THE MALAGASY SPECIES OF THE CREVICE WEAVER GENUS ANDOHARANO (ARANEAE: FILISTATIDAE)

2019·7.7 MB·English

Bulletin

of the
Museum of Comparative Zoology
Volume 162, Number 4 10 April, 2019
The Malagasy Species of the Crevice Weaver
Genus Andoharano (Araneae: Filistatidae)
Ivan L. F. Magalhaes and Cristian J. Grismado
US ISSN 0027-4100
MCZ Publications
Museum of Comparative Zoology
Harvard University
26 Oxford Street
Cambridge, MA 02138
[email protected]
© The President and Fellows of Harvard College 2019 HARVARD UNIVERSITY | CAMBRIDGE, MASSACHUSETTS, U.S.A.
BULLETIN OF THE
Museum of Comparative Zoology
BOARD OF EDITORS
Editor: Jonathan Losos
Managing Editor: Melissa Aja
Associate Editors: Andrew Biewener, Scott Edwards,
Brian Farrell, Gonzalo Giribet, James Hanken,
Hopi Hoekstra, George Lauder, James McCarthy,
Naomi Pierce, Stephanie Pierce, and Mansi Srivastava
Publications Issued or Distributed by the
Museum of Comparative Zoology
Harvard University
Bulletin 1863–
Breviora 1952–
Memoirs 1865–1938
Johnsonia, Department of Mollusks, 1941–1974
Occasional Papers on Mollusks, 1945–
General queries, questions about author guidelines, or permissions for
MCZ Publications should be directed to the editorial assistant:
MCZ Publications
Museum of Comparative Zoology
Harvard University
26 Oxford Street
Cambridge, MA 02138
[email protected]
EXCHANGES AND REPRINTS
All of our publications are offered for free on our website:
https://mcz.harvard.edu/publicationshome
To join our exchange program, please contact the
Ernst Mayr Library:
[email protected].
This publication has been printed on acid-free permanent paper stock. Photo on the front cover:
© The President and Fellows of Harvard College 2019. Andoharano decaryi, female lectotype, habitus, dorsal.
THE MALAGASY SPECIES OF THE CREVICE WEAVER GENUS
ANDOHARANO (ARANEAE: FILISTATIDAE)
IVANL.F.MAGALHAES1ANDCRISTIANJ.GRISMADO1
ABSTRACT. The Malagasy species of the filistatid er, it remains very poorly studied, as
spider genus Andoharano Lehtinen arerevised.The
virtually nothing is known about it except
four previously known species, all from caves, are
redescribed and have their type material illustrated. for the original descriptions of its species.
The paratypes of A. milloti Legendre are not Gray (1995) suggested Andoharano is a
conspecificwithitsholotypeandarehereindescribed
close relative of the mainland African
as A. rollardae sp. nov. The first epigean represen-
tativesofthegenusontheislandarerecorded,anda genus Afrofilistata Benoit, a relationship
further six species are described: A. simoni sp. nov., purportedly supported by characters of the
A.zonsteinisp.nov.,A.woodaesp.nov.,A.lehtineni
male palp. However, the poor taxonomy of
sp. nov., A. griswoldi sp. nov., and A. ramirezi sp.
nov. Interestingly, each of the species occurring in the genus and the lack of knowledge of its
cavesismorphologicallysimilartoadifferentepigean basic morphology hamper a more conclu-
species, suggestingrepeated invasions of the subter-
sive hypothesis for its phylogenetic place-
ranean realm. The fine morphology of the genus is
illustrated, including the first scanning electron ment.
microscopyimagesofgenitalia,spinnerets,andother Four of the known species of Andohar-
structures.
anocomefromMadagascar,all fromcaves.
Unfortunately, their original descriptions
Key words: Africa, Madagascar, New Species, Prithi-
nae,Spider,Taxonomy lack useful illustrations allowing their
recognition, and the type species of the
genus has never been illustrated (Simon,
INTRODUCTION
1901;Fage,1945;Legendre,1971). On the
The spider genus Andoharano Lehtinen
other hand, the examination of a series of
was erected to accommodate two Malagasy
collections of Malagasy spiders deposited
species then placed in the Eurasian genus
mainly in the California Academy of
Filistata Latreille (Lehtinen, 1967): F.
Sciences revealed the genus was not only
grandidieri Simon 1901 and F. decaryi
more diverse than previously known, but
Fage 1945. Later, additional species have
that it is also represented by several
been described from Madagascar (Legen-
hitherto undescribed epigean species.
dre,1971)andfromNamibia(Zonsteinand
Thus, the aims of this contribution are: 1)
Marusik, 2015), and currently the genus
to redescribe and illustrate the previously
includesfivespecies(WSC,2018).Uponits
known species of Malagasy Andoharano
originaldescription,Lehtinen(1967)noted
this was a ‘‘highly deviant genus.’’ Howev- based on the examination of type material,
2) describe new taxa, especially of the
1 Divisio´n Aracnolog´ıa, Museo Argentino de epigean fauna, and 3) provide data on the
Ciencias Naturales ‘‘Bernardino Rivadavia’’ – general morphology of the genus for a
CONICET, Av. A´ngel Gallardo 470, C1405DJR,
better understanding of its phylogenetic
Buenos Aires, Argentina. Author for correspon-
dence:[email protected] placement.
Bull. Mus. Comp. Zool., 162(4): 263–307, April, 2019 263
Downloaded From: https://bioone.org/journals/Bulletin-of-the-Museum-of-Comparative-Zoology on 11 Apr 2019
Terms of Use: https://bioone.org/terms-of-use Access provided by Innodata
264 Bulletin of the Museum of Comparative Zoology, Vol. 162, No. 4
MATERIALS AND METHODS arid, and semiarid regions (Gray, 1995;
Specimens for this study come from the Magalhaes and Ram´ırez, 2017; WSC,
following institutions: CAS—California 2018). Madagascar has a wide range of
Academy of Sciences (San Francisco, habitats,includingmorehumidhabitatsand
rainforests in the eastern side and a drier
USA); MCZ—Museum of Comparative
portion in the west, mainly due to the rain
Zoology, Harvard University (Cambridge,
shadow of the highlands in the central
Massachusetts, USA); MNHN—Muse´um
portion of the island. As a consequence,
National d’Histoire Naturelle (Paris,
Andoharano seems to be restricted to the
France);MRAC—Muse´eRoyaldel’Afrique
dry portions of the island, mainly in the
Centrale (Tervuren, Belgium). Complete
tropical dry forests in the north and the
voucher and locality data for all specimens
spiny forests and xeric shrublands in the
examined in this study are given in Supple-
mentary Table 1.2 southwest (Fig. 1). Their apparent absence
in the tropical dry forests in the western
The format of description, including
portion of the island (e.g., in Mahajanga) is
notation of leg macrosetae, follows Mag-
puzzlingandmaybeduetoasamplingbias.
alhaes and Ram´ırez (2017). All measure-
Interestingly, we here describe epigean
ments are in millimeters. Female internal
representatives that bear morphological
genitalia were examined after digestion
resemblance to the previously known cave
using a pancreatin solution. For imaging,
species. For instance, Andoharano decaryi,
theywereplacedinlactic acidintemporary
A. rollardae, and A. grandidieri are very
slides andphotographed under an Olympus
similar to A. simoni, A. griswoldi, and A.
BH2transmittedlightmicroscope;drawings
lehtineni, respectively, mainly in their gen-
weremadewithacameralucida.Malepalps
ital morphology. The cave species are
of some species were cleared in clove oil.
usually paler and have relatively longer
Measurements and photographs of habitus
appendages than their epigean counter-
and male palps were made with a Leica
parts, suggesting some degree of troglo-
M165C stereoscopic microscope using Lei-
morphism. Furthermore, if the
ca Application Suite 3.8. All stacked-focus
morphological resemblance was to indicate
imageswereprocessedinHeliconFocus6.8
that species in each of these three pairs are
(https://www.heliconsoft.com). Specimens
sister taxa, this would mean that Andohar-
were prepared for scanning electron mi-
ano invaded the subterranean habitat more
croscopy as described in Magalhaes and than oncein their evolutionaryhistory.This
Ram´ırez (2017), and all images were taken hypothesiscouldbeeventuallytestedwitha
in a Philips FEI XL 30 TMP scanning densely sampled phylogeny of the genus
electronmicroscopeatMuseoArgentinode that is beyond the scope of this paper.
Ciencias Naturales, Buenos Aires. The morphological details of representa-
tives of Andoharano unveiled during the
RESULTS AND DISCUSSION courseofthisstudy hintat thephylogenetic
placement of the genus. Gray (1995)
The study of the new specimens allowed
suggested Andoharano is closely related to
us to recognize at least seven undescribed
Afrofilistata due to the possession of a
species of Andoharano, elevating the total
‘‘lozenge-shaped cymbium.’’ The cymbium
from four to 11 species in Madagascar.
in Andoharano is indeed small when
Filistatids are more diverse in subtropical,
comparedwiththatofotherfilistatidgenera
and has a convex margin that could be
2 Supplementary material referenced in this
interpreted in that way. However, the
paper is available online at www.mcz.harvard.edu/
Publications/. cymbium in Afrofilistata is horseshoe-
Downloaded From: https://bioone.org/journals/Bulletin-of-the-Museum-of-Comparative-Zoology on 11 Apr 2019
Terms of Use: https://bioone.org/terms-of-use Access provided by Innodata
A REVISION OF MALAGASY ANDOHARANO (cid:2) Magalhaes and Grismado 265
Figure1. DistributionmapofAndoharanospeciesinMadagascar.Drawingsoffemalespermathecaeindorsalview,alltoscale.
shaped,andthisgenuslacksaciniformgland posterior end of the carapace and lacking
spigots (Magalhaes, unpublished data), submarginal bands, is also reminiscent of
characters that would instead ally it with that of Antilloides and Filistatoides F.O.
Pritha Lehtinen and its relatives. We here
Pickard-Cambridge.Thismightsuggestthat
show that Andoharano presents true feath-
Andoharano has affinities with the New
erysetaeandastrongretrolateralcondylein
World fauna, with puzzling biogeographic
the metatarsus II of males, characters
implications.
shared among the Prithinae only with New
World genera (e.g., Pikelinia Mello-Leita˜o
and Antilloides Brescovit, Sa´nchez-Ruiz & SYSTEMATICS
Alayo´n). The pattern of coloration of the
carapace, with a dark hourglass-shaped Filistatidae Ausserer, 1867
band extending from the eyes to the Andoharano Lehtinen, 1967
Downloaded From: https://bioone.org/journals/Bulletin-of-the-Museum-of-Comparative-Zoology on 11 Apr 2019
Terms of Use: https://bioone.org/terms-of-use Access provided by Innodata
266 Bulletin of the Museum of Comparative Zoology, Vol. 162, No. 4
Andoharano Lehtinen, 1967: 214. Type species carapace longer than wide, thoracic fovea
by original designation Filistata decaryi absent. Clypeus slightly notched in males.
Fage,1945 Eyes united in a low tubercle, anterior
median eye subequal to the anterior lateral
Diagnosis. This genus can be easily
eye. Eye apodemes absent (Fig. 2B).
distinguished from all Old World filistatids
Featherysetaepresent(Figs.3C,4F),white
by the presence of true feathery setae (Fig.
setae absent. Sternum slightly longer than
2A).Bothmalesandfemalesusuallypresent
wide; sigilla difficult to observe but detect-
an hourglass-shaped brown band extending
able under SEM, a single posterior pair
from the eye region to the posterior end of
(Fig. 2D). Cheliceral gland slightly raised
the carapace (Fig. 12; absent in some
(Figs. 3B, 4D). Female palp tarsal macro-
species).Malescanbefurtherdistinguished
setae absent. Leg formula 1423. Femora,
by the small cymbium with a straight
metatarsi, and tarsi macrosetae absent.
anterior margin and bearing a retrolateral
Tibiae macrosetae usually absent, but pres-
forelock of long setae that run along with
ent in the tibiae I of males of some species.
the bulb (Figs. 9E, 41D) and by the shape
Male tarsi entire. Trichobothria with ring-
of the sperm duct, which makes a strong
like socket present in tibiae and metatarsi,
backwards curve in the last section (Fig. 8).
in the last case, reaching the distal end of
Females possess a large fan-shaped seta in
the article (Fig. 4G). Male metatarsus II
the posterior median spinnerets (Figs. 6D,
bearing a strong retrolateral condyle, with
E, 7E) (convergent in some other prithine
modified metatarsus stopper, and apparent-
genera). The shape of the outer spermathe-
ly lacking a tarsal socket (Fig. 17F); in A.
cae (elongate, drop-shaped, widest distally,
ramirezi,witharetrolateralexcavation(Fig.
with pores restricted to apical portion; Figs.
41H). Calamistrum sessile, in some species
11D, 14) is typical of this genus, but some
with numerous setae, composed of three
species (A. ansieae, A. ramirezi) have
parallel rows; teeth present in all setae
rounded spermathecae. (Figs. 3F, G, 4I–K). Abdomen suboval.
Description. Size: small spiders, ranging Posterior respiratory system lacking lateral
from 1.82 (A. ramirezi male) to 6.7 mm (A. tracheae or transverse duct (Fig. 5). Anal
decaryi female) in total length (appendages tubercleunmodified.Spinnerets(Figs.6,7):
excluded). Color and pattern: carapace, cribellum divided, with each spinning field
sternum, labium, and appendages pale very wide, cribellum spigots strobilate.
yellowtoyellowishbrown.Carapaceusually Anterior lateral spinnerets (ALS) with
with dark brown hourglass-shaped band anterior row of setae, one major ampullate
extending from the eye region to the gland spigot on the anterior margin, and
posterior end of the carapace; submarginal about 15–35 piriform gland spigots. Poste-
bands absent; dark brown lining on sides of rior median spinnerets (PMS) flattened,
carapace present or absent. Sternum lightly with spatulate setae, a large fan-shaped seta
colored, occasionally with brown pigment medially, and one aciniform gland spigot,
on anterior margin. Chelicerae with an one minor ampullate gland spigot, and one
anterior brown patch. Legs usually with paracribellar gland spigot. Posterior lateral
brownannulations,threeonthefemoraand spinnerets (PLS) with around 10–20 acini-
two on the tibiaeandmetatarsi; annulations form gland spigots, and a longer spigot
sometimes lacking, especially in cave-dwell- probably serving the paracribellar gland.
ing species. Abdomen with well-defined Male genitalia (Figs. 8, 9): palpal femur
chevron-like pattern, usually with around straight,unarmed.Palpaltibiaincrassateor,
six markings; ventral side lightly colored, morecommonly,subconical,widestdistally;
with bands alongside spinnerets. Prosoma: in all species, except for A. ramirezi and A.
Downloaded From: https://bioone.org/journals/Bulletin-of-the-Museum-of-Comparative-Zoology on 11 Apr 2019
Terms of Use: https://bioone.org/terms-of-use Access provided by Innodata
A REVISION OF MALAGASY ANDOHARANO (cid:2) Magalhaes and Grismado 267
Figure2. Andoharanosimonisp.nov.,femalefromMadagascar,Toliara,Eloetse(CAS9014032),cephalothorax.A.Dorsal.
Insetshowingfeatheryseta.B.Eyeregion,dorsal.C.Lateral.D.Ventral.Insetshowingsternumsigillum.E.Anterior.F.Anterior,
detailoflabrumandleftserrula.Abbreviations:L¼labrum,S¼serrula.
Downloaded From: https://bioone.org/journals/Bulletin-of-the-Museum-of-Comparative-Zoology on 11 Apr 2019
Terms of Use: https://bioone.org/terms-of-use Access provided by Innodata
268 Bulletin of the Museum of Comparative Zoology, Vol. 162, No. 4
Figure 3. Andoharano simoni sp. nov., female from Madagascar, Toliara, Eloetse (CAS 9014032), appendages. A. Left
chelicera,anterior.B.Same,posterior.Insetsshowingcheliceraglandandrowofchemosensorysetae.C.Leftpalp,retrolateral.
D.LeftlegI,claws,retrolateral.InsetshowingtarsalorganIV,retrolateral.E.LeftlegIV,metatarsusstopper,dorsal.Insetshowing
trichobothriumonmetatarsusI.F.Leftcalamistrum,arrowstoeachofthethreerows.G.Calamistrumsetae,detail.Abbreviations:
CL¼apexofchelicerallamina,Ch¼chemosensoryseta,PL¼promarginallobe,SS¼slitsensilla.
Downloaded From: https://bioone.org/journals/Bulletin-of-the-Museum-of-Comparative-Zoology on 11 Apr 2019
Terms of Use: https://bioone.org/terms-of-use Access provided by Innodata
A REVISION OF MALAGASY ANDOHARANO (cid:2) Magalhaes and Grismado 269
Figure 4. Andoharano griswoldi sp. nov., female from Madagascar, Antsiranana, Montagne des Franc¸ais (CAS 9000850),
appendages.A.Leftchelicera,anterior.B.Same,posterior.C.Palpaltarsalorgan,retrolateral.D.Cheliceragland.E.Leftpalp,
retrolateral.F.MetatarsusI,dorsal,detailoftrichobothriaandfeatherysetae.G.LeftlegI,metatarsusstopper,dorsal.H.Tarsal
claws IV, retrolateral. I. Left calamistrum, dorsal. J. Same, retrolateral. K. Detail of calamistrum setae. Abbreviations: Ch¼
chemosensoryseta,MS¼metatarsusstopper,PL¼promarginallobe,SS¼slitsensilla,Tr¼trichobothria.
Downloaded From: https://bioone.org/journals/Bulletin-of-the-Museum-of-Comparative-Zoology on 11 Apr 2019
Terms of Use: https://bioone.org/terms-of-use Access provided by Innodata
270 Bulletin of the Museum of Comparative Zoology, Vol. 162, No. 4
Figure5. Andoharanosimonisp.nov.,femaleparatype(MCZ54669),clearedabdomenshowingspermathecaeandposterior
tracheae,dorsalview.Scalebars¼0.5mm(A)or0.1mm(B).
ansieae, with strong apical setae on the ansieae Zonstein & Marusik, 2015; A.
ventral side. Cymbium short, with straight griswoldi sp. nov.; A. lehtineni sp. nov.; A.
anterior margin, bearing a forelock of long ramirezi sp. nov.; A. rollardae sp. nov.; A.
setae that run parallel to the bulb. Bulb simoni sp. nov.; A. woodae sp. nov.; A.
drop-shaped to trapezoidal. Paraembolic zonsteini sp. nov.
lamina absent. Embolus straight to curved, Distribution. The 11 species treated in
retrolaterally bent; small slit present in the this contribution are endemic to the dry
retrolateral side; keel absent. Sperm duct portions of Madagascar. An additional
with a double coil, the second one not species has been described from Namibia
complete; a strong backwards curve is byZonsteinandMarusik(2015),anddetails
present in the last section. Fundus pointing of the male palp indicate it indeed belongs
ventrally. Basal bulb sclerite flattened; the to the Andoharano lineage (see ‘‘Internal
tendon of the claw flexor muscle (tM29) Relationships and Monophyly’’ section be-
attachesdorsally.Femalegenitalia(Figs.10, low). Because the filistatid fauna of main-
11): external region unsclerotized, with land Africa has never been revised and is
dark, strong pubescence. Interpulmonary severely undersampled, we would expect
fold with straight anterior margin and sub- additional species of the genus to be
squarish corners. Uterus externus membra- discovered in new localities in the future.
nous. Spermathecae: two pairs present Internal Relationships and Monophyly.
(except for A. ansieae, apparently with a Most Malagasy species of the genus have a
single spermatheca); the inner pair sub- rather uniform morphology and are similar
triangular to flattened apically, the outer to A. decaryi, the generotype. At least two
one usually elongate, drop-shaped (except character states are shared by these species
in A. ramirezi and A. ansieae, rounded), (and not by other filistatid genera): the
with pores restricted to the apical portion presenceof incrassate setae inthe venter of
on a slightly more sclerotized sector that themalepalpaltibia(Fig.27)andthedrop-
appears darker in light microscope shaped outer spermathecae with pores
Composition. Twelve species: Andohar- restricted to its apical portion (Fig. 11).
ano grandidieri (Simon, 1901); A. decaryi However, two species deviate from this
(Fage, 1945); A. milloti Legendre, 1971; generalpatternandlackthesetwocharacter
Andoharano monodi Legendre, 1971; A. states: A. ramirezi (from southern Mada-
Downloaded From: https://bioone.org/journals/Bulletin-of-the-Museum-of-Comparative-Zoology on 11 Apr 2019
Terms of Use: https://bioone.org/terms-of-use Access provided by Innodata