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The study pursues the description of covering setae across the whole family Gnaphosidae with using SEM. A detailed morphology of the setae of ground spiders (Araneae, Gnaphosidae) is presented. The six major types of covering setae recognized among gnaphosid spiders: squamose, plumose, lanceolate, pinnate, arborate, and sicate setae. Squamose setae are characteristic to Micaria lenzi and Nauhea tapa species. Plumose setae are more common in ground spiders and occur in genera Drassodes, Haplodrassus, Anagraphis, Nodocion, Zelotes, and species Berlandina caspica, Nomisia aussereri, Minosiella intermedia, Sosticus loricatus, Leptodrassus memorialis, Intruda signata, Parasyrisca caucasica, Scopoides catharius, Echemoides tofo, Zimiromus medius, Encoptarthria echemophthalma, Apodrassodes trancas, Apopyllus silvestri, Hemicloea sundevalli, Zelanda erebus, Orodrassus assimilis, Callilepis nocturna, and Synaphosus turanicus. Species Matua valida, Anzacia gemmea, Hypodrassodes maoricus, Homoeothele micans, and Scotophaeus blackwalli have lanceolate setae. Spiders of genus Gnaphosa have pinnate setae. Fedotovia uzbekistanica has arborate setae. Species Cesonia bilineata, Herpyllus propinquus, Litopyllus temporarius, Aphantaulax seminigra, and Kishidaia conspicua have sicate setae. Some genera, such as Drassodes and Synaphosus have a combination of different types of setae on their opistosoma, whereas others, like Eilica sp., Laronius erawan, Urozelotes rusticus, have no covering setae on their opistosoma at all. The study reveals the existence of different types of covering setae and provides a set of characteristics important for classification and phylogenetic analysis of the spider family Gnaphosidae.
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Arachnologische Mitteilungen 49: 34-46 Karlsruhe, Juni 2015
All ground spiders possess setae that cover their bo-
dies. Ovtsharenko (1983, 1985, 1989) and after that
Murphy (2007) recognized 10 dierent types of se-
tae on the cuticle of ground spiders. Setae have die-
rent morphologies and diverse functions, depending
on their location on the body (Ovtsharenko 1985,
1989). Aculeate setae are the most common type of
setae on the cuticle. ese setae are widely distribut-
ed over the spider’s body. e majority of these setae
are sensory organs or mechanoreceptors (Murphy
2007). Covering setae are located mostly on the ab-
domen, dorsally, ventrally and laterally, and may also
cover the cephalothorax, legs, pedipalps and spinne-
rets. Covering setae have no connection with sensory
receptor cells, they have no sensory function (Town-
send & Felgenhauer 1998a, 1998b, 1999, 2001, Foe-
lix 2011), and are identied by the following cha-
racteristics: (a) the covering setae rest in the shallow
depression or small elevation of the cuticle, (b) the
pedicel is bent at an obtuse angle where it emerges
from the socket, (c) the main axis of the setae is pa-
rallel to the surface of the spider’s body, and (d) the
trunk of the setae has appendages.
e covering setae are of special interest. Lehtinen
was among the rst who noted the importance of
these cuticular structures for spider classication
(Lehtinen 1967, 1975a, 1975b). us, he called the
covering setae ‘feathery hairs’ on the tibia of Micaria
pulicaria (Lehtinen 1975b). Hill (1979) and Town-
send & Felgenhauer (1998a, 1998b, 1999) made a
signicant contribution to our understanding of the
diversity of the shapes of these cuticular structures,
which they called ‘scales’. Studies of ground spiders
(Araneae: Gnaphosidae) demonstrate that they are
greatly varied in shape and are genus-specic (Ovts-
harenko 1983, 1985, 1989, Murphy 2007). e value
of scales for classication and establishing evoluti-
onary relationships were demonstrated for jumping
spiders (Hill 1979) and lynx spiders (Townsend &
Felgenhauer 2001). e wide diversity of scales and
their value for phylogenetic analysis were used for
the reconstruction of the phylogeny of araneomorph
spiders (Griswold et al. 2005, Ramírez 2014). How-
ever, knowledge about the diversity of the covering
setae among gnaphosid spiders is far from complete
and still has to be addressed. As Murphy (2007: 31)
said: “A serious study of spider setae may yet reveal
much of interest”.
The covering setae of ground spiders (Araneae: Gnaphosidae)
Boris Zakharov & Vladimir Ovtsharenko
doi: 10.5431/aramit4904
Abstract. Previous study of the setae covering the opisthosoma of ground spiders shows that the morphology of
the covering setae is genus-specic. The present study pursues the description of covering setae across the whole
Gnaphosidae family using SEM. A detailed morphology of the setae of ground spiders (Araneae, Gnaphosidae) is
present ed. The six major types of covering setae recognized among gnaphosid spiders are squamose, plumose,
lanceolate, pinnate, arborate and sicate setae. Squamose setae are characteristic for Micaria lenzi and Nauhea tapa.
Plumose setae are more common in ground spiders and occur in the genera Drassodes, Haplodrassus, Anagraphis,
Nodocion, Zelotes and the species Berlandina caspica, Nomisia aussereri, Minosiella intermedia, Sosticus loricatus, Lepto-
drassus memorialis, Intruda signata, Parasyrisca caucasica, Scopoides catharius, Echemoides tofo, Zimiromus medius, En-
coptarthria echemophthalma, Apodrassodes trancas, Apopyllus silvestri, Hemicloea sundevalli, Zelanda erebus, Orodras-
sus assimilis, Callilepis nocturna and Synaphosus turanicus. The species Matua valida, Anzacia gemmea, Hypodrassodes
maoricus, Homoeothele micans and Scotophaeus blackwalli have lanceolate setae. Spiders of the genus Gnaphosa
have pinnate setae. Fedotovia uzbekistanica has arborate setae. The species Cesonia bilineata, Herpyllus propinquus,
Litopyllus temporarius, Aphantaulax seminigra and Kishidaia conspicua have sicate setae. Some genera, such as Dras-
sodes and Synaphosus, have a combination of dierent types of setae on their opisthosoma, whereas others, like
Eilica sp., Laronius erawan, Urozelotes rusticus, have no covering setae on their opisthosoma at all. This study reveals
the existence of dierent types of covering setae and provides a set of characteristics important for the classication
and phylogenetic analysis of Gnaphosidae.
Keywords: arborate, lanceolate, pinnate, plumose, sicate, squamose setae
Boris ZAKHA ROV, Department of Natural Sciences, La Guardia
Community College of the City University of New York, New York, USA,
e-mail: bzakharov@lagcc.cuny.edu
Vladimir OVTS HARENKO, Department of Natural Sciences, Hostos
Community College of the City University of New York, New York, USA,
e-mail: vio@hostos.cuny.edu
submitted 12.11.2014, accepted 21.5.2015, online: 10.6.2015
Covering setae of ground spiders 35
In this study, we use the term ‘covering setae’
based on the following reasoning: 1. A scale is thin,
at plate, which is the most common shape for these
structures among jumping and lynx spiders. However,
this type of shape is comparatively rare among gna-
phosids. Ground spiders often have feather-shaped
setae, far from being a at plate. 2. e term feathery
hair was rst used by Lehtinen for these structures
(Lehtinen 1975b, Fig. 7, image 12). 3. In studies of
ground spiders (Araneae: Gnaphosidae) the term ‘se-
Fig. 1: External features of covering setae; squamose (c and d), sicate (a) and plumose (b, e and f) setae on abdomen of gnaphosid
spiders. A. Cesonia bilineata; B. Anagraphis sp.; C. Micaria lenzi; D. Nauhea tapa; E. Echemoides tofo; F. Drassodes lapidosus; A – appen-
dages, Ap – apex, Ped – pedicel, ISp – inferior spines, SSp – superior spines, Sh – shafts, So – socket, Tr – trunk.
36 B. Zakharov & V. Ovtsharenko
tae’ was traditionally used for these structures (Ovts-
harenko 1983, 1985, 1989, Ovtsharenko et al. 1994,
1995, Ovtsharenko & Platnick 1995, Platnick et al.
2001, Murphy 2007). us, in the article on spiders
of the genus Synaphosus the authors wrote that the
abdomen of the spider is “covered by thick, plumo-
se setae bearing 4-7 pairs of appendages originating
from ventral surface of setae (Figs 5, 6)” (Ovtsharen-
ko et al. 1994, p. 3).
Materials and methods
Specimens were examined with a Hitachi S-4700
Field Emission SEM at the American Museum of
Natural History (New York). e cut-o abdomen
was dehydrated in acetone, critically point dried in
carbon dioxide, mounted with double-sided sticky
carbon tape, and sputter coated using the necessary
materials. e resulting images were combined using
Photoshop.
Setae terminology follows Simon (1893), Ber-
land (1919), Lehtinen (1975a, 1975b), Platnick
(1975), Hill (1979), Ovtsharenko (1983, 1985,
1989), Townsend & Felgenhauer (2001) and Mur-
phy (2007). Collections examined: AMNH - Ameri-
can Museum of Natural History, New York, USA;
NMNZ - National Museum of New Zealand Te
Papa Tongarewa, Wellington, New Zealand; OMD -
Otago Museum, Dunedin, New Zealand; OPC - V.
Ovtsharenko personal collection; SAM - South Aus-
tralian Museum, Adelaide, Australia; ZDUC - Zoo-
logical Department of the University of Canterbury,
Christchurch, New Zealand; ZISP - Zoological In-
stitute Saint Petersburg, Russia.
e following species were studied (locality
and collection included): Anzacia gemmea (Dalmas,
1917) [NEW ZEALAND: Kaikoura, January, 1961;
OMD]; Anagraphis sp. [KAZAKSTAN: Atyrau
District, Ustyurt Natural Reserve, Ustyurt Plateau,
village Kemderlie, May 20, 1989, coll. I. I. Ibraev and
A. A. Zyuzin; OPC]; Apodrassodes trancas Platnick
& Shadab, 1983 [ARGENTINA: El Bolsón, Rio
Negro, September 1962, coll. A. Kovacs; AMNH];
Apopyllus silvestri (Simon, 1905) [ARGENTINA:
Epuyén, Chubut, June 12, 1962, coll. Andor Ko-
vacs; AMNH]; Berlandina caspica Ponomarev, 1979
[AZERBAIJAN: village Dubendy, May 21, 1977,
coll. Dunin; OPC]; Cesonia bilineata (Hentz, 1847)
[USA: Arkansas, Logan Co., mountain Magazine,
Mossback Ridge Blu, July 20, 1990, pitfall trap, coll.
B. Leary; AMNH]; Drassodes lapidosus (Walckenaer,
1802) [AZERBAIJAN: Pirgulu State Reserve, 1300
m, May 21, 1984, coll. D. Logunov; OPC]; Echem-
oides tofo Platnick & Shadab, 1979 [CHILE: Co-
quimbo, Llano de la Higuera, September 29, 1980,
coll. L. E. Péna; AMNH]; Encoptarthria echemoph-
thalma (Simon, 1908) [AUSTRALIA: Belair Natu-
ral Park, 300 m S Lower Waterfall, 35001’S, 138043’E,
pitfall, November 28-December 5, 1994, coll. E. G.
Matthews, J. A. Forrest; SAM]; Fedotovia uzbekista-
nica Charitonov, 1946 [KAZAKHSTAN, Muyun-
kum District, 51 km of highway Mirnyi-Khantau,
June 8-9, 1990, coll. A. A. Fedorov; ZISP]; Gnaphosa
muscorum (L. Koch, 1866) [RUSSIA: Chita Region,
Sokhondo Natural Reserve, river Upper Bukukun,
pitfall trap, July 21, 1990, coll. S. Danilov; OPC];
Gnaphosa taurica orell, 1875 [KIRGHIZSTAN,
Kirghiz-Ata gorge, northern slope, June 11, 1985,
coll. A. A. Zuzin, OPC]; Haplodrassus dalmatensis
(L. Koch, 1866) [AZERBAIJAN: Kashkachay vill.,
elev. 1000 m, June 24, 1977, coll. Dunin; OPC];
Haplodrassus soerenseni (Strand, 1900) [RUSSIA:
Altai, Turochak District, Altai Mountains, pine for-
est, pitfall trap, coll. S. B. Ivanov; OPC]; Homoeothele
micans Simon, 1908 [AUSTRALIA: 3.1 km WNW
Mount Lindsay, 27001’09”S, 129051’01”E, Pitjant-
jatjara lands Survey WAT 03, pitfall trap, October
1996; SAM, N 9061-2]; Hypodrassodes maoricus
Dalmas, 1917 [NEW ZEALAND: Wellington,
Karori, inside house, November 8, 1995, coll. C. Pal-
ma; NMNZ]; Intruda signata (Hogg, 1900) [NEW
ZEALAND: Auckland, Beechlands, 36°53’S,
174°46’E, Jan. 1951, coll. J. Campbell; OMD]; Lep-
todrassus memorialis Spassky, 1940 [RUSSIA: Rostov
Region, Zavetninskye district, 4 km S.-E. vil. Fe-
doseevka, clay riverbank, June 15, 1973, coll. Pon-
omarev; OPC]; Leptodrassus sp. [KAZAKHSTAN:
Atyrau, Ustyurt Natural Reserve, Usturt Plateau,
Baskorgan wells, May 28, 1989, coll. A.A. Raikhapov,
S. I. Ibraev; OPC]; Litopyllus temporarius Chamber-
lin, 1922 [USA: Black Rock Forest, Cornwall, NY,
41.42267°N, 74.03039°W, July 5, 2009, coll. V. Ovt-
sharenko, B. Zakharov; OPC]; Matua valida Forster,
1979 [NEW ZEALAND: Arrowtown, December
3, 1969, under stone, coll. R. R. Forster; OMD]; Mi-
caria lenzi Bösenberg, 1899 [RUSSIA: Magadan Re-
gion, Tenkinskye district, Sibit-Tiellakh village, May
15, 1983, coll. S. Buhkalo; OPC]; Minosiella inter-
media Denis, 1958 [TURKMENISTAN: Repetek,
April 6, 1981, coll. V. A. Krivohatsky; OPC]; Nauhea
tapa Forster, 1979 [NEW ZEALAND: Logan Burn,
Covering setae of ground spiders 37
11-23 February, 1983, 900 m, pitfall trap, coll. B. I. P.
Barratt; OMD]; Nodocion eclecticus Chamberlin, 1924
[USA: Tucson, Arizona, coll. O. Bryant; AMNH];
Nodocion mateonus Chamberlin, 1922 [USA: Lake
Co.: Albert Lake, July 2, 1961, coll. B. Malkin;
AMNH]; Nomisia aussereri (L. Koch, 1872) [AZ
ERBAIJAN: Kuba, Tenshalti, September 26, 1984,
coll. unknown; OPC]; Orodrassus assimilis (Banks,
Fig. 2: Plumose setae on the abdomen. A. Berlandina caspica, B. Nomisia aussereri, C. Haplodrassus dalmatensis, D. Sosticus loricatus,
E. Leptodrassus memorialis, F. Synaphosus turanicus
38 B. Zakharov & V. Ovtsharenko
1895) [USA: Oregon, Willow Creek Camp, Warner
Mountains, Lake Co., June 22, 1952, coll. B. Malkin;
AMNH]; Parasyrisca caucasica Ovtsharenko, Plat-
nick & Marusik, 1995 [RUSSIA: Caucasus, Krasnodar
Region, Caucasian Reservation, Mountain Mramor-
naia, 2500 m, July 27, 1975, coll. V. I. Ovtsharenko;
OPC]; Pterotricha sp. [UZBEKISTAN: Bukhara
region, 70 kn W of Utch-Kuduk, 30 km SW of the
village Minbulak, sand desert, coll. D. V. Logunov;
OPC]; Pterotricha strandi Spassky, 1936 [TURK
MENISTAN: Repetek, June 14, 1979, coll. V. A.
Krivokhatsky; OPC]; Scopoides catharius (Chamber-
lin, 1922) [USA: California, 1-2 mi W of Lane Pine,
Inyo Co., April 27, 1960, coll. W. J. Gertsch, Ivie and
Schrammel; AMNH]; Scotophaeus blackwalli (ore-
ll, 1871) [USA: California, Glendale, October 15,
1951, coll. Ted Tice; AMNH]; Sosticus loricatus (L.
Koch, 1866) [UZBEKISTAN: Tashkent district, vil-
lage Toitepa, June 18-30, 1981, coll. N. M. Kudrina;
OPC]; Synaphosus turanicus Ovtsharenko, Levy &
Platnick, 1994 [KAZAKHSTAN, Atyrau, District,
Ustyurt Natural Reserve, Ustyurt Plateau, Baskorgan
wells, May 25, 1989, coll. A. A. Raikhapov, S. Ibraev,
V. Ovtsharenko; OPC]; Zelanda erebus Foster, 1979
[NEW ZEALAND: Gainesville; ZDUC]; Zelotes
lasalanus Chamberlin, 1928 [USA: Arizona, Tucson,
coll. O. Bryant; AMNH]; Zimiromus medius (Key-
serling, 1891) [BRAZIL: S. Paulo, S. Bocaina, 1960
m, S. Jose Barreiro, November 1968, coll. M. Alvar-
enga; AMNH].
Results
Almost all ground spiders (Araneae: Gnaphosidae)
have covering setae. Some species possess a combina-
tion of dierent types of setae on their body. Density
of the setae varies on the spider’s body. Some species
have few setae loosely scattered over the dorsal side
of the abdomen. In other species, setae entirely cover
the opisthosoma, cephalothorax and legs, and create
the shingle-like overlapping coverage that may be
iridescent as in Nauhea tapa (Fig. 1D). e covering
setae rest in a shallow depression of the cuticle that
is slightly elevated above the integument surface or
located on small tubercle of the cuticle. e pedicel
of the seta is bent at an obtuse angle after it emer-
ges from the socket on the cuticle. As a result of this
bending, the main axis of the setae is parallel to the
surface of the spider’s body (Fig. 1A, B).
Setae have a comparatively short pedicel bent
on one side, and an apex on the other side. e
apex may be sharp, attened, or rounded (Fig. 1C,
D). In Gnaphosa the apex bifurcates (Fig. 6E). e
trunk of the setae consists of fused shafts. Often
there are three shafts that create ridges of the trunk
(Fig. 1A, B). Following Hill (1979), short triangular
outgrowths are dened as spines. ere are superior
spines (SSp) that are located on the upper surface
of the trunk, lateral spines (LSp) on both sides, and
inferior spines (ISp) on the side of the trunk that
faces the body surface (Fig. 1A, B). e inferior
spines often have a hook-like shape and, probably,
provide a connection between the seta and the body
surface (Murphy 2007). Setae may also have long
outgrowths called appendages (A) (Fig. 1A, B). De-
pending on the shape of the setae shaft and types
of accessory structures, there are six major types of
covering setae among gnaphosid spiders: squamose,
plumose, lanceolate, pinnate, arborate and sicate.
Squamose setae are at and broad. ey are
comparatively short. ey may bear some accessory
structures and may have a spinous apex. Usually, this
type of seta creates a dense, iridescent coverage of the
spider’s abdomen and cephalothorax (Fig. 1C, D).
Squamose setae are characteristic of small ground
spiders: Micaria and Nauhea. anks to the iridescent
squamose setae these spiders have a metallic blue to
green colour. Micaria demonstrates the diversity of
squamose setae, which take many forms and may or
may not have proximal appendages. e sides and
tips of the setae are serrated. Murphy called this type
“uncinate squamose setae”. ese setae show varia-
tion in size and shape. e side of some setae which
face towards the body bears hook-like inferior spines
(Murphy 2007). e tip of the setae may be serrated
or have a sharp spike (Fig. 1C). Murphy thought that
“uncinate squamose setae” are a characteristic feature
of Micaria (Murphy 2007). In the New Zealand spi-
der Nauhea tapa the pedicel of the seta proximally
bears one or two pairs of appendages. e tip of the
setae is wide, serrated, and with a sharp terminal
spike (Fig. 1D).
Plumose setae are bilateral and look like a feather.
Lehtinen (1967) called this type of seta a “feathery
hair”. Later he came to the conclusion that all types
of feathery setae are adaptations of a setal structure
that has appeared many times independently and
may be regarded as dierent modications of the
plumose seta type (Lehtinen 1975b, see Fig. 7.12).
us, we adopt the term “plumose setae” as the basic
name for all variations of this type of setae. ese
Covering setae of ground spiders 39
setae have a long narrow trunk, sharp at the apex. e
comparatively long appendages are branched from
both sides of the trunk. e appendages may be set
along the whole length of the trunk, or only on a part
of it, usually the proximal third of the trunk length.
e position and length of appendages are also great-
ly varied and may provide signicant features for the
spiders’ classication (Figs 2, 3, 4). e middle part
Fig. 3: Plumose setae on the abdomen (B, D, E, F), cephalothorax (C), and legs (A). A. Intruda signata, B. Zimiromus medius, C. Encopt-
arthria echemophthalma, D. Apodrassodes trancas, E. Apopyllus silvestri, F. Minosiella intermedia
40 B. Zakharov & V. Ovtsharenko
of the trunk is enlarged, whereas its tip is sharpened.
Appendages are set in two rows on the lateral sides of
the trunk. e total number of appendages is genus-
specic and varies from 2 to 44. e appendages may
be long, slender, and sharp at the tip, or enlarged in
their middle part, or clavate, i.e. enlarged at the tip.
As a rule, appendages are more or less of the same
length. However, there are exceptions. In these cases,
the longest appendages are situated at the base of the
trunk, and the shortest ones closer to its tip. e setae
are immovably attached to the body surface on the
cuticular elevation that has ne longitudinal stripes.
e trunk of the setae bends proximally. is position
and their immovable attachment are characteristic
for covering setae only. All other setae are at a right
angle or signicantly far away from the integument.
is suggests that plumose setae are mostly covering
structures, have a protective function for the spiders’
integumentary system, and do not have a sensory
function (Foelix 2011). ey are always present on
the dorsal side of opisthosoma. ey may create a
continuous coverage or may be loosely distributed
over the body surface (Ovtsharenko 1985).
e plumose setae of Berlandina, Nomisia, Mi-
nosiella and Pterotricha have lateral appendages along
almost their entire length. e upper part of the se-
tae that have no appendages may be only 1/4-1/5 of
its length as in Nomisia (Fig. 2B), Minosiella (Fig.
3F) and Pterotricha (Fig. 4D), or even less, as in Ber-
landina (Fig. 2A) (Ovtsharenko 1985). e number
of appendages is also dierent. Minosiella intermedia
has 11-18 appendages, Pterotricha strandi – 23-38. In
Nomisia, it is 10-20, whereas in Berlandina it is 30-
44 appendages or 15-22 pairs. e structure of the
setae may change, depending on the body part. e
abdominal setae of Berlandina, for example, have 30-
32 appendages; setae on the carapace may have up to
44 appendages. Nomisia has setae with 10-12 pairs of
appendages on its abdomen, 10-16 pairs on its cara-
pace, and 10-20 pairs on the legs. In all these genera,
the plumose setae create a dense coverage that com-
pletely covers all of the spider’s body (Ovtsharenko
1985).
Species of the genera Haplodrassus (Fig. 2C) and
Drassodes (Fig. 1F) have at the tip a clear part of their
plumose setae two times longer than its base, which
bears lateral appendages. e number of appendages
varies. Haplodrassus dalmatensis has 8-14 appendages
(Fig. 2C), Haplodrassus soerenseni has 8 appendages.
Haplodrassus signier has from 7 to 10 appendages
(Ovtsharenko 1985, 1989). e number of append-
ages in this species is dierent on both sides of the
setae’s trunk. Setae on one side may have one or even
three fewer appendages than on the other side. Of-
ten, if on one side it is even, then on the other it is
uneven (Ovtsharenko 1985). e plumose setae of
Drassodes vary in number from 8 to 14, but the most
common is 10 (Fig. 1F).
In Sosticus (Fig. 2D) and Leptodrassus (Fig. 2E)
the tip part is equal or only a little longer than
the base with its appendages attached. Sosticus has
8-14 appendages. ese appendages are of dierent
lengths. e longest appendages are in the middle
part of the trunk. Leptodrassus memorialis has 8-11
appendages. e tip of the setae is laterally serrated.
Intruda signata has plumose setae with 9 appendages
at the base of the trunk; 4 appendages on one side
and 5 appendages on the other side (Fig. 3A).
Plumose setae of Anagraphis have 12-15 append-
ages. e appendages are not organized into pairs.
eir number on opposite sides of the trunk is dif-
ferent. Usually, if on one side there are six append-
ages, the other side has eight. Beside that the setae of
Anagraphis spiders have two rows of superior spines
(SSp) and lateral spines (LSp) along the total length
of the trunk (Fig. 1B). Plumose setae on the abdomen
of Nodocion meteonus possess 12 long appendages, oc-
cupying almost half of the trunk, and 5-6 long spines
distributed on the distal part of the setae; all trunk
and appendage setae have ne, longitudinal ridges
(Fig. 4A). In Parasyrisca caucasica on the abdomen
there are plumose setae with 13 appendages; all ap-
pendages have dierent lengths (Fig. 4C). Plumose
setae on the abdomen of Scopoides catharius have 5-15
appendages, they occupy less than half of the proxi-
mal part of the trunk; the distal part of the trunk is
coved by short spines and look like scales (Fig. 4F).
Plumose setae of Echemoides tofo have 9-11 append-
ages. eir number on both sides may be dierent
and vary from 4 to 6. All of these appendages are
located at the proximal 1/3 of the trunk. e SSp are
distributed along the total length of the trunk from
the pedicel to the apex (Fig. 1E). Zimiromus medius
has plumose setae with 10 appendages. Appendages
are long and are attened in a dorso-ventral direc-
tion. e apex of the trunk is sharp. Superior and lat-
eral spines are also present (Fig. 3B). Encoptarthria
echemophthalma has plumose setae with 7-8 pairs of
proximal appendages, the distal part of the trunk is
short, less than 1/3 of the trunk (Fig. 3C). Apodrassodes
Covering setae of ground spiders 41
trancas has plumose setae with 4 pairs (total amount
of appendages 8) of appendages symmetrically set on
the proximal 1/3 of the trunk (Fig. 3D). Apopyllus sil-
vestri has plumose setae with 6-8 appendages on the
proximal 1/3 of the trunk. e distal 2/3 of the trunk is
attened (Fig. 3E). Hemicloea sundevalli has plumose
setae with 4-5 proximal appendages on the base of
the trunk. In Zelanda there are 4 pairs of proximally
situated appendages (Fig. 4E). In Orodrassus assimilis
the plumose setae possess 4 pairs of appendages, the
trunk of the setae is smooth, serrated apically (Fig.
4B). Some genera have a reduced number of covering
Fig. 4: Plumose setae on the abdomen (A, B, C, D, F) and cephalothorax (E). A. Nodocion meteonus, B. Orodrassus assimilis. C. Parasy-
risca caucasica, D. Pterotricha strandi, E. Zelanda erebus, F. Scopoides catharius
42 B. Zakharov & V. Ovtsharenko
setae. ese spiders may have covering setae only on
the abdomen and the legs, most commonly on their
femur. As a rule, the number of appendages on these
setae varies from 0 to 6. In the case that appendages
are absent the trunk remains present on the cuticle.
us, the plumose setae of the Palaearctic species
Callilepis nocturna have 4-6 appendages of dierent
lengths on their base (Ovtsharenko 1985). Its North
American relative Callilepis pluto has 5-8 appendages
(Platnick 1975). Species of two genera close to Cal-
lilepis, Eilica sp. and Laronius erawan, completely lost
the covering setae. Synaphosus (Fig. 2F) has plumose
setae with 6-7 pairs of appendages. e tip of the se-
tae is slightly enlarged. It is two times longer than it
is wide at its base. For species of the genus Zelotes the
number of appendages is characteristically decreased.
us, Zelotes subterraneus has 2-4 appendages on its
setae, Z. longipes has 2-3. e plumose setae in these
spiders are only on the dorsal side of the opisthosoma
(Ovtsharenko 1985). e setae of Drassyllus praecus
do not have appendages at all. e opisthosoma of
Urozelotes rusticus has no covering setae, only sensil-
lae.
Spiders of the genera Matua, Anzacia, Homoe-
othele, Hypodrassodes and Scotophaeus have lanceolate
setae (Fig. 5). e lanceolate setae are, as a rule, at.
An oval ridge ornaments them on both lateral sides.
e trunk is slightly bent. e tips of the setae are
serrated. is type of setae was found on the dor-
sal side of the opisthosoma, cephalothorax and legs;
setae can be modied depending on their location
on the body. e trunk of the setae of Anzacia gem-
mea is comparatively narrow and short, and also bears
4 long appendages in its proximal part, close to the
base of the setae. e remainder of the trunk is broad
and serrated with a sharp spike on the tip (Fig. 5C)
(Ovtsharenko & Platnick 1995). Lanceolate setae on
the opisthosoma of Hypodrassodes maoricus have one
pair of proximal appendages and strongly serrated
distal parts of the setae (Fig. 5D). Lanceolate setae
on the cephalothorax of Homoeothele micans have 4
pairs of appendages in the proximal part that reach
the middle of the seta; the distal part of the seta is at
and laterally serrated (Fig. 5E). e lanceolate setae
on the abdomen of Scotophaeus blackwalli are narrow,
smooth, with 3 pairs of long proximal appendages
and with 4-5 slender spines at the apical part of the
trunk (Fig. 6D). Modication of lanceolate setae oc-
curs in Matua valida, thus setae on the abdomen are
at, wide, proximally with 3 pairs of appendages and
distally on the tip with 3-4 spines on each side of
the seta (Fig. 5A). e lanceolate setae on the legs
of Matua valida are narrow, at, with 7-8 long ap-
pendages in the proximal part of the seta and slightly
serrated distal part of seta (Fig. 5B). Lanceolate setae
occur sometimes together with plumose setae, mostly
on the abdomen, for example in species of Drassodes
and Synaphosus.
Sicate setae have a broad curved trunk and the
appendages are all on one pro-curved side of the
trunk, which dierentiates them from the lanceolate
setae (Fig. 6A-D). ese setae were found mostly on
the dorsal side of the abdomen and carapace (Ovt-
sharenko 1985). e species of the genus Herpyllus
have sicate setae with 10 appendages of equal length
set in the middle part of the setae. On its tip there
are 1-2 short spines. In Herpyllus propinquus sicate
setae bear only two rows of short (almost spine-like)
6-8 appendages on the dorsal surface of the distal
part of the trunk (Fig. 6B). Beside sicate setae, spi-
ders of this species have plumose setae on the dorsal
side of their abdomen. In Aphantaulax seminigra and
Kishidaia conspicua the sicate setae have up to 25-30
appendages, distributed over the whole length of the
trunk (Ovtsharenko 1989). In Cesonia bilineata sicate
setae have 4-36 appendages located on the rim of the
trunk along the whole of its length. e tip of these
setae bears 8-9 spines dorsally (Fig. 1A, 6A). In Lito-
pyllus temporarius 15 short appendages are distribut-
ed along the total length of the trunk in two closely
spaced rows, and 4 pairs of long appendages are lo-
cated on the proximal part of the trunk (Fig. 6C).
Pinnate setae are characteristic of the spider ge-
nus Gnaphosa. ey are bilateral and folded along
the grooved longitudinal midline. e most pecu-
liar characteristic of this type of setae is a marginal
serration, created by short appendages, broad at the
base, along the whole length of the trunk. e apex
of the setae of Gnaphosa taurica bifurcates and has
the shape of a snake’s tongue (Fig. 6E). e setae of
Gnaphosa lugubris like G. taurica have a longitudinal
groove along the whole length of the trunk with lat-
eral boarders bent toward the outside. ese board-
ers have two rows of spines, broad at the base and
sharp apically. e pedicel of the setae is narrow and
is merged into the depression on the cuticle. us,
the major characteristics of the setae of these spiders
are the four rows of appendages, the grooved shape
of the trunk, and the way it merges into the cuticle
pedicel. Spiders of this genus have setae only on the
Covering setae of ground spiders 43
dorsal side of the abdomen, where they create a dense
coverage. Comparison of the nymphs of Gnaphosa
lugubris with adult spiders shows that their setae are
almost identical (Ovtsharenko 1985).
Arborate setae occur in the species Fedotovia uz-
bekistanica and this type of setae is the most unusual
covering seta among the ground spiders. ey look
like branches on the trunk of a tree. e trunk of
Fig. 5: Lanceolate setae on the abdomen (A, C, D), cephalothorax (E) and legs (B); sicate setae on the abdomen (F). A, B. Matua valida,
C. Anzacia gemmea, D. Hypodrassodes maoricus, E. Homoeothele micans, F. Cesonia bilineata
44 B. Zakharov & V. Ovtsharenko
these setae has a shaft with four longitudinal ridges.
e appendages are branched from these ridges in
four directions almost along the whole length of the
trunk (Fig. 6F).
Discussion and conclusions
More than a third of the total known genera of gna-
phosid spiders have been studied. e obtained data
allow us to make some generalizations. Almost all
gnaphosid spiders have covering setae on the dor-
sal side of their opisthosoma and additionally on the
cephalothorax, legs and spinnerets. Some groups of
ground spiders have very little (subfamily Zelotinae)
or have no covering setae on their abdomen at all
(subfamily Laroniinae). Setae demonstrate diverse
appearance depending on their location on the body.
Our data show the existence of stable characteristics
relating to setal morphology. Depending on the sha-
pe of the setae and their accessory structures, they
may be classied into six groups: squamose, plumose,
lanceolate, sicate, pinnate and arborate.
Lehtinen (1967, 1975a, 1975b) was the rst
who pointed out the value of setal morphology for
taxonomy and reconstruction of phylogenetic rela-
tionships among spiders. Galiano (1975) and Hill
(1979) considered the diagnostic value of salticid
scales. Ovtsharenko (1983, 1985, 1989) came to the
conclusion that morphology of gnaphosids’ covering
setae varies among the genera and provides additio-
nal characteristics for taxonomic analysis and recons-
truction of phylogenetic relationships among ground
spiders. Townsend & Felgenhauer (1998a, 1998b,
1999) studied these cuticular structures and showed
their usefulness for the taxonomy of oxyopid spiders.
Townsend & Felgenhauer (2001), Griswold et al.
(2005) and Ramírez (2014) show the importance of
these characters in the phylogenetic analysis of Oxy-
opidae, entelegyne, and dionychan spiders. Our study
supports previous observations made by Ovtsharen-
ko (1983, 1985, 1989) that setae in the family Gna-
phosidae demonstrate specic subfamilial and gene-
ric characteristics and provide valuable information
for taxonomy and phylogeny of these spiders.
e role of covering setae still remains unknown.
However, the type of seta attachment, the shape of
the setae, and the absence of a proven association of
the setae with sensory neurons (Townsend & Fel-
genhauer 1998a, 1998b, 1999, 2001, Foelix 2011)
allow us to suggest a protective function. Hill (1979)
noted that there are no veried experimental data
that may suggest a specic function of the spiders’
scales. At the same time, the scales’ shape and design
create a reective surface that refracts light and is re-
sponsible for the colourful body pattern of jumping
spiders that may be displayed during courtship (Hill
1979). Some gnaphosid spiders, such as Micaria and
Nauhea, also have a metallic coloured body created by
dense coverage of squamose setae.
Acknowledgements
e present study was supported in part by the Research
Foundation of the City University of New York providing
funds through the grant PSC-CUNY TRADA-43-582
and the American Museum of Natural History, New York.
e authors are grateful to the students Amanda Tsang
and Kseniia Rusinova. We are grateful to Henry Towbin
and Morgan Hill from the American Museum of Natural
History Microscopy and Imaging Facility for technical
support with the SEM, as well as Dr. Norman I. Platnick,
Curator of AMNH, for support of the study. We are very
grateful to Martin Ramírez from the Museo Argentino
de Ciencias Naturales, who generously provided us with
his beautiful photo of the setae of Gnaphosa taurica. e
authors are very grateful to Dr. Sandra Dickinson from the
LaGuardia Community College of the City University of
New York for her valuable help on the text preparation and
Don N. Cameron for his help with the proper naming of
setae. e authors are indebted to Dr. Martin Ramírez, Hay
Wijnhoven and eo Blick who reviewed this article and
gave generously their time and expertise.
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A phylogenetic analysis of the two-clawed spiders grouped in Dionycha is presented, with 166 representative species of 49 araneomorph families, scored for 393 characters documented through standardized imaging protocols. The study includes 44 outgroup representatives of the main clades of Araneomorphae, and a revision of the main morphological character systems. Novel terminology is proposed for stereotyped structures on the chelicerae, and the main types of setae and silk spigots are reviewed, summarizing their characteristics. Clear homologs of posterior book lungs are described for early instars of Filistatidae, and a novel type of respiratory structure, the epigastric median tracheae, is described for some terminals probably related with Anyphaenidae or Eutichuridae. A new type of crypsis mechanism is described for a clade of thomisids, which in addition to retaining soil particles, grow fungi on their cuticle. Generalized patterns of cheliceral setae and macrosetae are proposed as synapomorphies of the Divided Cribellum and RTA clades. Dionycha is here proposed as a member of the Oval Calamistrum clade among the lycosoid lineages, and Liocranoides, with three claws and claw tufts, is obtained as a plausible sister group of the dionychan lineage. The morphology of the claw tuft and scopula is examined in detail and scored for 14 characters highly informative for relationships. A kind of seta intermediate between tenent and plumose setae (the pseudotenent type) is found in several spider families, more often reconstructed as a derivation from true tenent setae rather than as a phylogenetic intermediate. Corinnidae is retrieved in a restricted sense, including only the subfamilies Corinninae and Castianeirinae, while the ‘‘corinnid’’ genera retaining the median apophysis in the copulatory bulb are not clearly affiliated to any of the established families. Miturgidae is redefined, including Zoridae as a junior synonym. The Eutichuridae is raised to family status, as well as the Trachelidae and Phrurolithidae. New synapomorphies are provided for Sparassidae, Philodromidae, and Trachelidae. Philodromidae is presented as a plausible sister group of Salticidae, and these sister to Thomisidae; an alternative resolution placing thomisids in Lycosoidea is also examined. The Oblique Median Tapetum (OMT) clade is proposed for a large group of families including gnaphosoids, trachelids, liocranids, and phrurolithids, all having the posterior median eye tapeta forming a 90u angle, used for navigation by means of the polarized light in the sky as an optical compass; prodidomines seem to have further enhanced the mechanism by incorporating the posterior lateral eyes to the system. The Teutamus group is recognized for members of the OMT clade that are usually included in Liocranidae, but not closely related to Liocranum or phrurolithids. The Claw Tuft Clasper (CTC) clade is proposed for a group of families within the OMT clade, all having a peculiar mechanism grasping the folded base of the claw tuft setae with a hook on the superior claws. The CTC clade includes Trachelidae, Phrurolithidae, and several gnaphosoids such as Ammoxenidae, Cithaeronidae, Gnaphosidae, and Prodidomidae. A remarkable syndrome involving the expansion of the anterior lateral spinnerets, often sexually dimorphic, is here reported for some Miturgidae and several members of the CTC clade, in addition to the known cases in Clubionidae and ‘‘Liocranidae.’’ The following genera are transferred from Miturgidae to Eutichuridae: Calamoneta, Calamopus, Cheiracanthium, Cheiramiona, Ericaella, Eutichurus, Macerio, Radulphius, Strotarchus, Summacanthium, and Tecution; Lessertina is transferred from Corinnidae to Eutichuridae. The following genera are transferred to Miturgidae: Argoctenus, Elassoctenus, Hestimodema, Hoedillus, Israzorides, Odomasta, Simonus, Thasyraea, Tuxoctenus, Voraptus, Xenoctenus, Zora, and Zoroides, from Zoridae; Odo and Paravulsor, from Ctenidae; Pseudoceto from Corinnidae. The following genera are transferred from Corinnidae to Trachelidae: Afroceto, Cetonana, Fuchiba, Fuchibotulus, Meriola, Metatrachelas, Paccius, Paratrachelas, Patelloceto, Planochelas, Poachelas, Spinotrachelas, Thysanina, Trachelas, Trachelopachys, and Utivarachna. The following genera are transferred from Corinnidae to Phrurolithidae: Abdosetae, Drassinella, Liophrurillus, Plynnon, Orthobula, Otacilia, Phonotimpus, Phrurolinillus, Phrurolithus, Phruronellus, Phrurotimpus, Piabuna, and Scotinella. Dorymetaecus is transferred from Clubionidae to Phrurolithidae. Oedignatha and Koppe are transferred from Corinnidae to Liocranidae. Ciniflella is transferred from Amaurobiidae to Tengellidae.
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