Table Of ContentPELEGRINA FRANGANILLO AND OTHER JUMPING SPIDERS
FORMERLY PLACED IN THE GENUS METAPHIDIPPUS
(ARANEAE: SALTICIDAE)
WAYNE P. MADDISON'
CONTENTS
Abstract 216 2. Pelegrina proxima (G. & E. Peckham,
Introduction 217 1901) 265
Acknowledgments 218 3. Pelegrina dithalea new species 268
Materials and Methods 219 4. Pelegrina edrilana new species 269
Explanation of Morphological and Behavioral 5. Pelegrina proterva (Walckenaer, 1837) __ 270
Terms 222 6. Pelegrina peckhamorum (Kaston,
The Subfamily Dendryphantinae 226 1973) 272
Relationships within the Dendryphantinae 231 7. Pelegrina neoleonis new species 273
The Bagheera Group of Genera 232 8. Pelegrina tristis new species 274
Two Genera That Have Exchanged Species 9. Pelegrina sabinema new species 275
with Metaphidippus: Dendryphantes 10. Pelegrina pervaga (G. & E. Packham,
and Beata 235 1909) 276
The Proper Placements of Metaphidippus 11. Pelegrina kastoni new species 277
Species 237 12. Pelegrinaflavipedes (G. & E. Peckham,
Phanias F. P.-Cambridge, 1901 238 1888) 278
Terralonus new genus 239 13. Pelegrinaflaviceps (Kaston, 1973) 279
Ghelna new genus 239 14. Pelegrina exigua (Banks, 1892) 281
The Genus Pelegrina Franganillo, 1930 240 15. Pelegrina montana (Emerton, 1891) 283
Phylogeny within Pelegrina 245 16. Pelegrina insignis (Banks, 1892) 284
Identifying Species of Pelegrina and the 17. Pelegrina chaimona new species 286
Metaphidippus niannii Group 247 18. Pelegrina clemata (Levi & Levi, 1951) .. 287
Key to the Males of All Species of Pelegrina 19. Pelegrina aeneola (Curtis, 1892) 288
and Those Metaphidippus mannii 20. Pelegrina halia new species 290
Group Species Occurring in the United 21. Pelegrina chalceola new species 291
States 248 22. Pelegrina jurcata (F. P.-Cambridge,
Key to the Female Pelegrina of the Eastern 1901) 292
United States and Canada (East of the 23. Pelegrina volcana new species 294
Mississippi River and Manitoba) 257 24. Pelegrina bicuspidata (F. P.-Cam-
Key to the Female Pelegrina of the Great bridge, 1901) 295
Plains (between the Rocky Mountains 25. Pelegrina ochracea (F. P.-Cambridge,
and the Mississippi River) 258 1901) 295
Key to the Female Pelegrina of the Pacific 26. Pelegrina morelos new species 296
Coast of the United States and Western 27. Pelegrina huachuca new species 296
Canada 258 28. Pelegrina arizonensis (G. & E. Peck-
Key to the Female Pelegrina and mannii ham, 1901) 297
Group of Arizona 259 29. Pelegrina helenae (Banks, 1921) 298
Key to the Pelegrina and Nagaina females 30. Pelegrina verecunda (Chamberlin &
of Mexico and Central America 260 Gertsch, 1930) 299
Descriptions of the Species of Pelegrina — 262 31. Pelegrina clavator new species 300
1. Pelegrina galathea (Walckenaer, 1837) 263 32. Pelegrina pallidata (F. P.-Cambridge,
1901) 301
33. Pelegrina variegata (F. P.-Cambridge,
' Department of Ecology and Evolutionary Biol- 1901) 302
ogy, University of Arizona, Tucson, Arizona 85721. 34. Pelegrina yucatecana new species 303
Bull. Mus. Comp. Zool., 154(4): 215-368, January, 1996 215
52
216 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4
35. Pelegrina sandaracina new species 304 cussed, and the harfordii group is transferred to the
36. Pelegrina tillandsiae (Kaston, 1973) 305 genusPhanias F. P.-Cambridge, whichappearstobe
37. Pelegrina bunites new species 306 relatively distantly related to most other dendry-
38. Pelegrina orestes new species 307 phantines. The following new combinations are es-
The Genus Nagaina G. & E. Peckham, 1896 ... 308 tablished: Phanias albeolus (ChamberHn & Ivie),
39. Nagaina incunda G. & E. Peckham, Phanias concoloratus (Chamberlin & Gertsch),
1896 308 Phanias dominatus (Chamberlin & Ivie), Phanias
Species of the mannii Group of the United furcifer (Gertsch), Phanias furcillatus (F. P.-Cam-
States and Canada 310 bridge), Phanias harfordii (G. & E. Peckham), Phan-
40. Metaphidippus mannii (G. & E. Peck- iasmonticola (Banks),Phaniasneomexicanus(Banks),
ham, 1888) 310 and Phanias ivatonus (Chamberlin & Ivie). Also re-
41. Metaphidippus diplacis (ChamberHn, moved from Metaphidippus are the mylothrus and
1924) 31 castaneus groups, for which the new genera Terra-
42. Metaphidippus tricolor ChamberUn & lonus and Ghelna are described, thus yielding the
Ivie, 1941 314 new combinations Terralonus californicus (G. & E.
43. Metaphidippus chera (ChamberHn, Peckham), Terralonus mylothrus(Chamberlin), Ter-
1924) 31 ralonus unicus (Chamberlin & Gertsch), Terralonus
44. Metaphidippus carmenensis (Chamber- shaferi (Gertsch & Riechert), Terralonus versicolor
Hn, 1924) 316 (G. & E. Peckham), Terralonusvittatus(Banks), Ter-
45. Metaphidippus emmiltus new species .... 317 ralonusfraternus (Banks), Ghelna castanea (Hentz),
Literature Cited 318 Ghelna barrotvsi (Kaston), Ghelna sexmaculata
Index 322 (Banks), and Ghelna canadensis (Banks). The new
combination Sassacus paiutus (Gertsch) is estab-
lished. Thevitis groupisretained in Metaphidippus.
Abstract. The genus Pelegrina FranganiUo con- The limits of the genera Dendryphantes C. L. Koch
tains 38 species of dendryphantine jumping spiders and Beata G. & E. Peckham are also reconsidered.
from NorthandCentral America that were formerly The combination Dendryphantes nigromaculatus
placed in the genus Metaphidippus F. O. Pickard- Keyserlingisrevivedandthefollowingcombinations
Cambridge. The close relatives of the Dendryphan- established: Beata hispida (G. & E. Peckham), Beata
tinae may include the Europhryinae and several inconcinna (G. & E. Peckham), Beata maccunii (G.
smaller groups, for they share an embolus that is & E. Peckham), and Beata rustica (G. & E. Peck-
coiledcounterclockwise (leftpalp)andseparatedfrom ham). Dryphias (G. & E. Peckham) is synonymized
thetegulumbyafullyexpandablehematodocha. The with Beata.
subfamily Dendryphantinae itselfisdelimitedbythe The largest group removed from Metaphidippus
derived conditions of a carina on the underside of is placed in the genus Pelegrina FranganiUo, 1930,
the male chelicera, the coil of the embolus folded whose species are, with some exceptions, distin-
back so as to be hidden behind the base of the em- guished from otherdendr\phantines b\ thepresence
bolus, and S-shaped epig\nal openings. oftwoterminalramiretrolateraltotheembolusopen-
Within the subfamily, generic relationships are ing, an embolic hematodocha that bulges distally,
poorly understood, but it is clear that the genus Me- wrinklesontheanteriormarginofthemalecheliceral
taphidippus is polyphyletic. The genus should in- fang, a distinct band of pale scaleson the side of the
clude at most a few species closely related to the face, and male courtship with the first legs held low
neotropical genera Messua G. & E. Peckham, Bag- and forward. The following species are moved into
heera G. & E. Peckham, and Gastromicans Mello- Pelegrina: P. aeneola (Curtis), P. arizonensis (G. &
Leitao. Gastromicansisremovedfromsynonymywith E. Peckham), P. bicuspidata (F. P.-Cambridge), P.
Beata G. & E. Peckham. The following new com- clemata (Levi & Levi),P.exigua (Banks),P.flaviceps
binations are established for species in this group: (Kaston), P.flavipedes (G. & E. Peckham), P.furcata
Bagheera prosper (G. & E. Peckham), Messua cen- (F. P.-Cambridge), P. galathea (Walckenaer), P. he-
tralis (G. & E. Peckham), Messua dentiger (F. P.- lenae (Banks), P. insignis (Banks), P. montana
Cambridge), Messua donalda (Kraus), Messua lata (Emerton), P. ochracea (F. P.-Cambridge), P. palli-
(Chickering), Messua laxa (Chickering), Messualim- data (F. P.-Cambridge), P. peckhamorum (Kaston),
bata(Banks),Messuamoma(F. P.-Cambridge),Mes- P. pervaga (G. & E. Peckham), P. proterva (Wal-
sua octonotata (F. P.-Cambridge), Messua pura ckenaer), P. proxima (G. & E. Peckham), P. tilland-
(Bryant), Messua tridentata (F. P.-Cambridge), Gas- siae (Kaston), P. variegata (F. P.-Cambridge), andP.
tromicans albopilosa (G. & E. Peckham), Gastromi- verecunda (Chamberlin & Gertsch). Pelegrina prox-
cans hondurensis (G. & E. Peckham), Gastromicans ima is shown to be a senior synonym of Pelegrina
levispina (F. P.-Cambridge), Gastromicans noxiosa geniciilata FranganiUo,thelatterbeingthetypesspe-
(Simon), and Gastromicans vigens (G. & E. Peck- cies of Pelegrina. A neotype is designated for Attus
ham). The combination Messua desidiosa G. & E. galathea Walckenaer. Seventeen species are de-
Peckhamisrevived. Theproperplacementofvarious scribed as new: P. balia, P. bunites, P. chaimona, P.
groups currently assigned to Metaphidippus is dis- chalceola, P. clavator, P. dithalea, P. edrilana, P.
Pelegrina Jumping Spidkrs • Maddison 217
huaclntca. P. kastoni, P. morelos, P. neoleonis, P. about half of the nearly 300 species of sal-
ovroelsctaenso.,Pa.ndsaPb.iynuecmaat,ecPa.nas.anEditaorpalciirnyas,IcPu.cotrpihsatiesa, CP. ticids occurring north of Mexico (accord-
L. Koch, Icius crassivcnfer Ke>serling, am) Meta- ing to the count of Richman and Cutler,
phidippus digitatus F. P.-Cambridge are newly syn- 1978). In the last three decades, increased
onymized with P. galathea; Dendrijphantes uteanus interest in the family has resulted in re-
poCphhhaarnmytbseescrohmnnicom&luosGrerBCtahsnackhmsbweiistrhhrenP.moawveientehdolPfa.r;ofamunrdscayDtneao.nndyrEmyuy-- PvihsiidoinsppoufsH(aEbdrwoanradtst,uisn p(rGerpiasrwaotlido,n)1,98a7n)d,
with P. proterva and considered asenior synonym of other genera (Proszynski, 1968, 1971a,
Sittaciis cursor Barrows, yielding the new combina- 1973a, 1980; Cutler, 1981a, 1987; Rich-
tion Sitticus concolor. Identification keys are pre- man, 1981, 1989). However, except for
sented for all Pelegrina males and for females from works by Kaston (1973) on some eastern
restricted geographical regions. All species are de-
scribed and illustrated. Male/female associationswere species and by Cutler and Jennings (1985)
achieved for all species north of Mexico. Courtship on the arizonensis group, Metaphidippus
behavior is described for 22 species of Pelegrina, has remained unrevised, perhaps because
karyotypes for 10 species, and habitat information its poorly defined limits have made the
MfoeriTahmpoehsigtedsnipupespciuPeses.lmeagnrniniaimgraoyupb,eNcalgoasierliyarealnadt/eodrtoErtihs.e sscoompee. oWfhaenny 1refvirisstiobnegpaontenttoiarlelvyisetrMouebtlae--
Nagaina incunda G. & E. Peckham is described and phidippus, I knew that I would have to
illustrated; Dendryphantes vegettts G. & E. Peck- restrict the revision to only some of the
ham, Meiaphidippusflavolineatus F. P.-Cambridge, disparate groups placed there. The largest
asnydnoMneyimaipzheiddiwpiptuhsNe.xpailnlciudnatduas.FT.hPe.-sCpaemcbiresidogfeatrhee groupplaced in Metaphidippus,including
mannii group (temporarily retained in Meiaphidip- the species most commonly collected in
pus)thatoccurintheUnitedStatesarealsodescribed northern and eastern North America, was
and illustrated; two new combinations, Meiaphidip- chosen for revision and is here moved to
pus chera (Chamberlin) and Meiaphidippus car- the genus Pelegrina Franganillo.
meriensis (Chamberlin), are established; one species,
Meiaphidippus emmilius, is described as new; Den- The jumping spiders placed in Pelegri-
dryphanies versicolor G. & E. Peckham is synony- na are medium-sized dendryphantines
mized with Meiaphidippus mannii (G. & E. Peck- distributed throughout North America,
ham),andMeiaphidippusfranciscanusSchenkelwith with some species extending as far south
Meiaphidippus diplacis (Chamberlin). asPanama. The38speciesincludethewell-
known P. galathea, P. proterva, P. fla-
INTRODUCTION vipedes, and P. aeneola. Males of Pele-
grina are generally brown with white
For about 50 years after the Peckham's stripes (Fig. 1), and most can be distin-
(1909) revision of the jumping spider spe- guished from other dendryphantines by
cies north of Mexico, taxonomic work on the wide embolus with two rami retrola-
North American representatives of this teral to the opening (Fig. 3). The spotted
large family consisted mostly of scattered females (Fig. 2) have large thickened flaps
species descriptions by Chamberlin, over the epigynal openings (Fig. 4). Al-
Gertsch, Ivie, and others. Some generic re- though the eastern species were well stud-
visions consolidating and clarifying the ied by Kaston (1973), most of the species
previous work began appearing in the occur in the western United States, Mex-
1950s (Gertsch and Ivie, 1955; Barnes, ico, and Central America, and they re-
1955, 1958), but most genera remained ceivedtheirlastcomprehensivetreatments
untouched, including the three largest by G. & E. Peckham (1909) and F. O.
genera, Habronattus* Phidippiis, and Pickard-Cambridge (1901). Many of the
Meiaphidippus, which together include western species have been inadequately
described and illustrated, often from only
one sex, making identification almost im-
Authorsofscientificnamesaregivenintheindex. possible by anyone other than an araneol-
218 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4
ogist familiar with the group. Many spe- encouragement that allowed me to grow
cies in the southwest were undescribed, on my own. My other friends and col-
and for most species there is Httle pub- leagues—at Harvard and in our spider lab-
Hshed information on natural history. The oratory Leticia Aviles, Anna Weitzman,
present revision has as its main goal to Mark Moffett,JonathanCoddington, Mark
makethe species known, by describing and Stowe, Jackie Palmer, Cecile Villars, John
illustrating them, their courtship displays, Hunter, Caty Sibble, ArdisJohnston, Laura
and their habitats. Although much prog- Leiben—sperger, Dee Woessner, and Ann
ress has been made in distinguishing spe- Blum made a very supportive commu-
cies and matching males to females, many nity in which to work. Leticia Aviles and
problems of geographical variation and Christopher Maddison gave me much aid
uncertain male-female matching remain and inspiration during the most difficult
for future work, especially among Mexi- stages of the project.
can Pelegrina. H. W. Levi, L. Aviles, D. Maddison, G.
In addition tothe Pelegrinaspecies,Na- B. Edwards, P. F. Stevens, and E. O. Wil-
gaina incunda and the U.S. species of the son read and commented on this paper.
Metaphidippus mannii group are de- Two anonymous reviewers gave useful
scribed because they could very well be suggestions. For discussion of jumping spi-
confused with speciesof Pelegrina and be- der issues, I thank B. Cutler, G. B. Ed-
causetheirtaxonomy is in needof revision. wards, D. B. Richman, J. Proszynski, F.
This work addresses the phylogeny of Wanless, and P. Wijesinghe. B. Cutler, G.
Pelegrina and the subfamily Dendry- B. Edwards, A. Moldenke, and D. B. Rich-
phantinae, but it has no pretensions of be- man have sent live specimens to me for
ing a comprehensive or modern phyloge- courtshipobservations. Brent Opell helped
netictreatment. My phylogeneticgoalsare withtrypsin clearing. H. D. Camerongave
toproposesomecharactersthat might pro- me helpful advice on the formation of
vide an outline of dendryphantine rela- names, though he is not to be held re-
tionships, focusing on the question of the sponsibleforanyillformed,especiallysince
monophyly of Pelegrina and a few other I did not alwaysfollow his nodoubt proper
groupsformerly placed in Metaphidippus. advice. Curatorsat various institutionslent
I hope that this and the basic exploratory, me specimens; a list can be seen under
species-level taxonomic work will provide Materials and Methods. I thank these cu-
the groundwork for future phylogenetic ratorsfortheirhelp. I wouldespecially like
treatments. to thank Luis F. de Armas of the Instituto
de Ecologia y Sistematica, Havana, Cuba,
ACKNOWLEDGMENTS
for sending the Franganillo collection of
This paper formed part of a Ph.D. thesis salticids.
for the Department of Organismic and This work was supported in part by a
Evolutionary Biology, Harvard Universi- NSERC (Canada) Postgraduate Fellow-
ty. Theworkdescribed in it hasroots many ship. For allowing me to participate in a
years deep, and during these years I re- major collecting trip to Mexico, I thank S.
ceived the help of many friends and col- B. Peck and R. S. Anderson. The trip was
leagues. As always, David Maddison has funded by NSERC and had assistance from
been foremost among my collaborators in the Universidad Nacional Autonoma de
formulating ideas, on collecting expedi- Mexico and Harvard University. The De-
tions, on behavioral observations, and in partment of Organismic and Evolutionary
many other activities. My thesis advisor, Biology of Harvard supported trips to Cal-
Herbert W. Levi, provided a well-orga- ifornia, Arizona, and Spain. Publication
nized laboratoryin which towork,amodel costs of this study were covered in part by
ofdedicationtoarachnology,andahelpful the Wetmore Colles fund.
Pelegrina Jumping Spiders • Maddison 219
MATERIALS AND METHODS
white silicone rubber (bathtub caulking).
Collections Examined. The taxonomic The silicone rubber is superior to paraffin
revision is based on specimens in the fol- for most purposes, for it can hold even
lowing collections. The abbreviation for minuten pins firmly and later heal, and it
the collection is followed by the name of offers the advantage over sand of allowing
the collection and the curator and others appendages to be pinned open. Palpi were
responsible for aiding in loaning the ma- mounted on Vaseline in an alcohol-filled
terial, to whom many thanks are due: depression slidewithacoverslipand drawn
at 100X and 200x under an Olympus
AMNH American Museum of Natural BH-2 compound microscope using inci-
History, New York (N. Platnick, dent fiber-optics illumination and a cam-
L. Sorkin) era lucida. Not only did the use of a com-
BMNH The Natural History Museum, pound microscope allow higher resolution,
London (P. Hillyard) but also the axial light path prevented the
CAS California Academy of Sci- drawing difficulties caused by the side-to-
ences, San Francisco (W. Pu- sideshiftingof the imagethatoccurswhen
lawski, D. Ubick) focusingon a stereodissecting microscope.
DU Darrel Ubick personal collec- For the external (ventral) view, epigyna
tion were examined using the same technique,
lESC Instituto de Ecologia y Siste- without clearing. Epigyna were dissected
matica, Havana (Luis F. de Ar- off of the specimen to allow for the small
mas) working distance of the compound micro-
MCZ Museum of Comparative Zool- scope. The Vaseline on which they were
ogy, Cambridge (H. Levi) mounted was madeopaqueby mixingwith
MSUW
Midwestern State University, chalk dust, in order to simulate the cream-
Wichita Falls, Texas (N. Hor- colored muscles and glands that would un-
ner) derlytheepigynum on an intactspecimen.
TXAM A&M
Texas University, College After examination of the specimen. Vas-
Station, Texas (A. Dean) eline was removed by a xylene rinse. The
UCB University of California, Berke- obliquedrawingsofthemalecarapaceand
ley (E. Schlinger, C. Griswold) chelicerae were made mostly under the
UWBM Burke Museum, University of compound microscope, at 40x Most
.
Washington, Seattle (R. Craw- drawings of the female abdomen were
ford) made under a Zeiss stereo dissecting mi-
WPM W. Maddison personal collec- croscope with a camera lucida. The draw-
tion ings of the male face and female abdomen
ZMB Zoologishes Museum Berlin (M. show the appearance in alcohol. Most
Moritz, S. Fischer) drawingsweredoneon coquille board with
ink, a Conte drawing pencil, and white
Notethat Canadian specimens are, in gen- paint. Small labels with my initials (WPM)
eral, underrepresented in this revision be- and the year drawn (e.g., 84) were placed
cause two major collections of Canadian in vials of specimens illustrated. Photo-
spiders, the Canadian National Collection graphsof livingspecimenswere made with
at the Biosystematics Research Centre, Ot- a standard 55-mm lens reversed on exten-
tawa, and the Royal Ontario Museum col- sion tubes to yield approximately 2.5x
lection, were not examined due to time magnification on Kodak Technical Pan or
limitations. Kodachrome film, using illumination by
Routine Examination and Illustra- flash. Measurements of carapace length,
tions. Specimens were examined in a glass carapacewidth,and body length (Galiano,
dish with a bottom layer of half black, half 1963; Wanless, 1978) were made from the
220 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4
dorsal view using an eyepiece reticle on a fixed, 80% ethanol fixed, and Kahle's fixed
Leitz stereo dissecting microscope. Gen- specimens were all used. Specimens not
erally only about five specimens of each fresh-killed were first rinsed in water for
sex were measured, because little reliance several hours before digestion. The body
on these measurements was made in this parts were separated to allow penetration
study. They are intended to give only a of the trypsin. The trypsin solution was
general idea as to the size and proportions made from about 1 ml purified trypsin
of the species. The results are presented as (Fisher #T-360) in 10 ml water, filtering
follows: minimum (median) maximum. after mixing. The trypsin was warmed un-
Thus, if five measurements for carapace der a light bulb during digestion. After
length for the females of one species are digestion,the parts, especially dark, heavi-
2.0, 2.1, 2.2, 2.3, and 2.3, these would be ly sclerotized palps or epigyna, were
reported as follows: 2.0(2.2)2.3, n = 5. bleached. They were first rinsed in water
Descriptions. A species description was and then 80% ethanol, then moved to a
originally written from a sample of five solutionof 1 partethanol: 1 part 10% Aero-
males and five females or more (if avail- solforaday,andthenforbleachingmoved
able). During subsequent identifications, into a solution of 1 part 30% hydrogen
the description was periodically checked peroxide: 1 partethanol:2 parts 10% Aero-
to ensure that it covered the range of vari- sol. Aerosol (Fisher) was added to the
ation within the species. Two characters bleaching solution to inhibit the formation
described that were less thoroughly sam- and buildup of bubbles, which otherwise
pled are the exact region of contact of the can fillthebody partand makeit unusable.
forehead band with the anterior median After about 1 day of bleaching, the body
eyes, which was examined in only four to part was moved to 80% ethanol and then
eight males, and the details of the internal transferred to 95-100% ethanol. The body
epigynal ducts of the female, which in part was then mounted temporarily in
many species was examined in only one to clove oil or permanently in Euparal. For
three females. most specimens, the transferral to Euparal
Clearing. Clearingwasusedfordetailed was accomplished by placing the body
examination of the integument, especially parts in Euparal thinned with either eth-
to observe external and internal structures anol or Euparal Essence and letting the
of the genitalia and mouthparts. The var- ethanoloressenceevaporateoff,thusgrad-
ious body parts were cleared by placing ually taking the specimen through a series
theminwarmtrypsinsolutionfor 1-2days ofstrongerEuparalsolutions. Thiswasmost
to digest internal tissues. When the tryp- easilydonedirectlyonthemicroscopeslide,
sin-clearing procedure is successful, it re- which was then let to dry partially in a
veals palp morphology to a level of detail dust-free area. Drying thickened the Eu-
not previously published (Figs. 3, 16-27). paral,allowingfinal positioningofthe parts
Trypsin was used instead of potassium or before adding the coverslip. After thecov-
sodium hydroxide because it damages the erslip is added, the body parts, especially
cuticle very little and has shown no ten- the palpus, may move. Thus, the palp was
dency to expand the palp except perhaps placed near the edge of the coverslip to
when hematodochae are tightly coiled as allow repositioning using a microneedle
in Ashtabula and Bagheera. If the tissues slipped under the coverslip.
of the specimen are well fixed and firm, Expansion ofPalps. Palps were best ex-
then digestion will be very slow. Hence, panded permanently by boiling the spec-
for best results the specimen should be imen alive and then fixing it in dilute
fresh-killed (not fixed) or "fixed" in a poor Kahle's fluid to harden it in an expanded
fixative with a low concentration of alco- position followed by gradual dehydraton
hol. Fresh-killed, dilute ethanol-triton to 80% alcohol. This technique is much
Pelecrima JiMi'iNG SPIDERS • MaddisoTi 221
liketl.-it describedby Sadana (1971). Palps back and forth at tips (n = 1), vigorously
sopreparedareresistanttocontractionand (ca. 5 c/s) (n = 1) but at low amplitude (n
can be critical-point-dried successfully = 5, 33)." The parenthetical comments in-
(Figs. 7, 9). A few palpi that were already dicate the number of observations (n) and
preserved in alcohol were expanded in a the number of males in which the feature
few minutes b\ placing them in a hot mix- was noted. The number of observations
ture of 15% hydrogen peroxide, 40% wa- was counted as follows: a male was ob-
ter, and 45% ethanol. served doing a bout of courtship display;
Chromosomes. Chromosomes were ex- any features of position or motion were
amined using the Feulgen technique as consideredtohavethusbeen observedonce
described by Maddison (1982). The results during this bout. If the male stopped dis-
are given under the description of each playingbecausethe female left or rejected
species examined. him, and if he began again later (perhaps
Courtship. Courtship observations were after I had moved them back together),
obtained for Pelegrina species and nu- then the next display was counted as a
merousotherdendryphantines. Specimens separate observation. While the more ob-
were examined within a few days after viousfeaturesofthedisplay may havebeen
collecting. A male and a female were observedmanytimes(forinstance,thatthe
placed on a cotton beating sheet, usually legs were flickered was observed 12 times
not in full sunlight, and manipulated until in six males in the preceding example),
the male faced the female and began dis- some of the more subtle details of the dis-
play. Behavior of the male was observed playwerenotnoticed in mostdisplaysand,
by eye, and notes and still photographs thus, would havebeen observed only a few
were taken. Female behavior was not re- times (for instance, that the legs flickered
corded. For most species, no filming or "alternately back and forth at tips" was
videotaping was done. As salticid behavior observed only once). Where there is vari-
can be fast, it is difficult to take notes ac- ation, the description lists each of the al-
curately, and there are likely errors in ob- ternatives with an indication of sample
servation, especially of subtle differences sizes. For instance, if the legs were usually
in timing and positions. Still, consistency flickered at low amplitude, but occasion-
of observations and videotape confirma- ally at high amplitude or not at all, the
tion of my own previously taken notes for description might read like this: "On each
Pelegrina dithalea, Metaphidippus man- series legs flickered (n = 9, 13) noticeably
nii, Metaphidippus chera, and Phanias (n = 1) or with fairly low amplitude (n =
watonus all indicate that the descriptions 7, 33) or perhaps not at all (n = 3, 13)."
should be generally accurate. There are a Sample sizes are in general small. It was
number of observations, such as the alter- felt that it is presently more important to
nate palp waving in the Pelegrina fla- obtain a broad survey, including many
vipedes group and the triangular crouch species, than a deep analysis of a few spe-
pose of Pelegrina aeneola, that have been cies. Explanationsoftermsusedtodescribe
repeated a number of times and in which courtship are given in the section Expla-
I have strong confidence. In the descrip- nation of Morphological and Behavioral
tions of the displays, the sample sizes for Termsand in thedescription of behavioral
the observations are indicated by listing characters (item 7) supporting the mono-
the number of observations for each fea- phyly of Pelegrina.
ture of the display. For instance, in the Phylogenetic Analysis. Though this
description of the courtshipof P. galathea, work is primarily concerned with the ge-
the following sentence occurs: "First legs nus Pelegrina, an attempt was made to
flicker (n = 12, 63) on each series (n = 4, outline the broader structure of the family
23) upanddown (n = 4,23) andalternately and the relationships of dendryphantines,
222 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4
partly for their own sake and partly to set vipedes,flaviceps, and exigua were main-
the context for the genus Pelegrina. The tained distinct despite apparent hybrid-
generaldiscussion of phylogeny within the ization because they differ in numerous
family is presented separately (Maddison, features. The two sympatric forms of ex-
1988, unpublished manuscript). My phy- igua were left under one specific name
logenetic proposals are presented in a nar- until they can be better studied.
rative discussion of groups and characters; Male/FemaleMatching. Carehadtobe
no numerical phylogenetic analysis was taken in proposing which males and fe-
done. Table 2 shows the distribution of males belonged together in the same spe-
some of the more important characters, cies, as in other spiders (Levi, 1985). The
but it will be noted that some of these do problem was especially bad in Pelegrina
not perfectly support thegroups proposed. species from the southwestern United
Problems with some of the characters are States, Mexico, and Central America,
noted in the phylogenetic discussion. Fur- where collecting has been limited. Despite
thermore, the presumption of ancestral the strong sexual dimorphism, similarity
state for a given character is usually not in body form and markings could often be
accompanied by rigorous outgroup anal- used as evidence. Other criteria used were
We
ysis (e.g., Maddison et al., 1984). are expected correlations in genitalia (e.g.,
still making only preliminary sketches of wide, robust embolus with strong epigynal
the phylogeneticstructureofthislargeand flaps), co-collecting in same geographical
poorly known family, and it would not be region, locality, or microhabitat, and sim-
productive to delay phylogenetic hypoth- ilarityofthemaleandfemaleeach tothose
eses until they can be rigorously defended. of another well-matched species. Com-
The suggestions made should have at least ments regarding evidence used to match
a glimmer of truth and will, I hope, stim- males and females are given in those spe-
ulate future work. cies descriptions where it seems needed.
Species Distinctions. Populations were Amongthelesscertain matchingsarethose
considered distinct species if several con- forPelegrinasandaracina,pallidata,chai-
sistent and discrete morphological differ- mona, huachuca, and morelos.
ences could be found among them, but
whentherewerefewdifferences,apparent EXPLANATION OF MORPHOLOGICAL
intergradation, or little material, the de- AND BEHAVIORAL TERMS
cision as to whether one or more species
Markingsin General. Colorpatternsare
arepresent wassometimesdifficult. In sev- generated by integument coloring and by
eral cases, only a single species was rec-
coveringofsetae,thoughpalesetaeusually
ognized despite geographical variation,
overlie pale integument and dark overlie
becausethegeographical variation wastoo
dark. The terms hair and scale are used to
confusing at present (furcata-mimus), or refer to a thin, more or less cylindrical seta
because the differences were slight and
and a broad, flattened seta, respectively
possibly not consistent {aeneola/uteanus,
northern and Floridian tillandsiae, north- (cf. Hill, 1979).
Markings of the Carapace. Male den-
ern and southern carmenensis, mannii/
dryphantines are commonly dark brown
versicolor) In other cases, allopatric pop-
. with bands of white to yellow scales, usu-
ulationsthataresimilarbut thatdiffercon-
sistently in a number of features were rec- ally including a major longitudinal band
on either side of the carapace and abdo-
ognized asdistinct (sabinema/pervaga,bi- men. The following names are used to re-
cuspidata/volcana, neoleonis/tristis, fer to the bands of pale scales on the car-
proxima/galathea/dithalea,chera/tricol-
or/diplacis), though in each of these cases apace (Fig. 1):
the decision was difficult. Pelegrina fla- side bands: Longitudinal bands on either
Pelegrina Jumping Spiders • Maddison 223
side of carapace, beginning beside the paired: the first pair just anterior to the
anterior lateral eyes (ALEs) and pro- muscle attachment of the second dorso-
ceeding just beneath the small eyes and ventral muscle (number 86viii -(- ix of
posterior eyes and beyond, onto the tho- Whitehead and Rempel, 1959), the second
rax. pairanteriortotheattachmentsofthethird
cheek band: Oblique band on the side of dorsoventral muscle, and the third, fourth,
the face, starting beneath the ALEs and fifth, and sixth pairs of spots behind this.
proceeding down and posteriorly to the The fifth and sixth pairs are very small.
carapace margin, in Pelegrina and the Often the fourth through sixth pairs are
mannii group. each connected medially and thus form
forehead band: A V-shaped marking on small chevrons. Between these pale spots
the dorsal cephalic area just behind the may be spots of dark brown, sometimes
anterior median eyes (AMEs). very dark (e.g.. Fig. 353).
marginal band: On the lower margin of Male Palpus(Figs. 3, 6-9). The descrip-
thesidesofthecarapace,often extended tions generally assume that the left palp is
from the cheek band in Pelegrina. Fe- being viewed from the ventral. The ad-
malesof most speciesshow none ofthese jectives basal and apical when unqualified
bands distinctly; the carapace is instead refer to the appearance in a contracted
often covered more or less uniformly palp (i.e., basal = toward the tibia and
with pale scales including a dense white apical = toward the tip of the cymbium).
covering on the clypeus. Incontrast,"anatomicallybasal"and "api-
cal" refer to the cymbium-embolus axis
Setae Surrounding Anterior Median of connections of the palp's bulb (ie., an-
Eyes. These are of various colors, from atomically basal = toward the basal he-
white to black. In the descriptions of Pe- matodocha and anatomically distal = to-
legrina males, the colors of setae around ward the tip of the embolus).
the circumference of the left anterior me- In the subfamily Dendryphantinae, the
dian eye (AME) are indicated using a no- shoe-shaped cymbium holds a bulb con-
tationderived from hourson aclock'sface. sisting of (from anatomically basal to api-
Usually, the colors on only the dorsal part cal) afullyexpandablebasalhematadocha,
of the AMEs are described, for those ven- a small subtegulum, a much reduced me-
trally are more variable. Thus, "white dian hematodocha, the large tegulum, a
forehead band contacts the AME dorsally fully expandable distal (embolic) hema-
10:30-12:30" meansthat asone looks from todocha, and the embolus. The basal he-
the front at the left AME there are white matodocha expands so as to give a clock-
setae from 10:30 o'clock to 12:30 o'clock wise rotation to the subtegulum and te-
thatarecontinuouswiththeforeheadband, gulum in ventral view of the left palp, as
and dark setae on either side of this. in other salticids. This has been observed
Markings of the Legs. The legs are yel- by artificial expansion on many dendry-
low to dark brown, but there are often phantines and during copulation of Den-
annulate markings (Fig. 1), especially in dryphantes nigromaculatus. The subte-
males, so that each leg has pale portions gulum is small and contains little more
covered with white scales alternating with than the basal portion of the sperm duct
darker portions lacking white. reservoir (Figs. 3, 8).
Markings of the Abdomen. The abdo- From the subtegulum, the sperm duct
men is usually brown above in males, loops up, down, and around the tegulum
ringed by white side bands (Fig. 1). The (Fig. 3) in a clockwise direction to the em-
dorsum of the abdomen often shows traces bolus. The distal retrolateral portion of the
of the paired pale spots seen in females tegulum, where the sperm duct enters the
(Fig. 2). These pale spots are central and tegulum from thesubtegulum, isgenerally
224 Bulletin Museum of Comparative Zoology, Vol. 154, No. 4
extended distally and contains a loop of bium), as indicatedby artificial expansions
the sperm duct pressed against its wall. (Figs. 7, 9). The counterclockwise move-
This portion I call the shoulder of the te- ment of the embolus itself probably ex-
gulum (Fig. 3). Just proximal to the em- plains why the counterclockwise-coiled
bolusandshoulderisafoldextendingacross embolus can still engage the epigynal
the surface of the tegulum, the tegular opening despite the clockwise movement
ledge (Figs. 3,6, 7), whichservesasa pock- of the tegulum (in salticids with the em-
et to hold the proximal part of the embolic bolus fixed to the tegulum, the embolus
base. Wanless (1984) suggested that the has a clockwise curve, which thus coin-
tegular ledge (his M3) is absent from sal- cides with the clockwise thrust of the te-
ticids other than spartaeines, in which he gulum). The embolus of dendryphantines
described it. However, the fold that I am usually consists of a basal portion, which
here calling the tegular ledge may be ho- is transversely directed, and an apical por-
mologous with his M3, for it appears as an tion, which is usually thin and erect and
extensionoftheembolichematodochacut- has the opening of the sperm duct at its
ting across the face of the tegulum and tip (Figs. 40-47; also Figs. 64b, d). The
occurs in many groups of salticids. The embolic base consists of a more or less
tegulumisfilledwith tegularglands,which sclerotized portion of the embolic hema-
empty into the sperm duct via a series of todocha that is exposed to the ventral sur-
poresin thesperm duct (seeOsterloh, 1922; faceandthatrestsbetweethetegularledge
Bhatnagar and Rempel, 1962). As in other and theembolus (Figs. 3, 6, 7). Its wrinkles
spiders, these pores are aligned intoa band suggest that it should be considered part
(Schult, 1980), which, when narrow, ap- ofthehematodocha. Asnoted,theembolus
pears as if it were a seam along the length is coiled counterclockwise, a feature much
ofthesperm duct. Alongthenarrowerpart more readily apparent in the euophryines
of the sperm duct toward the embolus, the and other subfamilies than it is in the den-
glandsareconnected tothesperm duct via dryphantines. Indendryphantines,thespi-
long ducts (Fig. 3; also known from other ral is hidden and best seen in expansions
spiders; Osterloh, 1922: figs. 20, 26). These or dissections (e.g., Fig. 35). The dendry-
pores, ducts, and glands have been largely phantine embolus arises prolaterally and
ignored in systematics but appear to be a moves across toward the retrolateral side
potential source of good systematic char- (the transverse basal portion of the em-
acters (for instance, in some salticids the bolus) and then folds back toward the pro-
pores are arranged in a broad ribbon, lateraland abruptlyrisesastheerectapical
whereas in others the band is very narrow, portion (Figs. 20-23). In many genera, the
and in Sitticus the pores are arranged into erect apical portion is fused against the
prominent craters). transverse portion so that there is no open,
The embolic hematodocha arises on the freely coiling spiral. A suture on the back
back side of the tegulum (Fig. 8), prolater- side of the embolus (the embolic suture),
al to the subtegulum. Its wrinkles sweep between the transverse portion and the
apically up toward the embolus. The exact erect portion,isoften presentand indicates
arrangement of the folds of the hemato- where the folded-back spiral has not com-
docha is probably of systematic value but pletely fused (Figs. 3, 8, 20-23, 31-35). In
is very difficult to untangle, for especially cleared palpi, a slight bend in the sperm
near the base of the embolus the folds are duct can be seen at this point. The coiling
twisted and confusing even in a well- oftheembolusand theembolic suturesug-
cleared palpus. During expansion, theem- gesting it are clearly visible in Dendry-
bolic hematodocha expands fully to move phantes, Eris, Pelegrina, Phidippus, Tu-
the embolus counterclockwise (i.e., pro- telina, Phanias, and many other genera.
laterally) and back (i.e., toward the cym- In Metaphidippuschera, thecoilingiseas-
