the systematics of neotropical orbweb spider 1863

[Begin Page: Page 1] rHE SYSTEMATICS OF NEOTROPICAL ORB-WEAVING SPIDERS INrHE GENUS METEPEIRA ARANEAE: ARANEIDAE... -Cambridge 5 ^ey to Female Metepeira 12 [ey to Male Metepeira 17 letep

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[Begin Page: Cover]

[Begin Page: Verso]

Orb-weaving Spiders in the

Genus Metepeira (Araneae: Araneidae)

JoHNSONiA, Department of Mollusks, 1941-1974

Occasional Papers on Mollusks,

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

Lyman, C P., and A R Dawe (eds.), 1960 Proceedings of the First ternational Symposium on Natural Mammalian Hibernation {Bulletin

In-of the M C Z, Vol 124.)

Orinthological Gazetteers of the Neotropics (1975-)

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

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

Proceedings of the Boston Society of Natural History.

Price hst and catalog of MCZ publications may be obtained from tions Office, Museum of Comparative Zoology, Harvard University, Cambridge,Massachusetts 02138, U.S.A

Publica-This publication has been printed on acid-free permanent paper stock

©The President and Fellows of Harvard College 2001

[Begin Page: Page 1]

rHE SYSTEMATICS OF NEOTROPICAL ORB-WEAVING SPIDERS INrHE GENUS METEPEIRA (ARANEAE: ARANEIDAE)

Collections Examined 3

Locality' Data Storage and Manipulation 4

Examination and Illustration 4

{etepeira F O P -Cambridge 5

^ey to Female Metepeira 12

[ey to Male Metepeira 17

letepeira foxi Group 19

1 Metepeira datona Chamberlin and Ivie 20

2 Metepeira desenderi Baert 21

3 Metepeira grandiosa grandiose

Chamberlin and Ivie 23

4 Metepeira grandiosa alpina

Chamberlin and Ivie 24

(etepeira vigilax Group 26

5 Metepeira cajahamba New Species _ 26

6 Metepeira gloinerabilis (Keyserling) _ 28

7 Metepeira vigilax (Keyserling) 30

8 Metepeira rectangula (Nicolet) 32

{etepeira labt/rinthea Group 33

9 Mete})eira spinipes F O P -Cambridge 34

10 Metepeira lacandon New Species 37

letepeira nigriventris Group 38

11 Metepeira nigriventris (Taczanowski) 38

12 Metepeira tarapaca New Species 40

13 Metepeira calamuehita New Species 42

14 Metepeira galatheae (Thorell) 43

15 Metepeira karkii (Tullgren) 46

(etepeira compsa Group 47

16 Metepeira compsa (Chamberlin) 48

17 Metepeira roraima New Species 53

18 Metepeira gressa (Keyserling) 54

(etepeira incrassata Group 56

19 Metepeira tnatja New Species 56

20 Metepeira inca New Species 58

' Museum of Cvomparative Zoology, Hanard

Uni-?rsity, Cambridge, Massachusetts 02138 Currentddress: Institute of Evolutionaiy and Ecological Sci-nces, Leiden University, 231 1 GP Leiden, Theletherlands; pielCa'rulsfb.leidenuniv.nl

21 Metepeira gosoga Chamberlin and Ivie

59

22 Metepeira ohnec New Species 60

23 Metepeira comanche Levi 62

24 Metepeira piinungan New Species 62

25 Metepeira triangularis (Franganillo) 63

26 Metepeira arizonica Chamberlin and

Ivie 66

27 Metepeira atascadero New Species 67

28 Metepeira incrassata F O

P.-Cambridge 68

Metepeira venttira Group 71

29 Metepeira ventura Chamberlin and

Ivie 71

30 Metepeira revillagigedo New Species 73

31 Metepeira celestun New Species 74

32 Metepeira uncata F O P-Cambridge 76

33 Metepeira crassipes Chamberlin and

35 Metepeira petatlan New Species 80

36 Metepeira minima Geitsch 82

37 Metepeira pacifica New Species 84

38 Metepeira jamaicensis Archer 86

Literature Cited 88

Index 91

Abstract Of the 39 species and three subspecies

of the orb-weaver genus Metepeira in the Americas,

36 species and two subspecies are knowni to occuroutside of the U.S and Canada Yet, despite theirconspicuous webs, diurnal foraging, and rclatixeK'common presence, the taxononi) of Metcj)iira ispoorly understood, probably because tlie genitalia aresmall and difficult to distinguish In fact, man\ namesfor species south of the U.S were, at some time, in-correctly synonymized with die name Metepeira la-bijrintliea In this paper, 14 new species are named(Metepeira atascadero, M cajabamba, M calamtichi-

ta, M celestun, M inca, M lacandon, M mai/a, M.olmec, M pacifica, M petatlan, M pinuingan, M re-villagigedo, M roraima M tarapaca); 11 new junior

Bull Mus Conip ZooL, 157(1): 1-92, June, 2001 1

[Begin Page: Page 2]

2 Bulletin Museum of Comparative Zoology, Vol 157, No 1

synonyms are reported (M acostai, M bani, M dom- [1989]), but this practice can lead tO

trou-inicana, M grinnelU, M latigijna, M perezi, M san- ^^^e 111 one case, the behavior of several

ta, M salei, M seditiosa, M vaurieorum, M virgi- ^.nr • ••j.-ii j.i-i

.so r 1 A ^ diiierent species was initially studied

un-nensis); hve cases oi erroneously synonymizeu names i i r i i i

are reversed; 22 species and two subspecies are re- der the false assumption that they all

be-described (M arizonica, M triangtdaris, M chilapae, longed to the same species (e.g., Uetz et

M Comanche, M compsa, M crassipes, M datona, ^L, 1982) Clearly, a strong taxonomic

M desenderi, M galatheae, M glomerabilis, M, go- foyj^^ation is important for further

biolog-saga, M grandiosa alpma, M grandiosa grandiosa, • i i

M gressa, M incrassata, M jamaicen.sis, M korkii, Icai WOlK.

M minima, M nigriventris, M rectangula, M spi- Ultimately, the relatively small,

indis-nipes, M uncata, M ventura, M vigilax); and a key tinct genitalia and the relatively

homoge-to all Metcpeira species is presented In addition, sev- ^^^^^ abdominal patterns are homoge-to blame for

eral ecological and life history observations are re- ,, i ^ i ii rn*

.AC- the weakness m our knowledge oi

Mete-ported tor vanous species r i

peira taxonomy Many or these species are

^-^1 ip-ri^Ki undoubtedly hard to distinguish, and this

fact has surely intimidated arachnologists

The absence of a comprehensive revi- from taking on the painful task of revising

sion of Neotropical Metepeira has left the the group In the absence of good

distin-taxonomy of this group in shambles Over guishing characteristics, the catalogs of

the years, a fair number of species have Bonnet (1957) and Roewer (1942)

synon-been named, particularly by A F Archer, ymized the names of many Neotropical

R V Chamberlin, and W Ivie However, species with the name Metepeira

labyrin-these efforts have been sporadic and, for then Levi's (1977) revision of Nearctic

the most part, scant For example, the de- species observes that M lahijhnthea is

ac-scription of Metepeira dominicana (Ar- tually limited to the eastern United States,

cher, 1965) provides little information oth- One task in this revision consists of

reas-er than "form typical of Metepeira in all sreas-erting the names of species that wreas-ere

im-respects," a few measurements, and two properly synonymized and clarifying the

unrecognizable figures Even when species diagnostic characters that are needed to

are properly described they have far less identify them,

taxonomic value when published alone, in

the absence of a full comparative revision ACKNOWLEDGMENTS

The poor understanding of Metepeira This paper is part of my Ph.D thesis for

taxonomy has persisted despite great eco- the Department of Organismic and logical and behavioral interest in this ge- lutionary Biology, Harvard University I amnus Indeed, many species are obligate or indebted to many people for their help,facultative social species and offer excel- assistance, and encouragement in this pro-lent models for investigating genetic and ject I am especially thankful for the ded-

Evo-environmental factors that influence colo- ication and support of my advisors,

Her-ny formation (e.g., Uetz and Cangialosi, bert W Levi and Edward O Wilson I am1986; Uetz et al., 1987) The monumental grateful that my colleagues in the Depart-work carried out over many years by G W ment of Invertebrate Zoology providedUetz has made great strides in our under- such a pleasant place to work: Edwardstanding of gregarious social behavior in Cutler, Ardis Johnston, Laura Leibensper-spiders and in risk-sensitive foraging the- ger, Damhnait McHugh, Diana Sherry,ory in general (e.g., Uetz, 1996) Still, in Van Wallach, and Dee Woessner, amongthe absence of solid taxonomic literature, others

behavioral ecologists have been forced to Field collecting and new specimen

ac-apply informal names to their study ani- quisitions were made possible with the

mals (e.g., Metepeira "atascadero" in Uetz help of Gita Bodner, Fundacion Capacitar,

[1989] or Metepeira "Species A" in Viera Tim Coonan (CINP), Fred Coyle, Dawn

[Begin Page: Page 3]

Metepeira • Piel

Fitzpatrick, Germania Jacome, Antdnia

Monteiro, Tila Perez, George Putnam,

Linda Rayor, Grace Smith (NAWF), and

George Uetz I am particularly indebted to

George Uetz for his assistance and

I am thankful for the comments by

those who read this paper — especially tothe members on my thesis committee: H

W Levi, N E Pierce, and E O Wilson

I am also indebted to Kathy Horton forher help in formatting and preparing themanuscript and to the Colles Fund for de-fraying the costs of publication Curators

at various institutions who lent me mens are listed in the Materials and Meth-ods section I cannot overstress the value

speci-of museum collections and expert curators,without which research in taxonomy wouldnot be possible Museum collections arethe most important tools available for un-derstanding biodiversity

MATERIALS AND METHODS

Collections Examined The taxonomicrevision was carried out on specimens bor-rowed from the following collections Theabbreviations correspond to those listedwith each record after every species de-scription I am grateful to the museums,curators, and staff that graciously loanedthe material

ADC A Dean, Texas A&M University,

College Station, Texas, United

States

AMNH American Museum of NaturalHistory, New York, United

States; N Platnick, L Sorldn

BMNH Natural History Museum, don, England; P Hillyard

Lon-CAS California Academy of Sciences,

San Francisco, California,

Ar-United States; G B Edwards

IRSNB Institut Royal des Sciences

Na-turelles de Belgique, Bnjssels,Belgium; L Baert

JAK J A Kochalka, Ciudad

Univer-sitaria, Paraguay

JEC J Carico, Lynchburg, Virginia,

United States

JMM J Maes, Leon, Nicaragua

MACN Museo Argentino de CienciasNaturales, Buenos Aires, Argen-tina; E A Maury, C L Scioscia

MCN Museu de Ciencias Naturais,Fundagao Zoobotanica do RioGrande do Sul, Porto Alegre,

Rio Grande do Sul, Brazil; E H.Buckup, M A L Marques

MCZ Museum of Comparative

Zool-ogy, Harvard University, bridge, Massachusetts, UnitedStates; H W Levi

Cam-MECN Museo Ecuatoriano de CienciasNaturales, Quito, Ecuador; Ger-

mania Estevez Jacome

MEG M E Galiano, Buenos Aires,Argentina

MLJC Maria Luisa Jimenez, Centre deInvestigaciones Bioldgicas del

Noroeste, La Paz, Mexico

MLP Museo de Universidad Nacional,

La Plata, Argentina; R F

MZSP Museu de Zoologia,

Unixersida-de Unixersida-de Sao Paulo, Sao Paulo, SP,

Brazil; P Vanzolini, | L Leme

MZUF Museo Zoologico de "La

Spe-cola" Universita di Firenze,

Florence, Italy; S Whitman

NRMS Naturhistoriska Riksmuseet,

Stockholm, Sweden; T

Krones-tedt

PAN Polska Akademia Nauk,

Warsza-[Begin Page: Page 4]

4 Bulletin Museum of Comparative Zoology, Vol 157, No 1

wa, Poland; J Proszynski, A

Slo-jewska, W B Jedryczkowski

REL R E Leech, Edmonton,

Alber-ta, Canada

SMF Forschungsinstitut

Sencken-berg, Frankfurt am Main,

Ger-many; M Grasshoff

SR Susan Riechert, Knoxville,

Ten-nessee, United States

USNM National Museum of NaturalHistory, Smithsonian Institution,

Washington, D.C., United

States; J Coddington, S F

Larcher

ZMB Zoologisches Museum der

Humboldt Universitat, Berlin,

Manipula-ing Claris FileMaker Pro® Geographiccoordinates were added to locality datathat lacked them using maps, USBGN gaz-etteers, and on-line databases (http://164.214.2.59/gns/html/ and http://mapping.usgs.gov/www/gnis/) Occasionally localityinformation was illegible or unknown orone of several homonymous sites In suchcases a reasonable, educated guess wasmade and a "[?]" designation was append-

ed to the locality In some cases the erary of a collector was reconstructed fromother known records, and the ambiguouslocality was assigned a coordinate halfwaybetween the previous and following knowncollection sites The locality databaseworked in concert with the mapping pro-gram Atlas Pro® to generate thematicmaps on the fly These maps helped in theprocess of delimiting species and discov-ering cryptic species

itin-Elevation (in meters) was estimated foreach locality that lacked this information

In some cases, elevation was estimated

us-ing contour maps, such as DMAAC ONCaeronautical maps; in most cases, elevation

was estimated using NOAA data with anon-line database server (http://phylogeny.harvard.edu/~piel/find.html).

The enhanced locality database was

used to reveal ecological and life historytraits Seasonality of species was expressed

by plotting a circular histogram showingthe relative amount of collecting activityper 5-day interval (Figs 300-337) Whilelocality dates alone cannot control for theseasonal activity of human collectors, thesedata at least provide an estimate of spiderseasonal abundance, if only approximate.Some sympatric species show incongruousseasonal abundance, which is at least someevidence that seasonality of spider collec-tors does not unduly overshadow the sea-sonality of the spiders themselves

Examination and Illustration

Speci-mens were examined under 80% ethanol

in a dish with light and dark sand grainsfor specimen support Digital photographs

of preserved specimens were taken

through a Nikon SMZ-10 scope using a Panasonic WV-CL320 CCDvideo camera, chosen for its high sensitiv-ity to light Video images were captured

photomicro-using a Quicklmage®24 digitizer and

ed-ited on a Quadra 700 Macintosh®

com-puter The comcom-puter allows relatively

in-expensive pictures to be printed rapidly on

a 1,200 dpi Xante® Accel-a-Writer 8200

laser printer Digital pictures were used to

help sort out individuals to species, to

cre-ate publishable pictures of gross dorsal

and ventral markings, and to aid in the

il-lustration of genitalia As an aid in

illustra-tion, the digital pictures functioned as a

camera lucida because they assured

accu-racy when drawing the proportions of

gen-ital parts and sclerites Usually a diggen-ital

picture was laid over carbon paper and an

outline of the genitalia was transferred to

coquille board underneath The

illustra-tion continued on the coquille board using

a Staedtler OmniChrom® pencil and a

drafting pen with India ink and then was

scanned at 600 dpi on a LaCie

Silverscan-[Begin Page: Page 5]

Metepeira • Piel

ner II® The resulting digital image was