few feet ofthe water's edge on a rocky ledge. When wounded,he claimed that they would take to the water.Stomach contents consisted of feathers and remainsof crabs.

The George Vanderbilt South Pacific Expeditionof 1937 stopped at Malpelo in February. Some fieldnotes by the Vanderbilt's collector Ronald Smithwere published by Fowler (1938).

Smith was confused in his taxonomy, since hebelieved that there were five species of lizardsroaming about. But from his descriptions it is ob-vious that there were only the two known forms.Sexual dimorphism, and perhaps distinctive sizeclasses led to his erroneous conclusion. Since wecan easily determine when Smith was referring toAnolis, and when to Celestus, we might mentiona few of his ecological observations. He claimedthat the Anolis was found throughout the island,all the way to the summits, but that the Celestuswas limited to the height of 150 meters. He alsoreported a habitat difference. The anoles wereseen mostly amongst loose rocks and exposed sur-faces, whereas the Celestus were found primarilyin small cave-like formations. The lizards weredescribed as fearless and quite bold.

The Vanderbilt expedition collected four speci-mens of Anolis and seven Celestus, and this servedas impetus for Dunn (1939) to reconsider thelizard fauna of the island. He made two majordecisions at the generic level, one that has stoodthe test of time, the other that has not. Dunnpointed out that the genus Celestus could not bemaintained as distinct from Diploglossus. He as-signed the large anguid to Diploglossus, the genericname that it still bears. More interestingly, hedemonstrated that the previously described C. mil-lepunctatus was not distinct from Slevin's C. han-cocki. Thus the specimen described by O'Shaugh-nessy in the mid-nineteenth century was almostcertainly obtained from Malpelo, although no re-cords of an expedition to the island have been

NUMBER 176 11

found. The second decision was to raise the anoleto a distinct genus (Mariguana). This decisionhas not been accepted (Etheridge, 1960).

Bond and deSchauensee (1938) presented anannotated checklist of the birds of Malpelo basedupon the observations and collections of the Van-derbilt expedition and incorporating data fromTownsend (1895). It is reprinted below in itsentirety.

ANNOTATED LIST OF THE BIRDSOF MALPELO ISLAND

1. Sula dactylatra granti Rothschild. (Sulidae, Pelicani-formes) Masked Booby. Two adult males were collectedFebruary 8, and an immature female February 9.

2. Fregata sp. (Fregatidae, Pelicaniformes). None collectedbut Smith notes: "Man o'War Birds 50; small colony onone of the outlying rocks on the southwest of the island."

3. Falco peregrinus anatum Bonaparte. (Falconidae, Fal-coniformes) Duck Hawk. Not collected but two birdsobserved twice by Smith on Malpelo Island.

4. Actitis macularia (Linnaeus). (Scolopacidae, Charadrii-formes) Spotted Sandpiper. Not collected but "ten seenin company with tattlers."

5. Heteroscelus incanus (Gmelin). (Scolopacidae, Charadrii-formes) 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 wellas a nestling were collected February 8 and 9. Smithrecords a colony of thirty birds on the island.

7. Anous stolidus ridgwayi Anthony. (Laridae, Charadrii-formes) Noddy. An adult female, the only specimenseen, was secured February 9. It was not in breedingcondition.

8. Anous minutus diamesus (Heller and Snodgrass). (Lari-dae, Charadriiformes) Black Noddy. An adult male col-lected February 9, was the only specimen seen. The birdwas not in breeding condition.

9. Hirundo rustica erthrogaster Boddaert. (Hirundinidae,Passeriformes) Barn Swallow. An immature female wassecured February 8. Four birds were seen flying out ofa cave.

10. Progne sp. Martin. (Hirundinidae, Passeriformes). Re-corded by Townsend but none secured. Smith did not seeany martins.

During March 1941, the schooner Askoy ap-proached Malpelo and Dr. Robert CushmanMurphy and another member of his party made alanding. His popular article (Murphy, 1945) pro-vides the best summary of the natural history ofthe island. He estimated the booby population atabout 25,000; added an additional species to the

bird list of Malpelo (Red Billed Tropic Bird,Phaethon aethereus); examined the stomach con-tents of a gull and found that it was eatingmarine crabs; and, most important, noted the pres-ence of a diversity of invertebrates includingspiders, pseudoscorpions, beetles, flies, ants, andother insects. Samples of Malpelo rocks were ob-tained by Murphy and a study of phosphatizationat 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 crea-tures with shells of a ghostly white. They may be quiteharmless toward human beings, yet they seem to eye a visitorwith an intent that grows uncomfortable. If you sit longenough the crabs will move up closer and closer, as if withwhetted appetites, and I have a feeling that they wouldmake life miserable for anyone who had to sleep ashore.

Garth (1948) provided important informationon this crab. He did not agree with Faxon (1893)that it was an endemic species, but considered itG. planatus Stimpson, a species widespread fromBaja California to Acapulco, that has successfullycolonized the Revillagigedo Islands and Clipper-ton, but that is notably absent from the Galapagos.

There have been additional landings on Mal-pelo, some documented by specimens. For example,the San Diego Society of Natural History haslizards collected in December 1931 and January1933, by Cyrus Perkins. There is, however, littleadditional published information about the ecol-ogy and natural history of the island or its inhab-itants. What inspired the present expedition? Whytravel hundreds of miles to visit a barren rock?The very harshness of the environment raises inter-esting questions. In addition to sea birds, and afew nonmarine birds that are very strong flyers butpresumably not residents, only three common ter-restrial residents, Anolis and Diploglossus (lizards)and Gecarcinus (a crab) had been described andidentified. What species of plants (if any) andinvertebrates would be found? What are their bio-geographic affinities? How does such a communitymaintain itself?

Our working assumption was that this was a ter-restrial ecosystem dependent upon primary pro-duction on the sea and not on the land itself.Presumably the birds which fed upon marine lifeprovided food resources for small terrestrial inver-


tebrates (parasites, carrion eaters, egg predators,etc.) which in turn provided sustenance for thelizards.

Several of us were specifically interested in theAnolis lizards. The genus is comprised largely oftropical arboreal species. They tend to be highlyterritorial, defending fixed perch sites with elabo-rate displays. Here was an environment devoid ofthe typical plant cover that we associate withAnolis. Did they have any special physiologicaladaptations that permitted life on warm, exposedrocks? Was there any modification of the socialstructure of populations in the absence of typicalperch sites that are associated with their terri-toriality?

These are the kinds of questions that were onour minds when we first discerned the form ofMalpelo Island, gray itself against the gray of theearly dawn on the sea on 29 February 1972. Ans-wers are what we sought when we braved the steepswells and jumped off the landing craft to beginserious exploration of the island and its life.

Literature Cited

Bond, J., and R. M. deShauensee1938. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part II: The Birds ofMalpelo Island, Colombia. Proceedings of the Acad-emy of Natural Science of Philadelphia, 90:155-157,plates 9-11.

Cieza de Leon, Pedro de1881. Guerras Civiles de Peru: Guerra de Chupas. Ma-

drid: Publicado por Dom Pedro de Novo y Colson.[English translation by Sir Clements Markham andreprinted 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 Lizardsof Malpelo Island, Colombia. Notulae Naturae ofthe 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 SkeletalMorphology. Unpublished Ph.D. Dissertation, Uni-versity of Michigan. Ann Arbor: University Micro-films Inc.

Faxon, W.1893. Preliminary Descriptions of New Species of Crus-

tacea. Part 6 in Reports on the Dredging Opera-tions off the West Coast of Central America to theGalapagos, to the West Coast of Mexico, and inthe Gulf of California, in Charge of AlexanderAgassiz, by the U.S. Fish Commissioner Steamer"Albatross," during 1891, Lieutenant CommanderZ. L. Tanner, U.S.N., Commanding. Bulletin of theMuseum of Comparatix>e Zoology at Harvard Col-lege, 24:149-220.

Fowler, H. W.1938. The Fishes of the George Vanderbilt South Pacific

Expedition, 1937. Monographs of the Academy ofNatural 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 Pan-ama Bight. Bulletin of the American Museum ofNatural History, 92(l):l-66, 4 figures.

Malaspina, A., and J. de Bustamante1885. Viaje politico—Cientifico alrededor del Mundo por

las corbietas "Descubierta" y "Atrevida": desde1789 a 1794. Madrid: Publicado por Dom Pedrode Nova y Colson.

McConnell, D.1943. Phosphatization at Malpelo Island, Colombia. Bul-

letin of the Geological Society of America, 54:707-716.


Slevin, J. R.1928. Description of a New Species of Lizard from Mal-

pelo Island. Proceedings of the California Academyof 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. Bulletinof 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 Museumof Comparative Zoology, 27:121-126, 1 plate.

Reconnaissance and Mappingof Malpelo Island

A. Ross Kiesterand Jeffrey A. Hoffman

ABSTRACT

Daily observations of the exploration party are re-ported. Early maps of Malpelo show it to have onepeak with a maximum elevation of about 300 m.The exploration party surveyed and mapped theisland and found that the island has twin peaks,which are both approximately the same height andabout 150 m higher than previous maps indicate.An improved map of Malpelo is given. It is be-lieved that the Smithsonian exploration team isthe first to reach the island's summit.

Introduction

An overflight of Malpelo Island, arrangedthrough the courtesy of the U. S. Army, in prepa-ration for the U. S. Navy-Smithsonian Expeditionto Malpelo showed serious defects in the officialmap based on a 1954 aerial survey. The publishedU. S. Navy Hydrographic Office map (No. 1685,North Pacific Ocean, Isla del Coco, Malpelo Islandinset) showed a single major peak of 260 meters(845 feet) on the island. Observers in the overflight

reported two distinct peaks of nearly the sameheight, read as roughly 390 meters (1200 feet) onthe plane's altimeter. Initial observations from theship on arrival at Malpelo did not reveal which ofthe peaks was higher. They were so close in heightthat whichever peak was closest to the ship ap-peared the highest. A previous expedition had re-

A. Ross Kiester, Museum of Comparative Zoology, HarvardUniversity, Cambridge, Massachusetts 02138. Jeffrey A. Hoff-man, Smithsonian Astrophysical Observatory, Harvard Uni-versity, Cambridge, Massachusetts 02138.

ported reaching the summit of the rock, but hadnot indicated which of the two summits they con-sidered this to be. Indeed very little generaldescription of the topography of the island existedin the literature, and no mention was made of theexistence of the two peaks. Because of this con-fusion, a general survey of the topography of theisland was one of the expedition's scientificobjectives.

In pursuit of this objective, two members of theterrestrial party of the expedition extensively ex-plored the entire island, while other members ofthe party carried on intensive studies at one site.In this report we give a general description of theisland as obtained by the survey party, and discussthe mapping of the island.

Mapping Technique

As the party traversed the island, measurementswere made of the horizontal and vertical anglesbetween the various summits and landmarks. Allmeasurements were made with a marine sextantand a Brunton compass. Further surveying wasdone from the ship. Sextant measurement of theangles between the various features of the islandcan give the height of these if the distance to theisland is known. This method, however, limitedthe survey to the various peaks and ridges visiblein profile from the ship. An accurate knowledgeof the location of each feature on the island wasnecessary for the reduction of sextant sights. Whilethe ship was circling the island, azimuth bearingswere taken with a pelorus at regular intervals onall of the main features, including the north and

13


south tips of the island. An attempt to use gun-ranging radar to fix the distance from the islandwas frustrated by an uncertainty in determiningthe radar reflection point on the island. Lackingthis distance measurement, the position of the shipwas fixed by the intersection of the azimuth bear-ings to the tips of the island and to the "SouthPeak," which position was shown on the old map.Further surveying had to accept the island outlineand scale of the published Hydrographic Officemap. But these are less likely to be in error thanthe photographically determined contour lines.The intersections of the azimuth bearings to eachof the features from the various ship positions lo-cated each feature on the map. Heights weredetermined from vertical sextant measurements ofthe angles between the features and the waterline,and horizontal angle measurements of the widthof the island. An azimuth bearing to the SouthPeak combined with the horizontal angle fixedthe ship's position during this phase of the survey-ing. The linear heights of the features were thenobtained by solving a simple proportional equationand correcting for the height of the observer abovethe waterline of the ship.

the southern end. Beside the peaks the most con-spicuous feature of the island is a large arm ex-tending out on the west side of the island and form-ing a small bay (Figure 4). The bay is narrow andthe cliffs surrounding it are very sheer, giving afjord-like appearance. Several sea caves and archeswere also seen on the circuit. Besides the mainisland itself, there are several other rocks and seastacks in the vicinity. The two largest groups ofthese rocks are clustered to the north and to thesouth of the island.

A total of three days were spent exploring themain island and one day was spent exploring thelargest of the offshore rocks.

After the circuit of the island, a landing sitewas chosen on the south side of the end of the spurwhich forms the fjord-like bay on the west side ofthe island. This site became the base for most ofthe exploration and the location of most of theintensive studies of the terrestrial biology of theisland. This landing and study site was designatedLanding Site 1 (Figure 4). After landing the en-tire party and its equipment, the exploration teamstarted climbing up the spur. The vista up the

Itinerary of the Exploration Party

A preliminary idea of the topography of theisland was gained the first morning by making acomplete circuit of the island close into shore.This survey also provided an opportunity to pickout possible landing sites. The island is quite steepalong the sides and has a rather flat-topped appear-ance. It has three distinct peaks (Figure 4). At thenorth end of the island is the smallest of the threewhich was termed "North Peak." This summit issharply set off by nearly vertical walls all of theway around it. The top, however, appears to berelatively flat. This peak is separated from the nextby an east-west fault. This next peak south is cy-lindrical and butte-like, and is much higher thanthe first. It appears to have equally as steep sidesand is also flat on top. This peak was referred toas "Plug." To the south of it, separated by a hugeeast-west fault which split the entire island is amore rounded peak which appeared to be of thesame height as Plug. This summit was called"South Peak." From it the island slopes clown tothe south until it reaches the rather sheer cliffs at

North Racks

5 Stools

.""Shoolt

FIGURE 4.—Reference map of Malpelo showing the variouslanding sites (1-4) mentioned in the text, the routes takenby the exploration party, and significant landmarks.

NUMBER 176 15

spur toward the saddle between Plug and SouthPeak was a rocky slope completely devoid of vege-tation. Climbing up the spur the party found sev-eral caves of various sizes, including one whichwent through the spur from one side to the other.The only vegetation that could be found was algaewhich was growing inside of the caves and creviceswhere some moisture was present. After somescrambling, the saddle was reached. Here a cairnwith a bottle containing the names of several Co-lombians from two mapping and survey expedi-tions was found. In the saddle ttte lava was shotthrough with holes of varying diameter, makingmovement slow. From here the party ascended tothe summit of South Peak. Once on the summitthe party had a view down the entire south end ofthe island. This view corresponded exactly to thephotograph taken from the summit by the Vander-bilt Expedition of 1937 (Bond and deSchauensee,1938, plate 10B). Thus it was concluded that theVanderbilt Expedition had in fact climbed SouthPeak.

As sighted from South Peak, the Plug had anelevation of 1° to 1.5° horizontal, showing it to bethe true summit of the Malpelo rock massif. Assurveyed from the ship, Plug was determined tobe approximately 6 meters higher than the SouthPeak. The root mean square errors for the mea-surements of the two peaks from various ship posi-tions are about equal to the difference in heightof the two peaks. The 1° to 1.5° elevation of thenorthern peak as sighted from the southern corre-sponds to a height difference of 6-9 meters, whichagrees with the difference as determined by sextantsights from the ship.

From South Peak the party returned to the sad-dle and climbed up to the base of Plug. From thispoint the climbing became more difficult. Aftersome searching, a chimney on the east side of Plugwas found that afforded a relatively easy, but none-theless technical (class 5), climb up to the top.No bottles or cairns were found on this summitplateau. None of the previous parties climbing onthe island had mentioned the necessity of technicalrock climbing for the routes they used. Thus, theSmithsonian party seems to have been the first toreach the summit plateau of Plug.

As with the southern summit, Plug was devoidof vegetation. However, both species of lizards(Anolis and Diploglossus) were common up to the

top of the two peaks, as were the land crabs (Gecar-cinus malpilensis). Blue-faced Boobies (Sula dacty-latra) were also omnipresent. At the top of bothpeaks a strong breeze was blowing continuously incontrast to the still conditions lower down on theleeward side of the island. From Plug the partyclimbed back down to the saddle and thence downto the landing site.

On the second day the party landed on thenorthwest edge of the island (Landing Site 2, Fig-ure 4). While two members of the party workednear the shore, two others climbed towards NorthPeak. Initially the peak was attempted by climbingaround the west side to its north face, which hadseveral large chimneys. Also in the area is a spec-tacular spire isolated from the main peak. Theroute was frustrated because the chimneys wereseen to contain formidable chock-stones. On theless steep slope to the northwest of the NorthPeak were some of the few patches of grass to befound on the island. The largest of these patcheswas not more than several square meters in area.Retracing their steps the party then climbed tobelow 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 wereobserved. One of these was perhaps one-half meterdeep and was the largest body of standing waterseen on the island. The pools contained someaquatic insects which unfortunately could not becollected. Continuing up the gully into the faultproved difficult and roped climbing was necessaryto scale a 25-meter lava wall. From the saddle itwas a relatively easy scramble up to North Peak.Again both species of lizards and numbers ofboobies were present all the way up to the summit,and the wind was blowing steadily in contrast tothe leeward side. From here the party climbeddown along the east side of Plug and over thesaddle south of Plug and down to Landing Site 1.

On the last day of exploration of the main is-land, a party of two landed on the east side of theisland (Landing Site 3, Figure 4). This landingsite was used by a Colombian expedition whichhad placed a bronze plaque to mark the site. Theparty made its way up to the saddle south of Plugand then up to South Peak. From there theyclimbed down to the southern end of the island.The area southwest of South Peak is relativelythe flattest part of the island. The slope leading


from South Peak down to the end of the islandwas a scree slope with small rocks cemented bysolidified lava. Lava bombs were strewn aboutliberally, and all over the lava had solidified intocubist architectural forms. The south end of theisland was extremely dry. Few lizards were seen onthe central dome of the south ridge, but bothspecies as well as the land crab were abundant onthe extreme south end. There was a small patchof grass on the southernmost cliff. From the southend the party hiked back up over South Peak andthen back down to Landing Site 1.

On the fourth day, the party visited the largestof the offshore rocks (Landing Site 4, Figure 4).This rock is in the South Rocks group and like theother rocks it is quite steep. A landing was difficultand roped climbing began from the tiny shelvesnear the water. Near the top of the rock (45m =130 ft) the slope decreased appreciably. Rathersurprisingly, a large patch of grass was found here.This was the largest and lushest stand of vegeta-tion found anywhere. Anolis were common in thegrass 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 alti-tudes and land contours based on the surveying of the STRIexploration party in March 1972.

numbers. No land crabs appeared to be on therock. Several small bushes and a fern were growingat the top of the rock. The bushes frequentlyshowed large trunk masses with only small greenshoots. This was suggestive of large seasonal varia-tions in rainfall.

A Red-footed Booby (Sula sula) was sightedperched in the middle of the bushes. This specieshabitually perches in trees in contrast to the Blue-faced Booby, and has not previously been reportedfrom Malpelo Island. Positive identification wasmade when one of the members of the party wasable to climb down to within a few meters of thebird before it flew, and secure several good photo-graphs showing the characteristic red feet of thisspecies.

As with the landing, departure from the rockwas somewhat difficult due, in part, to the nearbyshoals.

Conclusion

The survey undertaken by the exploratory partyresulted in the map shown in Figure 5. As men-tioned above, the outline of the island and thescale are taken from the old Hydrographic Officemap, but the topography and contour lines arethe result of the new survey. It should be borne inmind that the map may not be entirely accuratedue to the rugged nature of the terrain, and tothe limited time and equipment of the surveyparty.

Future visitors to Malpelo should be able tomake substantial improvements in the map, partic-ularly with regard to geological details. Any exten-sive on-site surveying, however, could not be accom-plished without a good deal of technical climbing.We hope that this map will be of some help to anyfuture visitors.

Finally we remark that the rather dry descrip-tion given here in no way does justice to the sus-tained surrealism of this unique and unusualisland.

Literature CitedBond, J., and R. M. deSchauensee

1938. Zoological Results of the George Vanderbilt SouthPacific Expedition of 1937, Part II: The Birds ofMalpelo Island, Colombia. Proceedings of the Acad-emy of Natural Science of Philadelphia, 90:155-157,plates 9-11.

Field Observations onthe Geology of Malpelo Island

Jeffrey A. Stead

ABSTRACT

Malpelo Island had a volcanic origin. Depth sound-ings around the island indicate that it was once8 to 10 times larger than its present size. Marineweathering has eroded the island and has formedsteep cliffs and sea caves along its sides. Severaltypes of igneous rocks and minerals occur on theisland. The distribution of major joints and faultsis shown.

Published accounts of the geology of MalpeloIsland include McConnell's (1943) analysis ofthree rocks from the island and the field observa-tions 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 phosphatecompounds derived from guano) while this processwas very advanced in another rock (phosphaterock). The specimen of augite andesite was onlyslightly altered by phosphates. Murphy reportedthat guano was not very concentrated on the islandand McConnell (1943) concluded that phosphati-zation occurs mainly in porous areas of the island(along faults and fractures) where guano could becarried and concentrated by rains.

Malpelo is volcanic in origin, the present islandis the remnant of a much larger structure. Most ofthe island's perimeter consists of nearly verticalrock faces which extend from 60 to 230 metersabove sea level (Figure 6).

Jeffrey A. Stead, Defense Mapping Agency, Hydrographic Cen-ter, Rodman Office, Box 2023, Rodman, Canal Zone. Presentaddress: Hahnemann Medical College, Philadelphia, Pennsyl-vania 19102.

Several types of igneous rocks are present onMalpelo. Field identifications are as follows: da-cite, trachyte, tuff, basalt, and andesite. Up to analtitude of 210 to 240 meters, the island appearsto be mostly trachyte, with lesser amounts ofdacite and tuff. This lower zone is intruded by atleast two sets of dikes made up of basic rocks. Thetop of the island is covered by an andesite cap.

Field identification of minerals is as follows:Specimens of dacite collected near the landing sitecontained amygdules of quartz and apatite. Oneamygdule contained fine, hairlike, zeolite crystalscoated with hematite. Small amounts of chrysocollawere found in a joint at an altitude of about 240meters. Guano occurs in limited amounts on allparts of the island.

Fresh water seepage was observed in severalplaces. Red streaks of iron oxide are deposited onthe rock faces where seepage has occurred. Someof these streaks represent principal areas of runofffollowing rainstorms rather than permanent seepage areas.

Steep cliffs around the perimeter are probablymaintained by the combined effects of wave action,which loosens and removes rock in low areas(Steers, 1953), rain, and alternate heating andcooling, which loosen the soft lavas (mostlytrachytes) above the normal splash zone until theyfall off along joints. The lava cap is a very resistantrock, and probably does not fall until the rockunderneath it has been weathered away.

Sea caves (Figure 6) are formed by the scouringaction of waves on brecciated rocks around faults(Guilcher, 1958). There are many small faults onMalpelo, most of which intersect the waterline atlarge (60°-90°) angles. The largest cave wasestimated to be 9 meters wide and 13 meters high

17


(above sea level) and to have a 10-fathorn waterdepth at its mouth. Some of the caves have a whiteformation on their ceilings which resembles flow-stone. This is probably composed of insoluble sub-stances from guano, washed down along the faultsand redeposited on the ceilings of the caves.

Small amounts of soil, consisting of variablemixtures of clay and animal feces, are found insheltered areas of the island. Soil is found nowherein large amounts, but increases in abundance withincreasing altitude, and rarely occurs below 90meters. The absence of soil in the lower levels canbe attributed to steep slopes, rain, and to waveaction which, during severe storms, can splashwater to considerable heights (Johnson, 1919).

Two major sets of joints and faults were meas-ured in the fjord area (Figure 7). Their averagemeasurements were as follows: strike 210°T, dip79°NW; strike 110°T, dip 82°SW. The easternwall of the fjord is the scarp of a major fault, repre-sented by a cliff over 60 meters high and half akilometer long. This fault, which is the most promi-

nent feature of the fjord, strikes 2O5°T and dips78°NW. The largest sea cave on the island is atthe apex of the fjord (Figure 7).

The U.S.S. York County made continuous depthsoundings around Malpedo Island. These sound-ings, plus those from a 1952 U.S. Navy sketch sur-vey, are shown in Figure 7.

Bottom depth falls off greatly to 50-60 fathomson all sides of the island, but the bottom gradientincreases rapidly at 60-70 fathoms (Figure 7).The area out to the 60-fathom curve may representa wave-cut plateau formed during the Pleistocene(Shepard, 1963). If this is assumed, the 60-fathomcurve would indicate the minimum size of the origi-nal volcano. A line has been drawn (Figure 7) toindicate a depression in the sea floor. This may bea fault. Its trend, 218°T, is close enough to the210±T average of one of the major joint sets onthe island to strongly indicate a structural relation-ship. The dashed line trends 136°T, and also mayindicate a fault, or it may mark the division be-tween two previously existing cones.

FICI'RE 6.—The southwestern tip of Malpelo Island showing the lava cap, the steep cliffs, andnarrow 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 sketchsurvey (boldface numbers), those from the T.S.S. York County (lightface numbers) and depthprofiles around the island. The location of two submarine faults (218°T and 136°T) areindicated.


ACKNOWLEDGMENTS.—I thank D. Meyer of theSmithsonian Tropical Research Institute, B. Ma-curda of the University of Michigan, and R. Ste-wart of the Panama Canal Company, EngineeringDivision, for reading this manuscript and offeringsuggestions. Macurda assisted me with field workand Steward helped me interpret photographs ofthe 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. Bul-letin of the Geological Society of America, 54:707-716, 2 plates.


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 foundamong the terrestrial invertebrates of Malpelo Is-land. Except for the birds and one of the lizards,almost all of the animals depend on a purely ter-restrial economy based on numerous algae, lichens,and mosses.

Studies on the biota of isolated islands andgroups of islands have produced exciting results,influencing and even initiating the theory of evo-lution. Almost all existing islands have now beensurveyed, at least to some extent, but until 1972Malpelo remained a notable exception. Except forthe vertebrates no collections had been made, pre-sumably because it is remote and almost inacces-sible. The location of the island, however, suggeststhat it may contain species that are highly interest-ing biogeographically, especially because introduc-tions of species by man are very unlikely. Althoughsome landings have been made (Fowler, 1938;Bond and deSchauensee, 1938), very few peoplehave ever set foot on the island.

A number of seabirds, a land crab, and twospecies of lizards are known from Malpelo, butstatements in the literature about plants and in-vertebrates are not very encouraging. It is reportedthat there is very little (Bond and deSchauensee,1938) or no (Slevin, 1928) plant life that couldsupport a community of animals. In fact, Slevin(1928) reports that he could not find "any lifewhatever" except for the species mentioned above.Such a situation seems a priori very unlikely. Evenvery 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 avariety of species and all kinds of animals are ableto reach islands more remote than Malpelo (Carl-quist, 1965). Moreover, one would expect at leastone of the lizards (Anolis agassizi) to feed pre-dominantly on insects. Indeed, Dunn (1939) found"two or three species of insects" in stomachs ofAnolis. Bond and deSchauensee (1938) report anabundance of "small black gnats or flies" and achirping, as if by crickets. The most encouragingstatement is that of Murphy (1945), who, in thestomachs of Anolis, found "remains of small crabs,spiders, pseudoscorpions, beetles, flies and fly lar-vae, ants and other insects." The problem to be

><?

South

Rock

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

21


investigated is what animals occur on the islandand how do they make a living. The near-absenceof plant life reported in the literature suggestedthat the economy of Malpelo is entirely based onthe birds bringing in food from the ocean, theirdroppings, and dead bodies.

In the first week of March 1972, I was able totest this idea. My collections and observations weremade in localities indicated on Figure 8 and Iexamined some material collected from the SouthRock.

The general impression one obtains from themain island is that it is bare rock (Figure 9), withno higher plants except for very few small patchesof a yet unidentified grass. On the South Rock thisgrass was fairly abundant and here also a shruboccurred, possibly a legume, and a fern, Pityro-

FIGURE 9.—General view of site 1.

gramma dealbata (Presl) Tryon. On the mainisland, however, lower forms of plant life wereabundant. On vertical rock faces moist with freshwater and under ledges, algae, mosses, and lichenscould be found everywhere (Figures 10, 11). A fewpieces of rock were collected and eight species oflichens were identified from them: Caloplacasp., Candelbria sp., ?A.spicilia sp., ?Diploschistes sp.,Lecidca sp., Graphidaceae sp., fBasidea sp. andPyxine cf. glebosa Tuck. Considering the smallnessof the sample, the number of sj>ecies present maybe much larger than this.

This vegetation is the basis for quite a complexcommunity of animals. In crevices and, especially,under rocks, a multitude of animals was discovered.My collection probably does not cover all thespecies present, but the fact that mine was aboutas good as the anoles (see below) suggests that Imay not have done too badly. I did not make ex-tractions from soil under the rocks and any micro-arthropods living there would have been missed.The abundance and diversity of the invertebratesobtained (Table 1) is rather impressive. At least17 species of insects were found, belonging to 8orders. The "black flies" mentioned by Bond anddeSchauensee (1938) represent a new genus ofChloropidae and the crickets they heard belongto the genus Hygronemobius, related to H. liuradescribed from British Guyana (Gurney, pers.comm.). These two species were among the mostabundant insects, together with a little black ant(Pheidole sp.) and staphylinid beetles. The twospecies of snails were fairly common in clustersunder rocks at locality 1. There was an extra-ordinary variety of spiders, some eleven species ofAraneida, none of which were webbuilders.Worms and isopods were abundant under rocks atlocality 2, far above the intertidal zone.

It is much too early to start speculating aboutthe origin of the species found, but there are indi-cations that some of them may have come from theIndo-Pacific. After a preliminary investigation ofthe centipeds, Crabill (pers. comm.) suggests thatthe closest relatives of some of these may not beSouth American species. Representatives of theopilionid family Assamidae seem to be mostlyliving in the Indo-Malayan region (Brues, Melan-der, and Carpenter, 1954). The land crab and allthree species of lizards (including the new gecko

NUMBER 176 23

described by Huey, pp. 44-46 herein) are all foundon Malpelo only, but how many of the otherspecies are endemic is not yet known.

To depict the structure of the ecosystem onMalpelo, i.e., to construct a foodweb, roles had tobe assigned to each of the species concerned. Theclassification of these species as a herbivore, a pred-ator, or a scavenger is based on what is knownabout relatives living elsewhere and on some ob-servations. The classification of the staphylinidsas herbivores, for instance, was made because theywere very abundant and found mostly amongmosses. The beetle, Enochrus, is classified as a pred-ator, although the adults may be omnivorous. Theresults are included in Table 1 and the tentativefoodweb is illustrated in Figure 12. The landcrabGecarcinus malpilcnsis, was very common and wasfrequently observed nibbling on stones, presum-ably feeding on lichens and algae. Remains of in-sects were often found in their feces. Many of thesemay have been scavenged or obtained while eatingthe feces of lizards, but the crabs can also act aspredators. On one occasion I saw two crabs thatwere holding a male Anolis, which was still verymuch alive, although one of its hindlegs was al-ready eaten. A third crab was eating the tail.These crabs also feed on dead birds. They areopportunistic omnivores, but probably mostlyscavengers.

I had the opportunity to examine the gut con-tents of 38 individuals of Anolis agossizi from the

main island, plus five from the South Rock. Thesewere kindly given to me by A. S. Rand. Virtuallyall the items listed in Table 1 were observed, in-cluding some juvenile crabs. The only obvious ex-ception were the earthworms, presumably becausethey are living in soil under rocks and thus inacces-sible. It is certainly not because they are distaste-ful. When I removed the rocks, several anoles camein immediately and feasted on the worms. Theanoles from the South Rock also had a variety ofarthropods in their guts, but differed from thosefrom the main island in that they had their rec-tums filled with grass seed.

Both individuals of the gecko, Phyllodactylus,had eaten crickets, a few ants (probably Strumi-genys) and a lepidopteran larva, possibly Ereun-etis. The number of species per individual wasmuch smaller than that observed in Anolis. Ants(Pheidole sp.) and staphylinid beetles, both ofwhich made up the bulk of the diet of anoles, wereabsent in the geckos. These differences could beaccidental, but are probably real, Anolis beingdiurnal and Phyllodactylus nocturnal.

Nematodes, probably several species, were foundin 60 percent of the anoles, averaging three perindividual (range 1 to 14). The geckos had 11 and12 nematodes, respectively.

The large lizard, Diploglossus millepunctatus,has a quite different way of life. The feces that Iexamined 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.


by boobies (Sula dactylata granti), on fish leftoversafter the boobies had fed their young, and on freshbird droppings. Slevin (1928) and Dunn (1939)found crab remains in Diptoglossus' stomachs andGarth (1948) suggested that this lizard feeds onboth crabs and bird eggs. The latter may be true,but I have no evidence to support this idea.

The foodweb (Figure 12) shows two interestingfeatures. The first relates to the rather large varietyof predators, some 24 species, including two lizards.In contrast I found only nine species of herbivoresand eight scavengers. This suggests that among the

species living on Malpelo, an unusually high per-centage are predators, many of which may be gen-eralists rather than specialists. Two of the herbi-vores, the snails, seem to be eaten exclusively bythe lizards, not by any of the invertebrate preda-tors. These predators are all eaten by Anolis, butthe bulk of their food consists of herbivores. Moreexhaustive collections, made at different times ofthe year, are needed to determine whether indeedthe predators make up such a large proportion ofthe fauna. If this is so, it might have some bearingon theoretical considerations which state that the

TABLE 1.—List of invertebrates found on Malpelo Island with estimate of theirfood habit (H = Herbivore; P = Predator; S = Scavenger; Par. = Parasite; O =Omnivore)

OligochaetaNematodaGastropodaDiplopodaGeophilomorphaScolopendromorpha

DecapodaIsopodaChelonethidaAraneida

••

OpilionidaSchizomidaAcarinaThysanuraCollembola

OrthopteraHemipteraColeoptera

Diptera

Lepidoptera"

Hymenoptera

ChilenophilidaeCryptopidae

Gecarcinidae

OecobiidaeSelenopidaeAgelenidaeLycosidaeSalticidaeSegestriidaePholridaeAssamidae

Entomobryidae"

GryllidaeEnicocephalidaeHydrophilidaeStaphylinidaeScydtnaenidaeEphydridae

"SciaridaeChloropidae?PyralidaeTineidaeFormicidae

1 (?)sp.spp.2 spp.sp.Ribautis sp.Scolopocryptops sp.Cryptops sp.Gecarcinus malpilensissp.sp.sp.Selenops sp.spp.spp.about 3 spp.PAriadna sp.sp.Paramitraceras sp.sp.sp.sp.Lepidocyrtus caprelsiLepidocyrtus 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.

SPar.HHPPPOSPPPPPPPPPPPar.HSSHPPHPS

ssHHSPHP

NUMBER 176 25

number of predator species can never exceed thenumber of prey species (cf. Levins, 1968), exceptin some special circumstances (Stewart and Levins,1973). If the phenomenon is real, it could be thatsome of the necessarily simplified assumptions onwhich such theories are based are violated. It couldbe that predation by the lizards on the inverte-brates, prey and predators alike, is so intense, atleast during some times of the year, that competi-tion between the invertebrate predators is greatlyreduced, thus enabling more of them to coexist.In that case, the problem boils down to what limitsthe numbers of lizards. We found no evidence ofpredation by birds on the lizards, nor do the

anoles seem to limit their density through behav-ioral means, as they are nonterritorial (see Rand,Gorman, and Rand, pp. 27-38 herein). Their gutcontents did not suggest that they were experienc-ing food shortage, but in other seasons this couldbe different.

The other interesting feature of the foodweb isthat my original idea, that the entire economy ofthe island was based on the sea, may be wrong.Only the birds and Diploglossus depend completely,and the landcrab partly, on income from theocean. All the other species seem to depend on apurely terrestrial economy. I would predict that ifthe birds, which provide the link between the

TERRESTRIAL

GRASSSOUTH ROCK

AALGAE

LICHENS

MOSSES

V

HERBIVORES9spp.

V

V

PREDATORS22spp.

AFECES

CORPSES

SCAVENGERS

8spp.

V ?

V

NEMATODES

ANOLISPHYLLODACTYLUS

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 questionmarks 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 nutrientsfor the plants. For simplicity, some relationships are omitted. These include lizards and crabsproviding food for the scavengers and ticks being eaten by Anolis.


ocean and the land, disappeared, Diploglossuswould soon be extinct and the land crab wouldreach lower densities, but that this would havelittle, if any, effect on the rest of the fauna. Theonly condition under which this would not be trueis when the weathering rock would not provideenough nutrients for the plants to grow, so thatfertilization (e.g., phosphates) by the birds is ofvital importance.

ACKNOWLEDGMENTS.—I wish to express my grati-tude to all those who identified the specimens:Drs. M. J. Dibben and T. Esslinger of Duke Uni-versity for the lichens, Dr. D. B. Lellinger of theNational Museum of Natural History, Smithson-ian Institution, for the fern, Dr. L. Abele of theSmithsonian 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. King-solver, 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, foridentification of the items listed in Table 1.

I also gratefully acknowledge Drs. N. G. Smithand J. B. Graham of the Smithsonian TropicalResearch Institute and G. C. Gorman of the Uni-versity of California at Los Angeles for criticallyreading the manuscript.

Literature Cited

Bond, J., and R. M. deSchauensee1938. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part II: The Birds ofMalpelo Island, Colombia. Proceedings of the Acad-emy of Natural Sciences of Philadelphia, 90:155-157, plates 9-11.

Brues, C. T., A. L. Melander, and F. M. Carpenter1954. 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 His-tory Press.

Clapp, R. B.

1972. The Natural History of Gardner Pinnacles, North-western Hawaiian Islands. Atoll Research Bulletin,163:1-25,7 figures.



Fowler, H. W.

1938. The Fishes of the George Vanderbilt South PacificExpedition, 1937. Monographs of the Academy ofNatural Sciences of Philadelphia, 2: v + 349 pages,10 plates. [Malpelo, pages 5-6.]


Remarks on Carcinological Collecting in the Pan-ama Bight. Bulletin of the American Museum ofNatural 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 fromMalpelo Island. Proceedings of the CaliforniaAcademy of Sciences, fourth series, 16:681-684,plates 25, 26.

Stewart, F. M., and R. Levins1973. Partitioning of Resources and the Outcome of

Interspecific Competition: A Model and SomeGeneral 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 Is-land, and in places is extremely abundant. In onestudy area we estimated one lizard per two squaremeters. Social behavior contrasts markedly withother anoles in the apparent lack of territoriality,infrequent aggressive encounters, and reduced dis-play repertoire. There is a strong tendency forsocial facilitation—the species is aptly described as"curious." Home ranges are large, and lizards tendto move long distances during the course of theday. Resources such as food, water, and perch sitesmay be shared by many individuals. There appearto be no special adaptations to temperature ex-tremes nor to desiccation. Reproduction is appar-ently highly seasonal, as evidenced by the completeabsence of small juveniles in museum collectionsmade in December and January, and in our col-lections of February and March. In our sampleabout half the females were carrying eggs. Thespecies is larger than most solitary anoles (adultmales often > 100 mm) and markedly sexuallydimorphic for size (males larger). Adult malesfall into two discrete reproductive categories thathave morphological correlates. Some have blackheads and permanently erect nuchal crests—thesehave active testes. Others lack the nuchal crestand have spotted heads like the females—thesehave 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 An-geles, California 90024. William M. Rand, Department ofNutrition, Massachusetts Institute of Technology, Cambridge,Massachusetts 02139.

Introduction

Almost all islands in the Caribbean have beencolonized by one or more stocks of Anolis lizards.Patterns of distribution, ecological relationships,and behavior have been extensively studied inthese West Indian anoles by Ruibal (1961), Rand(1964), Gorman (1968), Schoener (1969), Schoe-ner and Schoener (1971), Lazell (1972), Williams(1972) and many others. The Pacific waters adja-cent to the tropical American landmass are rela-tively impoverished in islands, and only two areinhabited by Anolis. One is Cocos Island, and thesecond is Malpelo. Malpelo is more isolated thanany of the West Indies, which have been naturallycolonized by anoles, although strictly in terms ofdistance it is within the limits known for coloniza-tions by Caribbean Anolis (Williams, 1969; Rand,1969). Malpelo is also more devoid of vegetationthan any West Indian island. Caribbean anolesare mostly arboreal, in the strictest sense, the ex-ceptions usually perch on other sorts of vegetation(bushes, grass, cactus, etc.). There are a few anoles

associated with rocks, such as A. rimanim on boul-ders in Hispaniola and A. bartschi on cliffs aroundcave entrances in Cuba; but none is from an envi-ronment so devoid of higher plants as is MalpeloIsland.

Previous to the expedition reported here, ourknowledge of Anolis agassizi was limited, but thevery nature of the habitat suggested that thisspecies might be rather different in behavior andecology from better known anoles of the Antilles.The species was first described by Stejneger (1900:161-163) with a detailed account of external mor-

27


phology. The paper included a superb watercolorplate of an adult male. Stejnegar also providedthe few field observations available: "Mr. CharlesH. Townsend, who collected these specimens inMalpelo, informs me that they were running overthe rocks near the water. The island was too steepto afford a landing, but the lizards were shot off orwhisked off the face of the cliffs, thus falling intothe water, whence they were secured by the col-lector." Slevin, in 1927, landed briefly on Malpeloand collected a series of A. agassizi, but he did notexplore the island.

Ronald Smith of the George Vanderbilt SouthPacific Expedition of 1937 did explore the islandand found anoles all the way to the summit. Hedescribed them as fearless and quite bold (Fowler,1938). Dunn (1939) utilized the Vanderbilt Expe-dition specimens to reconsider the status of thelizards of Malpelo. Because of peculiarities, par-ticularly a small throat fan, he erected a new,monotypic genus for agassizi. This he called Mari-guana. He reported that the agassizi stomachs con-tained small insects of two or three species. Heremarked that the unusual color pattern (see be-low) was similar to that of the other totally unre-lated lizard species on the island, Diploglossusmillepunctatus.

Etheridge (1960) considered the relationshipsof all available species of anoline lizards includingagassizi. He showed that this species fit squarelyinto the latifrons species group of Anolis, and hetherefore did not recognize the validity of Mari-guana. The latifrons series is an osteologicallyprimitive group of anoles otherwise restricted tothe southernmost of the Lesser Antilles and thetropical American mainland from Costa Ricasouthward. Anolis agassizi is virtually indistin-guishable from the Lesser Antillean members ofthe latifrons series on osteological characters alone,but genetic similarity evidence (Webster, p. 50herein) implies that it is not particularly close toeither the island or continental species examined.

Compared with West Indian anoles, the Malpelospecies was known to be unusual but not uniquein several aspects of its biology. Anolis agassizi islarger than most anole species that occur on is-lands without congeners (Schoener, 1969:389),but it is somewhat smaller than Anolis ferreus ofMarie Galante. Anolis agassizi has a very reduceddewlap 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, con-sisting of minute light spots on a very dark groundcolor, but many species of Anolis on small islandshave unique color patterns.

We were able to verify these attributes and todiscover additional peculiarities particularly inthe social behavior of A. agassizi during our visitsfrom 29 February to 3 March 1972.

ACKNOWLEDGMENTS.—We are grateful to Drs.Thomas A. Jenssen of the Virginia PolytechnicInstitute and Ernest E. Williams of Harvard Uni-versity for reading and commenting on the manu-script. Travel funds for G. C. Gorman were pro-vided by a grant from the National GeographicSociety; partial support for A. S. Rand was pro-vided by National Science Foundation grant No.B019801.

Distribution and Numbers

Anolis agassizi were found everywhere on Mal-pelo from sea level to the highest peaks, and alsoon the only one of the small rocks south of the is-land that was explored. They were rare on thesouth plateau and uncommon on the black rocksin the lowest 3 meters.

Anoles seemed less common on the large smoothrock faces than on the more irregular rocky areaswhich provided more hiding places and probablymore food and water. They certainly did not avoideither the vertical cliffs or the more nearly hori-zontal edges. The concept of structural niche asdefined by perch dimensions (Rand, 1964) failscompletely in an environment such as Malpelo.

Home Range Size and Population Density

Our home range data are based on marked in-dividuals. On the first afternoon we marked all ofthe anoles that we could catch in an area of per-haps 20 m long by 10 m high (Figure 13). Wevisited this area on three succeeding days and re-corded the numbers of anoles seen, as well as theiractivities.

We marked 29 anoles. Of these, we saw 21 againduring the next three days. Of the six large adultmales marked, five were resighted. The markedlizards 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. Onelizard was seen about 150 meters away from whereit had been marked; however, another marked liz-ard was seen at virtually the same spot on threesuccessive days.

Anoles gathered from distances as great as 10 to15 meters at concentrated resources, such as theoranges and jellied candy ("Chuckles") that weoffered, and at water seeps. Home ranges are largeand, even for adult males, are widely overlappingand seem little defended. The long distance movedby one anole and the fact that several marked ani-mals were not resighted suggests that there is afloating population like those postulated forother anoles (e.g., Philibosian, 1972).

During our observations, and when lizards con-gregated at sliced oranges we had set out (p. 31),we made periodic counts of the numbers presentand 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 Anolisagassizi 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 asof marked anoles, and assuming that of the 29marked anoles, 20 remained in the area, then thetotal number of anoles in our area would havebeen about 120. With this small sample and thesemany assumptions, we hesitate to calculate con-fidence intervals. We estimate, however, that anoleswere concentrating at the oranges from an area ofperhaps 500 to 1000 square meters. This wouldindicate a population density of about one anoleper 5 to 10 square meters. Although not all areasof the island are equally habitable by Anolis, theremust be at least 100,000 Anolis agassizi on theisland.

Measurements were taken from 45 animals,which were preserved and deposited in the Muse-um of Comparative Zoology. This series containedfour 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 themales had enlarged testes, dark heads, and well-developed nuchal crests (SV x 97.6 [89-114] SD7.3). The remaining 13 had small testes, female-like coloration, and small, if any, nuchal crests(SV x 84.2 [71-99] SD 7.9). (The latter malesare discussed on p. 32).

Weights of these animals just before preserva-tion were found to be highly correlated with snout-vent length in each sex (r = 0.% for males, r =0.85 for females). The lower correlation for femalesis due primarily to their smaller size range. Thelinear regression equations for weight (W) onsnout-vent length (SV) are: males, W = — 40 +0.62 SV; females, W = - 1 8 + 0.35 SV. Only asmall number (13.4 percent) of the lizards capturedhad unbroken tails, and an additional 6.8 percentwere broken in capture. The sexes do not differ inthe percentage of tails broken (p. 32). In the 18lizards with unbroken tails, the tail was slightlymore 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 specieswhich live on islands without congeners tend tofall into a restricted size range. The largest thirdof the males usually have a snout-vent length be-tween 55 and 90 mm, and the females between 40and 60 mm (for any given species, the female is


always significantly smaller). Anolis agassizi is anexception. The largest third of the males averages105.4 mm SV, the maximum is 114 mm. The com-parable female average is 85.2 mm SV with a re-corded maximum at 87 mm. Thus A. agassizi isabout the size of the Lesser Antillean giants A.richardi and A. bimaculatus, both of which aresympatric with smaller species.

Two hypotheses come to mind to explain thelarger than predicted sizes of A. agassizi: (1) Therewas a congener on the island that is now extinct.This is unknowable, but unlikely. The very re-moteness of the island makes the probability of asecond colonization extremely low; and insertionof a new colonist onto a small, already occupiedisland is the exception rather than the rule forWest Indian Anolis, where competitive exclusionseems to prevail (Williams, 1969; Gorman andBoos, 1972). (2) The average prey size is larger onMalpelo than in the Antilles. This is simply notthe case (see below).

Any number of additional hypotheses to explainthe body-size anomaly could be conjured up (e.g.,selection to resist predation; intense sexual selec-tion), but we have only our ignorance of the his-torical factors that have acted during the evolutionof this species with which to discuss such hypothe-ses. Thus we resign ourselves to the observationthat A. agassizi is large.

Food and Feeding

FIELD OBSERVATIONS AND ANALYSIS OF STOMACHCONTENTS.—The stomachs of 43 anoles were ex-amined and the contents identified by H. Wolda,who collected invertebrates on Malpelo (seeWolda's paper herein). Most of the species of in-vertebrates that he found on Malpelo, exceptearthworms, 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. Theinsects included beetles, flies, true bugs, caterpillars,ants, crickets, and Thysanurians. Many of theseanimals are cryptic. They were found under rocksand in crevices. Anoles certainly do enter crevicesand holes in the rocks where they probably catchsome prey.

The most common items in the stomachs wereants. These occurred in all guts and in larger num-

bers than any other prey. The next most commonitems were beetles, both larvae and adults, whichoccurred in 72 percent of all guts. Together, bee-tles and ants made up the majority of the foodeaten and probably more than half of the biomass.

The food items were small, nothing longerthan 20 mm, few over 10 mm. The majority of theitems and probably of the volume was in the 5 mmclass.

This range of prey corresponds with our sub-jective impressions of what was available. We sawalmost no invertebrates longer than 20 mm exceptearthworms, which were perhaps too well concealedto be captured, and the large land crabs, whichwere too big to be taken.

As previously pointed out, A. agassizi is largerthan might be predicted on the basis of Schoener's(1969) study of single species anole communities.The feeding data, however, do not support the hy-pothesis that A. agassizi is larger because it feedson unusually large prey. No precise comparisonscan be made, but the food of A. agassizi is certainlysmaller than that of the smaller mainland speciesA. polylepis (Andrews, 1971), and on the sameorder as that of the smaller A. roquet, which isalone on the island of Martinique (Schoener andGorman, 1968).

Anoles made repeated efforts to catch a hooklesstrout fly which we presented to them. They jumpedinto the air for it when the artificial fly was dan-gled overhead. One suspects that they may gatherat a bird carcass to feed on the invertebrates whichalso gather there. In this situation, they may leapinto the air to catch circling flies.

A small series of anoles was taken from one ofthe south rocks, where grass and bushes grew muchmore abundantly than on Malpelo itself. All ofthese anoles had their lower intestines packed withgrass 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 particu-larly attracted to the color orange. Thus, theyseemed especially interested in a Kodak film pack-age and in an orange screw-cap on a container ofsuntan lotion. This led us to perform two verysimple field experiments.

NUMBER 176

The Orange Experiment: On several occasionsin and near the study area, we set a half of anorange on a rock. Anoles from the surroundingareas gathered at the orange to lick repeatedly atthe cut surface, and to bite, pull off, chew, andswallow, bits of the pulp (Figure 14). They alsolicked the peel, but did not persist in this. Theanoles were intially attracted by the bright colorof the orange, and persisted because of its taste.

We saw lizards come to the orange from as faras 15 meters across the rock face. Five or ten indi-viduals would feed at the orange itself at one time,while another 10 to 20 remained within a 2 meterradius of it. There was almost no aggression shownat the orange though a male occasionally spreadhis small throat fan or bobbed.

The anoles seemed attracted to the area of theorange by the activities of the anoles already there,as well as by the orange itself. On one occasion wesaw a succession of anoles drinking (1 to 4 at atime) from a small wet spot on the rocks. Again wehad the impression that these lizards were attractedby the activities of other individuals.

Anoles did not stay at the orange for more thana few minutes though the same lizard might havereturned several times in the course of an hour ortwo.

The "Chuckles" Experiment: To test the possi-bility that the anoles actually showed a color pre-ference in making initial feeding choices, we usedthe jellied candy called "Chuckles." A package of

c . \ >

FIGURE 15.—Anolis agassizi making a choice between twoequal sized pieces of "Chuckles."

this candy contains five colors (flavors): red,green, black, orange, and yellow. Comparisons ofdifferent colored candy were made by setting outpairs of equal sized bits of candy close together inareas where we had not conditioned the lizards tooranges. We noted which bit of candy was selectedby the first lizard to approach (Figure 15).

The results (Table 2) show that yellow andorange are equivalent and that they both arechosen more frequently than the other colors (p<0.01). Of the remainder, red and green are aboutequal, and both preferable to black, though thedifference is not significant (p>0.05). We hastento add that this appears to be a color preferenceand not a taste preference, for once the less opti-mal candy was licked by the lizard, it was immedi-ately eaten.

We cannot separate hue from brightness inthese color choices, but all candies presented wereconspicuous.

TABLE 2.—Candy color preferences of Anolis agas-sizi (numerator is the number of times "Colorselected" was chosen before "Other color in testpair"; denominator is the total number of trials)

Color selected Other color in test pairYellow Red Green Black

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

OrangeYellowRedGreen

3/7 3/35/6--

6/64/62/6-

6/86/65/64/6


The preference for orange and yellow seemsmost peculiar because there are certainly no fruitsof this color on the island. The only objects thatwe could find in this color range were the largeland crabs, and the anoles showed not the leastinterest in these. One possible explanation is thatthe anoles are attracted to and feed on the yolks ofbroken seabird eggs. Many boobies nest on Mal-pelo and broken eggs may be a frequent occurrenceat certain times of year.

Predation

The most conspicuous potential anole predatorson the island are the large lizards, Diploglossus,the land crabs and the birds. The seabirds couldcertainly eat anoles, but Neal Smith (pers. comm.)says this is unlikely. The peregrine falcon (Falconperegrinus) has been recorded on the island (Bondand deSchauensee, 1938), but it is normally a birdpredator. The land crabs were everywhere. Wesaw a small group tearing apart and eating a stillliving anole (see p. 23). How it was caught is un-known for the crabs are relatively slow movinganimals.

The Diploglossus appear to depend largely onthe exuvia of boobies and on land crabs for food,and they gather where an adult booby is feedingits young. They do occasionally eat anoles, how-ever, for a specimen in the American Museum ofNatural History collections had one in its stomach(C. Meyers, pers. comm.). The Diploglossus couldnot be induced to eat oranges, but one big malerepeatedly approached an orange that had at-tracted a large group of anoles, and the Anolishurriedly fled at each approach. Though this Dip-loglossus was much swifter than the land crabs,the anoles were even faster and easily evaded himby keeping about a meter ahead of him on therocks.

There is strong circumstantial evidence thatpredation is important. Approximately 85 percentof the anoles that we examined had their tailsbroken and regenerated. Given the low level ofaggressiveness that we observed and the fact thatfemales showed as high a precentage of breakageas males, the implication is that the breakage wasnot caused by interaction with conspecifics, butrather by predation attempts. Pianka (1970) hasargued for a correlation between frequency of

tail breakage and intensity of predation in lizardsof the western United States.

If, as it seems likely, crabs and Diploglossus arethe important predators of anoles, it is likely thatthey catch more small than large ones. The colorpatterns of Anolis are highly cryptic on the rocks,suggesting selection by a visually hunting predator.

Adult males with their permanently erect nuchalcrests and black heads are very striking and aremore conspicuous than females and juveniles.Adult males of many Antillean Anolis also aremore conspicuous than females, presumably be-cause sexual and social selection favors conspicu-ousness in males and the larger size of the malesreduces their predation and consequently the selec-tion against being conspicuous.

Temperature

An open question in our minds, before seeingthe anoles of Malpelo in the field, concerned tem-perature adaptations. This species lives on barerock near the equator. Would A. agassizi preferhigh body temperatures? Would it be able to resisthigh environmental temperatures?

Our short-term, relatively simple studies indicateconclusively that with respect to temperature A.agassizi is a rather typical anoline lizard. Fortybody-temperature measurements of active animalswere taken in the field with a Schultheis thermo-meter. The mean temperature, representing pooledsamples of several mornings and early afternoonswas 30.6°C (24-38°C) with a standard deviationof 1.8. We believe that this approximates the pre-ferred body temperature as, quite obviously, thelizards could easily have had higher body tempera-tures by sunning themselves briefly on exposedrocks. Conversely, by spending more time in cre-vices, lower body temperatures could have beenmaintained. The A. agassizi temperatures werevery similar to the Lesser Antillean species A. rich-ardi and A. aeneus (Schoener and Gorman, 1968),and comparable to, but slightly lower than, A.homolechis on Cuba (Ruibal, 1961), a filteredsun animal.

Anolis agassizi does not show a temperatureadaptation to particularly high or low tempera-tures. This is what one might expect from thestructure of the island, which receives a great dealof insolation but also provides a great many hiding

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places to escape the sun. The most extensive slopesand cliffs orient north and south so that large partsof the island are in total shade for either themorning or the afternoon. This is probably themost peculiar aspect of this anole's thermal ecology.The A. agassizi were active during most of the dayin the shade and were seen basking only brieflywhen the sun first reached them. Since we workedon the west side, there were large areas where wesaw anoles active in the early morning and whichthey abandoned fairly shortly after the sun reachedthem (see p. 35).

The situation is quite unlike that which is usualfor a tree lizard who seldom needs to move morethan a few inches to get into the shade. An A.agassizi must move several meters if it is to exploitmany of the rocks and cliffs. In the absence of fly-ing predators it does not need to worry about keep-ing close to shelter.

Anoles that live in areas of high insolation tendto have either enlarged scales or black peritonealpigment. Anolis agassizi has the latter. The wallsof the body cavity are pigmented, most heavily onthe posterior half. There is some pigment on theheart and on the last 1 or 2 cm of the intestine. Insome examined A. agassizi the entire dorsal surfaceof the lower intestine is pigmented.

To determine adaptations to high temperatures,a simple heating experiment was performed onanimals brought to Panama from Malpelo. Theseexperiments were conducted immediately uponour return, and the lizards could not have accli-mated to the new climatic regime. Individualsrepresenting all size classes were tested. In theseexperiments we suspended a bright lamp above acage that measured approximately 30 X 30 X 45cm. The cages were constructed of wood with twoglass sides. Two parallel series of experiments wererun: in one, the lamp was placed at a height of30 cm above the cage top; in the second, at 42 cm.Each individual was dropped into the cage, thelight turned on, and the response observed for 3minutes.

It has been observed that when an anole isdropped into a novel situation its typical first re-sponse is to "freeze." As it warms up under theseexperimental conditions, it starts moving aboutwith the onset of discomfort; finally it opens itsmouth and begins panting (Ruibal, 1961).

All lizards moved within 50 seconds with the

light source at 30 cm, and in less than two minuteswith the light source at 42 cm. The temperature ofonset of panting was not correlated with body size,and only slightly with the distance between heatsource and cage. With the heat source at 30 cm,ten lizards began panting at a mean body tempera-ture (MBT) of 34.8°C; with the heat source at 42cm, 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 ani-mals opened the mouth soon after being droppedinto the cage. In all cases where mouth openingpreceded movement, the time of the trial was veryshort and the body temperature was well belowthat of all other trials. To confirm that movementwas a response to the heat, a transparent heat filterwas inserted between the light and the cage. Ineight of 10 cases the lizard did not move at all formore than 2.5 minutes (trial was discontinued at3 minutes).

This temperature at which panting begins(pooled mean, 34.2°C) is considerably lower thanpublished data for several Puerto Rican species ofAnolis (Heatwole et al., 1969), and is between datafor two Cuban species (Ruibal, 1961), A. allogusof deep shade (30.3°C) and A. homolechis of fil-tered sun (36.2°C).

Quite clearly, A. agassizi is not specificallyadapted to high body temperatures. Because of thenorth-south orientation of Malpelo and the greatcontrast between sun and shade on each part ofthe island with the passage of each day, some ofthe social organization discussed below may be re-lated to the lack of temperature adaptation. Ananimal cannot afford to defend a restricted terri-tory if it is unliveable for half of every day.

Water and Drinking

Many anoles that occur in very arid micro-habitats (A. poncensis in Puerto Rico, A. auratusin Panama) tend to have enlarged dorsal scales.Anolis agassizi does not. This reinforces our im-pression that Malpelo is not as dry as the lack ofvegetation suggests. Lying within the doldrums andbeing of some height, Malpelo probably receivesfrequent rain during most of the year. Water prob-ably does not stand very long on the exposed baresurfaces of cliffs and rocks. But fresh water wasfound spottily over most of the island during our


visit, even though it did not rain. The volcanicrock of which the island is made is porous andthere are many small seeps, springs, and rock pools.We saw anoles drinking together at a small seepand it may be that much of their water comes fromsuch limited and shared sources.

Initial tests done by Paul Licht of the Univer-sity of California at Berkeley on captive A. agassizisuggest that they lose water under drying condi-tions much as typical Antillean anoles do, andhave no special adaptations either to prevent desic-cation or to tolerate water loss. Several specimenscollected on Malpelo immediately excreted fluidurine, implying no special need for water con-servation.

Reproduction, Sex Ratio, and Size ClassDistribution

Our collection on Mapelo leads to the conclu-sion that the lizards must be highly seasonal (or atleast periodic) in their reproduction, or that wewere guilty of very biased sampling, or both (theyare not mutually exclusive). The smallest lizardcollected was 41 mm (SV). This is well above thehatching size of 25 to 30 mm (p. 35). Thus, wefailed to find evidence of any really young lizards.It is often the case that Anolis are more secretiveas young juveniles and special efforts are neededto collect them, but we had enough people spend-ing enough time in prime lizard habitat to makeus believe that hatchlings were truly scarce, if nottotally absent.

One of us (GCG) examined the two large muse-um series of A. agassizi: the Slevin specimens in theCalifornia Academy of Sciences, collected in Dec-ember 1927; and the Cyrus Perkins specimens inthe San Diego Society of Natural History, collectedin December 1931 and January 1933. Slevin (1928)wrote of his trip to Malpelo, whereas no informa-tion is available on the two journeys made byPerkins. Both these series then were made at ap-proximately the same time of year (December-January) and ours was made in the first days ofMarch. The smallest lizards in these collectionswere 64 mm (San Diego) and 56 mm (CaliforniaAcademy). Thus nothing is known about the timeof hatching or hatchling ecology in nature.

Studies on the reproductive cycles of tropicalAnolis (Licht and Gorman, 1970; Sexton et al.,

1971) indicated that within a population some fe-males were likely to be reproductive (with anoviducal egg or an enlarging follicle) throughoutthe year, the percentage seemingly affected by rain-fall. Thus in wet months 100 percent of the fe-males might be scored reproductive, but in exten-sive dry seasons, reproductivity could approachzero.

The reproductive condition of the females onMalpelo also hints at seasonality. In our collec-tion, five females had oviducal eggs, the smallestfemale being 72 mm SV; two additional femaleshad enlarging follicles, these were both large speci-mens (86 and 87 mm). Without adequate sampleswe cannot determine the size at sexual maturity.Since our smallest reproductive was 72 mm, let usconsider this our cut-off point. There were sixfemales larger than 71 mm which were completelynonreproductive. Thus, in early March, about 50percent of the collected female A. agassizi (6 of 13)were nonreproductive. In the museum samples(December-January) six of eight females (75 per-cent) larger than 71 mm were nonreproductive.To determine whether there is true seasonality orwhether there is merely a low level of reproduc-tive activity parceled out throughout the yearwould take further collections of A. agassizi.

The males presented the most aberrant and un-expected findings. They could be divided intothree groups. There were large males with theobviously developed secondary sexual characters,e.g., black head and erect nuchal crest. Theselizards ranged from about 90 to 114 mm SV andin weight from 13 to 34 grams. There were alsosmaller males (71 to 85 mm SV and 7 to 13 g) thatwere sexually mature, but lacked the impressivesecondary sexual characters of their larger neigh-bors. In other words, they looked like females.This is not particularly surprising, for in otherstudies of Anolis' reproductive cycles (Licht andGorman, 1970) it has been noted that males arephysiologically sexually mature long before theyare fully grown and, in fact, before they could becategorized as socially mature, i.e., assuming thesocial role of an adult male.

But there was a remaining class of animals thatpuzzled us in the field. Six male specimens (75, 85.86, 90, 94, and 100 SV) lacked male secondarysexual characters. Of interest is that three of thesewere 90 to 100 mm long and were as large as some

NUMBER 176 35

black-headed males, yet all six had completelyregressed gonads, to the extent that we at firstlabeled them "intersexes." Microscopic examina-tion led to the conclusion that these were fullyregressed males. Nothing like this has been seenin studies of Caribbean Anolis. Although mosttropical anole species show a moderate seasonalcycle for gonad size, the testes are never completelyregressed; they are always obviously testes, and inmost species they produce sperm throughout theyear. Thus there is a testes weight cycle in MalpeloAnolis. Furthermore, the population rarely showsa large variation for testes condition, i.e., the malesare generally in phase with one another. In con-trast, on Malpelo, we found some males that ap-peared fully reproductive and others that appearedfully regressed.

The museum collections also had specimenswith fully regressed gonads. In the San Diego col-lection there were putative males of 69, 78, 79, 82,and 83 mm SV that were fully regressed, yet malesas small as 74 that were reproductive. There is noway to determine whether this is a seasonal phe-nomenon or whether it is a population phenome-non. The general lack of aggression that weobserved (p. 36) might be related to the largenumber of nonreproductives in the population.

Male Anolis are often more conspicuous thanfemales and more likely to hold their groundagainst an approaching collector. Thus they havegreater likelihood of ending up in museum jars.Our collection consisted of 27 males and 14 fe-males, a sex ratio of almost 2:1. A pool of our sam-ple with the museum collections (and restrictingourselves to animals 72 mm and greater) shows asex ratio of 62:27, favoring males. We find it hardto believe that this is simply sampling artifact, butserious demographic study would have to be under-taken to resolve the problem.

Reproduction in Captivity

Four females returned to Panama laid eggs, oneat a time. Time intervals (n = 18) between suc-cessive eggs varied from 4 to 27 days (x = 13.6).Eggs were kept under ambient climatic conditionsin Panama (which must be close to those on Mal-pelo) and 13 hatched with an incubation time of48 to 69 clays (x = 57.8). The hatchlings mea-

sured 25 to 30 mm SV (x0.42 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 relativelack of aggressive encounters. There appears to bea very strong tendency towards social facilitation,i.e., when an anole took the initiative to approachan object, others in the area were likely to followsuit. They also appear to have a strong tendencyto examine any unusual object, making it very dif-ficult to perform unbiased observation without ablind. Thus the anoles frequently made active ap-proaches to the observers, on several occasionseven climbing on the camera and tripod set up tofilm their behavior. We could do little to avoidthem—backing off is not recommended on Malpe-lo's steep cliffs.

We interpret the social facilitation as an adapta-tion to patchy but superabundant food sources.Thus it is likely that the carcass of a bird mayoften serve as a focus for feeding of many anoles ina given area. In the West Indies, a well-establishedmethod for attracting large numbers of anoles isto break open a termite nest. Under such condi-tions, large numbers will often gather and feedactively with little aggressive interaction. The en-tire social organization of A. agassizi seems to bebased upon this type of situation.

Home ranges were large and activity was notrestricted to a few preferred perches. The follow-ing anecdotal observations support the conclusionthat home ranges overlap widely with residentsshowing little aggression. Up to 8 anoles simulta-neously would lick an orange, with up to 15 withina meter of it. One morning, approximately a halfdozen lizards that had been active on a smoothrock ledge moved laterally about 10 meters to acliff face. This occurred about a half hour after thesun reached the ledge. Presumably, they wereabandoning a foraging area that had become toohot. Many then gathered to try to catch a danglingtrout fly without showing any aggressive inter-action. On one accasion, a group of about a dozenanoles assembled at our portable radio and climbedall over it, though it provided neither food norwater. After some moments, all drifted away. This


is an example of the social facilitation referred toabove.

H. Wolda reported that as he turned over rockslooking for invertebrates, he was followed by agroup of anoles that fed on the food that he ex-posed. The A. agassizi in a walk-in cage in Panamagather around anyone who enters, as if expectingfood.

Occasionally we saw aggressive interactions be-tween individuals, including chasing and somedisplay. The instances involved crested vs. crestedmales, crested vs. noncrested males, and noncrestedvs. noncrested individuals of unknown sex. But weemphasize that we saw remarkably little display incomparison to our experiences with West Indiananoles. The only display that we saw was a verysimple multiple bob. This varied somewhat inamplitude and it was given sometimes with andsometimes without the dewlap being spread. Thedisplay was usually seen in seemingly aggressivesituations, but was also occasionally given in asomewhat less vigouous form when a lizard movedand stopped.

The captive anoles in Panama have shown littlein the way of additional kinds of display. We haveseen males jaw fencing, jaw locking and biting oneanother. More elaborate displays have not beenseen, nor any other simple displays. Males in cap-tivity that are courting approach the female withthe simple multiple bob, followed by pauses. Thisis similar to courtship displays of other Anolisspecies. Anolis agassizi has a display repertoire thatis more reduced than is typical for the genus andappears more like a sceloporine lizard. This is prob-ably a secondary simplification.

In a large outdoor cage in Panama, adult malesshow evidence of having developed a rather com-plex dominance hierarchy.

On Malpelo, the anoles exhibited direct compe-tition for food. Frequently, when an anole hadworried off a bit of orange pulp or picked up apiece of "Chuckle" too big to swallow, immediatelyit would run off with it. Sometimes it was pursuedby several other lizards, including much smallerindividuals that would try to seize the food fromits mouth. In these cases there was very little ag-gression shown.

No anoles attempted to monopolize the resourceby chasing or displaying at others. This was trueeven of the first individual to arrive at a resource.

There was not an initial territoriality that subse-quently broke down under repeated attacks as hasbeen described for gulls by Drury and Smith(1968).

At the orange, when a new lizard arrived with arush, one already there might run off. In general,large anoles supplanted smaller ones, but no or-ganized dominance hierarchy was evident.

Introducing anoles tethered to a long stick atvarious spots both close to and far from otheranoles produced a little interaction and in onecase an attack and bite. It also produced a littledisplay. The residents did not seem to be afraid ofthe intruder, nor of the way it was presented butin most cases they were just not very interested.

The social behavior in Anolis agassizi differsfrom that described for any other species of anole.This is most readily interpretable in terms of theatypical structural niche, viz., in place of discretedefensible perch sites (tree trunks, fence posts,etc.) there are large areas of cliff face that are ex-posed to complete shade or complete sun formany consecutive hours each day. In addition, foodresources are presumed to be much patchier thanin a typical tropical environment.

The basic difference in social organization ofthis species lies in the reduced overt aggressiveinteractions among individuals, and the high de-gree of social facilitation such that directionalmovements of one individual are likely to promptsimilar investigatory movements in several otherindividuals. In anthropomorphic terms, this speciesis extremely curious.

Both males and females seem to have overlap-ping home ranges. They do not defend territoriesor individual areas. There is a high degree of tol-erance for the close approach of conspeciflcs at acommon resource, be it food, water, or perch site.There are occasional overt aggressive interactionswhen "individual distance" appears to be viola-ted, but aggression is seemingly not associated withviolation of specific space (territory). Widely over-lapping home ranges without aggression have beenobserved 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 lesstolerant of members of the same sex.

Anolis agassizi not only has a reduced frequencyof display interactions but also seems to have a re-

NUMBER 176 37

duced display repertoire. This is in strong contrastto the anoline species in the West Indies (Gorman,1968) and to A. townsendi of Cocos Island (Car-penter, 1965). Our tentative explanation is: first,that, with reduced aggression and territorial de-fense, there are fewer bits of information to becommunicated; and second, that the striking, per-manently displayed color pattern and erect nuchalcrest of adult, reproductively active males assumemuch of the communication function otherwisetransmitted by complex movements of head, dew-lap, and tail (display action patterns). The onlyWest Indian anole known to have reduced displayis Anolis ferrens of Marie Galante which is like A.agassizi in being unexpectedly large and in havinga very striking permanently erect crest (in this caseon the tail) in adult males.

We would suggest that the reduced aggressionis a direct adaptation to living under conditionswhere resources are scattered so that a large homerange is necessary to include them. Furthermore,the resources are often so abundant that there isenough for all and consequently little advantagegained by attempting to monopolize them. Thereduced amount of display permits efficient utili-zation of common resources. The social facilitation("curiosity") aids in the discovery of these re-sources.

The social organization that we have describedfor A. agassizi is very different from that describedin 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 anolesto show a more or less similar pattern. In thesespecies the adult male maintains a small homerange centering on one or more preferred perches.The home range is defended against other adultmales as a territory and the resident gives frequentassertion displays. An adult female maintains astill smaller home range which she may defendagainst other females and against similarly sizedsubadult males. The territory of an adult maletypically overlaps those of one or more adult fe-males. All of the species showing this general pat-tern of social behavior have similarities in theirecologies. Evidence is accumulating from recentstudies 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 Anolisin social organization than had been realized. Wehave attempted to show in this paper that therather unusual social organization of A. agassiziis an adaptation to its unusual enviroment. Wepredict that A. agassizi will remain an extreme inthe continuum of social organization among anoles,but that intermediates between it and the morecommon types exist and, further, that the conti-nuum will be expanded in several quite differentdirections as species adapted to unusual ecologiesare 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 SouthPacific Expedition of 1937, Part II: The Birds ofMalpelo Island, Colombia. Proceedings of the Acad-emy of Natural Sciences of Philadelphia, 90:155-157,plates 9-11.

Carpenter, C. C.1965. The Display of the Cocos Island Anole. Herpeto-

logica, 21 (4):256-260.Drury, W. H., and W. J. Smith

1968. Defense of Feeding Areas by Adult Herring Gullsand Intrusion by Young. Evolution. 22 (I): 193-201.



Etheridge, R.1960. The Relationships of the Anoles (Reptilia: Sauria:

Iguanidae): An Interpretation Based on SkeletalMorphology. Unpublished Ph.D. Dissertation, Uni-versity of Michigan. Ann Arbor: University Micro-films Inc.

Evans, T. L.1938. Cuban Field Studies on Territoriality of the Lizard,

Anolis sagrei. Journal of Comparative Psychology,25:97-125.


Expedition, 1937. Monographs of the Academy ofNatural 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 Studyof Display Behavior. Breviora, 284:1-31.


Gorman, G., and J. Boos1972. Extinction of a Local Population of Anolis Lizards

through Competition with a Congener. SystematicZoology, 21:440-441.

Greenberg, B., and G. K. Noble1944. 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. Matta1969. 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 An-tilles. Bulletin of the Museum of ComparativeZoology, Harvard University, 143 (1): 1-115.

Licht, P., and G. C. Gorman1970. Reproductive and Fat Cycles in Caribbean Anolis

Lizards. University of California Publications inZoology, 95:1-52.

Pianka, E. R.1970. Comparative Autecology of the Lizard Cnemidoph-

orus tigris in Different Parts of Its GeographicRange. 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. Gorman1968. Some Niche Differences in Three Lesser Antillean

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

Schoener, T. W., and A. Schoener1971. 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. Licht1971. 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 CaliforniaAcademy 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 GenusAnolis from Cocos and Malpelo Islands. Bulletinof the Museum of Comparative Zoology, 34 (6): 161-164, 1 plate.

Williams, E. E.1969. The Ecology of Colonization as Seen in the Zoo-

geography of the Anoline Lizards on Small Islands.Quarterly Review of Biology, 44:345-389.

1972. The Origin of Faunas: Evolution of Lizard Con-geners in a Complex Island Fauna: A Trial Analy-sis. 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 ofDiploglossus millepunctatus

(Sauria: Anguidae)

A. Ross Kiester

ABSTRACT

The natural history of the large anguid lizardDiploglossus millepunclatus from Malpelo Islandis described. This species is characterized by itsabundance, flexible thermal regime, euryphagy,and curiosity.

Introduction

The large anguid lizard Diploglossus millepunc-tatus is one of the most conspicuous elements ofthe fauna of Malpelo Island. Because the islandis seldom visited little has been reported of itsnatural history. Here we begin to fill in some ofthe gaps in our knowledge of this species.

Diploglossus millepunctatus was first describedby O'Shaughnessy (1874) who listed the type-locality as "the north-west coast of America."Slevin (1928) described Celestus hancocki fromMalpelo Island, noting that it was similar to theunique type of O'Shaughnessy. Dunn (1939), whohad additional material, synonomized Slevin'sname on the basis of similarities of his series withthe type of millepunctatus and the fact that therewas some indirect historical evidence that the orig-inal specimen could indeed have been collectedon Malpelo. Dunn also placed the species in thegenus Diploglossus arguing that Celestus was notrecognizable. While Dunn's generic allocation ofthe species millepunctatus has prevailed, the issue

A. Ross Kiester, Museum of Comparative. Zoology, HarvardUniversity, Cambridge, Massachusetts 02138.

of the validity of the genus Celestus requires fur-ther attention (Myers, 1973).

Diploglossus millepunctatus is among the largestof the anguids, ranging up to 250 mm snout-ventlength with a tail length of 162 mm and a weightof 268 grams. It is rather heavy-bodied with shortlimbs. Older males possess the enlarged wide headsthat are characteristic of some other anguid generasuch as Gerrhonotus (Stebbins, 1954). The bodyhas a very shiny metallic appearance and like thatof other anguids is encased in a solid armor ofosteoderms. The tail tends to be rather blunt inadults but is more typically tapered in the young.The color pattern is dark black or brown-blackwith small yellow or yellow-white flecks in greatprofusion, as the specific name suggests (Figure16).

Fir.tRF. 16.—Diploglossus niillepurictatus aggregated about ayoung Blue-faced Booby (Sula dactylalra) which has justbeen fed.

39


ACKNOWLEDGMENTS.—I wish to thank T. L.Chorba of Oxford University, G. C. Gorman of theUniversity of California, Los Angeles, J. A. Hoff-man of the Smithsonian Astrophysical Observatory,A. S. Rand of the Smithsonian Tropical ResearchInstitute; and W. M. Rand of the MassachusettsInstitute of Technology for assistance in collectingdata, and R. B. Huey of the Museum of Compara-tive Zoology for aid in the preparation of thismanuscript.

Habitat

Diploglossus millepunctatus is found virtuallyeverywhere on the island from next to the water'sedge to the top of the summit (376.4 m). Lizardsusually did not occur far from the crevices intowhich they retreated. Otherwise, they scrambleover the bare rocks and cracks, sometimes movingwith agility in an almost serpentine fashion, butalso occasionally falling.

Although these lizards live next to the oceanand are capable of swimming, we were unable toget any individuals to enter the water voluntarilyas Slevin (1928) reported they would do.

Population Size and Structure

These animals are extremely common. No quan-titative measurements of population density wereattempted, but rough counts indicated that a den-sity of 1 per 10 square meters was perhaps a rea-sonable estimate. The population seems to consistmostly of adults. Of the literally hundreds of ani-mals that were observed only three young wereobserved. Except for these individuals, the entirepopulation fell into a size range of approximately180-260 mm snout-vent length. The juveniles wereapproximately 75 mm snout-vent length and noanimals of an intermediate size were seen. Thisstriking size distribution might possibly be due tothe fact that younger animals tended to remainhidden in crevices. However, the fact that someyoung were seen while no intermediate sized ani-mals were found seems to indicate that the obser-vations reflect a real phenomenon, namely thatreproduction is seasonal or erratic and that oursample was taken prior to a period of reproductiveactivity.

Activity and Thermal Relations

Although no nights were spent on the island, itseems likely that D. millepunctatus spends nightsinside crevices. Upon arriving on the island at 0800or earlier we observed few lizards out in the openbut saw many in crevices. Later, the lizards movedout to bask in the midmorning sun. By middaymost had retreated back into the crevices. Duringthe afternoon most remained in or adjacent tocrevices and only an occasional individual roamedin the open.

Overall mean body temperature (cloacal tem-perature ± se) of 12 D. millepunctatus duringactivity was 27.5 ± 0.64°C (range 24.8 to 32°C).However, body temperatures shifted upward be-tween morning and afternoon. The mean bodytemperature of six lizards active between 0838 and0910 was 25.8 ± 0.3°C, whereas the mean bodytemperature of six lizards active between 1324 and1342 was 29.3 ± 0.7°C. There was no overlap inbody temperature between these two samples, andthe difference between the samples is significant(p = 0.01, Kolmogorov-Smirnov test, Siegel, 1956:127).

Among anguids only Ophisaurus attenuatus isreported to have a higher body temperature (Fitch,1956:444). However, as with Anolis agassizi (p.33), D. millepunctatus appears in general to haveno special physiological adaptations to the ex-treme thermal environment in which it lives. Theready availability of the cooler crevices seems toafford D. millepunctatus a means of cooling itselfwhen the temperatures on the exposed parts of theisland are extreme. On the other hand, much ofthe island remains in the shade until the very latemorning, by virtue of the steep rock walls, andthis species can also be active at lower tempera-tures. Thus it appears to be flexible with regardto the temperatures at which it is active.

Drinking and Water Relations

Diploglossus millepunctatus drinks readily incaptivity. On Malpelo standing fresh water wasobserved in a few locations and water seepageswere seen in many of the crevices, although mostof the island was dry. It is impossible to know fromour observations what the seasonal pattern of wateravailability is, or whether there is water available

NUMBER 176 41

in the deeper crevices. Rock surfaces on Malpeloranged from very porous to extremely smooth andhard. In either case it seems that water would notstand on the surface of the island for any appreci-able time after a rain storm, either percolatingdown into the rock, running off rapidly, or evapo-rating. Although it did not rain during our visit,captive animals displayed an extremely rapid re-sponse to water. When kept in a large pen with noavailable water, the animals would respond to thefirst drops of water sprayed into the cage by run-ning immediately to the nearest puddle that wasforming. This response seemed to be about as fasta movement as the animals were capable of. Theyresponded by coming out of crevices or down offthe tops of large bricks that were in the cage. Thisrapid response appears to be a specific behavioraladaptation to a transiently available water supply.

Food and Feeding Behavior

Although this species will eat the usual kind ofprey items that are to be expected for a lizard ofthis size, such as insects and invertebrates, thebulk of the food taken by adults is quite out of theordinary for a lizard. There are small insects onMalpelo, and in captivity D. millepunctatus willreadily take crickets and mealworms. Actual feed-ing on insects on the island was not observed, how-ever, probably because the insects occur only insidethe crevices. Out in the open the lizards were ob-served to feed on a variety of bizarre foods. BothSlevin (1928) and Dunn (1939) reported finding

FICURE 17.—Diploglossus millepunctatus feeding on a fishregurgitated by the booby.

crabs in the stomachs of the animals they exam-ined, and Slevin also reported finding feathers. Weobserved this species feeding on carrion consistingof both dead crabs (Gecarcinus malpilensis) anda Blue-faced Booby (Sula dactylatra). In the caseof the booby, the lizards may have also been feed-ing on the insects attracted to the carcass.

Most of the feeding observed was connected withthe activities of living boobies. On two occasionsanimals partially consumed fresh booby feces. Themost spectacular feeding behavior occurs when anadult booby is feeding its young regurgitated fish.When an adult bird returns and alights near itsyoung, the young bird begins to squawk loudly.This noise attracts the attention of any nearby D.millepunctatus which immediately run to the vi-cinity of the two birds. If any of the fish is dropped,the waiting lizards immediately snatch the foodand drag it away. We observed this complete se-quence of behavior on two occasions, and sawlizards attracted to calling boobies on several others.In one case, more than seven adult lizards that hadaggregated about two birds successfully carried offand consumed an entire fish (Figures 16, 17). Dur-ing such activity the birds appear oblivious to thelizards. In captivity these lizards would readily eatcanned tuna or fish-flavored cat food.

It appears that the boobies provide a substantialamount of food for the lizards, at least during thenesting season. Obviously this source of food isnot available year round. The lizard's large stumpytail may possibly serve as a storage organ, andseveral of the individuals which we examined hadlarge fat bodies in the abdominal region. Presum-ably the young feed more on the insects that areavailable in the crevices and that may not be asseasonal as the supply of fish.

Social Behavior

In general this species does not appear to havea complex social behavior system. Individuals fre-quently ignore each other, although short chaseswere occasionally observed. The only activity thatcould be interpreted as a display is a particular"head-up" posture in which a lizard sits with itshead held back so that it points almost directlyupwards. Although two lizards within eyesight ofeach other could be observed to do this, no obviousreaction was observed. Further observations will


be needed to confirm that this is indeed a display.Social facilitation for feeding appears to be well

developed in this species. A running individualseems to attract the attention of others who fre-quently give chase. This reaction appears to be incontrast to the response elicited by a slowly movingindividual. When food or a source of food is pres-ent, it is not clear how much of the attraction isdue to the food itself and how much to the otherlizards; however, aggregation can be extremelyrapid. As mentioned above, seven or eight animalsaggregated about two boobies, and the aggregationtook place in a radius of over 5 meters in about 2minutes. Rapid aggregation in itself is evidence forsocial facilitation (Kiester and Slatkin, 1974), butthis point needs to be investigated experimentally.

One pair was observed copulating. They werefound in a crevice in the shade. The male held thehead of the female in his jaws, biting from theside. His body was positioned laterally and intro-mission took place from the side. The pair wasfound in copulation in the midmorning and re-mained in the same position for two and one-halfhours. The position is similar to that describedfor Gerrhonotus multicarinatus (Fitch, 1935:18).

Reproduction

A series of 18 specimens collected on a singleday during the visit (early March) showed markedreproductive activity. Females had many well-developed embryos, and males had large testes.Females brought to Panama and maintained incaptivity gave birth to live young two monthslater. Other females in this same group and somefemales taken to Cambridge, Massachusetts, whichinitially appeared gravid showed no embryos onexamination after their death some weeks later.Thus it may be possible that resorption of embryoscan occur.

We may conjecture that D. millepunctatus willhave its reproductive cycle tied to that of thebooby. From our collections we know that thelizards were in reproductive condition (and oftenfat) near the end of the fledging period of theboobies. If the amount of food available to theadult lizards is markedly increased by the appear-ance of nestling boobies which need to be fed, thenbreeding by the lizards might be expected to followthis period.

Conclusions

This species appears to be very opportunistic,ready to take advantage of a transiently availableresource on split second notice. Associated withthis opportunism is a high level of curiosity and atendency to respond to anything novel or disturb-ing in the environment. We found that individualsof D. millepunctatus would approach humanbeings or their artifacts with little regard for pos-sible danger. This curiosity seems linked to theunpredictable and meager food supply on MalpeloIsland where any disturbance may likely meanfood rather than danger for the lizards, since thereare no natural predators (with the possible excep-tion of migrating raptors; Bond and deSchauensee,1938). Thus the benefits of curiosity are greatwhile the disadvantages normally associated withthis behavior are reduced or absent. In this wayD. millepunctatus is typical of predators on remoteislands (Carlquist, 1965).

Literature Cited

Bond, J., and R. M. deSchauensee1938. Zoological Results of the George Vanderbilt South

Pacific Expedition of 1937, Part II: The Birds ofMalpelo Island, Colombia. Proceedings of theAcademy of Natural Sciences of Philadelphia, 90:155-157, plates 9-11.

Carlquist, S.

1965. Island Life. Garden City, New York: The NaturalHistory Press.



Fitch, H. S.

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

1956. Temperature Responses in Free-living Amphibiansand Reptiles of North-eastern Kansas. Universityof Kansas Publications of Museum of Natural His-tory, 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 Pan-

ama 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 Mal-

Collection of the British Museum. Annals and p e l o i s ia nd. Proceedings of the California AcademyMagazine of Natural History, fourth series, 13: of Sciences f o u r t h s e r i e s > 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 Amer-

ences. 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 fromMalpelo Island, Colombia, is characterized by ab-sence of tubercles on arms, legs, rear of head, andtail; absence of ear denticulation; absence of ab-dominal plaque; slight enlargement of dorsal tu-bercles, which form only two distinct paravertebralrows; terminal lamellae of digits distinctly widenedand truncate; 13 to 15 fourth toe lamellae; groundcolor medium gray-brown with dark chocolatebands. Source of this species is uncertain.

Introduction

The gekkonid genus Phyllodactylus is widelydistributed in the western hemisphere. Recentreviews by Dixon (1962, 1964a, 1964b) and Dixonand Huey (1970) have revised all New Worldmembers of the genus except species from theGalapagos Islands. Some 45 species are currentlyrecognized.

The collection of lizards from the Smithsonian-U. S. Navy Expedition to Malpelo Island, Colom-bia, included two specimens of Phyllodactylus, agenus not collected by the few previous expedi-tions to the island. Morphological characteristicsindicate distinctness of these specimens, which Ihere describe following the methodology of Dixon(1964a).

ACKNOWLEDGMENTS.—I thank J. R. Dixon ofTexas A & M University; G. C. Gorman of theUniversity of California at Los Angeles; J. B.Graham and A. S. Rand of the Smithsonian Tropi-cal Research Institute; A. E. Greer, A. R. Kiester,

Raymond B. Huey, Museum of Comparative Zoology, Har-vard University, Cambridge, Massachusetts 02138.

and E. E. Williams of the Museum of Compara-tive Zoology for helpful comments on the manu-script. I acknowledge the collectors of the type-series, A. S. Rand and H. Wolda of the Smithson-ian Tropical Research Institute.

Phyllodactylus Iransversalis (new species)

FIGURE 18

HOLOTYPE.—Adult female, Museum of Compara-tive Zoology (MCZ 130042), collected by A. StanleyRand from under a rock near the first collectingarea on Malpelo Island, Colombia, February 1972.

PARATYPE.—Immature specimen (MCZ 130043)collected from under a rock on Malpelo Island byHendrik Wolda, February 1972.

DIAGNOSIS.—Differs from the Central Americantuberculosus group, the Mexican delcampi group,and most South American groups by having smalland scattered dorsal tubercles forming only twoparavertebral rows; differs from unctus group inpossessing dorsal tubercles; differs from gerrhopy-gus group by absence of abdominal plaque; fromthe species microphyllus by having expanded ter-minal lamellae and a small nostril; and from in-aequalis in having dark chocolate dorsal bars, morefourth toe lamellae (13 to 15 vs. 10 to 12), smallerhead scales (third labial snout scales 32 vs. 20 to24; scales between eye and nostril 14 to 15 vs. 11to 12).

DESCRIPTION OF HOLOTYPE.—Rostral twice as wideas high, its dorsal edge almost straight with a dor-sal vertical groove one-half depth of rostral; 2internasals, somewhat rounded, their median edgesin broad contact, bordered posteriorly by 7 smallgranules and a postnasal on each side; nostril

44

NUMBER 176 45

surrounded by rostral, labial, internasal, and 2postnasals; first supralabial in narrow contactwith ventral edge of nostril; shallow depressionbetween internasals, slight depression in frontalregion; 15 scales on line between nostril and eye;scales in posterior loreal region slightly larger thanscales in mid-orbital region; 32 scales across snoutat level of third labial; 22 interorbital scales; eyelarge, its diameter contained in snout length about1.5 times; eyelid with 2 inner rows of granules, amedian row of larger scales, and 1 row of largescales on the edge, the last 8 to 9 are pointed; earlarge, its diameter contained in the eye diameter2.7 times; scales on anterior edge of ear opening

FIGURE 18.—Holotypc of Phyllodactylus transversalis, adultfemale, MCZ 130042, dorsal view.

flattened, overlapping, those on posterior edgesmaller and granular; rear of head granular with-out intermixed tubercles; 5 supralabials and 4 in-fralabials to a point below center of eye; mentalbell-shaped with posterior median edge sharplyangular, as wide as long, bordered posteriorly by2 postmentals; postmentals slightly longer thanwide, their median edges in broad contact, fol-lowed by transverse row of 5 flattened scales.

Dorsum with scattered, small, conical tubercles;only 2 paravertebral rows evident, extending ontoneck; paravertebral rows separated by 3 to 5 irreg-ular rows of small granules; 40 paravertebral tu-bercles between axilla and groin, separated by 0to 2 granules; remaining dorsal tubercles not indistinct rows; dorsal granules irregular; postanaltubercles 3 on each side, not distinct; enlargedabdominal plaque absent; venter with 27 scalesacross belly, from throat to vent 69.

Dorsal surface of upper arm with rounded,slightly elevated scales; forearm with slightlysmaller scales of similar shape; dorsal surface ofthigh and tibia with granular scales; claw long,visible from above and below; terminal toepadgreatly enlarged, longer than wide, truncate;fourth toe lamellae 13 to 14; tail partially regene-rated, original tail stub with a few wide medianscales on ventral surface, devoid of tubercles ondorsal surface, but with scattered, flattened tuber-cles at base.

Measurements (in mm): Snout-vent length 57,axilla-groin length 26, length of leg 25, length ofarm 17, length of tail 22 + 18, length of head 15.5,depth of head 5.7, width of head 11.2, length ofsnout 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-brownground color with white speckles; 8 distinct broadchocolate brown transverse bars from base of headto base of tail, somewhat broken along midline;width of bars slightly larger than ground inter-spaces; lateral area of trunk as dorsal ground; armsand legs with dark chocolate ground suffused withlight scales and small light spots, without definitepattern; top of head with suffusion of dark brownand small light blotches, no definite pattern; darkchocolate brown stripe from nostril to eye, blend-ing with head color posterior to eye, bordered be-low by a pale whitish line from first infralabialthrough ear; belly pale yellow.


VARIATION (based on paratype).—Similar toholotype except in the following characters: 14scales between nostril and eye; 24 interorbitalscales; 6 supralabial and 5 infralabial scales to apoint below center of eye; mental nearly triangu-lar; no contact between postmentals, followed pos-teriorly by transverse row of 6 flattened scales;venter with 30 scales across belly, from throat tovent 67; 39 tubercles in paravertebral row betweenaxilla and groin; fourth toe lamellae 15-14; tailregenerated 3 + 19; snout-vent length 36 mm.

NATURAL HISTORY.—Both geckos were collectedduring the day under rocks. Because of the topo-graphy of Malpelo, these geckos are necessarilyscansorial. Dixon and Huey (1970:66) have shownthat scansorial Phyllodactylus have expanded toe-pads, a characteristic of P. transversalis.

Two large diurnal lizards are also present onMalpelo (Diploglossus millepnnctatus and Anolisagassizi) and eat crustaceans as well as booby regur-gitations and feces. Ants and crickets were foundin the stomachs of P. transversalis (A. S. Rand,pers. comm.).

Accompanying papers in this volume describeMalpelo Island (Kiester and Hoffman, pp. 13-16)and the biology of the diurnal lizards (D. mille-punctatus, Kiester; A. agassizi, Rand, Gorman,and Rand).

ZOOGEOGRAPHIC RELATIONSHIPS.—The dorsal cross-banding and the two rows of dorsal tubercles ofP. transversalis are distinctive. These unique char-

acters suggest long isolation from source areasand hinder analysis of affinities.

Phyllodactylus transversalis shares many exter-nal morphological characters with the inaequalisgroup of Peru (Dixon and Huey, 1970:69). Afterexamining the type and paratype, however, J. R.Dixon (pers. comm.) suggested that P. transver-salis may be closer to certain Mexican species (P.paucituberculatus, P. unctus, and P. delcampi).There are no obvious affinities with Phyllodactylusfrom the Galapagos. The source of this gecko isthus equivocal.

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

Literature Cited

Dixon, J. R.1962. The Leaf-toed Geckos, Genus Phyllodactylus, of

Northeastern South America. Southwestern Natur-alist, 7:211-226.

1964a. Further Data on the Geckos (Phyllodactylus) ofIslands of the Extreme Southern Caribbean. South-western Naturalist, 9:203-206.

1964b. The Systematics and Distribution of the LizardGenus Phyllodactylus of North and Central Amer-ica. New Mexico State University Science Bulletin,64:1-139.

Dixon, J. R., and R. B. Huey1970. Systematics of the Lizards of the Gekkonid Genus

Phyllodactylus of Mainland South America. Con-tributions in Science of the Los Angeles CountyMuseum, 192:1-78, 14 figures.

Electrophoretic Estimates of GenieVariation in, and the Relationships of,

Anolis agassizi

T. Preston Webster

ABSTRACT

From data on proteins representing 30 genes, anaverage individual of Anolis agassizi is estimatedto be heterozygous at 2.1 percent of its loci. Thislow value is similar to that found in other solitaryanoles and in some anole populations of rela-tively recent origin, but it contrasts sharply withresults from some Greater Antillian species. Verylittle genetic similarity was found in the compari-son of A. agassizi to five congeners.

Introduction

An insular lizard population may have relativelylittle genie variation as a result of random pro-cesses and natural selection acting during its estab-lishment. The small size of the propagule, perhapsonly a single impregnated female, should limitvariation. This holds whether the alleles carriedby the founder or founders are a random or a se-lected sample of those in the parental population.Subsequently, random allelic loss (genetic drift)during the initial or secondary periods of smallpopulation size and selection would further reducevariation. A low level of heterozygosity could en-dure, but in a persistent population many newpolymorphisms might accumulate.

A considerable loss of genie variation may haveoccurred during the arrival and life of Anolisagassizi Stejneger on Malpelo Island. Malpelo mayhave received anoline immigrants only once and,as the source population is extinct, there certainly

T. Preston Webster, Museum of Comparative Zoology, Har-vard University, Cambridge, Massachusetts 02138.

have been no recent immigrants bearing new al-leles. On such a small island, secondary reductionsin population size are possible. The founders prob-ably encountered more extreme conditions onMalpelo than those to which they were adapted,even if the source population, like those anolesthat have been successful colonists in the Carib-bean (Williams, 1969), occurred in open, edgesituations. Anolis agassizi, however, is a long estab-lished population that has perhaps experiencedpressures and had time to reacquire heterozygosity.

In this study, genie variation in A. agassizi wasestimated by electrophoretic examination of enzy-matic and non-enzymatic proteins. Since there isno prediction to which this result can be compared,it is considered in the context of similar estimatesfrom other anole populations. Secondarily, manyof the same proteins were used to estimate thegenetic difference between A. agassizi and fivecongeners.

ACKNOWLEDGMENTS.—I thank the following fortheir contributions to this study: G. C. Gorman ofthe University of California at Los Angeles, T. A.Jenssen of Virginia Polytechnic Institute and StateUniversity, A. R. Kiester of the Museum of Com-parative Zoology, and A. S. Rand of the Smithson-ian Tropical Research Institute for collectingspecimens. J. B. Graham and A. S. Rand of theSmithsonian Tropical Research Institute, and E. E.Williams of the Museum of Comparative Zoologycommented on the manuscript. This study wasconducted in part while I was an NSF predoctoralfellow and was supported by NSF grant B 019801Xto E. E. Williams, Museum of ComparativeZoology.

47


Materials and Methods

Plasma and hemolysate were obtained from 32A. agassizi collected at the first landing site on thewest side of Malpelo. Six individuals were used forchromosome studies and preserved, and the re-maining 26 were homogenized in distilled water(1:2, W:V). Homogenates were centrifuged andstored at -80°C.

Electrostarch was used for horizontal starch gelelectrophoresis. Each protein was examined inas many electrophoretic environments as conven-ient and feasible. In several cases this multiplicityof observation increased the number of mobilitydifferences detected. Many of the buffer systemsand protein assays were those previously used instudies of Anolis (Webster, Selander, and Yang,1972; Webster and Burns, 1973). Additional tech-niques are those commonly used in studies of ver-tebrate proteins (Brewer, 1970; Shaw and Prasad,1970; Selander et al., 1971).

Twenty-eight enzymatic and nonenzymatic pro-teins were examined: leucine aminopeptidase, twopeptidases, fumarase, creatine kinase, phospho-hexose isomerase, eight dehydrogenases (lactate,two malate, two isocitrate, alcohol, a-glycerophos-phate, and 6-phosphogluconate), indophenoloxidase, two aspartate aminotransferases, two phos-phoglucomutases, an esterase, three "general pro-teins" (A,B,C), hemoglobin, transferrin, and threeplasma proteins. Like other anoles (Webster, Sel-ander, and Yang, 1972), A. agassizi appears to havefour or five genes encoding the polypeptides thatare subunits of hemoglobin, but it is treated hereas if it were under two-gene control. Lactate dehy-drogenase is encoded by two genes. Since each ofthe remaining proteins appears to be encoded bya single gene, the products of 30 genes have beenexamined.

Many of the proteins studied in A. agassizi werecompared with their apparent homologues in thesecongeners: 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 differ-entiation between members of the same speciesgroup, A. luciae was compared to A. extremus (St.Lucia: Vigie Beach, a recently introduced popula-

tion). Single individuals or pooled extracts wereused in these comparisons, but the variation ineach population has been studied and considered.

Genie Variation

In A. agassizi variation was detected at 4 of the30 loci. One of 26 individuals was heterozygousfor Protein B, 2 of 26 were heterozygous for one ofthe malate dehydrogenases, and 2 of 26 were heter-ozygous for one of the peptidases. Transferrin wasrepresented by 3 variants, with 14 of 32 individualsheterozygous. The remaining 26 genes appear tobe monomorphic. Thus the average individual ofthis population is estimated to be heterozygous at2.1 percent of its loci (H = 0.021).

This result can be evaluated within the contextof a broad range of estimates of H from anolepopulations (Table 3). Since a large proportionof protein polymorphism is expected (Lewontinand Hubby, 1966) and known (Boyer, 1972) to beundetectable by electrophoresis, such values aresurely underestimates of the actual extent of genievariation. Their comparison requires both that theelectrophoretically determined H be consideredas having some value as an index to actual genievariation among structural loci or within thegenome as a whole and that some standard erroror arbitrary criterion for difference be assigned.

From a joint consideration of morphologicalvariation and electrophoretically determined H,Soult et al. (1973) concluded that H has consid-erable power as an index to genie variation, evenwhen only 20 to 25 loci are examined. Two otherstudies support this relationship. Kluge and Ker-foot (1973) examined several sets of vertebratepopulations, finding that within each the averagevariability of a morphological character is corre-lated with the extent to which there has been diver-gence in its mean. Webster, Selander, and Yang,(1972) provided the complementary relationship,although their measure of morphological diver-gence was relatively crude. They found that amongAnolis species, those with higher values of H weredifferentiated into more geographic races. Thegenetic base for both morphological variation anddivergence probably involves many more genesthan the handful examined in the direct estima-tion of H. But whether these relationships aresimply 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 beit to other components of the genome is a matter considered indicative of less genie variation thanfor 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 ownlar 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. carolinensisSoutheastern U.S.

GREATER ANTILLEAN SPECIES

A. stratulusPuerto Rico

A. poncensisPuerto Rico

A. evermanniPuerto Rico

A. gundlachiPuerto Rico

A. krugiPuerto Rico

A. pulchellusPuerto Rico

A. porcatusHavana, Cuba

A. distichus dominicensis .Bon Repos, Haiti

SOLITARY SPECIES

A. extremusBarbados

A. roquetMartinique

A. luciaeSt. Lucia

A. marmoratusGuadeloupe

A. agassiziMalpelo

RELATIVELY RECENT POPULATIONS

A. carolinensisSouth Bimini

A. angusticepsSouth Bimini

A. distichusSouth Bimini

A. sagreiSouth Bimini

A. sagreiSwan 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.


Unfortunately, H has not been estimated forany of the South or Central American anoles.Whether A. carolinensis, a secondarily continentalspecies, has more genie variation than A. agassiziis questionable (Table 3).

Many of the Greater Antillean species representstocks long established on islands so large and bio-logically diverse that they probably have a conti-nental influence on genie variation. Estimates ofH have a broad range, but many of the values arehigher than those obtained from anoles in anyother evolutionary or ecological context. Sourceisland populations of two good colonists in theCaribbean, A. porcatus and A. distichus, appearto be more variable than A. agassizi.

Like A. agassizi, many of the species of the LesserAntilles have been long established and occur inthe absence of congeners. Their islands, however,tend to be larger than Malpelo and provide a moretypical anole environment. Estimates of H fromfour species are nonetheless similar to that fromA. agassizi.

The South Bimini and Swan Island popula-tions represent colonizations that are relatively re-cent (Williams, 1969). It is in such populationsthat any loss of heterozygosity during and soonafter founding should be most visible. The popula-tions of A. sagrei and A. angnsticeps on SouthBimini have little or no detectable genie variation,although the A. angusticeps have some esterasevariation that was refractory to interpretation andwas omitted from the reported H of 0.00. Else-where in the Bahamas and the West Indies, how-ever, these species have more variation (Webster,in prep.; Lister and Webster, in prep.; see Table3, A. sagrei, Swan Island). These results and thosefor insular populations of Peromyscus (Selanderet al., 1971) and cave populations of Astyanax(Avise and Selander, 1972) suggest that a majorloss of heterozygosity may frequently occur duringthe colonization of islands; however, the SouthBimini populations of A. distichus and A. caroli-nensis seem to have been less affected.

Since several Greater Antillean species appearto have as little genie variation as A. agassizi, thereis no compelling reason for concluding that it lostmuch during and subsequent to its colonization ofMalpelo. If it did, it is likely that both randomand selective processes played a role, but their rel-ative importance cannot be determined. It is

interesting to note that no variation was observedfor three esterases (only one of which was in-cluded in the estimate of H). These enzymes usu-ally show complex patterns of variation in anolepopulations, including some with little genievariation. That the estimates of H for A. agassiziand several other solitary anoles are fairly uniformsuggests that the net influence on genie variationmay be quite similar in these older populations.If this consistency is confirmed by further studiesof solitary species, it will be appropriate to con-sider selective explanations.

Genetic Differentiation

On the basis of a protein phenotype includingthe products of 24 genes, A. agassizi is quite distinctfrom all five of the species with which it was com-pared (Table 4). Two of the species are also mem-bers of the latifrons series of the alpha section(Etheridge, 1960): A. luciae (St. Lucia, LesserAntilles), like A. agassizi, has retained caudal auto-tomy, but A. squamulatus (Mainland, Venezuela)has lost it. Since the latifrons series is defined bycharacter states that are ancestral for the alphasection (Etheridge, 1960), it is not surprising thatthe species within it should have diverged to suchan extent. Even two members of the apparentlymonophyletic roquet species group (Gorman andAtkins, 1969), A. luciae and A. extremus (nativeto Barbados, Lesser Antilles), differ completely at17 of the 29 genes examined. Anolis singularis isas distinct from A. agassizi as are members of thelatifrons series; and the fourth alpha section species,A. marmoratus, has no shared electrophoretic mo-bilities. The beta section A. garmani is no moredistinct than some of the alpha section species.

These findings shed no new light on the affinitiesof A. agassizi. Except for very closely related popu-lations, electrophoretically determined genetic dif-ference is always an underestimate of genetic dis-tance. When the species compared are as distantlyrelated as these, it even becomes unreliable as anindex to genetic distance. Thus the small differ-ences in similarity that were observed in this studyshould be considered meaningless. As concludedpreviously (Webster, Selander, and Yang, 1972),when only 20 to 30 proteins are examined, electro-phoresis has little value in anoline taxonomy be-yond the species group.

NUMBER 176 51

TABLE 4.—Similarity (j*,; Nei, 1972) of eleclrophoretic mobilities of proteins incomparisons between Anolis agassizi and five congeners

Protein

aspartate aminotransferase-1phosphohexose isomeraseleucine aminopeptidaseisocitrate dehydrogenase-2lactatc dehydrogenase-1lactate dehydrogenase-2malate dehydrogenase-1malate dehydrogenasc-2ct-glycerophosphate

dehydrogenaseSum of Similarities

A.garmani

0.000.000.001.000.000.001.000.00

0.002.00

A.marmoratus

0.000.000.000.000.000.000.000.00

0.000.00

A.singular is

0.001.001.000.000.000.001.000.00

0.003.00

A.squamulatus

1.000.001.000.001.001.000.000.00

0.504.50

A.luciae

0.001.001.000.000.000.000.001.00

1.004.00

Values for the following are 0.00: 6-phosphogluconate dehydrogenase, aspartate aminotrans-ferase-2, peptidase, isocitrate dehydrogenase-1, two phosphoglucomutases, fumarase, indophenoloxidase, alcohol dehydrogenase, albumin, proteins A, B, and C.

Literature Cited

Avise, J. C, and R. K. Selander1972. Evolutionary Genetics of Cave-Dwelling Fishes of

the Genus Astyanax. Evolution, 26:1-19.Boyer, S. H.

1972. Extraordinary Incidence of Electrophoretically Si-lent 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 SkeletalMorphology. Unpublished Ph.D. dissertation, Uni-versity of Michigan. Ann Arbor: University Micro-films Inc.

Gorman, G. C, and L. Atkins1969. The Zoogeography of Lesser Antillean Anolis Liz-

ards: An Analysis Based upon Chromosomes andLactic Dehydrogenases. Bulletin of the Museum ofComparative Zoology, 138:53-80, 15 figures.

Kluge, A. G., and W. C. Kerfoot1973. The Predictability and Regularity of Character

Divergence. The American Naturalist, 107:426-442,10 figures.

Lewontin, R. C, and J. L. Hubby1966. A Molecular Approach to the Study of Genie

Heterozygosity in Naural Populations, II: Amountof Variation and Degree of Heterozygosity in Nat-ural Populations of Drosophila pseudoobscura.Genetics, 54:595-609.

Lister, B. C, and T. P. WebsterIn prep. Genie Morphologic Variation in Island Popu-

lations of the Lizard Anolis sagrei.Nei, M.

1972. Genetic Distance between Populations. The Ameri-can Naturalist, 106:283-292.

Selander, R. K., M. H. Smith, S. Y. Yang, W. E. Johnson, andJ. B. Gentry1971. Biochemical Polymorphism and Systematics in the

Genus Peromyscus, I: Variation in the Old-fieldMouse (Peromyscus polionotus). Studies in Genetics(University of Texas Publication 7103), 6:49-90.

Shaw, C. R., and R. Prasad1970. Starch Gel Electrophoresis of Enzymes: A Compila-

tion 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 VariationCorrelation. 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 ElectrophoreticallyDetected Sibling Species in a Haitian Lizard, Anolisbrevirostris. Evolution, 27:368-377.

Webster, T. P., R. K. Selander, and S. Y. Yang1972. 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 Zoo-geography of Anoline Lizards on Small Islands.Quarterly Rex'ieu' of Biology, 44:345-389, 17 figures.

Notes on the Chromosomes ofAnolis agassizi (Sauria: Iguanidae)

and Diploglossus millepunctatus(Sauria: Anguidae)

Brad Stamm and George C. Gorman

ABSTRACT

The karyotypes of Anolis agassizi and Diploglos-sus millepunctatus are very similar. When com-pared to other species in their respective groups,the karyotypes of the two Malpelo lizards are con-sidered primitive.

During the past ten years description of reptil-ian karyotypes has moved from relative novelty tovirtually routine. Chromosome preparations ofMalpelo lizards were made directly from testes fol-lowing methods described by Gorman, Atkins, andHolzinger (1967).

Both Anolis agassizi and Diploglossus millepunc-ctatus have kartotypes consisting of six pairs ofmetacentric macrochromosomes and 12 pairs ofmicrochromosomes, 2n = 36 (Figure \9a,b).

Among anguid lizards previously studied, thiskaryotype was found in Diploglossus costatus (theonly other Diploglossus studied to date) but notin the genera Gerrhonotus, Anguis, and Ophisau-rus (Bury, Gorman, and Lynch, 1969). McDowelland Bogert (1954) considered the diploglossinesthe most primitive of the anguids.

Similarly, representatives of the most primitivegroup of Anolis, the Lesser Antillean roquet group(alpha group, latifrons series; Etheridge, 1960),

Brad Stamm, Museum of Comparative Zoology, Harvard Uni-versity, Cambridge, Massachusetts 02138. George C. Gorman,Department of Biology, University of California, Los Angeles,California 90024.

have this same karyotype. Although many speciesgroups of Anolis show highly derived chromoso-mal morphology, a large section of the genus hasbeen conservative and retained the ancestral karyo-type (for review, see Gorman, 1973).

Anolis agassizi is also a member of the latifronsgroup. This species group has representatives onthe South American mainland as well as the LesserAntilles (the term roquet group follows Under-wood, 1959). The Malpelo anole and the roquetgroup all possess fracture planes in the caudal ver-tebrae, hence caudal autotomy is commonplace.This character appears to have been secondarilylost among living mainland forms. Thus the pre-sumed ancestors of A. agassizi and the Lesser Antil-lean latifrons anoles are extinct, and the livingSouth American anoles are more derived (Ethe-ridge, 1960).

Chromosomes tend to support this contention.Many members of the roquet group have a karyo-type virtually indistinguishable from A. agassizi.Only a single mainland latifrons member has beenpreviously studied, A. jacare, and it has a derivedkaryotype of 2n = 32, with a reduction in chromo-some number (Williams et al., 1970). For compari-son we add data on two other mainland latifronsspecies, A. frenatus from Panama, and A. squamu-latus from Rancho Grande, Venezuela. Webster,Hall, and Williams, (1972) argued convincinglythat within Anolis, 12 metacentric macrochromo-somes and 24 distinctly smaller microchromosomesis the primitive condition. Anolis squamulatus (notillustrated) 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 macrochromo-somes and 24 microchromosomes (pair 6 appears heteromorphic in this preparation, but we lacksufficient numbers of metaphase plates, and have no data on females to confirm this); B,Diploglossus millepunctatus $, 2n = 36, 12 ca. metacentric macrochromosomes and 24microchromosomes; c, Anolis frenatus $, 2n = 36, size break between pairs 4 and 5, macro-chromosomes do not break sharply from microchromosomes; pair 7 is intermediate in size.

saurian diploid number; but there is an enlargedmetacentric pair of "microchromosomes," and abreak in size between pairs four and five implyinga derived karyotype (Figure 19c).

The similarity between A. agassizi and theroquet group of Lesser Antillean Anolis in bothosteology and karyotype substantiates the beliefthat both retain primitive character states. We donot argue for common origin between these twogroups. The similarity in karyotype between theDiploglossus and Anolis of Malpelo is fortuitous.This chromosomal formula is common to repre-sentatives of numerous lizard families, often

among the most primitive representatives, as deter-mined osteologically (Gorman, 1973).

ACKNOWLEDGMENTS.—Travel to Malpelo by Gor-man was supported by a grant from the NationalGeographic Society. Laboratory work supported byNSF Grant B 019801X to Dr. E. E. Williams.

Literature Cited

Bury, R. B., G. C. Gorman, and J. F. Lynch1969. Karyotypic Data for Five Species of Anguid Lizards.

Experientia, 25:314-316.Etheridge, R.

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


Iguanidae): An Interpretation Based on SkeletalMorphology. Unpublished Ph.D. dissertation, Uni-versity of Michigan. Ann Arbor: University Micro-films Inc.

Gorman, G. C.1973. The Chromosomes of the Reptilia: A Cytotaxo-

nomic Interpretation. Chapter 12 in A. B. Chiarelliand E. Capanna, editors, Cytotaxonomy and Verte-brate EvohUion. New York: Academic Press.

Gorman, G. C, L. Atkins, and T. Holzinger1967. New Karyotypic Data on 15 Genera of Lizards in

the Family Iguanidae, with a Discussion of Taxo-nomic and Cytological Implications. Cytogenetics,6:286-299.

McDowell, S. B., and C. M. Bogert1954. The Systematic Position of Lanthanotus and the

Affinities of the Anguinomorphan Lizards. Bulletinof 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 Compara-tive Zoology), 121:191-226.

Webster, T. P., W. P. Hall, and E. E. Williams1972. 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 fromthe Andes of Venezuela. Brevoria (Museum ofComparative 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 describedin terms of the pattern of spatial coverage of sessileorganisms. The Malpelo subtidal is characterizedby a prevalence of large barnacles (Balanus penin-sularis) on steeply sloping rock walls and a highlystructured community of hermatypic corals ongradually sloping substrata, both communities ex-tending to a depth of about 30 meters. The rockwall communities of Malpelo have significantlymore free space available for recruitment (38.6%)than do comparable communities in continentalregions (7.3%). Size distribution suggests that re-cruitment is a greater risk, but conditions for adultsare better on Malpelo than in continental regions.Observations suggesting the importance of fishgrazing are discussed. An expanded depth rangeat Malpelo provides clearer zonation of the coralcommunity than along the Panamanian coast. Al-though coral growth is rich in terms of surfacecoverage and size of colonies, a true reef is notformed.

Introduction

The subtidal communities at Malpelo had neverbeen observed before our brief visit, and thus ourobjective was to describe these communities inquantative terms that would be useful for compari-sons to other areas and for any return visits tothe island.

We have utilized the relative surface area cover-age of macroinvertebrate species as the quantita-tive expression of community composition. Since

Charles Birkeland, David L. Meyer, James P. Stames, andCaryl L. Buford, Smithsonian Tropical Research Institute,P. O. Box 2072, Balboa, Canal Zone.

the Malpelo subtidal consists almost entirely ofvertical rock walls, surface area data were easilyacquired from underwater photographic transects,using color and black-and-white film. Relative sur-face area coverage was then estimated by tabula-tion of randomly located points on the projectedtransparencies and by planimetry (Tables 5,6,8).Photographs used for the quantitative survey areon file with C. Birkeland. We found that such amethod was most efficient for acquiring basic infor-mation from a large, unfamiliar, area within alimited time. Photographic transects have the fol-lowing additional advantages for primary survey ofan unexplored underwater area: First, size distri-bution of the predominant sessile organisms cansometimes be used to deduce aspects of the age orrecent history of a community, such as the fre-quency or heterogeneity of disruptive factors.Second, the extent of available unoccupied spacein a habitat is often an important limiting factorfor sessile organisms. Finally, the habitat or nur-sery of certain motile animals is often defined byspecific sessile organisms and, therefore, the sessileorganisms may provide the best definition of thecommunity as a whole (e.g., coral reef, oyster bed,kelp bed, sea grass bed, etc.).

Dives were made at each of eight stations (Fig-ure 20). At all stations except Station 6, the basal-tic rock wall sloped steeply, sometimes vertically,to a depth of about 37 meters on Malpelo itself orabout 49 meters on the offshore pinnacles (Stations4 and 5). Sometimes a talus slope of large bouldersprovided a more gentle slope in deeper water (be-low about 20 m; Stations 3, 7, and 8). Station 6 wasthe only area found with a gradual slope and itwas the only location of a rich community of her-matypic corals.

55


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 spongeyellow flat spongered flat sponge

•white flat spongewhite lumpy spongeblue spongegreenish gray spongegray-brown spongehydroidbryoz'oanLichenopora spPacifigorgia sppLophogorgia albagorgonaceanErrinopora pourtalesiiTubastraea aureaTubastraea aurea (dead)Pocillopora robustaPorites sppAstrangia spPavona variansOstrea irridescensOstrea irridescens (dead)Ostrea fisheriDidemnum candidumDidemnum spPolyandrocarpa tincta

2.9

3.3

500 350

1.117.8

5.316.822.111.64.9-

5.130.7

_-0.4_______

1.630.1

14.023.637.6

3.86.00.7

0.16.70.10.10.1__0.1____

4.634.0

5.88.7

14.51.05.21.2

0.81.11.60.51.60.30.5-_--_

5.224.6

1.86.68.4

18.82.2

_

6.00.2-6.2--

05--_

5.420.3

1.12.83.9

12.01.4

-

13.10.60.6

11.10.6--0.31.77.7

0.14.0___0.30.10.4__--0.1-7.20.4_

+10.00.6

0.32.40.50.99.0-0.53.0--1.73.3_

-2.6--7.60.60.2--1.4-6.4-0.85.41.40.8

—0.0-6.62.31.1-----8.0--2.6+_

• Predominant categories. An organism that occupies over 10 percent of the primary substratais arbitrarily denned as predominant.

+ indicates that the organism was present in photographs of the area, but it did not appearas 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 occupiedby macroscopic sessile organisms other than crustose coralline algae, comparingMalpelo Island and the continental coast of the eastern tropical Pacific (coverageestimated by number of rectangular coordinate points—taken from a randomnumber table—falling on the image of each category; 50 points tallied from eachquadrat photograph; several points in the continental photographs fell intoshadow so were not determined)

Region

Oceanic (Malpelo)Continental (Taboguilla Is.,

Panama and Playas delCoco, Costa Rica)

No. ofquadrats

56

Openspace

1081

213

Occupiedspace

1719

2705

Total no.of points

2800

2918

Mean percentof 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 duringdiving work at Malpelo by D. B. Macurda, Jr., ofthe University of Michigan and W. L. Smith ofthe State University of New York at Stony Brook.We thank D. Dexter of San Diego State University,and J. B. Graham of the Smithsonian TropicalResearch Institute for constructive review of themanuscript. The list of invertebrates was compiledwith the generous assistance of ten authorities.Maureen Downey of the National Museum of Nat-ural History, Smithsonian Institution, contributedher list of asteroids to the Appendix of this paper.John S. Garth of the Allan Hancock Foundationidentified the xanthid, dynomenid, and majid

crabs; and Janet Haig, also with the Allan Han-cock Foundation, identified the porcellanid anddiogenid crabs. L. G. Abele of Florida State Uni-versity completed the list of crabs and provided acomplete list for the decapods. Joseph Rosewaterof the National Museum of Natural History,Smithsonian Institution, confirmed and correctedthe lists of mollusks. Alan J. Kohn of the Univer-sity of Washington verified the species of Conusand Dora P. Henry, also from the University ofWashington, identified the barnacles. Peter W.Glynn of the Smithsonian Tropical Research Insti-tute confirmed and corrected our identification ofscleractinian corals. J. W. Wells of Cornell Uni-

TABLE 7.—Colony-polyp-number distributions in populations of Tubastraeaaurea at Malpelo compared with colony polyp number distributions in twoareas near the continental coast (counts were made from quadrat photographs)

Region

ATotal no.colonies

examined

BNo. solitary

polyps

CNo. colonies

with 10 ormore polyps

Mean no.polyps per

colony(±SE)»

Oceanic (Malpelo)Continental (Taboguilla Is.,

Panama and Playas delCoco, 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 3year sample of several mainland populations.


TABLE 8.—Zonation of the coral community of Malpelo Station 6: percent cov-erage by each species, bare rock, or dead coral substrata (data acquired by plani-meter measurements on tracings of projected photographs)

92 & 10.7Depth range (meters)

123 13.8 & 15.4 16.9 18.4 26.1 & 27.6

No. of planimeter samples

PERCENT OF SURFACE

Live hermatypic coralsDead coral substrata

Rock or sediment substrata

PERCENT OF LIVE CORAL SURFACE

Pocillopora robusta

Porites sppPavona clivosaAgariciella planulata

8 10 13

57.224.518.3

94.35.7_

69.327.6

3.1

80.519.5

_

74.822.7

2.5

14.477.3

8.3

89.210.8

—

2.443.254.4

58.629.811.6

80.519.5

42.718.038.4

47.4_

52.6

versity identified the hydrocoral and George Hech-tel of the State University of New York at StonyBrook identified the sponge.

500 Meters

o4

FIGURE 20.—Subtidal collection stations at Malpelo Island.

Rock Wall Communities

The steep rock walls (Figure 21) were predomi-nantly occupied by the large barnacle Balanuspeninsularis (Table 5). Sixty to 80 percent of theBalanus space was occupied by empty tests (Figure22) which remained on the walls after the death ofthe barnacles because their predators were snails(probably Thais planospira and possibly Purpurapansa) rather than sea stars. These tests provide amajor source of shelter or habitat for many poly-chaetes, mollusks, crustaceans, and fishes (Ento-macrodus chiostictus) and a substratum for hy-droids, sponges, and tunicates. The prevalence ofbarnacles is reflected in the white sand sedimentwhich encircles the island at the base of the rockwall. This sand is composed almost entirely of frag-ments of Balanus tests.

A clear zonation was observed on the verticalwalls (Table 5). To a depth of 3 meters, the largeblack sponge Polyfibrospongia sp. occupied over30 percent of the space. The brown alga Lobo-phora variegata was also prevalent above 3 meters.Both Polyfibrospongia and Lobophora decreasedrapidly in abundance between 3 and 6 meterswhere Balanus tests occupied nearly 37 percent ofthe 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 Pacifi-gorgia media (left), Porites sp. (right), and Balanus peninsularis.


very clear of "scuz" (filamentous algae, hydroids,etc.) and was covered mostly with crustose coral-line algae and scattered patches of anthozoans, i.e.,hermatypic corals (mostly Poriles spp.) an aher-matype (Tubastraea aurea), and gorgonaceans(Pacifigorgia spp.). Below 27 meters, filamentousalgae, small flat encrusting sponges, oysters (Ostreairridescens), and the violet branching hydrocorals(Errinopora pourtalesii) became prevalent. Al-though occupying only 7.6 percent of the substra-tum, the vivid violet hydrocorals appeared as a"field of Errinopora" below 27 meters. Certainmotile animals were also found only at depths of27 meters or below: the basket star, Astrodictyum

panamense, and the sea stars, Tamaria sp. andNarcissia gracilis.

The most striking aspect of the occupation ofthe subtidal substrata at Malpelo was the amountof "open" or "free" space. Table 6 compares theamount of open space in 59 permanent quadratphotographs from small Pacific islands less than14 km from the mainland in Costa Rica and Pana-ma with the amount in 56 survey photographsfrom Malpelo. Bare rock or space occupied bycrustose coralline algae is available as attachmentspace for other sessile or sedentary organisms. Be-tween the depths of 6 and 27 meters near the main-land, 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 = Balanuspeninsularis, 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 werecommon to continental and oceanic rock wall com-munities. The oceanic Malpelo populations forboth Tubastraea aurea and Lophogorgia alba ap-peared to be characterized by size distributions inwhich larger individual colonies were more preva-lent and less recruitment was observed in compari-son with the continental populations. The numberof polyps in each Tubastraea aurea colony in Mal-pelo photographs were counted and the polyp-number distribution (presumed to be a form ofage distribution) was compared with polyp-numberdistribution in Panama mainland quadrats (Table7). The distributions differ significantly by chisquare when the proportions of recruitment forsolitary polyps, colonies of 2 to 9 polyps, and colo-nies of 10 or more polyps are compared. The main-land data are from several areas through all sea-sons over a 3-year period while the Malpelo dataare from a single survey. The significant differencein age distribution could be the result of a particu-larly poor year, or time of year, in which the Mal-pelo survey was made, rather than to differencesbetween oceanic and continental populations. Ifthe solitary polyps are excluded from the analy-sis, the polyp-number distributions do not differsignificantly by Mest. Nevertheless, we predict thatmore data from further surveys would confirm ageneral trend towards proportionately less recruit-ment and longer adult life in oceanic populations.

A similar trend is evident for closely relatedspecies living on the island and mainland, respec-tively. For instance, Balanus trigonus (adults ca.0.8 cm basal diameter) is the prevalent species incontinental areas examined while Balanus penin-sularis (adults ca. 6 cm basal diameter) is the pre-valent species at Malpelo. The same generalizationcould be drawn comparing colonial tunicates ofthe genus Didemnum.

In summary, the oceanic rock wall communitiesat Malpelo had over five times as much open spaceas the continental rock wall communities. Sincespace is a limiting resource for sessile benthic or-ganisms on solid substrata, the abundant avail-ability of open space at Malpelo implies that re-cruitment is a more difficult problem for these pop-ulations. Once established, however, an individual

lives under less crowded conditions. In other words,an individual may be subjected to greater risk ini-tially, but may attain greater fecundity if it sur-vives until adulthood.

The reason for reduced recruitment success isunknown but two possible causes can be hypothe-sized: isolation from other populations and heavygrazing by fishes. Isolation may reduce recruitmentof shallow benthic organisms because most, per-haps 85 percent of the species (Thorson, 1961:456),have pelagic larval stages for which the mean dura-tion is about 2 weeks. Only about 30 percent haveplanktonic stages longer than 5 weeks (Thorson,1961:459). Malpelo is small, isolated, and sur-rounded by deep water. It seems likely that a largeportion of the larvae may be carried away beforesettlement can occur. The water moving past Mal-pelo comes predominately from the northeastwhere the nearest shallow water is around the Azu-ero Peninsula, 367 km away. The fauna of CobbSeamount, 430 km west of Washington State in thenorthern Pacific, has large populations of brood-ing species usually found only in the intertidal orshallow subtidal (Birkeland, 1971). Once a popu-lation of a brooding species is established on anisolated pinnacle, it may have a generally betterchance of maintaining a population than specieswith planktonic larvae. The emplacement of set-tling plates at Malpelo would provide particularlyinteresting quantitative data for comparison withdata from plates set nearer the mainland.

The sessile ciliate protozoan Folliculina wasfound on some of the Malpelo algae and it mayhave arrived by rafting on drifting algae. However,Scheltema (1973) has recently presented data show-ing that dispersal of this organism can also occurby attachment to the shells of veliger larvae. Fol-liculina is often found on the larger species ofveligers which are "long-distance larvae from shoal-water bottom-dwelling organisms of the shelf"(Scheltema 1973:123).

The effects of grazing fish in producing openspace is implied by some general observations con-cerning algae and is based on the assumption thatgrazers of algae also remove recently settled ben-thic animals. Algae and juvenile sessile organismswere very sparse on Malpelo (Tables 5, 6), yet theexceptionally clear water should provide favorablegrowth conditions for algae. As an example, onesmall sprig of Sargassum was found at 46 m. The


I

FIGURE 23.—"Shingle" formation of Porites sp. on the rock wall community.

NUMBER 176 63

substrata in shallower water presumably hadenough available light and other conditions forgrowth of Sargassum. The large schools of acanthu-rids may have kept the algae (and settling larvae)under heavy grazing pressure. This hypothesiscould be simply tested by the emplacement of fishexclusion cages for 4 to 12 weeks.

A species list from the collections and observa-tions made during the survey is given in the Ap-pendix. The most prevalent of all animals, Balanuspeninsularis (Table 5), is known from the Gala-pagos but not from the mainland. Since its testprovides a major source of shelter or habitat formotile animals and a substratum for sessile orga-nisms, the prevalence of B. peninsularis may be

the major factor contributing to the strong faunalsimilarities between Malpelo and the GalapagosIslands. Many familiar tropical eastern Pacificmainland species were conspicuous by their ab-sence: Muricea and antipatharians (the usuallypresent arborescent coelenterates), and the sea starsPharia and Phataria. Scarid fishes, hermit crabs,and gastropods were remarkably scarce.

Coral Communities

On the steeply sloping rock walk typical of theMalpelo subtidal, hermatypic corals occur onlyas a veneer interspersed with other growth (Figure22) or in some cases as shingled, overhanging

FIGURE 24.—Porites heads in the coral community; .Station 6, 18 m depth.


FIGURE 25.—Agariciella planulata; Station 6, SO m depth.

masses (Figure 23). Table 5 shows that the grandaverage surface area coverage by hermatypic coralson steep rock walls is only about 10 percent. Incontrast, a more gradual slope at Station 6 supportsan extensive coral community (Figure 24) extend-ing to 30 m depth. Table 8 shows that hermatypiccorals dominate the surface area coverage of thisslope. At 15 meters depth, a continuous growth ofPorites sp. measured 6.7 meters across by 2 metershigh. At 28 meters, a continuous growth of Agari-ciella planulata covered an area of 3 by 1.8 meters.Figure 25 illustrates that coral coverage at about30 meters is extensive and not merely confined toisolated colonies.

The rich development of hermatypic corals atStation 6 suggests that conditions for coral growthare highly favorable at Malpelo. The lack of coraldevelopment elsewhere around Malpelo may beprincipally due to two factors, both related to theprevalence of vertical rock walls around the island.Vertical surfaces are frequently shaded due to sunangle and substratal irregularities; thus, much ofthe light falling on them is reduced in intensity

depthin 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 separatingthem (r = Pocillopora robusta, s = Porites spp., c = Pavona clivosa, and v = Agariciellaplanulata).

NUMBER 176 65

FH.URE 27.—Nicks from fish grazing on Porites sp.

and quality by scattering or by passing obliquelythrough the water mass. By contrast, intensity onhorizontal surfaces at a similar depth would begreater. Calcification in corals is enhanced by thephotosynthesis of their symbiotic zooxanthellae(Goreau, 1959) and it is likely that the smallamount of coral noted along vertical walls at Mal-pelo is partly due to lower light intensities.

The second factor possibly contributing to thelack of extensive coral development at Malpelo issimply the lack of horizontal support on which

corals can build a reef. Figure 23 shows that coralsgrowing on the vertical faces tend to extend outinto the water mass to form a more horizontal sur-face which would better intercept light. This pro-cess can proceed only so far until erosion by bur-rowing animals causes collapse. Furthermore, sub-tidal talus piles and the eroding aspect of the cliffsabove suggest that rockfalls are a frequent occur-rence at Malpelo. Some talus appeared to be ofrecent origin, judging from the low numbers oflarge encrusting organisms. These rockfalls prob-


ably constitute a major disruptive phenomenonwhich prevents dense coral buildup.

Even at Station 6 the prolific coral growth wasnot developed above 10 meters depth, and it wasnot clear that the living corals were growing on apre-existing coral framework. We thus hesitate torefer to Station 6 as a coral reef. It is possible thatthe development of a true reef is prevented atMalpelo by other factors, one of which may betemperature. Surface temperatures were usuallyabout 26.5°C, but an abrupt thermocline wasformed at various depths. Temperatures below thethermocline were as low as 19.5°C and it has beensuggested that this is cool enough to limit coralgrowth (Glynn and Stewart, 1973).

To the casual observer, the pattern of coral zona-tion is more clearly expressed at Malpelo than incontinental eastern Pacific coral communities be-cause the greater depth range at Malpelo allowsthe lower zones to spread out over a larger area(Figure 26). For instance, Pocillopora spp., Pavo-na gigantea, and Pavona varians are all intermixedbetween 6 and 7 meters at Uva Island, near thecoast of Panama (cf. Figure 8 in Glynn, Stewart,and McCosker, 1972) while Pocillopora robusta,Pavona clivosa, and Agariciella planulata are sepa-rated by depth at Malpelo (Figure 26).

As on coral reefs in continental regions of thetropical eastern Pacific (Glynn, Stewart, and Mc-Cosker, 1972), signs of the effects of grazing byfishes are prevalent. The edges of the ridged exten-sions of the large Pavona clivosa colonies werelined with nicks scraped by browsing fishes. Fig-ure 11 in Glynn, Stewart, and McCosker (1972)is a typical illustration of this phenomenon. Oc-casionally, concentrations of nicks also occurredon sheets of Porites (Figure 27). Only three indi-viduals of the scarid fish, Scarus rubroviolaceus,were seen at Station 6 and none were observed atother stations. The tetraodontid, Arothron melea-gris, and balistids were common at all stations andprobably these fishes, rather than scarids, were re-sponsible for most of the damage to corals (Figure27).

Literature Cited

Birkeland, C.1971. Biological Observations on Cobb Seamount. North-

west 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 Condi-tions. Limnology and Oceanography, 18:367-379, 8figures.

Glynn, P. W., R. H. Stewart, and J. E. McCosker1972. Pacific Coral Reefs of Panama: Structure, Distribu-

tion and Predators. Sonderdruck aus der Geolo-gischen 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 CalciumDeposition 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 Wide-spread Geographical Distribution. NetherlandsJournal 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 byOcean Currents. In M. Sears, editor, Oceanography.American Association for Advancement of SciencePublication, 67:455-474, 3 figures.

Appendix

A List of Marine Invertebrates Identified fromMalpelo 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 VerrillPavona clivosa, VerrillPavona varians VerrillAgariciella 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 MenkeHipponix panamensis C. B. AdamsCrepidula aculeata (Gmelin)Capulus sp.Erato (?columbella Menke)Cypraea albuginosa GrayCypraea isabellamexicana StearnsCypraea teres pellucens MelvillSimnia aequalis (Sowerby)Cymatium pileare (Linnaeus)Colubraria ochsneri Hertlein & AllisonMuricanthns princeps (Broderip)Coralliophila sp.Coralliophila Qneriloides [Lamarck])Quoyula madreporarum (Sowerby)Thais planospira (Lamarck)Acanthina brevidentata (Wood)Purpura pansa GouldMorula lugubris (C. B. Adams)Anachis (?coronata [Sowerby])Mitrella sp. 1Mitrella sp. 2Fasciolaria princeps SowerbyConus diadema SowerbyConus dalli StearnsTylodina fungina GabbChromodoris sedna (Marcus)

Bivalvia

Area mutabilis (Sowerby)Barbatia reeveana (Orbigny)Septifer zeteki Hertlein and StrongLithophaga aristata (Dillwyn)Lithophaga plumula (Hanley)Isognomon recognitus (Mabille)Ostrea fisheri DallOstrea Qiridescens Hanley)

ARTHROPODA

Cirripedia

Balanus peninsularis PilsbryTetraclita stalactifer (Lamarck)Heteralepas quadrata (Aurivillius)Lepas anserifer Linnaeus

Decapoda

PALAEMONIDAE

Brachycarpus biunguiculatus (Lucas, 1849), circumtropicalHarpiliopsis depressus (Stimpson, 1860), Indo-Pacific-western

AmericaPseudocoutierea elegans Holthuis, 1951, western AmericaVeleronia laevifrons Holthuis, 1951, western AmericaPeridimenaeus hancochi Holthuis, 1951, western America

GNATHOPHYLLIDAE

Gnathophylloides mineri Schmitt, 1933, circumtropical (?)

ALPHEIDAE

Alpheus lottini Gue"rin, 1830, Indo-Pacific-western AmericaA. malleator Dana, 1852, western America, eastern & western

AtlanticA. grahami Abele, new species, endemic to Malpelo (see

paper by Abele in this volume)Synalpheus charon (Heller, 1861), Indo-Pacific-western

AmericaS. nobilii Coutiere, 1909, western AmericaS. digueli Coutiere, 1909, western AmericaS. biunguiculatus (Stimpson, 1860), Hawaii, Clipperton,

Malpelo5. 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, Mal-pelo

L. trisetacea (Heller, 1861), Indo-Pacific-offshore islandsPalinuridaePanulirus penicillatus (Oliver, 1791), Indo-Pacific-offshore is-

lands

PORCELLANIDAE

Petrolisthes haigae Chace, 1962, western AmericaP. glasselli Haig, 1957, western AmericaPachycheles biocellatus (Lockington, 1878), western AmericaClastotoechus diffractus (Haig, 1957), western AmericaPAGURIDAE

Pagurus sp., western America

DlOCENIDAE

Aniculus elegans Stimpson, 1858, western AmericaTrizopagurus 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 AmericaTrapezia digitalis Latreille, 1825, Indo-Pacific-western

AmericaT. ferruginea Latreille, 1825, Indo-Pacific-western AmericaMenippe obtusa Stimpson, 1859, western AmericaDomecia hispida Eydoux and Souleyet, 1842, Indo-Pacific-

western AmericaMedaeus spinulifer Rathbun, 1898, western AmericaPilumnus pyginaeus Boone, 1927, western AmericaGlobopilutnnus xantusii (Stimpson, 1859), western AmericaOzius perlatus Stimpson, 1859, western AmericaCarpilodes cinctimanus (White, 1847), Indo-Pacific-western

America


GRAPSIDAE

Grapsus grapsus (Linnaeus, 1758), western America, easternand western Atlantic

GECARCINIDAE

Gecarcinus mapilensis Faxon, 1893, endemic to Malpelo

MAJIDAE

Teleophrys cristulipes Stimpson, 1959, western AmericaMithrax pygmaeus Bell, 1835, western AmericaMicrophrys platysoma (Stimpson, 1859), western AmericaLissa tuberosa Rathbun, 1898, western America

ECHINODERMATA

ASTEROIDEA

Leiaster callipeplus FisherNarcissia gracilis malpeloensis, new subspeciesTamaria stria Downey, new species, endemic to Malpelo (see

paper by Downey in this volume)Mithrodia bradleyi VerrillNidorellia armata (Gray)

Asterope carinifera Miiller and Troschel

OPHIUROIDEA

Astrodictyum panamense (Verrill)Ophiocoma aethiops LiitkenOphiocoma alexandri LymanOphiactis savignyi (Miiller and Troschel)Ophiotheta mirablis Verrill

ECHINOIDEA

Diadetna mexicanum A. AgassizEucidaris thouarsii (L. Agassiz and Desor)Tripneustes depressus A. AgassizEchinometra sp.

CHORDATA

ASCIDIACEA

Polyandrocarpa (Eusynstyela) sp. aff. tincta (Van Name)Didemnum (Didemnum) sp. aff. candidum SavignyDidemnum (Didemnum) sp. aff. moseleyi (Herdman)

The Macruran Decapod Crustaceaof Malpelo Island

Lawrence G. Abele

ABSTRACT

Eighteen species of Macrura are reported fromMalpelo Island. Two new species of Alpheidae,Alpnens grahami and Synalpheus bannerorum, aredescribed and S. nobilii and S. digueti are rede-scribed. Notes are given on Periclimenaeus han-cocki, and Gnathophylloides mineri is recordedfrom the eastern Pacific.

Ecologically about half of the 43 decapods knownfrom Malpelo are commensals with other inverte-brates, mostly Pocillopora coral. Two species aresemiterrestrial or terrestrial and the remainingspecies occur in a variety of subtidal habitats.

Zoogeographically decapod fauna of Malpelo (43spp.) is most closely related to that of the westernAmerican mainland with 62 percent or 27 speciesoccurring there. Nine species (20%) occur in theIndo-Pacific region and seven of these reach thewestern American mainland. There are three en-demic species, two circumtropical species, and twospecies known only from offshore islands.

The very limited data available at present sug-gest a species-area effect on the shallow waterdecapods associated with oceanic islands.

Introduction

The first report of a decapod crustacean fromMalpelo Island was that of Faxon (1893) who de-scribed a land crab, Gecarcinus malpilensis, basedon material collected by the Albatross. This specieswas considered to be a synonym of G. planatus byRathbun (1918), but Turkay (1970) recognized

Lawrence G. Abele, Smithsonian Tropical Research Institute,P. O. Box 2072, Balboa, Canal Zone. Present address: Depart-ment of Biological Science, Florida State University, Talla-hassee, Florida 32306.

G. malpilensis as a species endemic to Malpelo. In1948, Garth reported on the Askoy collections,which included three species taken at Malpelo: anintertidal crab, Grapsus grapsus, the land crab,Gecarcinus malpilensis, and a pelagic portunid,Euphylax dovii, which was abundant in the watersaround the island. No additional material of deca-pod crustaceans appears to have been collected atMalpelo until the Smithsonian Tropical ResearchInstitute and U. S. Navy expedition visited theisland in 1972.

Several members of the expedition independ-ently made collections of decapods and sent themto various individuals for study. I was unaware thatboth Janet Haig and John S. Garth were studyingmaterial from Malpelo when I began studying thepresent collections. My study was then restrictedto the macrurans, the subject of the present report.A complete list of the decapods reported from Mal-pelo is included in the Birkeland et al. appendix(pp. 66-68, herein), and, therefore, it is unneces-

sary to repeat here.The macrurans collected at Malpelo Island form

a small but interesting collection of 18 species. Twospecies were previously undescribed, three speciesare recorded for the first time since the originaldescription, the genus Gnathophylloides is re-corded from the eastern Pacific for the first timeand many of the remaining species were previouslyknown from only a few records.

Collections of macrurans came primarily fromtwo localities on Malpelo Island referred to in thedescriptions as: "Malpelo coll. no. 3," which wasmade on the southeastern section of the islandamong coral in 10 m depth on 2 March 1972, and"Malpelo coll. no. 4," which was made on the westside of the island in 15 m depth on 3 March 1972.

69


"Malpelo coll." refers to general collecting aroundthe island from 29 February to 3 March 1972. Theabbreviation cl refers to carapace length excludingthe rostrum, tl to total length excluding the ros-trum unless otherwise specified and USNM to theNational Museum of Natural History, SmithsonianInstitution, Washington, D.C. All of the materialwill eventually be deposited in that institution.

ACKNOWLEDGMENTS.—I thank Jeffrey B. Grahamfor entrusting me with the study of this interestingcollection. Charles Birkeland supplied me with acollection of shrimps and the color slide of Panu-lirus penicillatus. The following three individualscontributed so much to this study that completionwould not have been possible without their aid:Dora M. Banner and Albert H. Banner of the Uni-versity of Hawaii, examined alpheid material andloaned material of Alpheus crockeri. Fenner A.Chace, Jr., of the Smithsonian Institution, hasonce again given his time and energy to the com-pletion of a study. He loaned material of Pericli-menaeus pearsei and compared material of Pseu-docontierea, Veleronia, and Gnathophylloides withtype and other material in the Smithsonian Insti-tution 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, 5ovigerous 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 22.7-28.7mm; ovigerous females tl 31.5-41.3 mm.

DISTRIBUTION.—This species is circumtropical indistribution occurring on most oceanic islands.

REMARKS.—All of the Malpelo material agreeswith the description and illustrations of B. biungu-iculatus, which has an extensive synonomy dis-cussed by Holthuis (1952). A second recognizedspecies of Brachycarpus, B. holthuisi, has beendescribed from deeper waters (30-60 m) off Brazilby Fausto Filho (1966). The present species occurs

in shallow waters in a wide variety of habitats: oncoral reefs, in tide pools, on grass flats, and amongsubtidal rocks (Chace, 1972).

Periclimenaeus hancocki Holthuis, 1951

FIGURE 28B-D

Periclimenaeus hancocki Holthuis, 1951:97, pi. 29a-k.

MATERIAL.—1 male, 1 ovigerous female, Malpelocoll.

MEASUREMENTS.—Male cl 6.0 mm, ovigerous fe-male cl 8.4 mm.

DISTRIBUTION.—The unique holotype was col-lected from Pinas Bay on the Pacific coast of Pana-ma. It is now reported from Malpelo Island.

REMARKS.—The male specimen agrees with thedescription of the unique male holotype given byHolthuis (1951). The lamella of the scaphocerite,however, does not extend as far beyond the lateraltooth as figured by Holthuis. The female specimendiffers in several respects. The rostrum is relativelyshorter than that of the male and it is armed withfour rather than five dorsal teeth. Holthuis hadonly the male holotype of P. hancocki availablewhen he noted certain differences between this

FICURE 28.—Periclimenaeus pearsei Schmitt: A, chela of firstpereiopod. Periclimenaeus hancocki Holthuis: B, chela of firstpereiopod; c, telson and left uropod of female; D, anteriorportion of carapace of ovigerous female. [Scale = 1 mm forA, B; 2 mm for c, D.]

NUMBER 176 71

species and the closely related species, P. pearseiSchmitt, 1932 from the western Atlantic and theseare listed in his key (1951:78). The present female,and to a lesser extent the male, bridge these differ-ences with one important exception: the form ofthe dactylus of the chela of the first pereiopods. InP. pearsei (Figure 28A), of which two males andtwo females (USNM 65113, 65114) were availablefor comparison, the dactylus is high and convexwhile in P. hancocki (Figure 28B) it is low and notnearly as wide. This distinct and so far constantdifference seems to warrant continued specific re-cognition of these two closely related species.

The eggs of both species are small (about 0.45X 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 ofAmerica from the Gulf of California to Colombia.

REMARKS.—Bruce (1970) examined the status ofPericlimenes pusillus Rathbun from Hawaii andconcluded that it represents a juvenile of Harpili-opsis depressus. Two color forms of this speciesoccur on Pocillopora coral heads in Panama andAustralia. Their status is being investigated by A.J. Bruce.

The translucent form of this species is one of themost abundant Pocillopora commensals in Pana-ma (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, 7ovigerous females, Malpelo coll.

MEASUREMENTS.—Males tl 7.1-7.3 mm, non-ovigerous 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, Galapa-gos Islands, Bindloe Island. It is now reported fromMalpelo Island.

REMARKS.—The present material differs in sev-eral respects from the description of P. elegansgiven by Holthuis (1951). All of the Malpelo speci-mens are smaller (tl 7.1-8.6 mm) than those listedby Holthuis (tl 9-16 mm). The lamella of the sca-phocerite is less produced in the Malpelo speci-mens than those figured by Holthuis. In the orig-inal description he notes that larger specimenstend to have the dactylus of the larger second pere-iopod robust with a large proximal tooth. Even thesmallest of the Malpelo specimens have this char-acteristic. The rostrum of the Malpelo materialseems to be shorter than Holthuis' material. TheMalpelo males lack a spine on the posterior mar-gin of the third pleuron, which is present in Hol-thuis' figure (see Chace, 1972:45). Dr. Fenner A.Chace, Jr., compared Malpelo material with para-types of P. elegans and noted that the Malpelospecimens seem to have the cornea set less obli-quely on the eyestalk and the general body formmore depressed.

Given, however, the relatively few specimensavailable from such widely scattered localities andthe variation known in other pontoniids, it seemsbest to refer the present material to P. elegansuntil more material is available.

Veleronia laevifrons Holthuis, 1951

Veleronia laevifrons Holthuis, 1951:199, pi. 63f-m.

MATERIAL.—3 males, 3 non-ovigerous females, 4ovigerous females, Malpelo coll.

MEASUREMENTS.—Males tl 4.2-4.9 mm, non-ovigerous females tl 6.4 mm, ovigerous females tl5.4-6.6 mm.

DISTRIBUTION.—Ecuador, off Cape San Franciscoand Santa Elena Bay, Galapagos Islands (HoodIsland). The species is now reported from MalpeloIsland.

REMARKS.—Dr. Fenner A. Chace, Jr., comparedthe present material with paratypes of V. laevi-frons from Hood Island and concluded they wereconspecific.

Schmitt (in Holthuis, 1951:200) gives color noteson specimens of this species that were found on agorgonian.


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 isnow reported from the eastern Pacific at MalpeloIsland.

REMARKS.—Dr. Fenner A. Chace, Jr., comparedthe Malpelo specimen with Atlantic material ofG. mineri and was unable to find any obvious dif-ferences such as those noted by Bruce (1973) inhis comparison of G. mineri with G. robustus.

In the Atlantic the species occurs under stonesor in coral, but it is usually found on the spines ofsea urchins (Tripneustes and Lytechinus). Aspecies of Tripneustes, T. depressus, is known tooccur at Malpelo Island (Birkeland et al., Appen-dix, 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, 8.2mm, juvenile cl 2.9 mm.

DISTRIBUTION.—Red Sea and South Africa, acrossthe Central Pacific to western America where itoccurs from Baja California to at least Panama.It is now reported from Malpelo.

REMARKS.—This species is a characteristic mem-ber of the Pocillopora coral community where itusually occurs in male-female pairs. Patton (1973)has dealt with aspects of the biology of this color-ful 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, non-ovigerous females cl 5.2-5.4 mm, ovigerous femalescl 7.8-11.4 mm.

DISTRIBUTION.—Western Atlantic: southern Flor-ida to Sao Paulo, Brazil. Eastern Atlantic: Senegalto the Congo. Eastern Pacific: Gulf of Californiato Ecuador, the Galapagos Islands, and now Mal-pelo Island.

REMARKS.—None.

Alpheus grahami new species

FIGURE 29A-C, F.-J

MATERIAL.—2 ovigerous females (smaller speci-men is holotype), 2 juveniles, Malpelo coll. no. 3.

MEASUREMENTS.—Ovigerous female holotype cl11.1 mm, paratypes, ovigerous female cl 12.3 mm,juveniles cl 3.7, 4.2 mm.

TYPE-LOCALITY.—Southeastern side of MalpeloIsland, Colombia in 10 m depth among coral.

DISTRIBUTION.—Known only from the type-locality.

DESCRIPTION.—The rostrum is short and triangu-lar with an acute apex; it extends to about the mid-dle of the basal antennular segment. The orbitalteeth are triangular, subacute, and extend almostto the apex of the rostrum.

The first three abdominal pleura are rounded;the fourth and fifth are bluntly angled and thesixth is subacute. The sixth segment is slightlylonger than the fifth and is about 0.7 of the lengthof the telson. The telson has the length about 1.8times the width. The distal lateral angles arearmed with a pair of spines, the inner one beinglonger and stronger. Two pairs of dorsal spines arepresent; the anterior pair is placed slightly lessthan half the distance from the anterior marginand the posterior pair is located about three-fourths the distance from the anterior margin. Theuropods extend well beyond the distal margin ofthe telson. The lateral uropod is armed with astrong movable spine at the distal lateral anglethat is adjacent to a smaller immovable spine.

NUMBER 176 73

The stylocerite extends distinctly beyond thedistal margin of the basal antennular segment. Thevisible portion of the basal segment is about three-fourths the length of the second segment. Thethird segment is about three-fourths the length ofthe second segment.

The basicerite is unarmed dorsally, the anglebeing blunt. The lateral spine is short and acute,extending to about half the length of the styloce-rite. The scaphocerite is widest proximally and

does not extend beyond the antennular peduncle.The third maxilliped extends beyond the carpo-

cerite by the distal third of the ultimate segment.The distal portion of the ultimate segment isarmed with strong setae but no spines.

The first pereiopods are strong and unequal.The ischium of the major cheliped is armed withabout four spinules on the inferior margin and asingle spine on a lobe on the distal superior mar-gin. 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, anteriorportion 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 majorchela of first pereiopod. [Scale -• 2 mm for A, B, D, E, I; 4 mm for c, F-H, J.]


inferior margin of the merus; the distal margin isunarmed. The major chela is about three times aslong as high. There are no superior notches but asulcus is present on the medial surface below thesuperior margin. It extends from the base of themovable finger to about the middle of the chelawhere it bifurcates; the upper portion extendsonly a short distance but the lower portion extendsdownward almost to the proximal margin of thechela. The lower margin of the chela is compressedproximal to the base of the immovable finger re-sulting in the formation of a weak inferior notchadjacent to two weak tubercles. The distal marginof the chela is armed with a pair of large, acuteteeth, one on either side of the movable finger. Be-low the lateral (outside) tooth, the margin of thechela leading to the immovable finger is high andcrenulate. The movable finger is high, slightly com-pressed, and blunt, although in the single chelaavailable the apex may be damaged. The ischiumof the minor chela is armed with about six spinuleson the inferior margin and a spinule on a lobe onthe distal superior margin. The merus is armedwith about 11 spinules on the inferior margin; thedistal margin is unarmed. The chela is about 1.7times as long as high. The movable finger is al-most 0.6 the length of the chela and is somewhatflattened dorsally. The distal margin of the palmis armed with two strong teeth, one on either sideof the movable finger. The tips of both fingers areacute; the hand is not balaeniceps in form. Thesecond pereiopods are subequal and extend beyondthe carpocerite by the distal three segments of thecarpus. The ischium is slightly longer than themerus and both are shorter than the carpus. Thecarpus is subdivided into five segments; the firstis the longest being slightly longer than the sec-ond and the fifth, which are subequal in length.The third and fourth segments are subequal inlength and are slightly less than half the length ofthe first. The movable finger is slightly more thanhalf the length of the chela. The fingers are un-armed. The third through the fifth pereiopods aresimilar in form. Three of the four available speci-mens have a small movable spinule at the base ofthe ischium of pereiopods three and four. Thisspinule is relatively large on the two small speci-mens (cl 3.7, 4.2 mm), but it is small and difficultto 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 thanhalf the length of the merus. The merus is slightlyless than five times as long as wide; it is about 1.5times the length of the carpus, about 1.3 times thelength of the propodus and about 5 times thelength of the dactylus. The propodus is armed withabout nine movable spines on the inferior margin.The dactylus has a minute accessory tooth, sur-rounded by setae, on the superior margin. Theeggs are small and numerous.

REMARKS.—This species seems to be most closelyrelated to Alphens crockeri (Armstrong, 1941)(= Crangon tuthilli Banner, 1953) known fromthe Central Pacific Ocean, Thailand (Banner,1953; Banner and Banner, 1966) and the easternAtlantic (Crosnier and Forest, 1966). Three speci-mens, through the courtesy of D. M. and A. H.Banner, of A. crockeri were available for compari-son: 1 male cl 8.0 mm, 2 non-ovigerous females cl5.0, 6.9 mm from Oahu, Hawaii. The two speciesare very similar, and were it not for the extremedifferences in the form of the major chela, otherdifferences would have been attributed to varia-tion. In A. crockeri (Figure 29u), the movable fin-ger is greatly compressed, with the superior marginbeing defined by a knife-like ridge; it is twistedin a horizontal plane and the apex is bulbous. Theimmovable finger has a large, high proximal toothpresent and is twisted in a horizontal plane. In A.grahami (Figure 29c, F), the movable finger isrobust and only slightly compressed; it is nottwisted and the width is the same throughout. Theimmovable finger lacks a proximal tooth and isnot twisted.

Other differences between the two species areslight but are constant in the material examined.In A. crockeri the stylocerite is narrow and extendsalmost to the distal margin of the basal antennu-lar segment, while in A. grahami the stylocerite ismore robust and extends distinctly beyond thebasal segment. The antennular peduncle of A.crockeri is slender with the second segment havingthe length about twice the width, while in A. gra-hami the length is distinctly less than twice thewidth. The scaphocerite of A. crockeri is slenderand extends to or beyond the distal margin of theantennular peduncle, while in A. grahami it isshorter than the antennular peduncle. The pereio-pods of A. crockeri are slender with the length ofthe 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 andBanner, 1966; Crosnier and Forest, 1966) itwould appear that the rostrum of A. crockeri islonger 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. gra-hami.

ETYMOLOGY.—The species is named for JeffreyB. Graham, who collected the specimens and en-trusted 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 MalpeloIsland.

REMARKS.—The Malpelo material all lack themajor chela making the identification somewhatuncertain. They agree in most respects with thedescription and illustrations of Banner (1953).The carpocerite in the Malpelo material is some-what longer than the spine of the scaphocerite, butthe 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), Malpelocoll. no. 3; 4 specimens, Malpelo coll. no. 4; 10specimens, Malpelo coll.

MEASUREMENTS.—Ovigerous females cl 4.1-4.9mm, other specimens cl 1.8-6.2 mm.

DISTRIBUTION.—St. Helena, Ecuador (Coutiere,1909), Galapagos Islands (Schmitt, 1924), Clipper-ton 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 an-tennular segment. It is on a higher level and ex-tends beyond the orbital teeth, which are smallacute extensions on the orbital hoods.

The third abdominal pleuron is obtuselyrounded, the fourth and fifth are bluntly angledand the sixth is acute. The fifth segment is slightlyshorter than the sixth, which is slightly more thanhalf the length of the telson. The telson is slightlylonger than its anterior width. It narrows distallyand has a distinct constriction proximally. Thetelson is armed with two pairs of spines at the dis-tal lateral angles; the lateral spine is about one-third the length of the medial. There are two pairsof dorsal spines present; the first pair is locatedslightly anterior of the middle and the second islocated about four-fifths of the distance from theanterior margin. The uropods extend well beyondthe distal margin of the telson. The lateral branchis armed with a strong movable spine situated be-tween two immovable spines at the level of thediaeresis.

The stylocerite is long and slender extending toabout the middle of the second antennular seg-ment. The visible portion of the basal segment isslightly longer than the second segment, which isdistinctly longer than the third segment.

The basicerite is armed with strong dorsal andlateral spines; the dorsal spine is about half thelength of the lateral. The lateral tooth of the scaph-ocerite extends beyond both the antennular pe-duncle and the carpocerite.

The third maxilliped has the apex armed withnine strong, dark-colored spines arranged in a cir-cle. The ultimate segment extends slightly beyondthe scaphocerite.

The first pereiopods are strong and unequal.The superior distal margin of the merus is armedwith a tooth. The major chela has the lengthslightly less than three times the height. The distalmargin of the palm is unarmed. A tubercle is pres-ent on the medial margin of the palm just belowthe distal margin. The immovable finger is armedwith two teeth proximally. The movable finger isslightly less than one-third the length of the chela.The superior distal margin of the merus of thesmaller chela is armed with a tooth. The smaller


chela has the length slightly more than three timesthe height. The distal margin of the palm is un-armed. The fingers are unarmed; the movable oneis about one-third the length of the chela. The sec-ond pereiopods are equal and extend beyond thescaphocerite by the distal segment of the carpus.The ischium is about three-fourths the length ofthe merus, which is slightly shorter than the car-pus. The carpus is subdivided into five segments;the first is slightly less than five times the lengthof the second, third, and fourth, which are sub-equal, and about twice the length of the fifth. Themovable finger is about half the length of the che-la. The third through the fifth pereiopods aresimilar. The merus of the third is about three

times as long as wide and is unarmed. It is abouttwice the length of the carpus and about 1.3 timesthe length of the propodus. The propodus is armedwith six movable spines, the sixth being paired,on the inferior surface. The carpus has a distalextension on the surperior margin and a pair ofspines on the distal inferior margin. The dactylusis bifid with a small ventral protuberance; the ven-tral tooth is wider than the dorsal and slightlyshorter. The eggs are small and numerous.

REMARKS.—Specimens of this species have beencollected from the rocky intertidal zone in the Bayof Panama. The material from Malpelo Islandcame from among coral and coral rubble in about10 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 firstpereiopod; 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, Malpelocoll.


DISTRIBUTION.—Baja California (Coutiere, 1909;Chace, 1937). The species is now reported fromMalpelo Island. Material from Taboguilla Island

and the Pearl Islands, Panama, was also examinedduring the present study.,

DESCRIPTION.—The rostrum is narrow, subacute,and short; extending to about one-fourth of thelength of the basal antennular segment. The orbit-al teeth arise from the anterior margin of the cara-pace; they are triangular, subacute, almost twiceas broad as the rostrum, and are subequal inlength to it.

The third through the fifth abdominal pleuraare bluntly angled; the sixth is acute. The fifthsegment is almost equal in length to the sixthwhich is slightly more than half the length of thetelson. The telson is longer than wide with twopairs of spines on the distal lateral angles; the

FIGURE 31.—Synalpheus digueti Coutiere: A, anterior portion of carapace; B, anterior portion ofcarapace of aberrant specimen; c, second pereiopod; D, minor chela of first pereiopod; E, thirdpereiopod; F, telson and left uropod; c, anterior portion of dactylus of third pereiopod; H, majorchela of first pereiopod; i, anterior portion of major chela of first pereiopod. [Scale = 2 mmfor A-F; 1 mm for c; 4 mm for H, I.]


lateral spine is slightly more than half the lengthof the medial. The distal margin of the telson isconvex and extends to about two-thirds the lengthof the medial spine. There are two pairs of dorsalspines; the first pair is located at about the middleof the telson and the posterior pair is locatedslightly more than two-thirds of the distance fromthe anterior margin. The uropods extend beyondthe distal margin of the telson. The lateral branchis armed with a strong movable spine situated be-tween two immovable spines at the level of thediaeresis.

The stylocerite is long and acute, extendingslightly beyond the middle of the second antennu-lar segment. The first antennular segment is about1.4 times the length of the second and about twicethe length of the third.

The basicerite is armed with dorsal and lateralspines; the dorsal spine extends to about one-fifthof the length of the lateral. The lateral tooth ofthe scaphocerite is long and acute and extendsbeyond the antennular peduncle. The carpoceriteis long and broad, extending well beyond the lateraltooth of the scaphocerite.

The first pereiopods are robust and unequal.The merus of the major cheliped is unarmed. Thechela is about 2.6 times as long as high. The distalmargin of the palm is armed with a downwardlydirected spine. There are two tubercles on thelateral margin of the palm at the insertion of themovable finger. The distal portion of the immov-able finger is compressed. The movable finger isslightly more than one-fourth the length of thechela. The merus of the smaller cheliped is un-armed, as is the chela. The length of the chelais about 2.5 times the height. The movable finger isabout half the length of the chela. The secondpereiopods are subequal. The ischium is unarmedand is about three-fourths the length of the merus.The merus is unarmed and is about four-fifths thelength of the carpus. The carpus is subdivided intofive segments; the first is over five times the lengthof the second, third, and fourth which are subequaland which are half the length of the fifth. Themovable finger is slightly more than half thelength of the chela. The third through the fifthpereiopods are similar. The ischium and merusare unarmed. The ischium is slightly less than one-third the length of the merus. The carpus has an

extension on the distal superior margin and a pairof spines on the distal inferior margin. The carpusis about half the length of the merus. The propo-dus is about four-fifths the length of the merusand is armed with seven movable spinules on theinferior margin; the last spinule is paired. Thedactylus is bifid; the superior tooth is slightlylonger than the inferior tooth and they are sub-equal 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 ofthe first pereiopods are red.

REMARKS.—The rostrum was somewhat variablein form often being wider than figured here. Onespecimen (Figure 31B) has a highly aberrant ros-trum but was normal in other features. Three ofthe 99 specimens examined lack a tooth on thedistal margin of the major chela. In other featuresthe species was not variable.

In the Gulf of Panama Synalpheus diguetioccurs as male-female pairs (the female is usuallyovigerous) 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.


DISTRIBUTION.—Red Sea across the Pacific toBaja California (Banner and Banner, 1966; Chace,1937). During the present study, material was alsoexamined from the Pearl Islands and TaboguillaIsland in the Gulf of Panama.

REMARKS.—This species is a characteristic mem-ber of the Pocillopora coral community, occurringdeep inside the coral at the base of the branches(Banner and Banner, 1964). It is common in thishabitat in the Gulf of Panama where it usuallyoccurs in male-ovigerous female pairs. The bodyand appendages of fresh material is an even redcolor.

NUMBER 176 79

Synalpheus bannerorum, new species

FIGURE 32

MATERIAL.—5 ovigerous females (holotype tl 11.1mm, cl 4.4 mm), 23 specimens, Malpelo coll. no. 3(paratypes); 2 specimens, Malpelo coll. no. 4.

MEASUREMENTS.—Ovigerous females cl 4.0-4.7mm, tl 9.9-11.1 mm, other specimens cl 3.3-5.0 mm,// 8.5-11.4 mm.

TYPE-LOCALITY.—Southeastern side of MalpeloIsland, Colombia, in 10 meters depth among coral.

DISTRIBUTION.—Known only from the type-locality.

DESCRIPTION.—The rostrum is narrow and acute;it does not extend to the distal margin of the basalantennular segment. It is on a higher level thanthe orbital teeth, which are small, acute, directedslightly medially, and do not extend to the apexof the rostrum.

The first five abdominal pleura are rounded; thesixth is subacute. The fifth segment is subequal inlength to the sixth, which is three-fourths thelength of the telson. The anterior width of the tel-son is greater than the length; it narrows distinctlyin the posterior half. The posterior margin is lessthan one-half the width of the anterior margin.The distal lateral angles of the posterior marginare acute and armed with a pair of spines; thelateral pair being shorter and more robust thanthe medial. Both extend well beyond the lateralangles of the telson. The dorsal surface of the tel-son is armed with a pair of strong spines; the firstpair is located about two-thirds of the distancefrom the posterior margin and the second pair islocated about one-third of the distance from theposterior margin. Both pairs are set in well fromthe lateral margins. The lateral angle of the outeruropod is armed with a long movable spine set

FIGURE 32.—Synalpheus bannerorum, new species: \, anterior portion of carapace; B, major chelaof first pereiopod; c, anterior portion of major chela; D, second pereiopod; E, anterior portion ofdactylus of third pereiopod; F, third pereiopod; c., telson and left uropod; H, minor first pereio-pod. [Scale = 2 mm for A-D, F, H; I mm for E, G.]


between a pair of smaller acute immovable spinesat the level of the diaeresis. The medial uropod islonger than the lateral, which is subequal in lengthto the telson.

The stylocerite is acute and extends to about themiddle of the second antennular segment. Thebasal antennular segment is longer than the secondand third, which are subequal in length.

The basicerite is armed with strong, acute dorsaland lateral spines; the dorsal spine extending toslightly over one-third of the length of the lateral.The scaphocerite is reduced in size but the lateralspine is strong, extending beyond both the anten-nular peduncle and the carpocerite. The carpoce-rite extends slightly beyond the antennular pe-ducle.

The third maxilliped has the tip of the apexarmed with six dark-colored spines. It extends be-yond the carpocerite by the distal third of theultimate segment.

The first pereiopods are unequal and robust.The merus of the major cheliped is armed with atooth on the distal superior margin. The carpusis armed with two acute extensions. The majorchela is about 2.3 times as long as high. The distalmargin of the palm is armed with a small down-wardly directed spine. There is a large tubercle atthe base of the immovable finger which is followedby a deep depression and then another tubercle.The other side of the finger is also depressed givinga pinched appearance to the base of the finger.The immovable finger is armed with two strong,acute teeth in the proximal portion. The movablefinger is more than one-third the length of thechela and has two distinct tubercles on the proxi-mal dorsal surface. There is a series of about fiveunequal tubercles on the medial surface of thepalm between the acute distal spine and the baseof the immovable finger. The merus of the minorcheliped is armed with an acute spine on the supe-rior distal margin. The carpus is cup-shaped. Thechela is unarmed; the movable finger is slightlyless than half the length of the chela. The secondpereiopods are subequal. The fingers are unarmedand the movable finger is slightly more than halfthe length of the chela. The carpus is subdividedinto five segments; the first is about six times thelength of the second, which is subequal in lengthto the third and fourth, which are about half thelength of the fifth. The ischium is about half the

length of the carpus and about three-fifths thelength of the merus. The third through the fifthpereiopods are similar and robust. The ischium isunarmed. The merus of the third is armed withfour to six movable spines. The length is slightlymore than three times the width. The carpus isarmed with two distal spines. The propodus isarmed with about nine movable spinules on theinferior margin. The dactylus has the superiortooth longer and slightly more than half the widthof the inferior. The merus of the fourth pereiopodis armed with three or four movable spines. Themerus of the fifth pereiopod is unarmed. The eggsare small and numerous.

REMARKS.—This species differs from all knowneastern Pacific species in the form of the meri ofthe third and fourth pereiopods and in the formof the major chela. It is the only western Americanspecies in which the meri of the third and fourthpereiopods are armed and the constriction of theimmovable finger of the major chela appears alsoto be unique. There is one east American speciesof Synalpheus (S. dominicensis Armstrong, 1949)from the Dominican Republic, which has the meriof pereiopods three and four armed. Synalpheusbannerorum can be distinguished from S. domini-censis by the form of the anterior portion of thecarapace, by the armature of the telson, by theform of the major chela and by the form of thedactyi of pereiopods three through five. In S. domi-nicensis the medial margins of the orbits meet therostrum at a sharp angle while they blend into thecarapace before they meet the rostrum in S. ban-nerorum. The dorsal spines of the telson are bothlocated in the posterior half of the segment in S.dominicensis, while in S. bannerorum only the pos-terior pair is located in the posterior half. Themajor chela of S. dominicensis is not compressedat the base of the immovable finger as it is in S.bannerorum. The dactylus of the third pereiopodin S. dominicensis has the lower tooth less diver-gent than that of S. bannerorum. Synalpheus ban-nerorum is also related to Indo-Pacific membersof the genus, but the form of the chela appears todistinguish it from all other species of the genus.

ETYMOLOGY.—This species is named for Dora M.Banner and Albert H. Banner who have contri-buted so much to the knowledge of the Alpheidaeand 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 Islandin the Gulf of Panama were also examined duringthe present study.

REMARKS.—The present material agrees with thedescription and illustrations of Chace (1937). Thespecies commonly occurs on Pocillopora coral inthe 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 Islandin the Galapagos Islands, and is now reported fromMalpelo Island.

REMARKS.—Both specimens agree in all respectswith the description and illustration of Schmitt(1924). The rostral formula of both specimenswas one postorbital, five preorbital and one sub-apical ventral teeth. The occurrence of this specieswith L. trisetacea offers additional support (Sch-mitt, 1939; Chace, 1962) for the specific distinct-ness 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 non-ovigerous females, 9 ovigerous females, Malpelocoll. 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 beenreported 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 tooccur on the western American mainland. Lysmatacalifornica (Stimpson, 1866) occurs in Californiaand is common in the Gulf of Panama while L.porteri (Rathbun, 1907) occurs in Chile. The re-cord of Sivertsen (1934) for L. intermedia (Kings-ley, 1878), a western Atlantic species, is based ona single specimen from the Galapagos Islands,which appears to be within the range of variationof L. californica (see Sivertsen, 1934, pi. 2: figs.9-15; and Limbaugh, Pederson, and Chace, 1961,fig. 7).

REMARKS.—Chace (1962) discussed variation inthis species and noted three differences betweenmaterial from Clipperton Island and material fromthe Hawaiian Islands. In some Hawaiian specimensthe posterior ventral rostral tooth is behind oropposite the anterior dorsal tooth while in mostof the Clipperton specimens both ventral teeth,when present, are well in advance of the anteriordorsal tooth. In larger adults from Hawaii thefused part of the upper antennular flagellum isnot much shorter than the free portion of theshorter branch while in large Clipperton speci-mens the fused portion is distinctly less than halfthe length of the free portion. The average num-ber of carpal segments in Hawaiian material (23)tended to be higher than that of Clipperton mate-rial (21).

These three characters were examined in theMalpelo material with the following results. Therostral formula was (number of specimens in pa-rentheses): 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 specimenswith two ventral teeth had them both placed wellin advance of the anterior dorsal tooth; those withone ventral tooth had it placed well in advance ofthe anterior dorsal tooth. The Malpelo materialagreed also with the Clipperton material regardingthe relative lengths of the fused and free portionsof the upper antennular flagellum. The numberof carpal segments in Malpelo material varied


from 21-26 distributed as follows (number of speci-mens in parentheses): 24 (5), 22 (3), 21 (2), 23(2), 25 (1), 26 (1). The average was slightly morethan 24 agreeing more with the Hawaiian materialthan with the Clipperton material. These differ-ences warrant further consideration when compar-ing material from throughout the range of L. tri-setacea as suggested by Chace (1962) and Holthuis(1947).

PALINURIDAE

Panulirus penicillatus (Olivier, 1791)

Astacus penicillatus Olivier, 1791:343.Panulirus penicillatus (Olivier).—Holthuis, 1946:125.—Hol-

thuis and Loesch, 1967:217. pi. 8.

MATERIAL.—None, the species was identified fromcolor slides.

MEASUREMENTS.—Cl 91-137 mm (Holthuis andLoesch, 1967).

DISTRIBUTION.—This species occurs in the Indo-West Pacific region: Red Sea, Southeast Africa toKorea, Formosa, and Polynesia. In the eastern Pa-cific it occurs at Revillagigedo Islands, ClippertonIsland, Cocos Island, Galapagos Island (Holthuisand Loesch, 1967), and now is reported from Mal-pelo Island. It has not been found on the westernAmerican mainland.

REMARKS.—Although no material of this specieswas collected, comparison of color slides taken byDr. Charles Birkeland with the color plate of thisspecies given by Holthuis and Loesch (1967) per-mitted identification.

Ecology

The short time available to members of the expe-dition did not permit detailed collecting notes onso many different groups of organisms (see Appen-dix in Birkeland et al., this volume). However,some observations are possible based on publishedinformation and my own experiences in Panama(Abele, 1972).

Perhaps the most interesting observation is thelarge percentage of decapods from Malpelo Islandwhich are commensals. Almost half of the 43species occur in close association with some otherinvertebrates. 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., Tra-pezia digitalis, T. ferruginea, Domecia hispida,Medaeus spimilifer, Carpilodes cinclimanus, Tel-eophrys cristulipes, and Mithrax pygmaeus. BothVeleronia laevifrons and Pseudocoutierea elegansare associated with gorgonians. Gnathophylloidesmineri occurs on the spines of sea urchins andPericlimenaeus hancocki probably occurs withsponges.

Two species, Grapsus grapsus and Gecarcinusmalpilensis, occur intertidally and supratidally,respectively.

The remaining species occur in a variety of sub-tidal habitats, mostly with rocks and coral rubble.

Distribution

The decapod fauna of Malpelo Island has affini-ties with both the western American mainland andIndo-Pacific decapod faunas (p. 67). Twenty-sevenspecies, 63 percent of the Malpelo fauna, occur onthe western American mainland. Nine species, 21percent of the fauna, occur in the Indo-Pacific re-gion and seven of these reach the western Ameri-can mainland. The other two, Lysmata trisetaceaand Panulirus penicillatus occur from the Indo-Pacific only to the offshore islands of the easternPacific. Three species, Gecarcinus malpilensis, Al-pheus grahami, new species, and Synalpheus ban-nerorum, new species, appear to be endemic toMalpelo Island. Two species, Brachycarpus biun-guiculatus and probably Gnathophylloides mineriare circumtropical in distribution. Lysmata galapa-gensis is only known from the Galapagos Islandsand Malpelo Island, while Synalpheus biunguicu-latus is known from Hawaii, Clipperton Island,and Malpelo Island.

The decapod fauna of Malpelo Island then hasits greatest affinities (63%) with that of the west-ern American mainland, next is the Indo-Pacificregion (21%), then endemic (7%), circumtropical(4%), and island species (4%).

Comparison with Other Islands

A detailed comparison, at the species level, ofthe decapod faunas of the offshore islands of the

NUMBER 176 83

fe 1.8.

I8 1.7

I.OC PERIMETER OF ISLAND

FIGURE 33.—The relationship between island perimeter andthe number of decapod crustacean species present at (1),Malpelo, (2) Clipperton, and (3) Cocos islands. Refer to textfor discussion.

eastern Pacific will have to await more data onother islands.

Some comparisons can be made, however, be-tween numbers of species of decapods associatedwith oceanic islands (Figure 33). There are 25species of decapods known from Saint Helena(Chace, 1966; 1968), an isolated island in thesouth Atlantic. Twenty percent, five species, areendemic. There are 58 species of decapods knownfrom Clipperton Island (Chace, 1962; Garth, 1965)in the eastern Pacific. None of these species isendemic. I have found about 100 speciesknown from Cocos Island in the eastern Pacific ofwhich at least six (6%) are endemic. Malpelo Is-land has a decapod fauna of 43 species with three(7%) of them being endemic.

Within a given region there generally exists anorderly relation between numbers of species andthe size of the area sampled. This relationship hasbeen worked out for a number of terrestrial organ-isms occurring on islands (MacArthur and Wilson,1967). In comparing shallow water organisms as-sociated with islands, it seems more appropriateto use island perimeter rather than island area. Acomparison between island perimeter and numbersof 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 strik-

ing, although the slope (z=0.56) is higher thanthat reported for other area-species curves (z =0.20-0.35; MacArthur and Wilson, 1967). Thepresent curve may not be representative beingbased on only three islands and is presented hereprimarily to stimulate interest in examining thisrelationship in other marine organisms.

Literature Cited

Abele, L. G.1972. Comparative Habitat Diversity and Faunal Rela-

tionships between the Pacific and Caribbean Pana-manian Decapod Crustacea: A Preliminary Reportwith Some Remarks on the Crustacean Fauna ofPanama. Pages 125-138 in M. L. Jones, editor. ThePanamic Biota: Some Observations Prior to a SeaLevel Canal. Bulletin of the Biological Society ofWashington, number 2.

Armstrong, J. C.1941. The Caridea and Stomatopoda of the Second Tem-

pleton Crocker-American Museum Expedition tothe Pacific Ocean. American Museum Novitates,1137:1-14, 4 figures.

1949. New Caridea from the Dominican Republic.American Museum Novitates, 1410:1-27, 9 figures.

Banner, A. H.

1953. The Crangonidae, or Snapping Shrimp of Hawaii.Pacific Science, 7 (1):2-144, 147, 50 figures, 1 frontis-piece.

1956. Collections from the Mariana Archipelago. Part Iin Contributions to the Knowledge of the AlpheidShrimp of the Pacific Ocean. Pacific Science, 10:318-373, 23 figures.

1958. On a Small Collection from Onotoa, Gilbert Is-lands. Part III in Contributions to the Knowledgeof the Alpheid Shrimp of the Pacific Ocean. PacificScience, 12(2):157-169, 4 figures.

Banner, A. H., and D. M. Banner

1964. Collections from the Phoenix and Line Islands.Part IX in Contributions to the Knowledge of theAlpheid Shrimp of the Pacific Ocean. PacificScience 18:83-100, 5 figures.

1966. The Alpheid Shrimp of Thailand: The AlpheidShrimp of the Gulf of Thailand and AdjacentWaters. The Siam Society Monograph Series, 3,vi + 168 pages, 168 figures.

Bruce, A. J.1970. On the Identity of Periclimenes pusillus Rathbun,

1906 (Decapoda, Pontoniinae). Crustaceana 19(3):306-310, 1 figure.

1973. Gnathophylloides robustus sp. nov., a New Com-mensal Gnathophyllid Shrimp from Western Aus-tralia, with the Designation of a New GenusLevicaris (Decapoda, Caridea). Crustaceana 24(1):17-32, 9 figures.


Chace, F. A., Jr.1937. Caridean Decapod Crustacea from the Gulf of Cali-

fornia and the West Coast of Lower California.Part VII in The Templeton Crocker Expedition.Zoologica (New York), 22(2):109-138, 9 figures.

1962. The Non-Brachyuran Decapod Crustaceans of Clip-perton Island. Proceedings of the United StatesNational Museum, 113(3466):605-635; 7 figures.

1966. Decapod Crustaceans from St. Helena Island, SouthAtlantic. Proceedings of the United States NationalMuseum, 118 (3536):623-661, 15 figures.

1968. A New Crab of the Genus Cycloes (Crustacea;Brachyura; Calappidae) from Saint Helena, SouthAtlantic Ocean. Proceedings of the Biological So-ciety of Washington, 81:605-612, 2 figures.

1972. The Shrimps of the Smithsonian-Bredin CaribbeanExpeditions, with a Summary of the West IndianShallow-water Species (Crustacea: Decapoda: Na-tantia). Smithsonian Contributions to Zoology, 98:1-179, 61 figures.

Coutiere, H.1909. The American Species of Snapping Shrimps of the

Genus Synalpheus. Proceedings of the UnitedStates National Museum, 36 (1659): 1-93, 54 figures.

Crosnier, A., and J. Forest1966. Crustaces Decapodes: Alpheidae. Part 19 in Cam-

pagnc de la Calypso dans le Golfe de Guinee etaux lies Principe, Sao Tome et Annobon (1956), etCampagne aux lies de Cap Vert (1959). Fascicle 7,volume 27, in Resultats scientifiques des Campagnesde la "Calypso". Annales de I'Institute Oceanogra-phique, Monaco, 44:199-314, 33 figures.

Dana, J. D.1852. Crustacea, part I. Volume 13 in United States Ex-

ploring Expedition, During the Years 1838, 1839,1840, 1841, 1842, Under the Command of CharlesWilkes, U.S.N. viii + 685 pages. Philadelphia: C.Sherman.

1855. Crustacea, Atlas. Volume 13 in United States Ex-ploring Expedition, During the Years 1838, 1839,1840, 1841, 1842, under the Command of CharlesWilkes, U.S.N. 27 pages, 96 plates. Philadelphia:C. Sherman,

de Man, J. G.1911. Family Alpheidae. Part II in The Decapoda of the

Siboga Expedition. Siboga Expedite, 39a1 (2): 113—327, 23 plates.

Fausto Filho, J.1966. Brachycarpus holthuisi, Nova Especie de Crustaceo

do Brasil (Decapoda PaJaemonidae). ArquivosEstacao de Biologia Marinha Universidade do Ceara,6 (2): 123-125, 11 figures.

Faxon, W.1893. Preliminary Descriptions of New Species of- Crus-

tacea. Part VI in Reports on the Dredging Opera-tions Off the West Coast of Central America to theGalapagos, to the West Coast of Mexico, and inthe Gulf of California, in Charge of AlexanderAgassiz, by the U.S. Fish Commission Steamer

"Albatross," during 1891, Lieutenant CommanderZ. L. Tanner, U.S.N., Commanding. Bulletin of theMuseum of Comparative Zoology at Harvard Col-lege 24:149-220.


Remarks on Carcinological Collecting in the Pan-ama Bight. Bulletin of the American Museum ofNatural History 92(1): 1-66, 4 figures.

1965. The Brachyuran Decapod Crustaceans of Clipper-ton Island. Proceedings of the California Academyof Sciences, fourth series, 33(1): 1-46, 26 figures.

GueYin-Meneville, F. E.1830. Crustaces, Arachnides ct Insectcs. Section 1 of part

2 in volume 2 (zoologie) in M. L. I. Duperrey,Voyage autour du monde, execute par Ordre duRoi, sur la Cornette de Sa Majeste, JM "Coquille,"pendant Us annees 1822, 1823, 1824, et 1825. 319pages, 24 plates. Paris.

Heller, C.1861. Synopsis der im rothcn Mccrc vorkommenden

Crustaccen. Verhandelinger Zoologische botanischeC.esellschaft (Wien) 11:3-32.

Holthuis, L. B.1946. The Stenopodidae, Nephropsidae, Scyllaridae and

Palinuridae. Part I in The Dccopoda Macrura ofthe Snellius Expedition. Part XIV in BiologicalResults of the Snellius Expedition. Temminckia, 7:1-178; 2 figures, 11 plates.

1947. The Decapoda of the "Siboga Expedition, IX: TheHippolytidae and Rhynchocinetidae Collected bythe Siboga and Snellius Expeditions with Remarkson Other Species. Volume 39a8 in Siboga-Expedite.Leiden: E. J. Brill.

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 GeneralRevision of the Palaemonidae (Crustacea DecapodaNatantia) of the Americas. Allan Hancock Founda-tion Occasional Papers, 12:1-396, 55 plates.

1958. Crustacea Decapoda from the northern Red Sea(Gulf of Agaba and Sinai Peninsula), I: Macrura.(Contributions to the Knowledge of the Red Sea,no. 8.) State of Israel Sea Fisheries Research StationBulletin, 17:1-40, 15 figures.

Holthuis, L. B., and H. Loesch1967. The Lobsters of the Galapagos Islands (Decapoda,

Palinuridea). Crustaceana 12 (2):2I4-222; 1 figure,plates 7-9.

Hult, J.1939. Crustacea Decapoda from the Galapagos Islands

Collected by Mr. Rolf Blomberg. Arkiv Zoologiske,3OA(5):1-18, 4 figures, 1 plate.

Kingsley, J. S.1878. Notes on the North American Caridea in the Mu-

seum of the Peabody Academy of Science at Salem,

NUMBER 176 85

Mass. Proceedings of the Academy of NaturalSciences of Philadelphia, 1878:89-98.

Lewis, J. B.1956. The Occurrence of the Macruran Gnathophylloides

mineri Schmitt on the Spines of the Edible SeaUrchin Tripneustes esculentus Leske in Barbados.Bulletin of Marine Science of the Gulf and Carib-bean, 6 (4):288-291, 2 figures.

Limbaugh, C, H. Pederson, and F. A. Chace, Jr.1961. Shrimps that Clean Fishes. Bulletin of Marine

Science of the Gulf and Caribbean, 11 (2):237-257,9 figures.

Lucas, H.1849. Crustaces, Arachnides, Myriopodes et Hexapodes.

In Exploration scientiftque de VAlgerie pendant lesannees 1840, 1814, 1842. Zoologie I in Sciencesphysiques. Volume I (403 pages, 8 plates) in His-toire naturelle des animaux articule.

MacArthur, R. H., and E. O. Wilson.1967. The Theory of Island Biogeography. xi + 203

pages. Princeton: Princeton University Press.Milne Edwards, H.

1837. Histoire Naturelle des Crustaces, ComprenantI'Anatomie, la Physiologie et al Classification de cesAnimaux. Volume 2, 532 pages; atlas, 32 pages,plates 1-14, 14 bis, 15-25, 25 bis, 26-42. Paris:Librarie Encydoped ique de Roret.

Olivier, A. G.1791. Ecrevisse. Astacus. Pages 327-349 of volume 6 in

A. G. Olivier, Encyclopedic methodique, Histoirenaturelle, Insects.

Patton, W. K.1973. Community Structure among the Animals Inhab-

iting the Coral, Pocillopora damicornis at HeronIsland, Australia. Pages 214-238 in W. Vernberg,editor, Symbiosis in the Sea. Columbia: Universityof South Carolina Press.

Randall, J. W.1839. Catalogue of the Crustacea Brought by Thomas

Nuttall and J. K. Townsend, from the West Coastof North America and the Sandwich Islands, withDescriptions of Such Species as are Apparently New,among Them Which are Included Several Speciesof Different Localities Previously Existing in theCollection of the Academy. Journal of the Academyof Natural Sciences Philadelphia, 8:106-147, plates3-7.

Rathbun, M. J.1906. The Brachyura and Macrura of the Hawaiian Is-

lands. Bulletin United States Fish Commission, 23:827-930, 79 figures, 24 plates.

1907. South American Crustacea. Revista Chile Historicnaturale, 11:45-50; 1 figure, 2 plates.

1918. The Grapsoid Crabs of America. Bulletin of theUnited States National Museum, 97, xxii + 461pages, 172 figures, 161 plates.

Schmitt, W. L.1924. The Macrura and Anomura Collected by the Wil-

liams Galapagos Expedition, 1923. Zoologica (NewYork), 5(15):161-171, figures 39-41.

1932. Coralliocaris pearsei. In Pearse, Inhabitants ofCertain Sponges at Dry Tortugas. (Papers of theTortugas Laboratory, volume 23.) Carnegie Institu-tion of Washington Publication, 435:123, 124, fig-ure 1.

1933. Four New Species of Decapod Crustaceans fromPorto Rico. American Museum Novitates, 662:1-9,4 figures.

1935. Crustacea Macrura and Anomura of Porto Rico andthe Virgin Islands. Pages 125-227 in part 2 ofvolume 15 in Scientific Survey of Porto Rico andthe Virgin Islands. 80 figures. New York: New YorkAcademy of Sciences.

1939. Decapod and Other Crustacea Collected on thePresidential Cruise of 1938 (with Introduction andStation Data). Smithsonian Miscellaneous Collec-tions, 98 (6): 1-29, 2 figures, 3 plates.

Sivertsen, E.1934. Littoral Crustacea Decapoda from the Galapagos

Islands. Part VII in The Norwegian ZoologicalExpedition to the Galapagos Islands, 1925, con-ducted by Alf Wollebaek. Meddelelser fra detZoologiske Museum (Oslo), 38:1-23, 1 figure, 4plates.

Stimpson, W.1860. Prodromus Descriptionis Animalium Evertebra-

torum, quae in Expeditione ad Oceanum PacificumSeptentrionalem, a Republica Federata missa,Cadivaladaro Ringgold et Johanne Rodgers Duci-bus, Observavit et Descripsit. Proceedings of theAcademy of Natural Sciences of Philadelphia, 1860:22-48.

1866. Descriptions of New Genera and Species of Macru-rons Crustacea from the Coasts of North America.Proceedings of the Chicago Academy of Sciences,1:46-48.

Turkay, M.1970. Die Gecarcinidae Amerikas: Mif einem Anhang

fiber Ucides Rathbun. Senckenbergiana biologica51 (5/6):333-354, 11 figures.

Asteroidea from Malpelo Islandwith a Description of a New Species

of the Genus Tamaria

Maureen E. Downey

ABSTRACT

One new species, Tamaria stria, and a new sub-species, Narcissia gracilis malpeloensis, are de-scribed from shallow waters of Malpelo Island. Acollection totaling six species of starfish is described,with notes on distribution.

Introduction

In the course of the faunal survey of the shallowwaters (0-50 m) of Malpelo conducted by C. Birke-land and his colleagues, only six species of starfishwere collected; however, all are species with inter-esting or unusual distribution patterns, and in-cluded in the collection are a new species of thegenus Tamaria and a new subspecies of Narcissisgracilis. The abbreviation R refers to the majorradius from center of disc to tip of arm; and r re-fers to the minor radius, from center of disc tointerradial margin.

ACKNOWLEDGMENTS.—I wish to thank CharlesBirkeland and David Meyer of the SmithsonianTropical Research Institute for the opportunity toexamine this collection, and Dr. David L. Pawson,Chairman, Department of Invertebrate Zoology,National Museum of Natural History, Smithson-ian Institution, for reading the manuscript andoffering helpful comments and suggestions.

Maureen E. Downey, Department of Invertebrate Zoology,National Museum of Natural History, Smithsonian Institu-tion, 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.—Steinbeckand 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 collec-tion by one specimen, quite large and heavy (R10 cm, r 6.5 cm, dry weight 177 grams). The mar-ginals, particularly the distal ones, are tremen-dously swollen and most bear 1 or 2 stout spines orlow tubercles. The much smaller principal platesof the abactinal surface usually bear 1 or 2 lowtubercles. The papular areas of the abactinal sur-face are covered with pedicellariae of the splitgranule type. This is a common shallow waterspecies of the eastern tropical Pacific, known fromlower 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, re-ported by Fisher from Maui, Lanai, Kauai, andBird Islands, in 58-124 meters. One fine specimen,collected at Malpelo in 49 meters at the base of arock wall (Station 5, Birkeland et al., Appendix,this volume) measures R 20 cm, r 2.9 cm. Leiastercallipeplus has also been collected near the centralAmerican mainland in the Gulf of Chiriqui, Pana-ma (Birkeland, pers. comra.).

Narcissia gracilis malpeloensis, new subspecies

Narcissia gracilis malpeloensis is represented inthis collection by six specimens. The species, de-scribed by A. H. Clark (1916:58) has previouslybeen collected from Baja California to the Gala-pagos Islands. The average size of these specimensis R 12 cm, r 2.5 cm. The Malpelo specimens repre-sent a new subspecies of N. gracilis, differing fromthe type-specimen and Clark's description in thefollowing respects:

Narcissia gracilis gracilis: Pedicellariae abundanton all surfaces, of 2 slender valves with expandedtoothed tips, lying in alveoli; granules hemispher-ical; subambulacral spines in 3 rows; madreporiteround; papulae single; and proximal marginalslonger than broad.

Narcissia gracilis malpeloensis: Few or no pedi-cellariae—when present, of 2 stout curved un-toothed valves of uniform thickness, not in alveoli;granules flat-topped, polygonal; subambulacralspines 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, andabundant pedicellariae versus few or no pedicel-lariae might be accounted for by the difference insize (holotype: R 54 mm, r 8.5 mm) between thesespecimens and the type (starfishes frequently havemany pedicellariae when young and few or nonewhen fully grown), the very dissimilar character ofthe pedicellariae, plus the other variations listedabove 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, adra-

dial, superomarginal, and inferomarginal platesdiamond-shaped, slightly overlapping distal cornerof preceding plate, connected transversely to adja-cent row of plates by small, irregular, secondaryplates. Six rows of papular areas, 6-10 pores perarea. Three rows of rounded actinal intermediateplates, becoming two distally. Entire body, up toadambulacral armature, covered with close uni-form coating of granules, those in papular areasslightly smaller than those on plates. Adambulac-ral furrow spines two, one small, acicular, the otherheavy, flared at tip, broad in plane of furrow. Sub-ambulacral spines single, heavy, thick, longer thanbroad. No pedicellariae noted. Madreporite small,round, covered with fine radiating gyri. Anussubcentral, inconspicuous. Ocular plates large,rounded, raised, mostly bare, with few granules.Color in life is reddish orange; dried, brightorange with blue papular areas.

MATERIAL EXAMINED.—Holotype, USNM El 1838,Station 4, Malpelo Island, 49 m, on rubble nearbase of vertical rock wall. Paratypes, USNMEl 1839, 4 specimens, Stations 4 and 5, MalpeloIsland, 36-49 m.

ETYMOLOGY.—The species name is the latinizedacronym for the Smithsonian Tropical ResearchInstitute.

DISCUSSION.—Only one other species of Tamaria,T. obstipa Ziesenhenne, has been described fromthe eastern tropical Pacific, Tamaria obstipa isknown from the type-locality, Cocos Island, CostaRica, and from James Island, Galapagos Islands.The present species agrees closely with T. obstipa,but differs in having secondary connecting platesnot only intermarginally but between all abacti-nal plate rows; in having a carinal series, lackingin T. obstipa; in the nature of the adambulacralarmature; and in the lack of pedicellariae. Thenumber 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:


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 col-lected at Malpelo at depths from 12 to 14 meters;the known distribution for this species is from BajaCalifornia to the Galapagos Islands. The Malpelospecimens have many more pedicellariae thanspecimens from Baja California in the collectionsof the National Museum of Natural History,Smithsonian Institution, but correspond well withmaterial 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 col-lected at Malpelo on a vertical rock wall at 36meters. It is distributed throughout the Indo-Pacific and the tropical eastern Pacific. This speciesis apparently attacked by the shrimp Hymenocera(Birkeland, pers. comra.), well known as a pre-

dator on Acanthaster plancii, the crown-of-thornsstarfish.

Literature Cited

Bell, F. J.

1884. Contributions to the Systematic Arrangement ofthe Asteroidea, II: The Species of Or easier. Pro-ceedings of the Zoological Society of London, 1884:57-87.

Boone, L.

1926. Echinoderms from the Gulf of California and thePerlas Islands. Bulletin of the Bingham Oceano-graphic Collection (New York), 2 (6): 1-14, 9 plates.

1933. Scientific Results of Cruises of the Yachts Eagleand Ara, 1921-1928, Win. K. Vanderbilt, Command-ing. Bulletin of the Vanderbilt Marine Museum(Huntington), 4:68-164, plates 25-102.

Caso, M. E.1943. Contribucion al conocimiento de los asteridos de

Mexico. 136 pages, 50 plates. Thesis, UniversidadNacional de Mexico, Faculte de Ciendas.

1944. Estudios sobre asteridos de Mexico. Algunasespecies interesantes de asteridos litorales. AnalesInstituto Biologico de Mexico, 15 (l):236-259.

1953. Estado actual de los conocimientos acerca de lafauna de los equinodermos de Mexico. Memoriadel Congreso Cientifico Mexicano, 7:209-222, 12figures.

1961. Los equinodermos de Mexico. 388 pages, 124 fig-ures, 20 plates. Universidad Nacional de Mexico,Faculte de Ciendas.

1962. Estado actual de los conocimiento acerca de losequinodermos de Mexico. 388 pages, 124 figures, 20plates. Universidad Nadonal de Mexico, Facultede Ciencias.

1963. Estudios sobre equinodermos de Mexico. Contri-bucion al conocimiento de los equinodermos de lasislas Revillagigedo. Anales del Instituto Biologicode Universidad de Mexico, 33:293-330, 9 plates.

Clark, A. H.1916. Six new Starfishes from the Gulf of California and

Adjacent Waters. Proceedings of the BiologicalSociety of Washington, 29:51-62.

1946. Echinoderms from the Pearl Islands. SmithsonianMiscellaneous Collections, 106 (5): 11.

Clark, A. M.1967. Echinoderms from the Red Sea, Part 2: Crinoids,

Ophiuroids, Echinoids and More Asteroids. (Num-ber 21 in Reports of the Israel South Red Sea Ex-pedition 1962.) Sea Fisheries Research Station Bulle-tin (Haifa), 41:26-58, 5 figures.

Clark, H. L.1910. The Echinoderms of Peru. Bulletin of the Museum

of Comparative Zoology at Harvard, 52(17):321-358, 14 plates.

1940. Notes on Echinoderms from the West Coast ofCentral America. Zoological Society, 25 (3a):331-352,2 plates.

1958. Estrellas de Mar (Asteroidea). Biota, 2:85-105, 13figures.

Doderlein, L.1916. Ueber die Gattung Oreaster und Verwandte, Zool-

ogische Jahrbuch (Systematischen), 40:409-440.1936. Die Asteriden der Siboga-Expedition, III: Die

Unterfamilie Oreasterinae. Pages 295-369 of volume46c in Siboga-Expeditie. Plates 21-32. Leiden: E. J.Brill.

Ely, C. A.1945. Shallow Water Asteroidea and Ophiuroidea of

Hawaii. Bulletin of the Bernice A. Bishop Museum,176(1942):l-63, 8 figures, 13 plates.

Engel, H., D. D. John, and G. Cherbonnier1949. The Genus Mithrodia Gray, 1940. Zoologisch Ver-

handelungen (Leiden), 2 (1949): 1-39, 12 figures.


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. Bulletinof the Bernice P. Bishop Museum, 27:63-87, plates5-8.

1928. Sea Stars from the Arcturus Oceanographic Expedi-tion. Zoologica, 8:487:493.

Gray, J. E.1840. A Synopsis of the Genera and Species of the Class

Hypostomata (Asterias, Linnaeus). Annals andMagazine of Natural History, series 1, 6:175-184,275-290.

1866. Synopsis of the Species of Starfish in the BritishMuseum. 17 pages, 16 plates. London: John VanVoorst.

Ives, J. E.1889. Catalogue of the Asteroidea and Ophiuroidea in

the Collection of the Academy of Natural Scienceof Philadelphia.

Lamarck, J. B.1816. Histoire naturelle des animaux san vertebres. Edi-

tion 1, volume 2, 568 pages. Paris.Leipoldt, F.

1895. Asteroidea der "Vettor-Pisani" Expedition (1882-1885). Mit Anhang: Die von F. Orsini im rothenMeere gesammelten Asteroideen. Zeitschrift wissen-schaften 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: Steller-ides. Memoire Societe physique Histoire naturelleGeneve, 29 (4): 1-84, plates 7-22.

Lutken, C.1859. Bidrag til Kundskab om de ved Kysterne af Mel-

lemog Syd-Amerika levende Arter af Sdstjerner.Videnskabelige Meddelelser fra den naturhistoriskeForening i Kjovenhavn, pages 25-96.

1864. Kritiske Bemaerkninger om forskjellige Sostjerner(Asterider), med Beskrivelse af nogle nye Arter.Videnskabelige Meddelelser fra den naturhistoriskeForening 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 Ma-rine and Freshwater Research, 2:712-715.

Mobius, K.1859. Neue Seesterne des Hamburger und Kieler Mu-

seums. 14 pages, 4 plates.Mueller, J., and F. H. Troschel

1840. [Not seen, title unknown.] Akademie Wissen-shaften. Monatsberichtc der Koniglichen Preuss.

1842. System der Asteriden. 134 pages, 12 plates. Braun-schweig: Friedrich Viewig und Sohn.

Perrier, E.1875. Revision de la collection de stellerides du Museum

d'Histoire Naturelle de Paris (Astcriidae, F.chinas-teridae, Ophidiastcridae). Archives de Zoologie Ex-perimentale et Generate, 4, 5: 384 pages.

1876. Revision dc la collection de Stellerides du Museumd'Histoire Naturelle de Paris. Archives de ZoologieExperimental et Ginerale, 5:1-104, 203-309.

1878. £tude sur la repartition geographiquc des Astcrides.Nouvelles Archives du Museum, (2): 1-108.

Sladen, P.1889. Report on the Asteroidea collected by H.M.S. Chal-

lenger. Pages 1-893 of volume 30 in Report on theScientific Results of HM.S. Challenger, Zoology. 117plates.

Stanek, V. J.1955. La Beaute de la nature. 373 pages, illustrated.

Prague: Artia.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 ConnecticutAcademy of Arts and Sciences, 1 (2):247-613, plates4-10.

Viguier, C.1878. Anatomie comparee du squelette des Stellerides.

Archives de Zoologie Experimentale de Genirale,7:33-250, 11 plates.

Ziesenhenne, F.1937. The Templeton Crocker Expedition: Echinoderms

from the West Coast of Lower California, the Gulfof California and Clarion Island. Zoologica, 22 (15):209-239, figure 2.

Fishes Collected at Malpelo Island

John E. McCoskerand Richard H. Rosenblatt

ABSTRACT

Seventy species of shore fishes are reported fromMalpelo Island, the majority of them previouslyunrecorded from that locality. Four species areundescribed and may be endemic to Malpelo. Nonew taxa are described. On the basis of these col-lections, it appears that the fauna is primarilyeastern Pacific mainland in composition, althoughfive species are shared only with the Galapagos andCocos islands.

The fish collections made by the expedition con-tain the first shore fishes taken by rotenone ichthy-ocides and diving at Malpelo. Previous collectionsof Malpelo fishes were limited to those caught byhook and line (Fowler, 1938, 1944) or dipnettedbeneath surface night lights (Clemens, 1957; Cle-mens and Nowell, 1963). Important material repre-senting several new species and many new localityrecords was obtained by the STRI expedition. Thecollections, however, were limited in scope and arenot sufficiently representative of the probabletotal ichthyofauna to allow any definitive state-ments concerning the faunal composition.

Malpelo occupies an important zoogeographiclocation in that it arises from deep water beyondthe continental shelf, and could provide a"stepping-stone" to the Galapagos Archipelago forshore fishes. The known fauna is mainly easternPacific mainland in origin. However, there arecertain similarities to both the Galapagos andCocos islands. This is evidenced by the presence

John E. McCosker, Steinhart Aquarium, California Academyof Sciences, Golden Gate Park, San Francisco, California94118. Richard H. Rosenblatt, Scripps Institution of Ocean-ography, La Jolla, California 92037.

of Mycteroperca olfax, Apogon atradorsatus,Eupomacentrus arcifrons, and E. beebei, speciesotherwise known only from those localities. Otherspecimens collected included fishes previouslythough to be Galapagos endemics, e.g., Labrisomusdendriticus and possibly a deep water species ofLythrypnus. The presence of certain reef-associatedIndo-Pacific species, e.g., Gymnothorax flavimar-ginatus, Zanclus canescens, and Acanthurus glau-copareius, is relatable to hydrographic and ecologi-cal conditions found on other eastern tropical Pa-cific offshore islands (Rosenblatt, McCosker, andRubinofF, 1972). Included in the Malpelo collec-tions were undescribed, and apparently endemic,species of Chriolepis, Acanthemblemaria and Axo-clinus, and an undescribed genus and species oftripterygiid.

The following annotated list of fishes from Mal-pelo Island is based primarily on those collectedby J. B. Graham and party at two localities, andunderwater photographs and sight records madeby C. Birkeland (pers. comm.). Fishes from thefirst locality (SIO 72-96) were collected along asloping basalt and sparsely covered Pocilloporabottom between the surface and 6 meters along thewestern 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 20meters. Fishes and photographs are deposited inthe Marine Vertebrates Collection of the ScrippsInstitution of Oceanography (SIO), University ofCalifornia at San Diego. Verified records fromhook and line and night-light stations are also in-cluded. The basis of identification is indicated bythe following abbreviations: * specimens collectedand deposited at sio, s sight record by STRI divingparty, p photographed underwater by C. Birke-land, 1 recorded by Fowler (1938), 2 recorded by

91


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. malpeloensisC. leucas (Muller and Henle) STriaenodon obesus Ruppell S

SPHYRNIDAE

Sphyrna sp. S

MURAENIDAE

Gymnothorax flavimarginatus (Riippell).P Abundant in theWest Indo-Pacific and Hawaii, and several eastern Pacificoffshore and nearshore localities (Rosenblatt, et al., 1972).

G. castaneus Jordan and Gilbert PG. dovii (Gunther)PMuraena lentiginosa Jenyns P

HOLOCENTRIDAE

Myripristis leiognathus Valenciennes • 3Holocentrus suborbitalis (Gill) S

KUHLIIDAE

Kuhlia taeniura (Cuvier) S.1,2

APOGONIDAE

Apogon atradorsatus Heller and Snodgrass.* Previouslyknown only from the Galapagos and Cocos islands.

SERRANIDAE

Epinephelus dermatolepis Boulenger •E. labriformis (Jenyns) •Paranthias furcifer (Valenciennes) *1Mycteroperca olfax (Jenyns).l Previously known from the

Galapagos, Malpelo, and Cocos islands. Phillip C. Heemstrahad kindly reexamined Fowler's specimens (ANSP 89121)and found their gill rakers and dorsal spine profile toagree with Galapagos specimens of M. olfax.

MULLIDAE

Pseudupeneus grandisquamis (Gill).4 Based on surface dip-netted prejuvenile specimens.

Mulloidichthys dentatus (Gill) S

CARANGIDAE

Selar crumenophthalmus (Bloch) 1Caranx caballus GiintherSC. vinctus Jordan and Gilbert 3

C. tnelampygus Cuvier and Valenciennes PElagatis bipinnulatis (Quoy and Gaimard) 2

LUTJANIDAE

Lutjanus Jordan; (Gilbert) l.PLutjanus 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 Gala-

pagos and Cocos islands.Microspathodon dorsalis (Gill) *

LABRIDAE

Thalassoma lucasanum (Gill) •Bodianus diplotaenia (Gill) !»•

SCARIDAE

Scar us rubroviolaceus BleekerS

KYPHOSIDAE

Sectator ocyurus (Jordan and Gilbert) SKyphosus sp. S

CHAETODONTIDAE

Holacanthus passer Valenciennes I.*Heniochus nigrirostris (Gill) PPomacanthus 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 fromthe Galapagos Islands, and an undescribed species fromPanama 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 SPrionurus laticlavius (Valenciennes) SZanclus canescens (Linnaeus) S

CANTHIGASTERIDAE

Canthigaster punctatissima (Giinther) •

TETRAODONTIDAE

Arothron meleagris (Bloch and Schneider) •

BALISTIDAE

Sufflamen verves (Gilbert and Starks) 1Melichthys niger (Bloch).l.P As M. buiva in Fowler, 1938.Canthidermis maculatus (Bloch). As C. sp. in Clemens, 1957.Cantherhines dumerilii (Hollard) SAlutera 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 TunaCruises, 1952 through 1959. California Fish andGame, 49:240-264.


Expedition, 1937. Monographs of the Academy ofNatural 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 NaturalSciences of Philadelphia, 6:57-529.

Rosenblatt, R. H., J. E. McCosker, and I. Rubinoff1972. Indo-West Pacific Fishes from the Gulf of Chiriqui,

Panama. Contributions in Science of the Los An-geles 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 thegobiid fish genus Chriolepis were collected duringthe 1972 Smithsonian Institution-U.S. Navy Expe-dition to Malpelo Island. The new species is hereindescribed, and its taxonomic relationships arebriefly discussed.

Introduction

METHODS.—Measurements were made with dialcalipers to the nearest 0.1 mm, and are expressedas percent of standard length (SL) in Table 9.Counts and measurements requiring explanationare as follows: Head length is distance from ante-rior margin of upper lip to upper attachment ofopercular membrane. Head depth is depth of headat the vertical, posterior margin of preoperculum.Head width is the maximum width between cheeks.Depth at anal fin origin is shortest distance be-tween base of anal spine and dorsal fin base. Post-orbital distance is the distance from posterior rimof eye to upper attachment of opercular mem-brane. Transverse scale rows is the number of scalerows, counted upward and forward, between thelast anal ray and dorsal fin base. The last dorsaland anal rays, branched to their bases, werecounted as one.

The holotype and paratype (the only knownspecimens) are deposited in the National Museumof Natural History (USNM), Smithsonian Institu-tion, Washington, D. C.

ACKNOWLEDGMENTS.—Thanks are due the follow-

Lloyd Talbott Findley, Department of Biological Sciences,University of Arizona, Tucson, Arizona 85721.

ing persons: C. E. Dawson of the Gulf Coast Re-search Laboratory, Mississippi, J. E. McCosker ofthe California Academy of Sciences, and D. A.Thomson of the University of Arizona for readingand improving the manuscript; J. B. Graham ofthe Smithsonian Tropical Research Institute forproviding collection data and editorial advice; andJenean Thomson of the University of Arizonafor the illustration. I thank C. E. Dawson and D. F.Hoese of the Australian Museum for encourage-ment, and R. J. Lavenberg of the Los AngelesCounty Museum of Natural History for examina-tion of the type-species. This paper is a contribu-tion from the University of Arizona Marine Bio-logy Program.

Chriolepis lepidotus, new species

FIGURE 35

HOLOTYPE.—USNM 211456, adult male, 30.0 mmSL, Malpelo Island, Colombia, east side of island,depth ca. 10 m, above the coral zone, cobble bot-tom with some calcareous sand and boulders, verysparse algal growth; collected by J. B. Grahamusing rotenone ichthyocide and SCUBA, 2-3 March1972.

PARATYPE.—USNM 211457, adult male, 29.1 mmSL; collected with holotype.

DESCRIPTION.— (See Table 9 for meristic and mor-phometric data). A robust species. Head plump,with wide cheeks, more or less rounded; depth atpreoperculum 0.71 (holotype) and 0.68 (paratype)of head width. Mouth oblique, forming an angleabout 40°-45° with body axis, the posterior anglereaching a point approximately under rear marginof pupil. Interorbital narrow, about one-third or

94

NUMBER 176 95

less of eye length. Pectoral base broad. Gill slitnarrow; opercular membrane attachments at upperand lower edges of pectoral base in advance of in-sertion of fin rays.

Gill rakers short on first arch, those on secondand following arches shorter; 3+8=11 on outerface of first arch; 9 on lower limb of second arch.Pseudobranchiae in four small tufts.

Head without pores or barbels. Sensory papillaeon head relatively large; in rows below eye, oncheek, at edge of preoperculum, on operculum, onsnout, and on dorsal and ventral surfaces of head

TABLE 9.—Counts and measurements of the holo-type and paratype of Ghriolepis lepidotus

Characters

Sex

Standard Length (mm)

COUNTS

Dorsal fin raysAnal fin raysPectoral fin rays (left/right)Pelvic fin raysSegmented caudal fin raysBranched caudal fin raysProcurrent caudal fin rays

(upper/lower)Gill rakers (outer face of first arch)Longitudinal scale rowsTransverse scale rows

PROPORTIONS (% of SL)

Head lengthHead depthHead widthBase of second dorsal finDepth at anal fin originPectoral fin lengthPelvic fin lengthCaudal fin lengthCaudal peduncle lengthCaudal peduncle depthEye lengthSnout lengthMouth lengthWidth between mandible tipsInterorbital widthWidth between posterior nostrilsLength between anterior &

posterior nostrilsPostorbital distance

Holotype(USNM211456)

30.0

VII-1,101,920/201,5

1715

5/511ca.3413i/2

33.315.521.828.516.827.320524.716.013.7

6.77.6

13.410.22.44.1

2.919.6

Paratype(USNM211457)

$29.1

VII-1,101,920/20131714

6/511

ca.3513i/2

32.215.623.527517524.721.023.513.613.86.26.9

125

10.11.95.1

2.319.1

including mandibular area, and similar to otherspecies in the genus (Figure 35). A low, shallowlyV-shaped, transverse, fleshy crest on head behindeyes. A low, short, fleshy "keel" extending mediallybetween nostrils. Nostrils tubular; anterior tubenarrow, its length about equal to pupil diameter;posterior tube wider, its length about one-half thatof anterior tube. Tongue tip truncate.

Teeth in rows in both jaws, relatively large,sharp, mostly caniniform, slightly to distinctly re-curved, mostly depressible and with faint yellowishapical 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 toanterior portion of jaw; (2) behind outer row an-teriorly is a patch of many small (about % to 14length of outer row teeth) closely set, less recurvedteeth in irregular rows, becoming fewer in number,more widely spaced, and confined to one row poste-riorly at side of jaw; (3) inner row of large, slender,recurved, well-separated teeth anteriorly, becom-ing more widely spaced and stouter posteriorly,until 2 to 3 much enlarged and recurved "fanglike"teeth at side of jaw, followed by a few, muchsmaller, closely set teeth. Upper jaw with (1) outerrow of large, slender, recurved, well-separatedteeth anteriorly and at side of jaw (not absentposteriorly as in outer row of lower jaw); (2) be-hind outer row anteriorly is a patch of many small,closely set, less recurved teeth in irregular rows,becoming fewer in number posteriorly; (3) innerrow of many small, closely set teeth, becoming morewidely spaced posteriorly. No teeth on vomer orpalatines.

First dorsal fin with seven flexible spines; basesof first five spines equally spaced; the last twospines widely separated from the first five, theirbases widely spaced. Third and fourth spines slight-ly the longest, but no spine notably longer thanany other, and none filamentous.

Second dorsal fin separated from first dorsal finby distance slightly more than eye length. Firstelement of both second dorsal and anal fins a flexi-ble spine; remaining elements are subequal, seg-mented, branched rays, with the last branched totheir bases.

Pectoral fins when adpressed reach past tips ofpelvic fins to level of end of spinous dorsal; allpectoral rays branched.


Origin of pelvic fins in advance of pectoral base.Pelvic fins short, not reaching anus; separated anddiverging, not united by a membrane into an ad-hesive disc as in most gobiids; interspinal mem-brane absent, but pelvic bases slightly connectedby a very low, inconspicuous frenum. Pelvic spineflexible; segmented pelvic rays well developed andbranched, most with three main branches, theinnermost ray branching less than others, but notreduced or splinted to the fourth. Fourth rayslightly the longest, then the third, then the fifth(which is almost equal to the third). Caudal finshort, rounded.

Urogenital papilla of male long, slender, atten-uate; with a few scattered melanophores.

SQUAMATION.—Four enlarged strongly ctenoidbasicaudal scales (some lost from both specimens);the outer two (at upper and lower angles of caudalfin base) slightly narrower and longer than theinner two. Body extensively scaled. Scales alongmidline extending forward from hypural to nearaxil, beneath origin of spinous dorsal fin (Figure35), in about 34-35 more or less irregular longitu-dinal rows; difficult to enumerate (some scales lostfrom both specimens, but scale pockets usuallydiscernible). Fewer scales in anterior rows; in awedge-shaped pattern as approaching axil. Trans-verse scales difficult to enumerate, about 13i/£in an irregular row (first scale at base of last analray very narrow in comparison to scales above it,counted as i/2 scale).

Posterior scales large, with prominent ctenii; be-

coming progressively smaller, thinner, more em-bedded, and with fewer and shorter ctenii, untilcycloid anteriorly. Ctenii becoming progressivelyrestricted to central portion of posterior edge ofeach scale proceeding anteriorly, until completelylost at about twelfth scale along midline in ad-vance of hypural. Anteriormost scales cycloid,minute, embedded, and difficult to see.

A naked area beginning near base of sixth dor-sal spine, equivalent to one scale width from baseof sixth spine to below base of fifth spine, to twoscale widths below fourth spine, becoming widerand sloping downward and forward to near uppermargin of pectoral base. A lower naked area fromunder pectoral base sloping downward and back-ward to near midline of venter, continuing as anarrow strip around anus to about base of fourthanal ray. Head, nape, chest, and venter scaleless.

COLORATION IN ETHYL ALCOHOL.—Backgroundcolor of head and body brownish yellow. Headwith a number of prominent black spots, especiallylaterally and ventrally. An elongate spot at mid-posterior edge of eye (diffuse in paratype); onebehind eye at dorsal edge of cheek near end of lowtransverse dorsal crest; one at dorsal margin ofpreopercular-opercular junction; one at midante-rior edge of operculum which begins a row ofabout 9 or 10 widely spaced spots lying in pre-opercular-opercular groove, following curvature ofgroove downward and forward and onto the imme-diately adjacent branchiostegal membrane, thencontinuing 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. Therow 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 bya shorter row of about 5 spots beginning at lowerposterior edge of preoperculum, running forwardbelow lower jaw to near symphysis, the spots lyingclose to and opposite the spots of row describedabove. About 5-8 widely spaced spots scattered overcheek (more prominent on paratype). A spot (onparatype) at posterior end of interorbital, fol-lowed by profuse minute melanophores on the lowtransverse dorsal crest. A spot at posterior dorsalcorner of operculum. A prominent spot on upperand lower angles of outer pectoral fin base; a moredistal irregular spot at bases of fifth to eighth pec-toral fin rays (slightly larger on paratype). A smallspot on inner pectoral fin base near upper margin.

A black bar below anteroventral edge of eye,passing obliquely downward across suborbital andjaws, terminating in anteriormost ventral spot ofthe series described above. A fainter dark bar ex-tends from midventral edge of eye downward acrosssuborbital region, and curves around end of lowerjaw. Snout, nostrils, upper jaw, and anterior por-tion of lower jaw densely peppered with minutemelanophores.

Two wide, transverse, light brown bars acrossdorsal surface of head, the posterior is across thenape and bisected at either side by pectoral fin.Five wide, vertical, light brown bands on body,most noticeable dorsally as dark saddles, becomingmore diffuse ventrally, and terminating near mid-ventral line. Brownish yellow interspaces betweenbands with a few scattered melanophores. Eachband with one or two adjacent, prominent, blackspots at midline. Anteriormost body band crossesdorsum between first and fifth dorsal spines, andpasses downward behind pectoral fin; second bandbetween sixth spine and end of spinous dorsal fin;third band between second and fifth dorsal rays;fourth band between seventh and ninth rays; fifthband behind end of second dorsal fin on anteriorportion of caudal peduncle. Dorsal segments ofbands 3 and 4 appearing as darker blocks offsetposteriorly from their ventral counterparts belowmidline, with dorsal block of band 3 most notice-able and lying over the interspace between thirdand fourth ventral blocks. An irregular dark blotchbehind fifth band at dorsal edge of caudal pe-duncle, adjacent to first procurrent ray. A narrowvertical bar of darker pigment underlying basicau-dal row of specialized ctenoid scales.

Dorsal fins with about four or five oblique rowsof dark lines, noticeable as prominent black spotson spines and rays (membranes torn in both speci-mens). A dark horizontal bar on anal fin mem-brane, running the length of fin near its edge. Afew faint diffuse spots on caudal rays suggest threeor four vertical lines. Pectoral fins slightly duskyon interadial membranes. Pelvic fins dark.

Chest and venter whitish, with numerous minutemelanophores, appearing "peppered" on micro-scopic inspection. Inside of gill chamber with adark horizontal stripe ventrally, in advance oflower pectoral base; a few melanophores dorsallyin area surrounding pseudobranchial tufts.

RELATIONSHIP.—The general head shape, rela-tively high number of fin rays, and general colorpattern suggest a relationship with a species com-plex of Chriolepis, most of whose members in theeastern tropical Pacific are undescribed. Amongthe described species, C. lepidotus is perhaps clos-est to C. minutillus Gilbert, 1891, and C. tagusGinsburg, 1953 (judging by original descriptiononly). Complete discussion of relationships withinthe genus Chriolepis and allied genera of seven-spined gobiids with separate pelvic fins is deferredpending completion of a revision in progress bythe author of all known species in the eastern trop-ical Pacific.

COMPARISONS.—Chriolepis lepidotus differs fromother described congeners in the eastern tropicalPacific in the extent of its squamation; its scalesextend forward to beneath the origin of the spi-nous dorsal fin. Among the other described species,C. minutillus Gilbert, 1891 (Gulf of California),most closely approximates this condition; its scalesextend forward to beneath or slightly in advanceof midspinous dorsal. From the description of C.tagus Ginsburg, 1953 (Galapagos Islands), and inC. zebra Ginsburg, 1938 (Gulf of California),scales only extend forward to beneath the end ofthe spinous dorsal, or slightly posterior to this posi-tion in the latter species.

The new species also differs from other describedeastern tropical Pacific congeners in having thehighest number of pectoral fin rays (20 vs. 19 inC. tagus; 17 or 18 in C. zebra; 15 or 16 in C.minutillus.) The prominent black spots on thehead further serve to distinguish C. lepidotus fromthe other species described from these waters.

Chriolepis lepidotus further differs from C. min-


utillus in having a larger mouth (posterior angleextends to below posterior margin of pupil vs. tobelow midpupil in C. minutillus), shorter pelvicfins (when adpressed, not reaching anus vs. reach-ing or almost reaching anus in C. minutillus), andmore 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 inhaving a rounded rather than a depressed head, alarger mouth (posterior angle extends only to be-low midpupil in C. zebra), and more second dorsalfin rays (11 vs. 10 in C. zebra).

Chriolepis lepidotus differs from the descriptionof C. tagus in having fewer second dorsal fin rays(12 in C. tagus), and fewer anal fin rays (10 vs. 11in C. tagus).

DISTRIBUTION.—Chriolepis lepidotus is presentlyknown only from Malpelo Island, Colombia.

ETYMOLOGY.—From the Greek lepidotos (scaly),referring to its squamation, the most extensivelyknown in the genus.

REMARKS.—The genus Chriolepis was erected byGilbert (1891:557-558) for the species C. minutil-lus, which he described as being totally scaleless onthe basis of a single specimen trawled by the Alba-tross in the Gulf of California. His description wasin error, since the holotype retains a single basi-caudal scale, as mentioned by Ginsburg (1938:111;1953:21), and confirmed by examination of thetype by D. F. Hoese and R. J. Lavenberg (pers.comm.). Recently collected material of C. minutil-lus under study by the author shows that, indeed,squamation is extensive in this species, less so thanin C. lepidotus, but more so than in C. zebra, forexample.

Species of Chriolepis are sublittoral in habitatand secretive, hiding under rocks and in crevices.Until recently, few specimens were known, butwith the introduction of SCUBA and rotenone ich-thyocide collecting techniques by diving ichthyo-logists, specimens have begun to accumulate. Thesewill allow for a comprehensive systematic, zoogeo-graphic, and evolutionary treatment of the genusand its allies (briefly treated by Bohlke and Ro-bins, 1968:129-131) in the eastern tropical Pacific.There remain about five undescribed species ofChriolepis in these waters.

Literature Cited

Bohlke, J. E., and C. R. Robins1968. Western Atlantic Sevcn-spined Gobies, with De-

scriptions of Ten New Species and a New Genus,and Comments on Pacific Relatives. Proceedings ofthe 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 FishesCollected in 1888 and 1889, Principally among theSanta Barbara Islands and in the Gulf of Cali-fornia. Proceedings of the United States NationalMuseum, 14 (880):539-566.

Ginsburg, I.1938. Eight New Species of Gobioid Fishes from the

American Pacific Coast. Allan Hancock PacificExpeditions, 2(7):109-121.

1953. Ten New American Gobioid Fishes in the UnitedStates National Museum, Including Additions to aRevision of Gobionellus. Journal of the WashingtonAcademy of Sciences, 43(1): 18-26.

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The Biological Investigation of Malpelo Island, Colombia

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