Resu ec ion o Synema globosum cana iense Dahl,
1907 (A aneae: Thomisidae) wi h commen s on o he
synonyms wi hin his genus
Au ho s: Lissne , Jø gen, and Suá ez, Daniel
Sou ce: A achnologische Mi eilungen: A achnology Le e s, 65(1) : 18-
26
Published By: A achnologische Gesellscha e.V.
URL: h ps://doi.o g/10.30963/a ami 6505
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A achnologische Mi eilungen / A achnology Le e s 65: 18-26 Ka ls uhe, Augus 2023
Synema globosum (Fab icius, 1775) is a c ab spide ha am-
bushes insec s on lea es and on lowe s o he bs and sh ubs.
Being a p eda o o lo i o es and o he he bi o es i may
lowe he bi o e damage on some plan species, as shown o
E ophaca bae ica (L.), Boiss. (Can e bu y 2019). C ab spide s
using lowe s as cap u e si es may also al e he s uc u e o
communi ies o lowe - isi ing insec s, and he eby in luence
plan pollina ion (Ma e o e al. 2015). Taxonomy is c ucial
o s udying such ecological in e ac ions because i clea ly
iden i ies he species in ol ed. As in he case o many o he
gene a, ou iew o Synema has e ol ed o e ime wi h species
being added, synonymized, o ans e ed. Se e al subspecies
ha e been e ec ed, bu nea ly all we e la e synonymized wi h
he nomino ypical subspecies (WSC 2023). The genus cu -
en ly holds 124 accep ed species including S. globosum, which
also is he ype species (WSC 2023).
Acco ding o he cu en unde s anding Synema globosum
is widely dis ibu ed in he Palaea c ic ealm, anging om
he Cana y Islands o Eas e n Asia (Nen wig e al. 2023, Ono
1988, WSC 2023, Wunde lich 1987). In Eu ope i is ound
in Sou he n Eu ope, Wes e n Eu ope, and he sou he n hal
o Eas e n Eu ope (Nen wig e al. 2023). I is pa icula ly
common in he Medi e anean egion, which also includes
Tunisia, Alge ia, and Mo occo (Nen wig e al. 2023). I is no
known whe he he species has become es ablished in No h-
e n Eu ope. In ecen yea s i has been eco ded se e al imes
in England, bu he e is as ye no e idence o es ablished pop-
ula ions (Bee e al. 2020).
A well-known ea u e o S. globosum is he a ia ion in
colou among specimens, being much mo e p onounced in
emales han in males (Fig. 1). The opis hosoma ( olium ex-
cep ed) can be colou ed in shades o ei he whi e, yellow, o -
ange, o ed. Some specimens a e bicolou ed, bu wi h one
colou domina ing (Fig. 1: F4, F5, M1). The do sum o he
opis hosoma has a b oad, oliaceous, black pa e n ( a ely
b own), and he an e io pa esembles he ou line o a bi-
co ne ha . Hence, he e nacula name o his species is ‘Na-
poleon Spide ’. The head is b own o da k b own, and he
on 1s and 2nd pai s o legs a e da k b own o blackish wi h
pale annula ions dis ally. The 3 d and 4 h pai s o legs a e bo h
sho e and pale han he on legs. The species exhibi s sex-
ual dimo phism, males a e smalle and da ke and wi h less
colou a ia ion compa ed o emales.
Acco ding o he Cana y Islands Go e nmen (2022),
S.globosum is he only Synema species lis ed om he islands,
wi h eco ds om G an Cana ia, Tene i e, La Gome a and
La Palma. In 2011, Synema specimens we e bea en om
bushes in he pine o es bel on he sou he n slopes o Ten-
e i e. Due o hei gene al appea ance i seemed unlikely
ha hese specimens belonged o S. globosum, as hei col-
ou ma kings di e ed signi ican ly om hose o specimens
om he Eu opean mainland. La e , mo e specimens om
he islands o La Gome a and G an Cana ia we e collec ed,
again none o which looked like Eu opean S. globosum. In
ac , he habi us o he Cana ian specimens is a much be -
e ma ch wi h ha o S. imi a ix (Pa esi, 1883) om he
A ican con inen when compa ed o images o he la e
species in Dippenaa -Schoeman e al. (2020). Wi h hese
obse a ions in mind, i was hypo hesized ha he syn-
onymiza ion o S. g. cana iense wi h S. globosum is in alid.
A mo phological compa ison o he wo subspecies was
pe o med o es his hypo hesis. Also, ba code sequenc-
es o Cana ian specimens we e compa ed o da a a ail-
able a BOLD and GenBank o p o ide addi ional con ex .
Abb e ia ions
COI = cy och ome c oxidase I (gene)
CJL = Collec ion Jø gen Lissne (e en ually deposi ed a
Na u al His o y Museum o Denma k)
Resu ec ion o Synema globosum cana ienseDahl, 1907 (A aneae: Thomisidae)
wi h commen s on o he synonyms wi hin his genus
Jø gen Lissne & Daniel Suá ez
doi: 10.30963/a ami 6505
Abs ac . The homisid spide Synema globosum (Fab icius, 1775) is widely dis ibu ed in he Palaea c ic ealm and is pa icula ly com-
mon in sou he n pa s o zones wi h a empe a e clima e. Se en subspecies ha e been desc ibed; he widely dis ibu ed nomino ypical
subspecies, S. g. globosum and six wi h much mo e es ic ed anges. As o now, jus wo subspecies a e s ill ecognised as alid while
he o he s ha e been synonymized wi h he nomina e subspecies. This s udy now e alida es S. g. cana iense Dahl, 1907 (subsp. e al.)
based on indings om molecula and compa a i e mo phological analysis. We also a gue ha he synonymiza ion o he subspecies S.
g. daghes anicum U ochkin, 1960 and he species S. japonicum Ka sch, 1879 wi h S. globosum in Ono (1988) is o mally co ec , bu needs
u he in es iga ion wi h mode n me hods.
Keywo ds: Cana y Islands, c ab spide , napoleon spide , Spain, subspecies, axonomy
Zusammen assung. Re alidie ung on Synema globosum cana ienseDahl, 1907 (A aneae: Thomisidae) mi Kommen a en zu an-
de en Synonymisie ungen inne halb de Ga ung. Die K abbenspinne Synema globosum (Fab icius, 1775) is wei e b ei e inne halb
de Paläa k is und is besonde s häu ig in südlichen Zonen mi gemäßig em Klima. Sieben Un e a en wu den besch ieben; die wei e -
b ei e e nomino ypische Un e a S. g. globosum und sechs wei e e mi s ä ke eingesch änk en Ve b ei ungsgebie en. Heu zu age sind
nu noch zwei diese Un e a en ane kann wäh end die ande en mi de nomino ypischen Un e a synonymisie wu den. Diese S udie
e alidie nun S. g. cana iense Dahl, 1907 (subsp. e al.) basie end au eine gene ischen sowie e gleichend-mo phologischen Analyse.
Des Wei e en wi d a gumen ie , dass die Synonymisie ung de Un e a en S. g. daghes anicum U ochkin, 1960 und de A S. japonicum
Ka sch, 1879 mi S. globosum du ch Ono (1988) o mal ko ek is , abe wei e e Un e suchungen mi mode nen Me hoden benö ig .
Jø gen Lissne , Na u al His o y Museum Aa hus, Wilhelm Meye s Allé 10, Aa hus,
Denma k; e-mail: [email p o ec ed]
Daniel Suá ez, Island Ecology and E olu ion Resea ch G oup, Ins i u o de P oduc os
Na u ales y Ag obiología (IPNA-CSIC), A enida As o ísico F ancisco Sánchez, 3. La
Laguna, Spain; E-mail: [email p o ec ed]om
Academic edi o : Tobias Baue
submi ed 15.2.2023, accep ed 2.6.2023, online 4.8.2023
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Resu ec ion o Synema globosum cana ienseDahl, 1907 19
Fig. 1: Synema globosum globosum (Fab icius, 1775), habi us o li e specimens. F = Female, M = Male. F1. Nages e Solo gues, F ance; F2. Osani, Co -
sica, F ance; F3. Cala Mesquida, Majo ca, Spain; F4. Campane , Majo ca, Spain; F5. Meuilley, Dijon, F ance; F6. Pó oa Dão, Po ugal; M1. Sain -Rémy-de-
P o ence, F ance; M2. Viei a do Minho, Po ugal. © Jø gen Lissne , excep F1 © Jean-Philippe Tabe le . No e a e ac s due o lash e lec ions om ing lash
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20 J. Lissne & D. Suá ez
Ma e ial and me hods
Mo phological compa ison o habi us was made using colou
pho og aphs o li e specimens (Figs 1-2). The s e na o p e-
se ed specimens we e pho og aphed using a Leica M165C
s e eomic oscope i ed wi h a Leica MC 190 HD digi al
came a. The s e eomic oscope was connec ed o a compu e
wi h Leica Applica ion Sui e V. 4.12 and Ze ene S acke
so wa e V. 1.04. Specimens used o DNA ba coding a e
deposi ed in he in e eb a e collec ion o he Ins i u o de
P oduc os Na u ales y Ag obiología (IPNA-CSIC). DNA
was ex ac ed om a selec ion o se en specimens ollowing a
Chelex p o ocol (Casque e al. 2012). The 5’ egion (658 bp)
o he m DNA COI gene was ampli ied using LCO1490 and
HCO2198 p ime s (Folme e al. 1994). PCR eac ion condi-
ions we e as ollows: ini ial dena u a ion a 95°C o 2min,
ollowed by 40 cycles o 94°C o 30 s, 42°C o 35 s, and
72°C o 45 s, and a inal ex ension o 72°C o 5 min. 2μL
o dilu ed (1/10) DNA ex ac was ampli ied wi h 23 μL o
PCR mix ( o a o al olume o 25 μL), comp ised o 14.4μL
o wa e , 2.5 μL o 10x NH4 bu e (Bioline), 1.5 μL o
50mM MgCl2 (Bioline), 2 μL o 2.5 mM dNTPs (Bioline),
0.5 μL o BSA (20 mg/ml), 1 μL o each p ime (10 μM),
and 0.1μL o Taq polyme ase (BIOTAQ). PCR p oduc s
we e sequenced using he Sange DNA sequencing se ice o
Mac ogen (www.mac ogen.com). Sequences we e hen edi ed
in Geneious 2021.1.1 (www.geneious.com) and compa ed o
he “nucleo ide” da abase o GenBank (www.ncbi.nlm.nih.
go /genbank/) using BLAST (Al schul e al. 1990). A da ase
was assembled including ou sequenced ba codes, all a ail-
able ba code sequences wi hin GenBank o S. globosum and
a selec ion o close ela ed species (ou g oups). Unco ec ed
p-dis ances among sequences we e ob ained om Geneious
2021.1.1 (www.geneious.com). A maximum-likelihood ee
was cons uc ed wi h RAxML 8.2. on he CIPRES Science
Ga eway (Mille e al. 2010) using a GTR+G model o e o-
lu ion. The bes sco ing ML ee was selec ed among 100
sea ches on he o iginal alignmen wi h di e en andomised
pa simony s a ing ees. Suppo alues we e ob ained wi h
1000 boo s ap eplica es (Felsens ein 1985).
Ma e ial examined: Synema globosum globosum (Fab icius
1775). AUSTRIA: Bisambe g (48.3239°N, 16.3622°E)
300m, 09.V.2005, 1), Jan Pede sen leg., CJL-7158; Pe onell-
Ca nun um (48.1180°N, 16.8567°E) 150 m, 08.V.2008, 1),
Jan Pede sen leg., CJL-7226. FRANCE: Co sica, Po o
(42.2656°N, 8.7059°E) 0-250 m, 11.VII.2013, 1)1(, Jø -
gen Lissne leg., CJL-9079; Co sica, Osani (42.3214°N,
8.6365°E) 120-180 m, ock s eppe wi h bushes, 12.VII.2013,
1(, Jø gen Lissne leg., CJL-9008; Sain Remy-de-P o ence
(43.7891°N, 4.8317°E) 60 m, pa k, 05.IV.2007, 1), Jø gen
Lissne leg., CJL-4087; A cenan (47.1478°N, 4.8278°E)
465 m, ock s eppe wi h bushes, 21.VI.2004, 1(, Jø gen
Lissne leg., CJL-1693; Meuilley (47.1373°N, 4.8884°E)
290 m, all pas u e, 24.VI.2004, 2))2((, Jø gen Lissne leg.,
CJL-1926. GREECE: Laconia, Pa non O os (37.2207°N,
22.6287°E) 1700 m, 08.VII.1982, 1), Bja ne Skule leg., CJL-
2604; Olympia (37.6380°N, 21.6300°E) 70 m, 24.VI.1964,
1(, Johan Thomas Sko gaa d leg., CJL-2649. ITALY: Finale
Ligu e (44.1739°N, 8.3420°E) 320 m, 06.VII.2015, 1(, Jan
Dolansky leg., CJL-10742. SPAIN: Majo ca, Puig Randa
(39.5243°N, 2.9182°E) 380 m, oak o es s, 08.IV.2016, 1),
Jø gen Lissne leg., CJL-11459; Majo ca, Be lem, Cala
Camps (39.7493°N, 3.3015°E) 5 m, ock s eppe wi h bush-
es, 30.IV.2019, 1), Jø gen Lissne leg., CJL-3245; Majo ca,
San ua i de Lluc (39.8231°N, 2.8834°E) 480 m, mixed o -
es , 21.IV.2013, 1), Jø gen Lissne leg., CJL-8824; Majo ca,
Cala Romàn ica (39.5195°N, 3.3069°E) 16 m, acan lo
wi h bushes, 23.IV.2019, 1), Jø gen Lissne leg., CJL-2852;
Majo ca, Es Cas ell (39.7882°N, 2.9269°E) 170 m, mixed
o es , 20.IV.2018, 1), Jø gen Lissne leg., CJL-12331; Ma-
jo ca, Cap de ses Salines (39.2650°N, 3.0561°E) 44318 m,
ock s eppe wi h bushes, 27.IV.2018, 2)), Jø gen Lissne
leg., CJL-12413; Majo ca, La T apa (39.5999°N, 2.3609°E)
270 m, ock s eppe wi h bushes, 24.IV.2014, 1(, Jø gen Liss-
ne leg., CJL-9694; Majo ca, Dunes es T énc (39.3491°N,
2.9865°E) 2 m, sand dune wi h bushes, 25.IV.2014, 1(,
Jø gen Lissne leg., CJL-9719; Majo ca, Dunes es T é-
nc (39.3491°N, 2.9865°E) 2 m, sand dune wi h bushes,
27.IV.2018, 1(, Jø gen Lissne leg., CJL-12423; Majo ca,
Cala Mesquida (39.7398°N, 3.4304°E) 22 m, pine o es ,
30.IV.2019, 1(, Jø gen Lissne leg., CJL-2987; Majo ca,
Cala Mesquida (39.7423°N, 3.4328°E) 25 m, sand dune wi h
bushes, 05.IV.2016, 1(, Jø gen Lissne leg., CJL-11370; Ma-
jo ca, Es Cas ell (39.7882°N, 2.9269°E) 170 m, mixed o -
es , 27.IV.2019, 1(, Jø gen Lissne leg., CJL-3357; Majo ca,
To en de San Miquel (39.7934°N, 2.9652°E) 48 m, s eam
bank wi h bushes, 03.V.2019, 1)1(, Jø gen Lissne leg., CJL-
3077; Majo ca, Pò ol (39.6303°N, 2.7646°E) 215 m, mixed
o es , 25.IV.2018, 1)1(, Jø gen Lissne leg., CJL-12389;
Majo ca, Cala Mesquida (39.7423°N, 3.4328°E) 25 m, sand
dune wi h bushes, 04.V.2019, 3((, Jø gen Lissne leg., CJL-
2945. PORTUGAL: Pó oa Dão (40.5488°N, 7.9436°W) 225
m, 31.V.2006, 1)1(, Jo ge Alexand e Mo a de Almeida leg.,
CJL-3594; Rua da Fon e da Po ca (41.6447°N, 8.2073°W)
330 m, sh ubs, 02.IIX.2017, 1), Jø gen Lissne leg., CJL-3475.
Synema globosum cana iense Dahl, 1907. SPAIN: Tene i e,
Mon e del Agua (28.3303°N, 16.8385°W) 800 m, lau el
o es , 18.IV.2011, ju enile, Jø gen Lissne leg., CJL-7635;
La Gome a, Las Poya as (28.1488°N, 17.1930°W) 295 m,
xe ophylic zone, 03.IV.2017, 1), Jø gen Lissne leg., CJL-
11865; G an Cana ia, Calde a de Bandama (28.0332°N,
15.4557°W) 220 m, xe ophylic zone, 06.IV.2017, ju enile,
Daniel Suá ez leg., CJL-12596; G an Cana ia, El B ezal
(28.1072°N, 15.6016°W) 470 m, wax my le - ee hea h,
07.IV.2017, ju enile, Daniel Suá ez leg., CJL-12595; Ten-
e i e, Las Me cedes (28.5224°N, 16.2941°W) 670 m, xe o-
phylic zone, 07.IV.2017, 1(, Jø gen Lissne leg., CJL-11922;
G an Cana ia, Te o (28.0353°N, 15.5523°W) 800 m, ga -
den, 09.XI.2018, 1(, Jø gen Lissne leg., CJL-1389; G an
Cana ia, Ten eniguada (27.9758°N, 15.5289°W) 800 m, u-
al landscape, 14.XI.2016, 1(, Daniel Suá ez leg., CJL-12594;
Tene i e, Taucho (28.1535°N, 16.7198°W) 970-1060 m, pine
o es , 13.IV.2011, 1)1(, Jø gen Lissne leg., CJL-7549; La
Gome a, Los Tela es (28.1508°N, 17.1978°W) 240 m, xe o-
phylic zone, 22.IV.2015, 1)1(, Jø gen Lissne leg., CJL-10592.
Resul s
Ba coding
A agmen o he 5’-COI was success ully ampli ied and
sequenced (GenBank accession numbe s OM807203-
OM807209) o se en specimens om he Cana y Islands.
By conduc ing a maximum-likelihood analysis including
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Resu ec ion o Synema globosum cana ienseDahl, 1907 21
hose se en sequences as well as sequences ob ained in U e
e al. (2021) and sequences om BOLD and GenBank, ou
main clades we e eco e ed (Fig. 4). All se en sequences om
he Cana ian specimens we e iden ical, o ming a monophyl-
e ic sis e clade wi h indi iduals om mainland Spain, Po u-
gal, and I aly (‘g een clade’ in U e e al. 2021). Unco ec ed
p-dis ances be ween he Cana ian clade and he ‘g een clade’
ange be ween 0.9-1.6% while dis ances o he emaining
clades anges be ween 2.2-3.2% (‘pu ple clade’ in U e e al.
2021) and 6.1-6.9% (‘blue clade’ in U e e al. 2021).
Mo phological compa ison
Synema globosum cana ienseDahl, 1907 was e ec ed due o
colou di e ences be ween Cana ian specimens and hose
o Wes e n Eu ope (Dahl 1907). Dahl deno ed he con i-
nen al popula ion wi h a inomen, Synema globosum globo-
sum, o poin ou he nomino ypical subspecies. Acco ding
o he key in Dahl (1907), S. g. cana iense di e s om S. g.
globosum by ha ing a di e en colou ed p osoma: “Au dem
Cephalo ho ax is de hin e e Teil des Kop es und auch de
Sei en and imme helle als die dazwischen liegenden Teile”
[au ho ansla ion: he back pa o he head and also he
side ma gins a e always ligh e han he pa s in be ween].
Dahl (1907) desc ibed he colou a ion o he cephalo ho ax
in S. g. globosum as: “Au dem Cephalo ho ax is höchs ens de
hin e e Teil des Kop es hell” [au ho ansla ion: on he ceph-
alo ho ax, a mos he back o he head is pale]. La e , Syn-
ema g. cana iensewas synonymized wi h S. globosum by Wun-
de lich (1987:252). Wunde lich (1987) did no p o ide any
a gumen s o his synonymiza ion, bu i was ne e heless
Fig. 2: Synema globosum cana ienseDahl, 1907, habi us o li e specimens. GC = G an Cana ia, T = Tene i e; LG = La Gome a. © Jø gen Lissne . No e b igh
gla e a e ac s due o lash e lec ions, pa icula ly on he glossy cephalo ho axes
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22 J. Lissne & D. Suá ez
accep ed by he WSC (2023). In he p esen s udy we com-
pa ed se e al mo phological cha ac e s o assess he alidi y
o he synonymiza ion (Tab. 1). Mos o he mo phological
di e ences lis ed we e ound o be dis inc and no clinal wi h
only a ew cha ac e s sligh ly o e lapping. The di e ences
ound apply o all indi iduals examined wi hin each subspe-
cies ange. The esul s ag ee wi h Dahl (1907), bu wi h some
de ia ions. The colou di e ences o p osomas a e ep oduced
in his s udy o only emales and ju enile males. In hese
specimens, he da k ma king o he cephalo ho ax o ms a “U”
in emales and ju enile males o S. g. cana iense (Fig. 2). How-
e e , in examined adul males only he back pa o he head is
ligh in Synema g. cana iense (Fig. 2) while in S.g. globosum i
is uni o mly colou ed, b own o blackish b own (Fig. 1), no
in ag eemen wi h Dahl (1907), a leas no in li e specimens.
The da k/ligh a eas become much less con as ing in alcohol
p ese ed specimens. In emales o S. g. globosum he e is a
sligh ly pale a ea a he back o he head which o ms a wide
“ ” poin ing owa ds he udimen a y o ea and hey lack he
da k “U”-ma king (Fig. 1). Colou a ion o he s e na is shown
in Fig. 3. Di e ences a e mino , ne e heless dis inc among
he wo subspecies (Tab. 1). Opis hosomal ma kings we e no
men ioned by Dahl (1907), bu hey also show dis inc i e di -
e ences (Tab. 1.).
Discussion
Ba coding
U e e al. (2021) s udied molecula and mo phological a i-
a ion in 72 adul specimens o S. globosum collec ed ac oss he
Eu opean con inen and Tu key. The s udy did no include
ma e ial om he dis ibu ional anges o he subspecies, S.g.
cana iense and S. g. nig i en e o om anges o he possi-
ble subspecies, S. g. daghes anicum and S. g. japonicum. Thus,
only specimens om he ange o he nomino ypical subspe-
cies we e in es iga ed. They ound a he la ge (up o ≈ 6 %)
gene ic dis ance in he COI ba code, bu wi hou any bio-
geog aphical pa e n. Soma ic and geni al cha ac e s in bo h
sexes and mo phome ics o he male palp exhibi ed high in-
aspeci ic a ia ion ha did no ma ch wi h he gene ic da a.
Acco ding o U e e al. (2021), one possible explana ion is
ha S. g. globosum could be highly polymo phic bo h in e ms
o i s m DNA and mo phology. Di e ences in bo h mo pho-
logical ea u es compa ed o mainland specimens as well as
he monophyle ic posi ion o he Cana ian specimens a e su -
icien a gumen s o esu ec he subspecies S. g. cana iense.
Di e en s udies dealing wi h spide species delimi a ion had
eco e ed a wide a ie y o di e gence h eshold. Robison e
al. (2009) es ablished a di e gence se a 6.8%, based on he
mean di e gence o di e en spide amilies. This s udy in-
cludes wo Thomisidae gene a, Misumenops and Xys icus wi h
alues o di e gence o 4.9% and 6.85%, espec i ely. A s udy
add essing he phylogene ic ela ionship o he ibe Co ia -
achnini (B ei ling 2019) eco e ed in e speci ic di e gences
anging om 3.5% (genus Mecaphesa) o 9.6% (genus Xys i-
cus). In eg a i e axonomic s udies om non- homisid spide s
had eco e ed a wide ange o in e speci ic/in aspeci ic di-
e gence such as 4–5%/0.6–2.7% (genus Mica ia, Gnaphosi-
dae; Mus e & Michalik 2020), 3.8–13.0%/0.6–2.5% (genus
Dysde a, Dysde idae; Řezáč e al. 2014), 5.2–10.2%/0.2–3.9%
(genus Aphonopelma, The aphosidae; Hamil on e al. 2014) o
10.6–17.2%/1.4–4.6% (genus Loxosceles, Sica iidae; Planas &
Ribe a 2014). A s udy comp ising up o 371 mo phospecies
om 39 di e en amilies eco e ed a mean in aspeci ic di-
e gence o 2.07%, while mean in e speci ic di e gence was
9.73% (Domènech e al. 2021). Al hough he e a e mo pho-
logical di e ences as well as a clea physical ba ie be ween
mainland and insula popula ions, he low di e gence ound
be ween he Cana ian clade and i s sis e clade (0.9–1.6%)
p e en s ele a ion o he subspecies S. g. cana iense o species
le el. Gene ic s uc u ing in ou main clades and he g ea
Fig. 3: Synema globosum globosum (Fab icius, 1775) (uppe ow) and Synema globosum cana ienseDahl, 1907 (lowe ow), s e na o p ese ed specimens.
F = Female, M = Male, EU = Eu ope, CNY = Cana y Islands. B igh whi e spo s a e gla e a e ac s om LED illumina ion o subjec . F1- EU. Po o, Co sica,
F ance; F2- EU. Cala Mesquida, Majo ca, Spain; M1- EU. Pò ol, Majo ca, Spain; M2- EU. Rua da Fon e da Po ca, Po ugal; F1-CNY. Te o , G an Cana ia; F2-
CNY. Las Me cedes, Tene i e; M1- CNY. Los Tela es, La Gome a; M2- CNY. Taucho, Tene i e
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Resu ec ion o Synema globosum cana ienseDahl, 1907 23
di e gences ound among hem sugges he po en ial o high
c yp ic di e si y wi hin S. globosum. An in eg a i e axonomic
app oach is sugges ed o shed ligh on he di e si y o S. glo-
bosum and, po en ially, o delimi new subspecies, p obably
i ing wi h each clade eco e ed by U e e al. (2021). Fu u e
gene ic s udies o Synema should also include ma e ial om
o he subspecies and p e e en ially also o he Synema spe-
cies, and analyses o he nuclea genome o m DNA lineages
in sympa y. Such nuclea genomic analyses would enable
es ing o po en ial ep oduc i e isola ion associa ed wi h
m DNA lineages, hus e ealing i hese di e gences ep e-
sen c yp ic species, o an unusually high le el o in aspeci ic
m DNA a ia ion (e.g., Pé ez-Delgado e al. 2022). Se e al
Synema species ha e been poo ly desc ibed, some o hem no
e en being illus a ed, poin ing o he need o a e ision o
Synema.
Compa a i e mo phological analysis
May (1963) de ined subspecies as an “agg ega e o local pop-
ula ions o a species inhabi ing a geog aphic subdi ision o
he ange o he species and di e ing axonomically om o h-
e popula ions o he species”. This was elabo a ed by Mon oe
(1982), de ining subspecies as e e ing o “one o wo o mo e
popula ions o a species li ing in di e en , isola ed subdi i-
sions o he species’ ange and a ying om one ano he by
mo phological cha ac e is ics ha a e no mino o clinal and
wi h hese condi ions applying o all indi iduals wi hin he
subspecies’ ange”. Pe haps a mo e mode n way o subdi id-
ing poly ypic spide species in o subspecies is he concep o
supe species wi h allospecies as subuni s (K aus 2002). The
unde lying concep in hese de ini ions is he biospecies com-
p ised o wo o mo e subspecies wi h allopa ic specia ion
as he p ima y mechanism o specia ion. Based on esul s
om he abo e men ioned molecula and compa a i e mo -
phological analysis i ollows ha all condi ions o Mon oe
(1982) a e me o ecognizing S. g. cana iense as a subspecies.
Thus we he e e alida e S. g. cana iense Dahl, 1907 (subsp.
e al.), ejec ing Wunde lich (1987: 252). The i le page o
his book on he spide s o he Cana ian Islands and Madei a
(Wunde lich 1987) shows a pho o o a yellow emale, a com-
mon colou mo ph o he nomino ypical subspecies o he
con inen . Howe e , his pho o is also ound in a book wi h
colou images o Eu opean spide s (Saue & Wunde lich
1982), indica ing ha he pho o p obably depic s a Eu opean
specimen. Bo h subspecies a e geog aphically isola ed as he
Cana ian Islands a e sepa a ed om A ica by he 100 km
wide Lanza o e Passage, wi h an addi ional 50 km o G an
Cana ia which is he eas e nmos island wi h known p esence
o Synema. Pho os o Cana ian Synema on iNa u alis (e.g.
h ps://www.ina u alis .o g/obse a ions/115217436) indi-
ca e he p esence o specimens wi h da ke oliaceous ma k
han epo ed in his s udy, bu appa en ly no comple ely je
black as in mos Eu opean specimens.
Synema g. globosum om he Eu opean mainland may a -
i e in he Cana y Islands acciden ally by human induced
in oduc ion. The e a e se e al examples o his species be-
ing in oduced o new a eas ou side i s na u al ange, such as
o England (Bee e al. 2020). A i al o Synema g. globosum
o he Cana y Islands could b eak ep oduc i e isola ion and
esul in in e g ada ion wi h Synema g. cana iense. Hope ully,
his can be a oided. The spide auna o he Cana y Islands
is cha ac e ised by an e olu iona y his o y in ol ing chance
e en s o a i al and es ablishmen , o en ollowed by spe-
cia ion e en s ha may ha e occu ed wi hou equen in-
e g ada ion. Subspecies o local geog aphic aces ep esen
conse a ion uni s o special in e es , and in ou opinion, hey
should ha e he same ocus in conse a ion e o s as species
since hey ha e he po en ial o e ol e in o new species. The
pale colou o Cana ian Synema could be an adap a ion o
highe sola adia ion o he mo e sou he ly si ua ed Cana -
ian Islands. In e es ingly, he habi us o Synema g. cana iense is
nea ly iden ical o some colou mo phs o S. imi a ix (Pa esi,
1883) acco ding o colou images in Dippenaa -Schoeman e
al. (2020). Howe e , he la e species also encompass speci-
mens wi h b igh g een ca apaces which a e ne e seen in S.
g. cana iense.
Commen s on o he synonymies in Synema
As men ioned abo e, mos subspecies o S. globosum a e no
cu en ly accep ed. Fi e ou o he se en desc ibed subspe-
cies o S. globosum ha e been synonymized wi h he nomi-
no ypical subspecies by B ei ling e al. (2016), Ono (1988)
Tab. 1. Colou di e ences be ween Synema g. globosum (Fab icius, 1775) and Synema g. cana iense (Dahl, 1907)
Cha ac e Synema g. globosum (Fab icius, 1775) Synema g. cana iense (Dahl, 1907)
Male ca apace Uni o mly colou ed, b own o blackish b own (Fig. 1) B own o da k b own wi h pale o pale pos e io pa o
head (Fig. 2), some specimens wi h da k b own “U”-ma k
and pale ma gin as in emales
emale (and ju enile)
ca apace Yellow-b own o da k b own wi h back o head sligh ly
pale (Fig. 1) Pale wi h a la ge, da k b own “U”-ma k and pale ma gin
(Fig. 2)
S e num Uni o mly colou ed b own o blackish b own some imes
pale medially, ne e pale an e ola e ally (Fig. 3) Bicolou ed (whi ish ma ks an e ola e ally, howe e only
aguely disce nible in males, no pale medially (Fig. 3)
Colou polymo phy A dis inc ea u e o emales, indis inc in males. Female
opis hosoma whi e, g eyish whi e, c eam, g eenish whi e
o ligh g een, g eenish yellow, yellow b own, golden,
b igh yellow, o ange, ed o da k ed, males whi e (Fig. 1)
A mos udimen a y and hen only males. Female
opis hosoma gene ally pale, whi ish, ne e b igh yellow,
o ange o ed, males whi e o di y whi e (Fig. 2)
Opis hosoma
(no including
oliaceous ma k)
O en bicolou ed ed/yellow (Fig. 1 F4), o ange/yellow
(Fig. 1 F5), whi e/o ange (Fig. 1 M1) No bicolou ed in all examined ma e ial (Fig. 2)
Foliaceous ma k Usually je black (Fig. 1), some imes b own wi h da ke
edges (Fig. 1F2) Ligh b own wi h dis inc ly da ke edges o spo s, some-
imes blackish, no je black (Fig. 2)
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24 J. Lissne & D. Suá ez
and Wunde lich (1987) acco ding o he WSC (2023). This
lea es Synema globosum nig i en e Kulczyński, 1901 om
Sou h Sibe ia as he only cu en ly s ill accep ed subspecies o
Synema globosum (WSC 2023) beside om he nomino ypi-
cal subspecies. The h ee synonymiza ions o B ei ling e al.
(2016) a e jus i ied as F anganillo e ec ed sepa a e subspe-
cies wi hin he polymo phic S. globosum based on some o
he abo e-men ioned colou mo phs: Synema globosum cla um
F anganillo, 1913 = Synema globosum (Fab icius, 1775) (B ei-
ling e al. 2016: 55), Synema globosum la um F anganillo,
1913 = Synema globosum (Fab icius, 1775) (B ei ling e al.
2016: 56) and Synema globosum pulchellum F anganillo, 1926
= Synema globosum (Fab icius, 1775) (B ei ling e al., 2016:
56). Acco ding o he WSC (2023), Ono (1988) s a ed he
ollowing synonymiza ions in his e isional s udy o Japanese
homisids: Synema globosum daghes anicum U ochkin, 1960
= Synema globosum (Fab icius, 1775) (Ono, 1988: 147) and
Synema japonicum Ka sch, 1879 = Synema globosum (Fab icius,
1775) (Ono, 1988: 147). Synema japonicum Ka sch, 1879 has
been he subjec o se e al axonomic changes. Dahl (1907)
Fig. 4: Maximum-likelihood ee showing he phylogene ic placemen o Synema globosum cana iense. Boo s ap alues a e p esen ed only o main
clades. Coun ies a e indica ed a e GenBank accession numbe . Sequences a e colou ed ollowing U e e al. (2021) classi ica ion, i.e. blue o ‘blue clade’,
pu ple o ‘pu ple clade’ and g een o ‘g een clade’. Red: specimens sequenced in his s udy. Black: sequences no included in U e e al. (2021)
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Resu ec ion o Synema globosum cana ienseDahl, 1907 25
ega ded i a subspecies o S. globosum. U ochkin (1960)
seems o ha e been unawa e o i s subspecies s a us as he s ill
conside ed S. japonicum a alid species. Howe e , Ono (1988)
decided he would ea his species as S. globosum which was
accep ed as a synonymiza ion by he WSC (2023). He a -
gued ha 1) di e ences in colou a ion and ma kings could
no be used o iden i y subspecies, 2) he e we e only li le o
no a ia ion in he geni al pa s, 3) he ega ded he ma e ial
a ailable o him as “no su icien o de e mining he subspe-
cies di ision o he species”. None o he a gumen s o Ono
(1988) con adic s he de ini ions o a subspecies (e.g. Mon-
oe 1982). Thus, his decision “I p e e o pu all he desc ibed
subspecies and a ie ies in he complex o S. globosum in he
b oades sense“ is no logically deducible om his easoning.
Hence, al hough o mally co ec , he synonymiza ions o he
subspecies S. g. daghes anicum U ochkin, 1960 and he species
S. japonicum Ka sch, 1879 wi h S. g. globosum in Ono (1988)
a e ques ionable. These axa a e in u gen need o being in es-
iga ed o pheno ypic and gene ical di e ences o ecognize
hei p ope axonomic anks.
Acknowledgemen s
We wish o hank Jean-Philippe Tabe le , Nages e Solo gues,
F ance o allowing us o use one o his images o Synema globo-
sum. We also wish o hank B en Cha les Eme son o p o iding
a e ision o he languague s yle. Daniel Suá ez was unded by he
‘Minis e io de Ciencia e Inno ación’ h ough an FPI PhD ellowship
(PRE2018-083230). This wo k was suppo ed by p ojec s CGL2017-
85718-P ( unded by MCIN/AEI/10.13039/501100011033, Spain
and EDRF, EU), PID2020-116788GB-I00 ( unded by MCIN/
AEI/10.13039/501100011033, Spain), and P oID2021010093
( unded by he Cana y Islands Go e nmen , Spain and EDRF, EU)
awa ded o B en Cha les Eme son. Fieldwo k was suppo ed by he
collec ing pe mi s 246285, Nº 5059 (23/10/2020), A/EST-001/2020,
and AFF 193/21 kindly p o ided by he Cana y Islands Go e nmen ,
Cabildo de La Gome a, Cabildo de La Palma and Cabildo de Te-
ne i e, espec i ely.
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