Revision of the Genus Trechalea Thorell (Araneae, Trechaleidae) with a Review of the Taxonomy of the Trechaleidae and Pisauridae of the Western Hemisphere

32 Pages·1993·33.2 MB·English

by  James E Carico

1993. The Journal ofArachnology 21:226=257
REVISION OF THE GENUS TRECHALEA THORELL
(ARANEAE, TRECHALEIDAE) WITH A REVIEW OF THE
TAXONOMY OF THE TRECHALEIDAE AND PISAURIDAE
OF THE WESTERN HEMISPHERE
James E. Carico: Department ofBiology, Lynchburg College, Lynchburg, Virginia
24501 USA
ABSTRACT. The spider family Trechaleidae, introduced by Simon in 1898 but abandoned immediately
thereafter, isdefined. Charactersproposedfortrechaleidsbyrecentauthorsforthisfamilyand somecharacters
usedtoseparateitfromthePisauridaeandLycosidaearereviewed.GeneraincludedinthefamilyTrechaleidae,
all ofwhich were previously placed in the Pisauridae, are: Trechalea, Hesydrus, Syntrechalea, Dossenus, Par-
adossenus, Dyrines and Enna. Familial placement ofall other genera ofthe western hemisphere previously
placed in the Pisauridae are considered.
The genus Trechalea, with eleven species, is revised. Redescriptions ofeight known species are given: T.
longitarsis (C. L. Koch), T. cezariana Mello-Leitao, T. macconnelli Pocock, T. paucispina di Caporiacco, T.
connexa (O. Pickard-Cambridge), T. extensa (O. Pickard-Cambridge), T. gertschi Carico & Minch and T.
amazonica F. Pickard-Cambridge. Descriptions ofthree new species: T. boliviensis, T. lomalinda and T. trin-
idadensis are presented. The holotype ofthe type species of Trechalea, T. longitarsis, is lost. The specimen
whichwaspreviouslyregardedastheholotypeismislabelledandbelongsactuallytoanother,unnamed,trechaleid
genus. All other relevant species names not synonymized with any ofthe above and not previously placed in
Trechaleaare removed to otherknown genera orwill be laterincluded in newgenera.
Intheearly 1970’sIbeganaseriesofrevisions (1897). Trechalea Thorell (1869) was described
oftheAmericanPisauridaewithPisaurina(Car- approximately thirty years earlier.
ico 1972)andDolomedes(Carico 1973).Theim- Simon apparently realized that he was study-
mediategoalwas to clarifythe taxonomy, atthe ing a group of unique spiders when he, rather
genus and species levels, which would establish cryptically, inserted a two-line notation of the
the required base for a later analysis at higher newfamilyTrechaleidaeinhissummaryof1890.
levels. Thetraditional family Pisauridaewas re- Nodescriptionwasofferedandonlythereference
tained as a matter ofconvenience with deferral to two genera, Dendrolycosa and the type genus
until later of the re-examination of the family Trechalea, wasgiven. Heneveragainreferredto
taxon itself. this family and continued (1898) to name new
While working on a group ofprimarily Neo- species ofthe genus Trechalea in his old family
tropical genera, including Trechalea, I became Pisauridae.
awarethattheyrepresentedamonophyleticclade DuringameetingoftheInternationalCongress
which challenged the limits of the traditional ofArachnology, IreintroducedtheTrechaleidae
family Pisauridae. This group of genera was (Carico 1986), erroneouslyreferringtothisfam-
therefore set aside for special consideration and ily name as a nomen oblitum. At that time I
wasusedasabasisforastudyofthefamily-level presented a provisional cladogram showing its
taxonomy. relationship to the families Pisauridae and Ly-
Ofthe seven genera placed here in the family cosidae, with the Pisauridaeas the probable sis-
Trechaleidae Simon, six were described around ter group.
theturnofthecenturybySimonandthePickard- ThecomplexityoftheTrechaleidaecombined
Cambridges. These genera include Hesydrus with the rather close relationship ofthe genera
Simon (1898a), Syntrechalea F. Pickard-Cam- madeithighlydesirablethatallpossiblematerial
bridge (1902), Dossenus Simon (1898b), Para- beexaminedbeforeformalpublicationofafam-
dossenus F. Pickard-Cambridge (1903), Dyrines ilyanalysiswasattempted.Onlyafterexamining
Simon (1903) and Enna O. Pickard-Cambridge all available related type specimens and collec-
226
CARICO-THE GENUS TRECHALEA (TRECHALEIDAE) 227
tionsscatteredinnumerousmuseumswasIable To studyepigyna, thesofttissuewasremoved
tosortoutthegeneraandspecies. Itthenseemed by a combination ofdissection and immersion
KOH
necessary to revise the entire family at once be- in or a proteolytic enzyme. Since the in-
fore any generic revision could be published to ternalpartswereoftendenseanddark,theywere
providewhatwasbelievedtobetheproperfoun- immersed in Chlorox (sodium hypochlorite so-
dation for a secure set offamily apomorphies. lution), as per the method ofGriswold (1993),
Asaresult, muchtimewasrequiredtobringthe until only the internal lining of the ducts was
project to completion. uncleared.Inaddition,totracebetterthetubules,
In the meantime, various authors have ex- the cleared, alcohol-saturated structure was al-
amined specimens ofthe family and have gen- lowed to air dry completely, which causes the
erally also concluded that the Trechaleidae rep- silvery trace of the lumen to appear when the
resents a distinct group and is probably a valid structure was reimmersed in alcohol.
family. Sierwald (1990), in her efforts to work Drawings were made with pen and ink, using
out the higher taxonomy of the American Pi- a Bausch & Lomb Stereozoom® Microscope
sauridae, has shown through an excellent study equipped with eyepiece drawing grid and mi-
ofthe male genitalia that the "'Trechalea genus- crometer.Therightpalpwasillustratedfromthe
group” represents a distinct taxon from the Pi- ventral and retrolateral positions. SEM micro-
sauridae and that the Lycosidae is the probable graphsofthe medianapophysisandtibialretro-
sister group of her ""Trechalea genus-group.” lateral apophysis at ventral view are provided
Griswold(1993)confirmed,byapreliminarycla- for comparison and diagnosis. Epigyna were
disticanalysisoftheLycosoidea,thedistinctness drawnfromtheventralviewwithsetaeremoved,
and monophyly ofthe Trechaleidae and placed and from the dorsal view, uncleared, with the
the family as the sister group ofthe Lycosidae. soft tissue removed.
Dondale (1986) listed family level synapomor- Measurements are in mm. As an index to the
phies for the Lycosidae, and further noted that size ofthe body, only the length ofthe carapace
the family shared some ofthese characters with is given because ofvariability in the condition
Trechalea. ofthe abdomen. Generally, the length oftheab-
To date no complete treatment ofthe family domen is approximately equal to the carapace
has been presented nor have any ofits included lengthinmaleswhiletheabdomenmaybesome-
genera been revised. It is my intention in this what longer in females. Eyes or eye group mea-
papertorevisethegenusTrechaleaandtoreview surementsweremadewiththesurfaceplaneper-
the taxonomy ofthe family Trechaleidae in ref- pendiculartotheaxisofsight.Abbreviationsand
erencetotherelatedfamiliesPisauridaeandLy- additional notes pertaining to eye group mea-
cosidae. Later,revisionsoftheremainingsixde- surements are in Table 1.
scribed genera and new genera will follow. A Museum abbreviations.—AMNH~American
summaryoftheirphylogeneticrelationshipsand Museum ofNatural History; BMNH—TheNat-
akeytotheiridentificationwillalsobepresented ural History Museum, London; CAS-—Califor-
at the appropriate time. nia Academy of Sciences; EXPE—Exline-Peck
(now in CAS); FMNH-=-Field Museum ofNat-
METHODS
ural History; INPA-—Institute National de Pes-
Careful dissections ofunexpanded palps were quisas Amazonia; JAK—J. A. Kochalka; JEC—
MACN—
conducted to learn the relative positions ofthe J. E.Carico; MuseoArgentinedeCien-
MCN—
variouscomponents,asintheapproachofDon- cias Naturales, Buenos Aires; Museu de
dale (1986). This method develops a good un- CienciasNaturals, FundagaoZoobotanicadoRio
derstanding ofpalpal structure that makes pos- Grande do Sul; MECN—Museu Equitoriano de
siblecomparisonsamongdifferentgenerabecause Ciencias Naturales, Ecuador; MEG—M. E. Ga-
all palpi in this unaltered condition are more liano; MCZ—Museum ofComparativeZoology;
likely to be similar. Expanded palpi were also MZUCR—Museo de Zoologia Universitaria,
studied, butexpansionbyitsverynature, causes Costa Rica; REL—R. E. Leech; USNM—Na-
distortion in positional relationships which in- tional Museum ofNatural History.
evitably results because ofdifficulty in control- Abbreviations ofanatomical terms for genita-
ling the degree ofexpansion. Comparisons be- lia.—Many anatomical terms and abbreviations
tweenspecimensofdifferentgeneraweretherefore ofgenitalia were adopted primarily from Sier-
much more difficult on palpi in this state. wald (1989, 1990) while others were coined for
228 THEJOURNALOFARACHNOLOGY
purposes of this paper: <2/?—accessory bulb of ofthediscoideggsac(Fig. 6), transportofyoung
epigynum; (^/—anterior field of the epigynum; on the empty egg sac, and non-reattachment of
bs—base of spermatheca; c—conductor; cd— dislodged egg sac.
copulatory duct; cdd-~copu\aXoty duct divertic- Description.—Very large (body length up to
ulum; CO—copulatory opening; cy—cymbium; 21 mm, leg span up to 167 mm; Trechalea ce~
o'o'—dorsal division of median apophysis; c— zariana)to small (bodylengthdownto 3.5 mm,
embolus; cZ)—embolic base; ecd—ee\a\ division legspandownto 18 mm; undescribedgenusand
ofthe retrolateral apophysis ofthe male palpal species), entelegyne, araneomorph, ecribellate,
tibia; cp/—epigynal fold; c^—embolic groove; lycosoid. Carapacelowtomoderatelylow,about
end-~en\a\ division ofthe retrolateral apophysis as wide as long with cephalic area moderately
ofthe male palpal tibia; c/?—epigynal p\aXe\fd— distincttoindistinct, longitudinalfoveadistinct.
fertilization duct; guide (part of mo); hs^ Eyes viewed from above in two rows, PE row
head of spermatheca; //o—internal fold of an- recurved, subequal in size, approximately equi-
terior field; //—lateral lobes; mo—median distantand separated byabout an eye diameter,
apophysis; m/—middle field of the epigynum; AErowstraightornearlyso,alwayssmallerthan
/?—petiolus; pmo—posterior margin ofanterior PE, ALE smaller than AME; ocular quadrangle
fieldofepigynum; r/o—retrolateraltibial apoph- wider above, about as high as wide. Clypeus
ysis; 5—spermatheca; sperm duct; 55—stalk height variable. Sternum about as wide as long,
ofspermatheca;5/—subtegulum; /—tegulum; //— truncated anteriorly and acute posteriorly. La-
tibia; vcm—ventral cymbio-tibial membrane of bium free, length-width ratio may be greater or
the palpal tibia; v^/—ventral division ofmedian lesser than 1.0. Chelicerae vertical, base robust,
apophysis; vp—ventrodistalprotuberanceofmale often enlarged anteriorly on males, promarginal
palpal tibia; vr—ventrodistal rim ofmale palpal teeth three, equidistant with middle one largest,
tibia. retromarginal teeth variable between three and
five in number varying in size and distribution.
Family Trechaleidae Simon
Endites longer than wide, parallel.
Trechaleidae Simon, 1890:82, type genus Trechalea Abdomen oval, low, somewhat flattened ven-
Thorell, 1869. Bonnet, 1955-1959:4680; Carico, trally,moderatelycoveredwithsetae,oftenpatch
1986:305; Coddington & Levi, 1991:22, (dado- ofsetae present ventrally and anterior to spin-
gram); Griswold, 1993:1-39. nerets, no plumose setae found. Six spinnerets
Dolomedeae (in part), Simon, 1898a:301. andcoluluspresent. Trachealspiraclelocatedjust
Dolomedidae (in part), Lehtinen, 1967:372.
anterior ofcolulus.
Diagnosis.—The family Trechaleidae can be Maleandfemalegenitaliaaredescribedbelow.
distinguishedfromtheLycosidaebythepresence Legsgenerallylong,slender,oftenwithflexible
of: a retrolateral apophysis and a ventrodistal tarsi(Fig. 1), and, insomespecies, flexiblemeta-
refoldedrimonthemalepalpaltibia(Fig. 7),the tarsi. Coxaenotchedventrally.Thirdlegsalways
posterior eye row (PE) in a single recurved row, shortest but relative lengths ofother legs vary.
and eyes in two rows. Large macrosetae present on most segments,
In only the Pisauridae, in comparison with found in varying numbers of pairs on ventral
LycosidaeandTrechaleidae:thefemaleproduces surfacesoftibiaeandmetatarsi.Longsetae,often
anurseryweb(amatrixofirregularwebbingsur- curvedattips(Fig. 1), presentmainlyonventral
roundingthe egg sac anduponwhichthe female surfaceoflegs. Trichobothria: tarsiwithtworows
sits and defends against intruders), in some spe- and in alternating position, metatarsi with one
cies the male binds the female’s front two pairs row, tibia with cluster near proximal end with
oflegswith silkduringcopulation, and aunique few others scattered distally, femur none. Both-
tubular retreat with associated resting behavior rium(Fig. 2)withhooddistinct,angledlaterally,
is constructed by some species. notridgedorembedded.Tarsiandmetatarsimay
The Trechaleidae can be distinguished from be scopulate (Fig. 1) with one median and two
both the Lycosidae and Pisauridae by: the male lateral claws, all usually dentate. Female palpus
palpuswhichhasalargedistallysituatedmedian with single, dentate claw.
apophysisequippedwithadorsalembolicgroove Distribution.—The family is found entirely
thatextendsdistallyintoanapicalguide(Fig. 7), withintheregionfromtheGilaRiverofArizona,
the presence ofa discoid egg sac carried only on United States southward to northern Argentina.
thespinnerets, presence ofa“skirt” onthe seam Natural history.—The included species have
CARICO-THE GENUS TRECHALEA (TRECHALEIDAE) 229
Figures 1-5."Scanningelectron micrographs ofstructures oftrechaleids: 1,2, Trechaleagertschi; 1, tarsus,
retrolateral view showing flexible integument and scopula; 2, bothrium; 3-5, Hesydrus habilis; 3, spinnerets (a
= anal pore, c = colulus); 4, enlargement of3 with emphasis on posterior median spinnerets, mesally located
pairs ofminor amputate gland spigots with silk emerging from three ofthem; 5, attachment offour “carrying
threads” to top valve ofegg sac. Magnifications: 1, 60x; 2, 2000x. Scales: 3, lOO/u; 4, lOju; 5, 100^.
a wide size range, and one would expect a wide No nursery web is constructed. The female
divergence ofhabitat preferencesto exist. Infor- carries the egg sac containing eggs or first instar
mationisavailable,however,mainlyforthelarg- spiderlings by the spinnerets, specifically the
er species. The general impression from collec- paired minor amputate gland spigots on each
tion data and from personal observation is that posteriormedianspinneret(Figs. 3, 4, identified
the preferred habitat is around the margins of by J. Coddington, pers. comm.) in the manner
bodies offreshwater. Some moderate-sized spe- oflycosids. The egg sac has a unique structure,
cies have been taken from trees and other veg- described as hemispherical by Simon (1898a),
etation away from water. with a generally flattened shape and with a dis-
No species is known to make a snare ofany tinct fringe or “skirt” at the seam between the
kindandallareapparentlyentirelycursorial.The two valves (Fig. 6). As the eggs hatch and the
most aquatic species are very adept at walking spiderlings occupy more ofthe internal space of
on thewatersurface and crawlingunderwaterin the egg sac, the “skirt” is obliterated by stretch-
themannerfamiliartotheaquatic speciesofthe ing. The young emerge from the weakjunction
pisaurid genus Dolomedes (Simon 1898a; pers. at this seam presumably without aid from the
obs.). The feature offlexible tarsi is common in mother. After emergence, the young are carried
the family. The flexibility is due to a general upon the egg sac mostly on the uppervalve but,
softnessofthe cuticleratherthantheoccurrence with crowding above, also sometimes below.
of pseudosegmentation. This flexibility, which Youngmayalsobefoundontheabdomenofthe
may be quite extensive, could be an advantage mother, butapparentlyonlyasaconsequenceof
inthe supportand locomotion onthewatersur- insufficient space on the egg sacand then only if
face. they are in contact with sac-bome spiderlings. I
230 THEJOURNALOFARACHNOLOGY
have not observed young only on the abdomen;
whenonlyafewyoungarepresent, theyareonly
on the egg sac. The mother persists in carrying
theeggsacandhasbeenobservedcarryingitwell
afterspiderlingshaveleft. Onlyfourstronglines
are attachedfrom the posteriormedian spinner-
etstothecenterornearcenteroftheuppervalve,
leavinga distinctsingle scar(Fig. 5). The female
will not reattach the egg sac at any stage ifit is Figure6.—Diagramofatrechaleideggsacwith“car-
dislodged. Both upper and lower valves are
ryingthreads” attached.
smooth on the surface, the upperusually darker
and thicker while the lower is lighter, thinner,
and flatter (Simon 1898a; Sierwald 1990; pers. with a pin. Only a single entire leg is still at-
obs.) with the eggs or young often visible. tached;othersareshattered,(b)Thevialcontains
Statusofthetypegenus.—Byreferenceto Tre- atotaloffivelabelswhichareasfollows:“2006”,
chalea as the type genus ofthe family Trechal- “longitarsis Koch?”, “Brazilien Lgsof’, “Ty-
eidae (Simon 1890), it follows that the family pus”,andamodemlabelwhichhasthenumber,
name therefore hinges on the type of its type species name, etc., and the notation, ^\sensu
species, Trechalealongitarsis(C.L.Koch).How- Karsch)”. The firstthreeoftheselabelshavepin
ever, the correctidentity ofthis species presents perforations (two ofwhich were mentioned by
a problem. Karsch) and were presumably pinned originally
C. L. Koch (1848) described a female from with the specimen. The “Typus” and modem
what is now known as Colombia with a body label, which were not perforated, were appar-
length of IVi lines (= 15.7 mm, H. Levi pers. entlyaddedwiththespecimenafteritwasplaced
comm.) and gave it the name ofTriclaria longi- into alcohol storage.
tarsis. Some important distinctive features list- After careful consideration ofthe evidence, I
ed in his description included the pattern ofthe have concluded thatthis specimen, hereafterre-
eyes(whichwerefigured)andthe“sickle-shaped” ferredto as “2006,” isnotthetype specimenfor
tarsi, both ofwhich are important identifiers of Trechalea longitarsis (Koch). My conclusion is
what has traditionally been considered Trechal- based on the following observations: (1) Koch
ea. Trechalea was the name latergiven as a no- indicatedthathisspecimenwasa“female”,pre-
men novum by Thorell (1869) to the genus be- sumably meaning it was adult. “2006” is a ju-
cause the name Triclaria was preoccupied by a venile.(However,theseauthors,includingSimon,
bird genus. often assumed that juveniles were females and
Later, Karsch (1879) found a dried spider in didnotlookforanepigynum[Levi,pers.comm.].)
theBerlinMuseum,no. 2006,whichheregarded (2) Koch stated that the length ofhis specimen
as the specimen Koch used when making the is “7V2'"”, (or 15.7 mm). “2006” is 12.2 mm,
description of Triclaria longitarsis because of whichissignificantlyshorter.(3)Anexamination
Koch’s reference to the “sickle-shaped” tarsi. ofthe external characters of“2006” shows it to
Karschmentionedtwolabelswiththespecimen, be a member ofa new genus (which will be de-
one ofwhich states “Brasilien, Langsdorff” and scribed in a later publication) with a distinctive
the other simply as “longitarsis?”. Noting the carapace shape, and offset fifth retromarginal
discrepancyofthepublishedlocalityas“Colom- cheliceral tooth. As will be shown later, this ge-
bia” and the label contained with the specimen nus is found only in the southernmost states of
as “Brazil,” he dismissed it merely as a change Brazilinthe Rio delaPlatadrainage system, far
made presumably in curating. Kraus (1955) from Colombia. Therefore, “2006” is not from
agreed with Karsch’s conclusion. Colombia, andthe specimenlabelwhichhasthe
I have examinedthe specimenno. 2006 men- locality “Brasilien” is probably correct. (4) The
tioned by Karsch and found the following: (a) labelinthevialwith“2006”,“longitarsisKoch?,”
The specimen is ajuvenile with the body length placed after Koch’s description and before
of 12.2 mm, presently preserved in alcohol, but Karsch’sdiscovery,indicatesthatthepersonwho
was apparently originally mounted dry. The pinnedthelabeltothespecimenhaddoubtsabout
presenceofaholethroughthecarapaceandster- it, asindicatedbythe questionmark. Therefore,
num indicates that it was probably perforated uncertainty about the specimen apparently pre-
CARICO^THE GENUS TRECHALEA (TRECHALEIDAE) 231
dates Karsch. (5) The “Typus” label was added ofthese genera based on an examination ofthe
after Koch, because it lacks the perforation and available types.
does not add proofthat Koch made this desig- TunaboChamberlin(1916)isbasedonanim-
nation himself about this particular specimen. maturemaleholotypeandisalycosid(confirmed
The conclusion that this specimen was the type by H. Levi pers. comm.) but was regarded as a
could have been based on the assumptions of zoridbyLehtinen(1967)andwasthereforelisted
Karsch and may have actually been added, in- in the latter family by Brignoli (1983) and Plat-
correctly, by him. (6) Karsch’s conclusion was nick (1989). Sisenna Simon (1898a) is ajunior
apparently promoted by the lack of other ma- synonym of Architis and is treated elsewhere
terialavailabletohimforcomparisoncombined (Carico 1993). Two species described by Key-
with an overemphasis on the distinct eye and serling in the genus Tetragonopthalma Karsch
tarsuscharactersthatledhim to makethe state- (1878), T. granadensis (1877) and T. obscura
ment that “2006” agrees exactly with the de- (1891), are in the lycosid genus Porrimosa (Ca-
scription. pocasale 1982). Aglaoctenus Tullgren 1905 is a
Thepartiallyinsect-eaten, dried, andpresum- junior synonym ofPorrimosa (NEW SYNON-
ably faded specimen after 30 years must have YMY), a decision based on an examination of
lostmuchofthedetail, especiallypattern, ofthe theholotypeofAglaoctenusbifasciatusTullgren.
fresh specimen. Thus, the conclusion ofKarsch The genus Dyrinoides Badcock (1932) was de-
may have been influenced by the combination scribedfortwo specieswhichare representedby
of a process of elimination, misleading labels, tiny, unidentifiable spiderlings; however, they
and an overemphasis on the few distinct char- may be trechaleids. Thaumasia Perty (1833) is
acters remaining on the specimen. a pisaurid.
Theaboveevidenceleadsmetoconcludethat The Mello-Leitao genera Xingusiella (1940a)
“2006”isnottheholotypespecimenofTriclaria and Demolodos (1943) (Demelodos [5/c] Mello-
longitarsis designated by Koch. With this con- Leitao 1943;Demolodes[5ic]Roewer 1954;Brig-
clusion, one is still left with the problem about noli 1983; Platnick 1989) are each represented
the status ofthe nomenclature ofthe genus and byasingle species, andtheirtypesarelostinthe
thus ofthe family. In order to provide stability Rio de Janeiro museum and are not available
ofthe names, I make certain assumptionsbased for examination.
on theexamination ofaconsiderableamount of A species ascribed to Nilus (O. Pickard-Cam-
material. bridge 1876), N. amazonicus, was described by
I assume that the type specimen is lost and Simon (1898b) from the Amazon region ofBra-
that Colombia was the actual type locality as zil. The type is a large, recently moltedjuvenile
statedby Koch. Furthermore, I assume that the of unknown sex which is unlike any known
mostprobablepartofthecountrywherethespec- American pisauridand is notatrechaleid. Since
imenwastakenisthatareaofnorthernColombia the genus is otherwise known only from Africa
extendingfrom theAtlanticcoastto Bogota, the eastwardthroughAustraliaandSimondescribed
mostlikelyareathata Europeancollectormight manyspidersfromothercontinentsinhis 1898b
visit. Thus, the presumed type locality is in the publication, it is likely that this specimen was
coastal drainage region rather than in the Am- from another area and its locality is in error.
azonian drainage area. As explained later, these Two species ofa primarily African and Asian
two drainageshavedistinctspecies ofTrechalea genus,HygropodaThorell(1895),weredescribed
which fall into the size range stated by Koch. by Simon (1898a) from South America. Exam-
Therefore, the Trechalea species found in the ination ofthe types ofthese species, H. andina
coastal area, is assumed to be T. longitarsis. andH. venezuelana, showsthattheyactuallybe-
Status of unrevised “pisaurid” genera in the long to the genus Paradossenus.
Western Hemisphere.—Since this work brings AncylometesBertkau(1880)bearssomectenid
about some reassignments ofa major group of anatomical characteristics, but it is probably a
generafromthe Pisauridae (orDolomedidae)to pisaurid because ofits clearly pisaurid behavior
the Trechaleidae, it is appropriate now to sum- (Merrett 1988).Astudyofitsrelationshipsshould
marize the familial placement ofthe remaining be conducted, but is beyond the scope of this
unrevised “pisaurid” genera in the Western paper.
Hemisphere. The remainder of this section is Status of various pisaurid species assigned to
devoted to my determination ofthe disposition revised genera of the Western Hemisphere.—
232 THEJOURNALOFARACHNOLOGY
During this study, no specimen of the widely Lehtinen’s (1967) attempt to place these gen-
distributedgenusDolomedeshasbeenfoundfrom era into subfamilies and into new families re-
anywhere south ofthe Yucatan Peninsula. This sultedinthefollowing: Trechalea(“..Lycosoidea
confirms the opinion ofCarico (1973) that the incertae sedis; probably related to Zoridae, es-
genus has only a Nearctic distribution in the peciallyNeoctenus.'"), Dossenus(Dolomedidae),
WesternHemispherewiththeexceptionofareas Dyrines (Dolomedidae), Hesydrus (Dolomedi-
ofsouthern Mexico. There are, however, a few dae), Paradossenus (Dolomedidae), Syntrechal-
species in the older literature described from ea (“Position obscure, but evidently a represen-
South America which I have considered. Gie- tative of Lycosoidea.”). Additionally, genera
beTs (1863) description of the eyes of his Do- whichwerepreviouslyassignedtoPisauridaeare
lomedesintermedinsfromColombiaclearlysug- assigned as follows: Xingusiella (Amaurobiidae:
gests that it is not a Dolomedes. I cannot Rhoicininae), Aglaoctenus (Dolomedidae) and
determine its genus, however, because the ho- Dyrinoides (Lycosoidea incertae sedis).
lotype has not been found. The holotype ofDo-
STRUCTURE OF THE GENITALIA IN
lomedes albicoxa Bertkau (1880) is lost but a TRECHALEA
readingofthedescriptionindicatesthatthisspe-
cies is not a Dolomedes because ofthe strongly Malepalpus.—Thebasicstructureofthemale
procurved anterioreye row and relative sizes of palpus of Trechalea was described by Sierwald
the eyes. It may be a Porrimosa or a species of (1990) in her survey ofvarious pisaurid genera.
Architis. Although the holotype of Dolomedes Manyofherconclusionsandabbreviationscon-
pullatus Nicolet (1849) is missing, the descrip- cerning structure are used here; a more detailed
tion ofthe color pattern suggests to me that this description of the anatomy is available in her
species may be a Thaumasia. The holotype of paper. Additional details were pointed out by
Dolomedes elegens Taczanowski (1873) is miss- Dondale (pers. comm.).
ing,butthedescriptionofthecolorpatterngives Basically, there are four, major, articulating,
very good indication that it is a Thaumasia. sclerotized elements which make up the bulb of
ThanatidiusspinipesF.O.Pickard-Cambridge themalepalpus, eachseparatedbymembrane(s)
(1903:156, 157)isajuniorsynonymofStaberius (Fig. 7). Beginning at the base is the 1) subte-
spinipes(Taczanowski).NEWSYNONYMY. All gulum(st),attachedtothecymbium(c)byalarge
of the species of Thanatidius listed by Bonnet membrane, the basal hematodocha (bearing
(1955“1959)havebeensynonymized. Thegenus within its wall the sclerotized petiolus). Next,
wassynonymizedearlierby Carico (1972) to the moving distally, one encounters the ring-like 2)
entirely Nearctic genus Pisaurina. tegulum{t),describedbySierwald(1990)ashav-
Previous attempts at higher taxonomy.—Re- ing several loops or “switchbacks” ofthe sperm
movalofthese sevengenerafromthe Pisauridae duct. Attached to the membrane distal to the
{sensu Simon) {Trechalea, Hesydrus, Syntre- tegulum is the large, conspicuous, 3) median
chalea,Dossenus,Paradossenus,Dyrines,Enna) apophysis (ma), and the 4) embolus (e) which is
into a monophyletic family is accompanied be- athin,curvedstructurewithabroad, subdivided
low by an historical examination of previous base {embolic base, eb; [perhaps incorporating
schemes ofhigher taxonomy. At least three au- the terminal apophysis]). The embolic base is
thors have made a significant attempt to name partly subdivided by a narrow membrane. Of
familiesorsubfamiliesinwhichthesegenerawere these, only the embolus is not usually visible
included. when viewed ventrally in the unexpanded state.
Simon’s(1898a)placementofTrechalea,Hes- The median apophysis is a large, complex
ydrus, Dossenus, Drances (= Dyrines), and Hy- structure that appears to have a unique form in
gropoda into ‘"Dolomedese” (along with other the trechaleids (Sierwald 1990) and is located
genera), indicates his concept that these trechal- distally on the bulb where it characteristically
eidgeneraarecloselyrelated. However, onlythe occupies about a fourth to a third of the bulb
American species ofHygropoda are trechaleids mass. On its dorsal side is a deep, narrow, lon-
as indicated above. gitudinalgroove,theembolicgroove(eg),inwhich
Roewer (1954) assigned the genera Dossenus, the thin part ofthe embolus rests and is appar-
Dyrines, Hesydrus, Paradossenus, Syntrechalea entlycomparableto thatfoundin some lycosids
and Trechalea to his Thaumasiinae. However, (Dondale & Redner 1983b; Roth 1985 [key]).
he assigned Enna to his Pisaurinae. Thisgroove (independentlydiscoveredby Don-
CARICO~THE GENUS TRECHALEA (TRECHALEIDAE) 233
The size and shape ofthe retrolateral apophysis
isgenus- and species-specific. In addition to the
retrolateral apophysis, there is a feature ofthe
ventrodistalmarginofthetibiainwhichthebor-
derfolds down upon itselfforming a depression
down into the distal end of the tibia. This pit
thusformedhasbeenusedas ataxonomicchar-
acterofsomeimportancebysomeworkers(Sier-
wald 1990; Griswold 1993). See the section be-
low on character analysis for further discussion
ofthisfeatureandadditionalfeaturesofthetibia,
ventrodistal rim (vr), and ventrodistalprotuber-
ance {vp), and ventral cymbio-tibial membrane
{vcm).
Sierwald (1990) identified in the palpal bulbs
ofspecies of Trechalea and Paradossenus addi-
tional structures such as a conductor, terminal
apophysis, etc., which are not used here for di-
agnostic purposes. Refer to her paper for a dis-
cussion ofthese details.
Female epigynum.—A thorough study ofthe
homologies ofthe structures ofthe female gen-
italia is beyondthe scope ofthe present analysis
and the terminology used here is adapted from
the work of Sierwald (1989) on American pi-
sauridgenera. Emphasisisheredirectedtoiden-
tification ofmajor components and unique fea-
Figure7.-=TrechaleidpalpusanatomybasedonTre- tures of the genitalia of only Trechalea for
chalealongitarsis,ventralview.Emboluswithitslarge diagnostic purposes. A preliminary survey of
embolicbasein shadingbehind(dorsad) and showing various trechaleid genera reveals a considerable
thenarrowpartcontainedintheembolicgrooveofthe range ofvariation, and each one will be treated
medianapophysis. Abbreviationsin methodssection.
individually later. Thus, broad generalizations
concerningthe family orany assortment ofgen-
dale,pers.comm.)continuesdistallyintoacurved era within it are left to work in progress. For
orhooked projection arising from the dorsaldi- descriptivepurposes, termsarecoinedforusein
vision {dd), which probably serves as aguide{g) Trechalea and may or may not be applicable to
for the embolus during intromission (as is the other members ofthe family.
caseinsomelycosids[Dondale&Redner 1978]) The epigynum is generally heavily sclerotized
butseems notto supportorprotecttheembolus {T.trinidadensisexcepted),darkandopaque,with
atrestaswith,respectively,thefulcrumandcon- some components fused together, which makes
ductorfoundin, e. g., Dolomedes. Inadditionto detailed observation difficult without initial
thedorsaldivision,thereistheconspicuoussub- clearing. Theexternalepigynalplate{ep) (Fig. 8)
divisionontheventralside{ventraldivision, {vd) is composed offour regional elements, two lat-
which varies in size and shape. The shape and eral lobes (//) which comprise the elevated por-
positionoftheprojectionsarisingfromthesedi- tions on either side, an anteriorfield {aj) which
visionsarehighlygenus-andspecies-specificand occupies most ofthe anteriorhalfofthe epigyn-
aregivenspecialattentioninthedescriptionand um and is usually continuous with, but not al-
figures for each genus and species. waysdistinctfrom, thelaterallobes. The middle
Thepalpal tibia {ti) bears a distal retrolateral field {mf) is a distinct, posterior, median com-
apophysis {rta), which in some genera may be ponent set off from the other components by
subdividedintotwoparts:theentaldivision{end) furrows and/or the posterior margin ofanterior
which is surrounded by membrane (defined be- field{pma). (The “external, outerlateral margin
low), and the ectaldivision {ecd) usually located oftheepigynalfold”describedbySierwald(1989)
laterally and proximally to the ental division. forpisauridsappears in trechaleidsto be round-
234 THEJOURNALOFARACHNOLOGY
ed, less distinct and to lie out of sight under
[dorsad to] the mf.) The relative shapes and di-
mensions ofthese components tend to be spe-
cies-specific and are illustrated.
Internally (Figs. 9, 10), on each side, a gen-
erallylarge,voluminouscopulatoryduct(cd)arises
from the anterior portion of the epigynalfold
{epf) at the copulatory opening {co) and curves
inward and posteriorly where it gives rise to a
spermathecum {s) composed of a terminal en-
largement {head ofspermatheca, hs) at the end
ofastalk{stalkofspermatheca,ss),mostofwhich
lies fused against the copulatory duct. A second
diverticulum, calledthe copulatoryductdivertic-
ulum {cdd) arises secondarily from the copula-
toryduct.Althoughthefunctionofthisstructure FiguresS~10.—Trechaleaepigynumanatomybased
isunknown,itspresencehasbeennotedbyGris- on T. longitarsis: 8, epigynal plate, ventral view; 9,
wold (1993; ‘Tobate spermathecal base”, char- internalstructures,dorsalview; 10,diagramofrelative
acter #39). My interpretation ofits relative po- positionsofinternalstructures.Abbreviationsinmeth-
sition(between andfd),however,departsfrom ods section.
his. I have been unable to find this structure in
alltrechaleids,except,e.g.,aspeciesofDossenus. ofsynapomorphies forthat family and proceed-
Afertilizationduct(fd)continuesfrom thisjunc- ed further to define a number ofsubfamilies.
tion to meet the oviduct {o) on its ventral side. Griswold(1993) presentedawide-rangingbut
An internalfold ofthe anteriorfield {ifa) in the preliminary analysis ofthe Lycosoidea and in-
shape ofa thin flap (called “wing” by Sierwald) cluded as exemplars in his cladogram members
occursinvariousshapesandismostconspicuous ofthe Pisauridae {Dolomedes tenebrosus Hentz,
laterally where itjoins the cd. Pisaura mirabilis (Clerck)), Lycosidae {Lycosa
helluoWalck., SosippusplacidusBrady), andthe
CRITICAL REVIEW OF SOME
TAXONOMIC CHARACTERS USED IN Trechaleidae {Trechalea sp.). The controversial
genusRhoicinus(Rhoicininae),consideredapos-
RELATED FAMILIES
sibletrechaleidbySierwald(1990),wasincluded.
Since I reintroduced the family Trechaleidae In his cladogram, a total of 20 apomorphies
(Carico 1986), ithasbeentheindependentopin- emergedtodistinguishthesixexemplarsofthese
ion of Sierwald (1989, 1990), and Griswold three families.
(1993), that the Trechaleidae (orthe "'Trechalea Both Sierwald (1990) and Griswold (1993)
genus-group”) deserves family status and has as considered rhoicinines to be related to the tre-
its nearest relatives the families Pisauridae and chaleids. The latter author included an unde-
Lycosidae, with Lycosidae the closer. Codding- scribed Rhoicinus in his cladogram which
ton 8l Levi (1991) adopted the conclusions of emergedas a sistergroupto hisgenus Trechalea
Griswold and used the family name Trechalei- sp. andthereinimpliedthetwocouldbeconsid-
dae. Additionally, Dondale (1986) emphasized ered members of the Trechaleidae. Because of
in Trechalea its unique and lycosid-like features the problems historically in assigning a family
in his description ofthe subfamilies ofthe Ly- for rhoicinines (Platnick 1979), the perceived
cosidae. weakness of Griswold’s synapomorphies (dis-
The Pisauridae, as currently constituted, ap- cussed below), and the diversity of characters
pears to be a very complex group and may not among the various genera currently assigned to
bemonophyletic.Sierwald,inherexcellentstud- the group (Exline 1960), I do not include rhoi-
ies ofthe American pisaurid female copulatory cinines in the Trechaleidae. However, I do not
organs (1989) and male palpal organs (1990), excludethepossibilityofthefamilybeingbroad-
statedthattheknowngeneraareapparentlypoly- enedlaterto includerhoicinines aswell as other
phyletic,aconclusionwithwhichIagree. Incon- genera, e. g., Shinobius ofYaginuma (1991).
trast, however, monophylywas assumedforLy- Inanycase, theconclusionsheremustbecon-
cosidaebyDondale(1986),whoproposedagroup sidered provisional pending the outcome ofon-
CARICO--THE GENUS TRECHALEA (TRECHALEIDAE) 235
going studies by various workers who are con- Trechaleatogetherinhiscladogramwiththeim-
tinuingtorevisethegenerawithinthesefamilies. plicationthatthe formergenuswas atrechaleid.
Such work is required to reveal consistent char- An examination of Rhoicinus, however, re-
acters which contribute to greater confidence in veals that there is indeed a “ring” present, but
whateverconclusionisreachedaboutsystematic the cuticle enclosed is actually sclerotized and
relationships. Therefore, no cladistic analysis is thewhole structureis situatedonthe sclerotized
offered here. portion ofthe tibia. This ring and its enclosed
Following is a discussion of the taxonomic cuticle are not synonymous with the “membra-
characters used to distinguish the families Tre- nous pit” in the trechaleids as reported by these
chaleidae relative to Pisauridae and Lycosidae latter workers and is actually homologous with
usedbyrecentworkers(Dondale 1986; Sierwald theretrolateralapophysis{rtd)oftrechaleidsand
1989, 1990; Griswold 1993) including some of- pisaurids.
fered here for the first time. In the Trechaleidae (as here defined), another
PLEsituated behindPMEtoformathirdrow type ofpit (not found in Rhoicinus) is actually
ofeyes.—Thischaracterisatraditionaloneused formedfromtheventralcymbio-tibialmembrane
toseparatelycosidsfromthepisaurids.Itismen- {vcm, Fig, 7)ofthemalepalpaltibia. Thismem-
tioned by Dondale (1986) and Sierwald (1990), brane, found in all spiders, may be broad and
and Griswold (1993). flexible to permit a wide arc of flexion at the
Retrolateral apophysis on the male palpal tib- articulation between the cymbium and tibia. It
ia.—This is also a long-standing character used isnotuncommonto note aconcavityofvarying
to separate pisaurids and lycosids and is men- degrees in this soft membrane in spiders ofall
tioned by Dondale (1986), Sierwald (1990) and three families. Since this concavity results from
Griswold(1993). Separately, Dondale(1986)as- varyingconditions ofpreservation, itisthus not
sumed that the loss of a retrolateral apophysis taxonomically significant. The pit of Trechalea
in the lycosids is a derived state. It is present in referredtobythepreviously-mentionedauthors,
Trechaleidae and Pisauridae. however, is due less to a condition ofthe mem-
Median apophysis position on bulb.—This brane itself, but ratherto the morphology ofthe
character is offered here for the first time and adjoining sclerotized ventrodistal rim (vr) (Fig.
refers to the position ofthe median apophysis 7) ofthe tibia. In the trechaleids and, to a lesser
on the palpal bulb as viewed from the ventral degree, the pisaurids, there is a folding back
side. The position is either somewhere on the downward ofthe rim into the inside ofthe cy-
ventral face, e. g., Lycosidae and Pisauridae, or, lindrical tibia, forming a noticeable depression
inthecaseoftrechaleids,enlargedandoccupying in the ventral and retroventral end ofthe tibia
thedistalfourthorthirdportionofthebulbmass. because the vcm typically arises from this deep
Folded ventrodistal rim of the male tibia.— recess. In these same two families, the rim is
This is a reinterpretation of a so-called “pit” usuallyalso moldedintoa ventrodistalprotuber-
locatedventrallyattheapicalendofmalepalpal ance (vp) and thus forms a characteristic shape
tibia reported by others. Attention to a pit per- tothe tibialrim from ventral view. No suchrim
hapsbeganwith afeature notedinRhoicinusby or protuberance is found in American lycosids.
Exline (1960) who described it as an “unscler- Whatmakesthe so-calledpitmorenoticeablein
otized pit, surrounded by a fairly high, rebor- the trechaleids is that the distal, gaping opening
dered, chitinous ring.” A “membranous pitthat ofthe cylindrical tibia is larger, and the dorso-
accompaniesthetibialapophysis”wasnotedlat- ventral elongation is accentuated by the vp.
erby Sierwald (1990)also in Trechalea, and she Method of egg sac transport.—I depart from
proposed that this may be synapomorphous for other workers in the interpretation and use of
the two genera. Recently Yaginuma (1991, pp. thischaracter.Thetraditionalapproachhasbeen
2, 4)noteda similarfeatureinaJapanese spider to state the choice between “carrying egg sac in
Shinobius orientalis (Yaginuma) and thus as- chelicerae” or “carrying egg sac on spinnerets”.
signed this spider to the Rhoicininae in the Pi- This is similar to Griswold’s (1993) character
sauridaewithreservation. Griswold(1993) used #68 and is mentioned by Sierwald (1990) and
a similar character in his analysis of lycosoids Dondale (1986). This seems an oversimplifica-
andnameditas,“Malepalpaltibiawithretroap- tion and inaccuracybecause in all three families
ical cuticle unsclerotized.” He used this as an thefemalescarrytheeggsacattachedtothespin-
important character to link both Rhoicinus and nerets.Therefore,thepisaurids,whichaloneem-