Tarsal Scopula Division in Theraphosinae (Araneae, Theraphosidae): Its Systematic Significance

8 Pages·1994·6.9 MB·English

by  Fernando Pérez-Miles

1994. TheJournal ofArachnology 22:46-53
TARSAL SCOPULA DIVISION IN THERAPHOSINAE
(ARANEAE, THERAPHOSIDAE): ITS
SYSTEMATIC SIGNIFICANCE
Fernando Perez-Miles: Division Zoologia Experimental, Institute de Investigaciones
Biologicas C. Estable, Av. Italia 3318; Montevideo, Uruguay.
ABSTRACT. A study ofentire/divided tarsal scopulae distribution was carried out on 28 genera ofThera-
phosinae. Although this character has been considered an important taxonomic tool for more than a century,
present findings show that the scopula condition is related to spider size. These results call into question the
systematic value oftarsal scopula division. Fine structure oftheraphosid scopula is described, being different
fromthatpreviouslydescribedforAraneomorphae.Functionandphytogenyofscopulaconditionarediscussed.
RESUMEN. Se realize un estudio de la distribucion del caracter escopulas tarsales enteras/divididas en 28
generos de Theraphosinae. A pesar de que dicho caracter fue considerado importante en taxonomia durante
mas de un siglo, los resultados mostraron que la condicion dela escopula estarelacionadacon el tamano de la
arana. Estosresultados permitencuestionarel valorsistematicode lacondicionde lasescopulas. Sedescribe la
estructurade laescopulade lasterafosidas, encontrandosediferenciascon lasescopulasdescriptaspreviamente
para las Araneomorphae. Se discute la funcion y filogenia de los tipos de escopula.
The structure and function oftarsal scopulae becoming entire in adults of some subfamilies
have been studied extensively in araneomorph (such as Theraphosinae). Raven (1985) used en-
spiders (Homann 1957; Foelix & Chu-Wang tire/dividedscopulatoseparategeneraandgroups
1975; Rovner 1978; Miller et al. 1988; Roscoe in Theraphosidae and also studiedanterior-pos-
& Walker, 1991), and less thoroughly in Myga- terior gradation of the character. Smith (1988)
lomorphae. The structure of tarsal scopulae in disregardedthescopuladivision,consideringthat
Theraphosidae was discussed by Foelix & Chu- inmanytheraphosidsitonlymeansimmaturity,
Wang(1975)inDugesiellaPocock 1901 andmore butlater(Smith 1990)considereditanimportant
recently studied by Roscoe & Walker (1991) in taxonomic character for Ischnocolinae and Eu-
Sericopelma rubronitens Ausserer 1875 and by menophorinae. Perez-Miles (1992) polarized this
Prentice (1992) in Aphonopelma paloma Pren- character on the basis of ontogeny, concluding
tice 1992. Despite our limited knowledge, the that an entire scopulacould be a synapomorphy
tarsal scopulae conditionhasbeen usedin Ther- ofsome genera ofTheraphosinae.
aphosidae systematics for a long time. A studyofentire/dividedscopuladistribution
For more than a century the character “tarsal was carried out on most (28) genera ofThera-
scopulaeentireordivided”,hasbeenwidelyused phosinae, fourofAviculariinaeandfourofHar-
in taxonomy ofTheraphosidae to diagnose gen- pactirinae. Finestructureoftarsalscopulaoffive
era and subfamilies. Entire tarsal scopulae have Uruguayan species from different genera is de-
homogeneous adhesive hairs analogous to those scribed and compared. The results ofthis study
describedasTypeAbyRovner(1978)inLycosa call into question the systematic and phyloge-
spp. Divided scopulae have a longitudinal band netic significance ofthe character. The function
oflong thick setae similar to Type B ofRovner and phylogeny oftarsal scopulae in Theraphos-
(1978). Beginning with Ausserer (1871), and inae are discussed.
Simon(1889, 1892)tothepresent,thischaracter METHODS
hasbeenconsideredanimportanttaxonomictool.
G.dePikelin&Schiapelli(1973),followingAus- Takinginto accountthat width oftarsal scop-
serer(1871), considered it a key character ofIs- uladivisionincreasesinhindlegs(Raven 1985),
chnocolinae and pointed out that scopulae di- legs IV were studied to avoid ambiguity with
visioniscommoninjuvenilesofalltheraphosids. respect to entire or divided condition. Few iso-
46
PEREZ-^MILES=~SCOPULACONDITION IN THERAPHOSID SYSTEMATICS 47
Figures 1-6.—Structuresofthetarsalscopula. \.Acanthoscurriasuina,tarsusIV,ventralview,showingentire
scopula, TypeAhairs; 2. Homoeomma uruguayense, tarsusIV, ventralview, showingdividedscopulae. Arrow
showsthebandofTypeBscopulahairs,withTypeAhairslaterally; 3. TypeAhairsontarsusIVofCeropelma
longisternale, showingthecylindrical stalkandtheclub-shapeddistalzone. Theentirehairiscoveredby small
spineswhich are denserin the distal portion; 4. Tips ofType A hairs ofCeropelma longisternale, showing the
small spines covering the hairs; 5. Type B hairs on tarsus IV ofHomoeomma uruguayense, showing their
subconicalshape; 6. TypeBhairtipontarsusIVofCeropelmalongisternale, showingthesmallspinescovering
it. Scales: Figs. 1,2 = 2 mm; Figs. 3, 5 = 0.2 mm; Figs. 4, 6 = 0.02 mm.
lated long and fine hairs (not line ordered) in microscope submersing the specimens in 80%
tarsal scopulae were not considered the divided alcohol. Light microscope studies were carried
condition. out to observe and measure scopula hairs. To
Observations were made with a stereoscopic estimate spider size, carapace length (CL) was
1
THEJOURNALOFARACHNOLOGY
48
Table 1.—Tarsal scopula condition (TS) and carapace length (CL) ofholotypes oftype species, ofnon type
species(*)orcongeners(**)ofTheraphosinaeandsomeAviculariinaeandHarpactirinae.Abb=abbreviatures;
LOC = location ofexamined material: BMNH = British Museum ofNatural History, London; IB = Institute
Butantan, Sao Paulo; MACN = Museo Argentino de Ciencias Naturales, Buenos Aires; MHNP = Museum
National d'Histoire Naturelle, Paris; MHNM = Museo Nacional de Historia Natural, Montevideo; MZSP =
Museu de Zoologia, Sao Paulo; NMV = Naturhistorisches Museum, Vienna; SI Smithsonian Institution,
Washington; SAM = South African Museum, Capetown; ZMB = Zoologisches Museum, Berlin; NE = not
examined, data from literature. Eighteen types ofGrammostola spp. (all with entire scopulae) are omitteddue
to space reasons.
Abb CL TS LOC
Theraphosinae
Acanthoscurriageniculata(Koch 1842) Ag 25.5 E BMNH
Acanthoscurriasuina (Pocock 1903) As 18.5 E BMNH*
Aphonopelmaseemani(Cambridge 1897) Ase 20.0 E NE
Aphonopelma crinita(Pocock 1901) — 13.0 D BMNH*
Aphonopelmapaioma Prentice 1992 — 5.7 D NE*
Brachypelma emilia (White 1856) Be 21.5 E MHNP
Ceropelma iwM/areMello-Leitao 1923 Ci 7.8 D MZSP
Ceropelmaflavohirtus(Simon 1889) 6.0 D MHNP*
Ceropelma longisternaleSch. & Ger. 1942 __ 7.5 D MACN*
Ceropelmasemiauranticum (Simon 1897) 6.2 D MHNP*
Crypsidromus isabeUinusAusserer 1871 Cis 12.0 D NMV
Citharacanthus longipes(Cambridge 1897) Cl 26,0 E BMNH
Cyriocosmussellatus(Simon 1889) Cs 4.3 D MHNP
Cyrtopholiscursor(Ausserer 1875) Cc 13.0 E BMNH
CyclosternumschmardaeAusserer 1871 Csc 17.8 D MHNP
Cyclosternum rufohirta(Simon 1889) Cr 14.0 D MHNP*
Dryptopelmajanthina (Simon 1888) Dj 14.4 D MHNP
EupalaestruscampestratusSimon 1891 Ec 20.0 E MHNP
Eupalaestrus weijenberghi(Thorell 1894) Ew 14.6 E BMNH*
EupalaestrustarsicrassusBucherl 1947 __ 17.6 E IB*
Euathlustruculentus (Ausserer 1875) Et 20.0 E BMNH
Grammostola mollicoma (Ausserer 1875) Gm 19.0 E BMNH*
Hapalotremusalbipes Simon 1903 Ha 9.7 D MHNP
HapalopusformosusAusserer 1875 Hf 9.0 D NMV
Holothelesericeus(Simon 1903) Hs 7.2 D MHNP*
HomoeommastradiingiCambridge 1881 Hs 17.5 E BMNH
Homoeomma uruguayense(Mello-Leitao 1946) Hu 7.0 D MHNM*
Lasiodora klugii(Koch 1842) Lk 27.0 E BMNH
MygalarachnebrevipesAusserer 1871 Mb 17.0 E NMV
Megaphobema robusta (Ausserer 1875) Mr 28.0 E BMNH
Phormictopuscancerides(Latreille 1806) Pc 20.7 E MHNP
Paraphysa manicata Simon 1892 Pm 20.7 E MHNP
Pamphobeteus nigricolor{Amserer 1875) Pn 26.5 E BMNH
Phrixotrichus roseus(Guerin 1838) Pr 28.2 E MHNP
Sphaerobothria hojfmaniKarsch 1879 Sh 17.5 E BMHN**
Schizopelma macropus(Ausserer 1875) Sm 12.0 D BMNH*
Theraphosa blondi(Latreille 1804) Tb 34.2 E MHNP
Xenesthis immanis(Ausserer 1875) Xi 29.0 E BMNH
Aviculariinae
Avicularia sp. __ 22.0 E MHNM**
Ephebopus murinus (Walckenaer, 1837) 15.3 D BMNH
Iridopelma hirsutum Pocock 1901 16.5 D BMNH
Pachistopeima rufronigrum Pocock 190 ” 12.5 D BMNH
PEREZ-MILES~=-SCOPULA CONDITION IN THERAPHOSID SYSTEMATICS 49
Table L—Continued.
Abb CL TS LOC
Harpactirinae
CeratogryusdarlingiiPocock 1897 23.5 E BMNH
CoelogeniumpillansiPurcell 1902 — 24.0 E SAM
EucratosceiuslongipesPocock 1898 — 17.0 E BMNH
Harpactirella treleaveniPurcell 1902 - 6.8 D SAM
measuredwithanocuiarmicrometer.Finestruc- portion (Fig. 3). No differences in hair mor-
ture ofscopulae was documented using a scan- phology among genera were detected but the
ning electronic microscope. lengthvariesrelativewithspidersize.Totallength
Holotypes and additional material examined ofType A hairs is 0.49 mm (± 0.023) inAcan-
mm
are referenced in Table 1. InAphonopelma see- thoscurria suina (Pocock 1903); 0.66
manni (Cambridge 1897), CL and scopula con- (±0.027) in Grammostola molUcoma (Ausserer
mm
dition weretaken from Valerio (1980) and inA. 1875); 0.43 (±0.044) in Eupalaestrus wei-
mm
paloma from Prentice (1992). jenberghi(Thorell 1894); 0.35 (± 0.032) in
I followed Raven (1985) in classification and CeropelmalongisternaleSchiapelliyGerschman
nomenclaturalaspects,withthefollowingexcep- 1942; and 0.30 mm (±0.029) in Homoeomma
tions: a)Aphonopelma Pocock 1901 precedence uruguayense (Mello-Leitao 1946). All possible
over Rhechostica Simon 1892 (ICZN 1991, mean comparisons among hair length of these
Opinion 1637); b) Pterinopelma Pocock 1901 fiveUruguayanspeciesresultedinsignificantdif-
transferredtoEupalaestrusVqcoc]l 1901 (Perez- ferences {P < 0.00001). However, distal zone
Miles 1992); c) Brachypelma Simon 1891 re- length (approx. 0.096 mm) did not show signif-
stored (Schmidt 1991); d) Oligoxystre Vellard icantdifferencesinmostcomparisons,exceptbe-
1924 and Stichoplastus Simon 1889 removed tween suinawith C. longisternaleandwithH.
from Theraphosinae (Perez-Miles 1992); e) uruguayense. Hairs are covered by small spines
Ephebopus Simon 1892 transferred from Ther- oriented at an angle of30° with the axis ofthe
aphosinae to Aviculariinae (Lucas et al. 1991). hair and with tips oriented toward the apical
Statistical analyses were carried out with the region (Fig. 4); no triangular end feet were ob-
PrestaP-C,V1.1, packagedevelopedattheCen- served in such spines. These spines are distrib-
tro Ramon y Cajal, Spain. Mean comparisons uted in an orderly fashion and are more dense
were done by Student ?-test, with restrictions to in the distal region (Figs. 3, 4). Type A hairs are
variance (Snedecor’s F). moreorlessverticalorslightlyinclined(80°with
respect to the surface oftarsus) with tips orien-
RESULTS
tated distally.
Structure of scopulae.—Two types of tarsal Type B hairs are subconical (Fig. 5). They are
mm
scopulae were found within the Theraphosinae: longer (0.607 in H. uruguayenseand 0.596
entire and divided. A third type ofscopula (di- mm in C. longisternale) and thicker than Type
vided by strong spines) was present inAcantho- A; no significant differences in hair mean length
pelma Cambridge 1897 but is not considered werefoundbetweenthesetwospecies{P=0.866).
here. The entire scopula (Fig. 1) has homoge- The surface is also covered by small spines, re-
neous adhesive hairs on the ventral surface of sembling those ofType A, but less dense (Fig.
the tarsus. Following Rovner’s (1978) nomen- 6). Type B hairs have angles ofapproximately
clatureonlycosids, andwithoutestablishingany 30°withrespectto theventral surface ofthe tar-
homology (only positional criteria), I call them sus with their tips orientated distally.
Type A hairs. Character distribution.—A close relation be-
Divided scopula (Fig. 2) has another type of tweensizeandtarsalscopulaconditionwasfound
hairs (called Type B) arranged on a medial lon- (Fig. 7, Table 1). Tarsal scopulae are divided in
gitudinal band which lies between the two ven- the small species ofTheraphosinaeandentire in
tro-lateral bands ofType A hairs. large ones. Size/scopular condition relationship
Type A hairs are cylindrical in theirproximal wasobservedbetween adults ofdifferentspecies
portion (stalk) and club-shaped in their distal of differing sizes and between adults (or large
8
50 THEJOURNALOFARACHNOLOGY
7
W
/iri carapace
length
entire scopula
Bi
30 divided scopula
Figure 7.—Tarsal scopulaconditionand carapace length (in mm) in Theraphosinae species(abbreviationsas
in Table 1).
juveniles)andsmalljuvenilesofthesamespecies example, in Homoeomma (described by Simon
(which have entire scopula as adults). 1892 as having entire scopula but diagnosed by
Divided tarsal scopulae were found in type Gerschman & Schiapelli 1972 ashavingdivided
species of seven genera ofTheraphosinae, plus scopulae), large species {H. stradlingi (Cam-
Holothele and Schizopelma (non type species); bridge 1881), carapace length 17.5 mm) have
entire tarsal scopulae were found in type species entire scopulae (Fig. 8) whereas small ones {H.
of 19 genera ofTheraphosinae (Table 1). uruguayense,CL= 7mm)havedividedscopulae
Size/scopular condition relationship was also (Fig. 2). A similarphenomenonwas observedin
found between species within same genus. For Aphonopelma: A. crinita (CL = 13 mm) and A.
paloma (CL = 5.7 mm) have divided scopula
while A. seemanni (Cambridge 1897) (CL = 20
mm) has entire scopulae. Only two species con-
8 tradicting the size/scopular condition relation
were found: Cyrthopholiscursor{hussQVQv 1875)
and Cyclosternum schmardae Ausserer 1871.
Examined theraphosid species with divided
scopula have a mean CL = 10.66 mm (SD =
3.50), while species with entire scopula present
a mean CL = 22.1 1 mm (SD = 5.27). A com-
parison ofCL means showed highly significant
differencesbetween spiderswith entire ordivid-
edscopulae{t= 6,285,P=0.0000014).Asshown
mm
in Fig. 7, Theraphosinae speciesupto 12 of
carapacelengthexhibiteddividedscopulaeonly;
mm
Figure%.~Homoeommastradingli(Holotypemale), speciesbetween 12 and 18 hadeitherentire
ventralviewoftarsusIV,showingentirescopula(Type or divided scopulae, and species larger than 1
mm
A hairs). exhibited entire scopulae only. This trend
PEREZ-MILES=~SCOPULACONDITION IN THERAPHOSID SYSTEMATICS 51
wasalsoobservedintypespeciesofsomegenera abilitiestomove(burrowersandnonburrowers)
ofAviculariinae and Harpactirinae (Table 1). did not show differences in scopulae morphol-
Size/scopulardivisionrelationshipisalsohold ogy, whichwas conflictingwith the incidence of
between smalljuvenilesandlarge onesoradults adaptive pressures on locomotion. It also seems
ofthe same species (which have entire scopulae possiblethattheraphosidswithdifferentscopula
as adults). Some evidence ofa shiftinthis char- condition and CL live in similar habitats. Field
acterconditionrelatedtotheindividual’sgrowth observations in Uruguay showed that thera-
wasfound. AstudyofjuvenilesofG. mollicoma phosidsofsimilaradultsizesdonotoccurinthe
showeddividedscopulaewhenCLis 10.40mm, samehabitat, withtheexceptionofA. suinaand
scopulae proximally divided and distally entire E. weijenberghi.
when CLis 15.60, and entire scopulae when CL The exceptions observed in C. cursor and C.
mm
is 16.00 (clearlybefore maturity moltwhen schmardae could reveal possible phylogenetic
CL is more than 19.00 mm). constrains to the character.
Ontogeny and scopula condition.—All Thera-
phosidaehavedividedscopulaeinjuvenilestates,
but they become entire (Type B hairs absent) in
DISCUSSION adults of several species (Gerschman & Schia-
Character distribution and phylogenetic sig- pelli 1973). Present findings in G. mollicoma
nificance.—Since Ausserer (1871), “entire/di- showed that the entire scopula condition is
vided scopula” has been extensively considered reached much before the adult state. This sug-
as an important taxonomic tool in Theraphosi- gests that scopular condition is more related to
dae. Recently Perez-Miles (1992) polarized this size and weight than to maturity. On the other
character by ontogeny, to apply it in a prelimi- hand, small species retain divided scopulae
nary phylogenetic analysis of Theraphosinae; throughout their lives.
concluding that entire is apomorphic. Present Structureandfunction.—Tarsalscopulaehairs
results, however, question the use ofthis char- (Type A) observed here in theraphosid spiders
acter in phylogenetic systematics. At least havedifferent morphologyfrom thosedescribed
Aphonopelma and Homoeomma are polythetic for Philodromus aureolus (Foelix & Chu-Wang
for this character, although there would be sev- 1975; Foelix 1982) and from those ofLycosidae
eral synapomorphies supportingthe monophyly (Rovner 1978; Milleretal. 1988). The maindif-
ofeach genus (e. g., the digitiform apophysis of ferenceisthattriangularend-feetwerenotfound
the bulb in Homoeomma). Both states are pres- inspinesofTypeAhairsofTheraphosidae,which
entwithintheTheraphosinaeanditssistergroup, agreeswith the observationofRoscoe & Walker
{sensu Lucas et al. 1991, the Aviculariinae) and (1991) in Sericopelma rubronitens. Differences
in the Harpactirinae. A close relation between inTypeAhairsizewerefoundwithPhilodromus
body size and scopula condition in these taxa andwithinsomegeneraofTheraphosinae,prob-
andalsowithinAphonopelmaandHomoeomma ably due to a correlation with spider size. In
suggests a functional adaptation or a develop- Theraphosinae, this correlation is associated
mental effect. Influence ofenvironment and in- mainlywith hairstalk(with high variability)but
teraction with phylogeny in this character re- notwith distal zone (lessvariable). Type B hairs
mainsobscure.Consideringcharacterdistribution weresimilartothosedescribedbyRovner(1978)
andsince sizecouldchangeindependentlyin di- andMilleretal. (1988)inLycosidae;butjudging
verse taxa, similarity in scopulae condition will from their photographs, those of theraphosids
not clearly reveal phylogenetic relationship in have a more spine-like appearance.
Theraphosinae. Acanthoscurria suina and Eu- Differences in hair fine structure between
palaestrus weijenberghi, both with entire scop- TheraphosidaeandPhilodromidaemayrelateto
ulae and large CL, live in similar environments differences in adhesive requirements between
(holes in meadows), which would be interpreted both taxa: Philodromidae are small sized, light
as convergence. However, Grammostola molli- weight(withtrachea);whereasTheraphosidaeare
comaandAviculariasp., alsowithentirescopula large and heavy (with booklungs instead oftra-
andlargeCL, liveindifferentenvironments(the chea). Alsodifferencesinhairstructurewouldbe
former under stones in hills and the latter ar- related to differences in hairdistribution, which
boreal). Similarly Miller et al. (1988) observed is restricted to claw tufts in Philodromus. The
that lycosid groups with different tendencies or resemblancebetweenTypeBhairsofLycosaspp.
52 THEJOURNALOFARACHNOLOGY
andTheraphosinaewouldbeacaseoffunctional LITERATURECITED
convergence:thetractionroleduringlocomotion Ausserer, A. 1871. BeitragezukenntnisderArachn-
established by Rovner (1978) in the formerand idien familie der Territelariae Thorell (Mygalidae
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alongwiththeincreaseofbandwidthinposterior El genero Homoeomma Ausserer, 1871 (Araneae:
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arboreal) adhesive requirements probably pre- La subfamilia Ischnocolinae (Araneae: Theraphos-
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International Commission on Zoological Nomencla-
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ture. 1991. Opinion 1637, Bull. Zool. Nomencla-
uultaedcobnyditRioosnc.oeMe&chaWnailksemrs (o1f9a9d1h)estoiosncoatpturliabe- Luctausr,e,S.4,8P:.1I6.6d-a16S7il.va&R.Bertani. 1991. Thegenus
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smoothsurfaces, themainroleattributedto sco- Miller, G. L., P. R. Miller & A. Brady. 1988. Ad-
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