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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.195.2342" ID-GBIF-Dataset="4882baaf-3747-4865-87e1-49e725955118" ID-PMC="PMC3361087" ID-Pensoft-Pub="1313-2970-195-1" ID-PubMed="22679386" ModsDocAuthor="" ModsDocDate="2012" ModsDocID="1313-2970-195-1" ModsDocOrigin="ZooKeys 195" ModsDocTitle="The velvet spiders: an atlas of the Eresidae (Arachnida, Araneae)" checkinTime="1451249010182" checkinUser="pensoft" docAuthor="Miller, Jeremy A., Griswold, Charles E., Scharff, Nikolaj, Řezac, Milan, Szűts, Tamas &amp; Marhabaie, Mohammad" docDate="2012" docId="B97386714EABCE1B7A9F2A94D5FBD963" docLanguage="en" docName="ZooKeys 195: 1-144" docOrigin="ZooKeys 195" docSource="http://dx.doi.org/10.3897/zookeys.195.2342" docTitle="Gandanameno Lehtinen" docType="treatment" docVersion="3" lastPageNumber="58" masterDocId="FFABFFD09B65FF5EFFA6FFE4FFE1FFC2" masterDocTitle="The velvet spiders: an atlas of the Eresidae (Arachnida, Araneae)" masterLastPageNumber="144" masterPageNumber="1" pageNumber="47" updateTime="1668153919305" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>The velvet spiders: an atlas of the Eresidae (Arachnida, Araneae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Miller, Jeremy A.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Griswold, Charles E.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Scharff, Nikolaj</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Řezac, Milan</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Szűts, Tamas</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Marhabaie, Mohammad</mods:namePart>
</mods:name>
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<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
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<mods:part>
<mods:date>2012</mods:date>
<mods:detail type="volume">
<mods:number>195</mods:number>
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<mods:start>1</mods:start>
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<mods:url>http://dx.doi.org/10.3897/zookeys.195.2342</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.195.2342</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-195-1</mods:identifier>
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<treatment ID-GBIF-Taxon="152035665" LSID="urn:lsid:plazi:treatment:B97386714EABCE1B7A9F2A94D5FBD963" httpUri="http://treatment.plazi.org/id/B97386714EABCE1B7A9F2A94D5FBD963" lastPageId="58" lastPageNumber="58" pageId="46" pageNumber="47">
<subSubSection pageId="46" pageNumber="47" type="nomenclature">
<paragraph pageId="46" pageNumber="47">
<taxonomicName LSID="http://species-id.net/wiki/Gandanameno" authority="Lehtinen" class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="46" pageNumber="47" phylum="Arthropoda" rank="genus">Gandanameno Lehtinen</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="46" pageNumber="47" type="reference_group">
<paragraph pageId="46" pageNumber="47">
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="46" pageNumber="47" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
Lehtinen, 1967: 235. Type species
<taxonomicName class="Arachnida" family="Eresidae" genus="Eresus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Eresus spenceri" order="Araneae" pageId="46" pageNumber="47" phylum="Arthropoda" rank="species" species="spenceri">Eresus spenceri</taxonomicName>
Pocock, 1900.
</paragraph>
</subSubSection>
<subSubSection lastPageId="47" lastPageNumber="48" pageId="46" pageNumber="47" type="note">
<paragraph pageId="46" pageNumber="47">Note.</paragraph>
<paragraph lastPageId="47" lastPageNumber="48" pageId="46" pageNumber="47">
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="46" pageNumber="47" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
contains five recognized species from eastern and southern Africa. We examined the collection holdings of several museums and most primary type speci
<pageBreakToken pageId="47" pageNumber="48" start="start">mens</pageBreakToken>
. The oldest available name,
<taxonomicName class="Arachnida" family="Eresidae" genus="Eresus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Eresus fumosus" order="Araneae" pageId="47" pageNumber="48" phylum="Arthropoda" rank="species" species="fumosus">Eresus fumosus</taxonomicName>
C. L. Koch, 1837, apparently lacks any type specimen (
<bibRefCitation author="Lehtinen, PT" journalOrPublisher="Annales Zoologici Fennici" pageId="95" pageNumber="96" pagination="199 - 468" title="Classification of the cribellate spiders and some allied families." volume="4" year="1967">Lehtinen 1967</bibRefCitation>
: 235). We evaluated morphological variation and analyzed DNA sequences from 24 individuals based on previously published and new data. Descriptions are based on a female specimen from Tanzania (ZMUC 19970530, ZMUC) and a male from Zimbabwe (AcAT 2005/123, NCA) supplemented by collections mostly from South Africa.
</paragraph>
</subSubSection>
<subSubSection lastPageId="50" lastPageNumber="51" pageId="47" pageNumber="48" type="diagnosis">
<paragraph pageId="47" pageNumber="48">Diagnosis.</paragraph>
<paragraph lastPageId="50" lastPageNumber="51" pageId="47" pageNumber="48">
Distinguished from other eresids except
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="47" pageNumber="48" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
by the position of the PLE which are both advanced (&lt;0.28) and widely spaced (PER/AER&gt; 0.95; Fig. 9F, H); other eresids with advanced PLE (e.g.,
<taxonomicName class="Arachnida" family="Eresidae" genus="Stegodyphus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Stegodyphus" order="Araneae" pageId="47" pageNumber="48" phylum="Arthropoda" rank="genus">Stegodyphus</taxonomicName>
, some
<taxonomicName class="Arachnida" family="Eresidae" genus="Paradonea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paradonea" order="Araneae" pageId="47" pageNumber="48" phylum="Arthropoda" rank="genus">Paradonea</taxonomicName>
) have them closer together (PER/AER &lt;0.90) than the ALE (e.g., Figs 10B, 11F). Male further
<pageBreakToken pageId="48" pageNumber="49" start="start">distinguished</pageBreakToken>
from other eresids except
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="48" pageNumber="49" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
by the more or less ventral-dorsal axis of the palpal bulb with the embolus encircling the ventral part (Figs 13D, E, 48
<normalizedToken originalValue="AC">A-C</normalizedToken>
; proximal-ventral in other eresids with the embolus encircling the distal part, e.g., Fig. 20
<normalizedToken originalValue="AF">A-F</normalizedToken>
) and conductor arising from the center of the tegulum with opposing projections covering much of the palpal bulb (Fig. 55C); distinguished from
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="48" pageNumber="49" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
by the lack of prominent tubercles bearing the ALE (Fig. 9F; compare with Fig. 8J) and the fringed conductor (Fig. 55C, E; smooth in
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="48" pageNumber="49" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
, Fig. 34D). Female further distinguished from other eresids except
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="48" pageNumber="49" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
and
<taxonomicName class="Arachnida" family="Eresidae" genus="Eresus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Eresus walckenaeri" order="Araneae" pageId="48" pageNumber="49" phylum="Arthropoda" rank="species" species="walckenaeri">Eresus walckenaeri</taxonomicName>
by the copulatory openings, which are broadly separated by hirsute cuticle (Figs 17
<normalizedToken originalValue="AC">A-C</normalizedToken>
, 59A; separated by a gla
<pageBreakToken pageId="49" pageNumber="50" start="start">brous</pageBreakToken>
median lobe in other eresids, e.g., Figs 16B, 29C);
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="49" pageNumber="50" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
and
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="49" pageNumber="50" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
together distinguished from other eresids including
<taxonomicName class="Arachnida" family="Eresidae" genus="Eresus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Eresus walckenaeri" order="Araneae" pageId="49" pageNumber="50" phylum="Arthropoda" rank="species" species="walckenaeri">Eresus walckenaeri</taxonomicName>
by the vulva, which have the spermatheca extending anterior of the spermathecal heads, together subtended by helical copulatory ducts (Figs 17
<normalizedToken originalValue="DF">D-F</normalizedToken>
, 59C; other eresids have the spermathecal head anterior, spermatheca posterior, and copulatory ducts other than helical, e.g., Fig. 29D), and by the subdivided PMS (Fig. 57C; entire in other eresids) with numerous short, conical, cylindrical gland spigots (Fig. 58E; cylindrical gland spigots absent or difficult to distinguish from aciniform gland spigots in other eresids); distinguished
<pageBreakToken pageId="50" pageNumber="51" start="start">from</pageBreakToken>
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="50" pageNumber="51" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
by the somewhat more prominent paired atria (Figs 17
<normalizedToken originalValue="AC">A-C</normalizedToken>
, 59A; compare with Figs 16C, 37D) and possibly by having three loops of the copulatory duct (Figs 17
<normalizedToken originalValue="DF">D-F</normalizedToken>
, 59C; fewer in
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="50" pageNumber="51" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
, although this character has been investigated in few
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="50" pageNumber="51" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
species, Figs 16F, 37E). Both sexes usually distinguished from
<taxonomicName class="Arachnida" family="Eresidae" genus="Dresserus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dresserus" order="Araneae" pageId="50" pageNumber="51" phylum="Arthropoda" rank="genus">Dresserus</taxonomicName>
by the two part cribellum (Figs 57E, 60E, compare with Fig. 36F), although signs of the distinctive 4-part cribellum are sometimes apparent in
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="50" pageNumber="51" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="57" lastPageNumber="58" pageId="50" pageNumber="51" type="distinguishing species">
<paragraph pageId="50" pageNumber="51">Distinguishing species.</paragraph>
<paragraph lastPageId="53" lastPageNumber="54" pageId="50" pageNumber="51">
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="50" pageNumber="51" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
are variable in several conspicuous characteristics. In the male, these include the presence or absence of a cheliceral boss (Fig. 56
<normalizedToken originalValue="CF">C-F</normalizedToken>
), palp size, details of the conductor shape, and the curvature and position of the tegular sperm duct (Figs 48B,
<normalizedToken originalValue="DF">D-F</normalizedToken>
, S2
<normalizedToken originalValue="AJ">A-J</normalizedToken>
). In the female, these include the height of the cephalic region, and the presence of cuspules of various weights on the carapace, sternum, and basal segments of legs I, II, and sometimes III (Figs 50, 52
<normalizedToken originalValue="AF">A-F</normalizedToken>
, 53
<normalizedToken originalValue="AF">A-F</normalizedToken>
, 54
<normalizedToken originalValue="AF">A-F</normalizedToken>
), also details of the epigynum shape (Figs 17
<normalizedToken originalValue="AC">A-C</normalizedToken>
, S2K, L, S3
<normalizedToken originalValue="AF">A-F</normalizedToken>
,
<normalizedToken originalValue="HL">H-L</normalizedToken>
). It has been noted previously that multiple forms may occur sympatrically and that intermediate combinations make species determination problematic (
<bibRefCitation author="Tucker, RWE" journalOrPublisher="Annals of the South African Museum" pageId="96" pageNumber="97" pagination="439 - 488" title="Contributions to the South African Arachnid Fauna. II. On some new South African spiders of the families Barychelidae, Dipluridae, Eresidae, Zodariidae, Heracliidae, Urocteidae, Clubionidae." volume="17" year="1920">Tucker 1920</bibRefCitation>
). Indeed, in most regions where several specimens are known, a wide range of variation can be
<pageBreakToken pageId="51" pageNumber="52" start="start">found</pageBreakToken>
(Fig. 50). We attempted to sequence DNA from several museum specimens. Unfortunately, most of our successes were from specimens at one end of the range of variation, i.e., specimens with a moderately to strongly raised cephalic region and abundant, heavy cuspules on the carapace, sternum, and anterior legs. The three con
<pageBreakToken pageId="52" pageNumber="53" start="start">trasting</pageBreakToken>
female specimens (13-10: KwaZulu-Natal; 18-04: AcAT 2002/181; 14517) did not group together. Instead, relationships appear to be more strongly tied to geography than to morphology (Fig. 51). The genitalia of 23 out of 24
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="52" pageNumber="53" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
<pageBreakToken pageId="53" pageNumber="54" start="start">specimens</pageBreakToken>
included in the molecular phylogenetic analysis are photo-documented in the electronic supplementary materials (Figs S2
<normalizedToken originalValue="AL">A-L</normalizedToken>
, S3
<normalizedToken originalValue="AL">A-L</normalizedToken>
).
</paragraph>
<paragraph lastPageId="55" lastPageNumber="56" pageId="53" pageNumber="54">
We looked for patterns of variation in adult female morphology. Adult female specimens are much more abundant in collections than males. We defined eight geo
<pageBreakToken pageId="54" pageNumber="55" start="start">graphic</pageBreakToken>
regions based on two criteria: 1) concentration of specimens available to us and/or 2) type localities of
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
species (Figs 49, 50). We assessed a series of characters: carapace length, carapace width, presence and strength of cuspules on the ventral surface of femur I and II, prosoma, and sternum for all specimens available
<pageBreakToken pageId="55" pageNumber="56" start="start">from</pageBreakToken>
these regions. When cuspules are present on the prosoma, they always occur on the femurs, but the reverse is not always true. So the set of specimens with cuspules on the femurs is larger than and contains the set of specimens with cuspules on the prosoma. There appears to be a general trend towards greater spinulation and a higher carapace with increasing size (carapace length), but the spinulation data are fairly noisy (Fig. 50). No morphological pattern emerged to segregate specimens by region. Not all degrees of spinulation were observed in every geographic region.
</paragraph>
<paragraph lastPageId="57" lastPageNumber="58" pageId="56" pageNumber="57">
<pageBreakToken pageId="56" pageNumber="57" start="start">Based</pageBreakToken>
on the combination of genetic, morphometric, and spinulation data, we see no evidence that the characters traditionally used to discriminate
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
species are valid. We speculate that ontogenetic factors are responsible for the morphological variation in this genus (perhaps juvenile nutrition or post adult molting). However, we judge that the synonymy of all
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
species is premature, particularly in light of the phylogeographic signal suggested by the molecular data and the limited avai
<pageBreakToken pageId="57" pageNumber="58" start="start">lability</pageBreakToken>
of specimens from beyond the Republic of South Africa. We therefore identify our specimens simply as
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="57" pageNumber="58" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
sp. We encourage additional investigations that might further elucidate the systematics of
<taxonomicName class="Arachnida" family="Eresidae" genus="Gandanameno" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gandanameno" order="Araneae" pageId="57" pageNumber="58" phylum="Arthropoda" rank="genus">Gandanameno</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="58" lastPageNumber="59" pageId="57" pageNumber="58" type="natural history">
<paragraph pageId="57" pageNumber="58">Natural history.</paragraph>
<paragraph lastPageId="58" lastPageNumber="59" pageId="57" pageNumber="58">
Associated with crevices in or under rocks or tree bark in savanna, parks, and gardens. They build a silken tube with a widened entrance; the tube of an adult female under bark can be up to 1 m long (Martin Forman, personal observation). Also found under tree bark well above the ground in dense aggregations (Nikolaj Scharff, Jere
<pageBreakToken pageId="58" pageNumber="59" start="start">my</pageBreakToken>
Miller, Iringa, Tanzania). Prey remnants are placed at the bottom of the tube. Clutches consist of tens of juveniles (Martin Forman, personal observation). Juveniles do not feed on their
<normalizedToken originalValue="mothers">mother's</normalizedToken>
corpse and adult females can produce a number of sequential clutches. Males mature in less than one year, females take longer (Martin Forman, personal observation).
</paragraph>
</subSubSection>
</treatment>
</document>