Sixty years after "A review of the genera of the family Tydeidae" (Acariformes: Tydeoidea: Tydeidae)

Recei ed 18 Decembe 2024
Accep ed 25 Feb ua y 2025
Published 21 Ma ch 2025
Co esponding au ho
Hen i M. And é:
[email p o ec ed]
Academic edi o
Akashi He nandes, Fabio
h ps://doi.o g/10.24349/l5 b-767l
ISSN 0044-586X (p in )
ISSN 2107-7207 (elec onic)
And é H. M.
Licensed unde
C ea i e Commons CC-BY 4.0
Six y yea s a e “A e iew o he
gene a o he amily Tydeidae”
(Aca i o mes: Tydeoidea: Tydeidae)
Hen i M. And éa,b,c
aRe i ed, cu en ly, 22, ou e de la Cô e, CH-1615 Bossonnens, Swi ze land.
bMusée oyal de l’A ique cen ale, B-3080 Te u en, Belgium.
cUni e si é Ca holique de Lou ain, B-1348 Lou ain-la-Neu e, Belgium.
O iginal esea ch
ABSTRACT
The goal o his pape is o p esen an o e iew o he gene a o he p os igma ic mi e
amily Tydeidae 60 yea s a e he appea ance o E.W. Bake ’s seminal pape on he amily
(1965), and o de ine in sys ema ic and scien i ic e ms he gene a wi hin he supe amily
Tydeoidea. Cumula i e cu es a e gi en o gene a, species and names. The numbe o
mo phospecies is es ima ed o be a ound 11,600 o which 3.72% ha e been desc ibed.
E olu iona y plas ici y and a new concep o he amily a e explo ed. The e ised amily
is conside ed he e o con ain 31 gene a and 3 non-nominal subgene a. Nomencla u al
changes a e no ed in amilies, gene a, subgene a and species: Bake lasio ydaeus n. gen.
(mo ed o Iolinidae, Tydaeolinae); Neoa o ydeus n. gen. (Tydeidae); Neohomeo ydeus n.
gen. ( eplaces Homeo ydeus no in acco dance wi h ICZN); A o ydeus, junio synonym
o Tydeus (n. syn.); Pa alo yia, junio synonym o Calo ydeus (n. syn.); Pe ydeus,
subgenus o Tydeus ( es . comb.); Quad o ydeus, subgenus o B achy ydeus ( es . comb.),
Eo ydeus, subgenus o B achy ydeus (n. comb.), Bake lasio ydaeus. k an zi n. comb.
(Iolinidae); Tydeus nieuke keni (jus i ied emenda ion) and Homeo ydeus o mosus (gende
ag eemen ). A able allowing he iden i ica ion o emale ydeid gene a and subgene a is
p esen ed.
Keywo ds Iolinidae; dac yly; e olu ion; Tydeinae; P e ydeinae; Aus ali ydaeinae
Zoobank h p://zoobank.o g/F3176DAB-5A34-4299-8730-8174F468233C
述而不作 信而好古
I ansmi bu do no inno a e. Con ucius, Analec s 7: 1.
Tex and ansla ion by Slinge land (2003).
In oduc ion
Tydeidae was e ec ed by K ame (1877: 224) unde he name Tydidae, a name no cong uen
wi h he guidelines o he In e na ional Code o Zoological Nomencla u e (he ea e e e ed
o as ICZN, o he Code) as a sub amily o P os igma a p oposed o accommoda e he single
genus Tydeus e ec ed by Koch (1836). P e iously, he genus Tydeus was placed in Eupodidae
(Koch, 1850: 70). La e , Tydeus was s ill associa ed wi h eupodid mi es by Canes ini (1892:
575) and Be lese (1925: 94). A i s monog aph ea ing he Tydeidae as a amily including 14
gene a, 4 subgene a, 69 de e mined and 13 unde e mined species was p o ided by Tho (1933:
12) who placed hem in Eupodoidea ( oge he wi h E eyne idae). Bake , in his seminal pape
on he gene a o he amily Tydeidae, kep he sense used by Tho (Tho ian sense), de e mined
15 gene a and admi ed ha he amily was “di icul o cha ac e ize” (Bake , 1965: 96). He
How o ci e his a icle And é H. M. (2025), Six y yea s a e “A e iew o he gene a o he amily Tydeidae”
(Aca i o mes: Tydeoidea: Tydeidae). Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l
di ided he ydeids “in o wo g oups, hose wi h [se a] L[La e al]2in he la e al posi ion, and
hose wi h L2in line o nea ly in line wi h he D[o sal] se ae” (Bake , 1965: 98).
And é (1979) di ided he ydeids in o se en sub amilies bu kep he adi ional Tho ian
sense. Subsequen ly and a e nume ous a icles on species iden i ica ions, Kaźmie ski (1996a,
1998a) p esen ed he i s app oaches based on a phylogene ic analysis o Tydeinae and
P e ydeinae.
And é and Fain (2000) a ibu ed a new meaning o he amily hey included in he
supe amily Tydeoidea. This app oach is adop ed he e. The supe amily included he
E eyne idae, Iolinidae, Tydeidae and T ioph ydeidae. This classi ica ion wi h 4 amilies
is also adop ed. A diach onic iew was gi en by And é (2021, ig. 1).
Ecologically speaking, “ he Tydeidae is pe haps he mos common o all olia mi e amilies,
and almos any ee, sh ub, o liana ha has no been ea ed by pes icides will ha e hese mi es
on i s lea es, s em, and ba k” (Wal e and Behan-Pelle ie , 1999: 9). The ela ionship be ween
ydeid mi es and plan s was ecognized as ea ly as he 1800’s, especially by bo anis s. One
o he oldes eco ds o his associa ion is Tydeus olio um Sch ank, which was collec ed in
I aly on he unde sides o lea es, in moss, and in hay deb is (Canes ini, 1896a: 233), no h o
Finland whe e i was equen ly ound in hay and g ass (No denskiöld, 1900: 35). Lunds öm
(1887: 9), Heinsius (1891: 197) and Ross (1904: 17) e e ed o he p esence o T. olio um in
plan doma ia. Howe e , Abliz e al. (2014) men ioned hem in passing in hei publica ion o
he Encyclopedia o UNESCO. Tydeids a e pionee s in inne An a c ica (Jane schek, 1967) and
a e li ing in he S alba d a chipelago (Tho , 1933; Seniczak e al. 2020). They we e collec ed
om he sea le el (Willmann, 1952) o he Himalayas (Momen and Solhøy; 1996; Chand a e
al., 2018).
In ag oecosys ems, Te anychidae and E iophyidae a e amous due o he plan eeding
p oblems hey pose which we e consequen ly cla i ied in compila ions by Lindquis e al.
(1996) and Uecke mann (2010). This is no he case o Tydeidae which o ms a mino
ag icul u al mi e g oup e en i Banks (1904) desc ibing a new ydeid i led his publica ion as
“Fou new species o inju ious mi es” and s a ed ha hey we e all “o conside able economic
impo ance”. Subsequen ly, Tydeidae we e ea ed as po en ial pes s in majo ex books such as
he “Mi es Inju ious o Economic Plan ” by Jeppson e al. (1975), he “Ag icul u al Aca ology:
In oduc ion o In eg a ed Mi e Managemen ” by Hoy (2011), o he “Handbook o Mi es o
Economic Plan s” by Vacan e (2016). A amous excep ion is he species “Lo yia o mosa”
desc ibed by Coo eman (1958) (see he sec ion on Inadequa ely desc ibed axa). Howe e , he
s a us and unc ion o Tydeidae as pes s, bene icials, p eda o s o al e na e p ey, a e beyond he
scope o his pape .
In ood and s o ed p oduc s, Tydeidae we e no e en illus a ed by Smiley (1987) and only
3 ydeid species we e desc ibed om s o ed g ain by Momen and Sinha (1991). Howe e , hey
may be locally signi ican as in a chocola e p oduc ion sys em wi h 55% o he mi es collec ed
(Sil a e al., 2024).
In medical and e e ina y sciences, Tydeidae a e also unimpo an and no ci ed in he
e iews by Mullen and OConno (2019) and Lynn e al. (2024). A no able excep ion is Tydeus
moles us desc ibed by Moniez (1889, 1894) in he ga den o a Belgian a m and hei impac
on people. Analogous de mopa hies p oduced by his mi e o a simila species we e desc ibed
in an I alian school by P incipa o e al. (2008) and P incipa o and Sc iboni (2009).
The sys ema ics o he amily a e some imes con using, especially o beginne s. E en
a ecen species such as Quad o ydeus sleipne i desc ibed by Momen and Lundquis (1996)
has been assigned o wo o he gene a: Lo yia (Kaźmie ski, 1998a: 327) and B achy ydeus
(Khaus o and Khaus o , 2023: 268). The beginne s a e s ill mo e emba assed by he double
assigna ion, in he same publica ion, o sleipne i o Quad o ydeus and o B achy ydeus (Sil a
e al., 2016: 25, 34). This double assigna ion ecalls ha Dugès’ cauda us was simul aneously
ea ed as a Tenuipalpidae and as a Tydeidae (And é, 2011). “Neglec ” o li e a u e, di e gen
in e p e a ions o con adic o y obse a ions also lead o con usion: A o ydeus was e ec ed
by Bake (1970) as subgenus o Tydeus o accommoda e h ee species, muns e i, kenyensis,
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 150
and meye ae, which exhibi h ee di e en leg chae o axies. The wo ld ca alog published by
Sil a e al. (2016) ea ed a genus, Pseudolo yia, p e iously synonymized wi h Calo ydeus by
And é (2005: 995), and did no include he genus Calo ydeus. The gene a e ec ed by Be lese
a e named a e he name wi h “a” a e he “d” (Lasio ydaeus,Melano ydaeus) con a ily o
S ylo ydeus e ec ed by Tho , Be lese e en w o e Tydaeus ins ead o Tydeus in his publica ions
and in his collec ion. This misspelling is some imes ound in ecen a icles. A las example
conce ns he coexis ence o se e al designa ion sys ems based on loca ion and a no a ion
sys em ounded on idionymy such as ha de eloped by And é (1980), which can be con using
as deplo ed by De Vis e al. (2024: 1097).
Qui e simply, his pape aims a p esen ing an o e iew o ydeid gene a six y yea s a e
he seminal pape on he amily Tydeidae p esen ed by Bake (1965) and a de ining hem in
a sys ema ic and scien i ic way wi hin he supe amily Tydeoidea, e en i he supe amily is
conside ed pa aphyle ic (Szuda ek-T ep o e al. 2022).
Ma e ials and me hod
This a icle ea s ex an mo phospecies (i.e. species based on mo phological c i e ia) desc ibed
un il 30 June 2024. Tydeid species desc ibed la e in 2024 a e no included (e.g. B achy ydeus
u kiyensis desc ibed by Al unç e al., 2024). The no a ion sys em p oposed by And é (1981b)
is used only o apo eles and a si, a designa ion sys em is applied elsewhe e.
A i s da abase (1760 lines x 12 columns) was p o ided by he In eg a ed Taxonomic
In o ma ion Sys em (ITIS), namely by Da id Nicolson and Geo ey D. Owe who we e
de eloping a web c awle o Tydeoidea. Thei basic da abase was upda ed and comple ed,
e.g. he numbe o combina ions.
O he da a a e essen ially based on pe sonal da a and bibliog aphical e e ences (books,
a icles, e iew pape s) on Tydeoidea axonomy and on he da abase in he Wikispecies pla o m
(And é, 2021). Pe inen wo ks we e also consul ed, such as monog aphs, books and ca alog:
Tho (1933), Bake and Wha on (1952), Bake (1965), And é (1980), Kaźmie ski (1998a) and
Sil a e al. (2016).
To a oid con usion, special a en ion is gi en o gene a and hei ype species. Fo ins ance,
Lo yia cumb ensis Bake , 1944 is he ype species o he genus Pa alo yia e ec ed by Bake
(1965). And é (1980: 119) placed i in a new genus he called Homeo ydeus, a nomencla u al ac
no in con o mi y wi h ICZN. Simila ly, And é (1980: 128) placed Tydeus muns e i, he ype
species o A o ydeus, in a new genus he called O ho ydeus ( ype species: T. goe zi). Again,
his ails o mee he ules o ICZN.
A he gene ic le el, only me is ic cha ac e s we e conside ed, i.e. coun s o disc e e se ially
homologous s uc u es (Lawing e al. 2008). Roughly, his co esponds o he p esence/absence
o idionymic cha ac e s ou lined by G andjean (1952). Diagnoses based on me is ic cha ac e s
we e qui e unusual be o e Bake ’s (1965) seminal pape and some gene a we e de ined mainly
by o namen a ion. Al hough o namen a ion may be a key cha ac e o delinea e species (see
igs 25-27 by Momen and Lundq is , 1996), cu en ly i does no p o ide adequa e suppo a
he gene ic le el.
To de e mine he numbe o gene a and species, nomina no ae and eplacemen names we e
no e ained. Simila ly, misspelled names (e.g. Aus alo ydeus ins ead o Aus alo ydaeus) and
speci ic epi he s wi h no gende ag eemen ( o mosa ins ead o o mosus) we e no included
in his publica ion. New combina ions and e ised diagnoses we e likewise no included.
Species wi h ag onomical in e es s we e conside ed so when hey we e desc ibed om an
ag oecosys em oliage o unk, a s a emen such as “ om soil samples o mango o cha ds”
was no su icien . Ince ae sedes (e.g. species in which he phane o axy is no desc ibed in he
o iginal publica ion and in subsequen publica ions) we e also coun ed.
The slopes, m, indica ed in igu e 3 we e calcula ed he usual way: m= (y2-y1)/(x2-x1)
whe e yand xa e he coo dina es o he wo end poin s o he line segmen .
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 151
Resul s
Lis o gene a excluded om Tydeidae
Gene a ini ially desc ibed in Tydeidae bu ans e ed la e o o he amilies include (in
alphabe ical o de ):
Genus ans e ed o Pa a ydeidae: Scolo ydaeus.
Gene a ans e ed o T ioph ydeidae: Apo ioph ydeus,Edbake ella,Meye ella,P e io-
ph ydeus,Pseudo ioph ydeus,S enipedis,Tele ioph ydeus and T ioph ydeus.
Genus ans e ed o E eyne idae: Pseudo ydeus.
Gene a ans e ed o Iolinidae: Aes he ydeus,And e ydaeolus,Apop onema us,Cocco-
ydaeolus,Coleo ydeus,Homeop onema us,Me ap onema us,Me a ydaeolus,Mic o ydeus,
Naudea,Neonaudea,Neop onema us,Oak illea,Pa ap onema us,Pa a ioph ydeus,Pausia,
P imo ydeus,P oc o ydaeus,P onecupula us,P onema ulus,P onema us,Pseudop onema u-
lus,Recki ydeus,Tydaeolus and Tynda eus.
Lis o gene a and subgene a o Tydeinae
In he Linnaean sys em o nomencla u e, he gene ic name is pa o he La in binomial
a ibu ed o all species, and he e o e appea s in e e y scien i ic pape dealing wi h li ing
beings; i also gi es his name an impo an ole in sys ema ics, ce ainly much mo e impo an
han ha o he names o highe axa (Dubois, 1988: 15-16).
A pa ial and diach onic iew o ydeine gene a and subgene a is gi en in Figu e 1. The
cu en numbe o gene ic names amoun s o 45 among which wo a e eplacemen names.
In case o homonymy, eplacemen names a e sys ema ically appended o he eplaced name
(Kuzne so ydides o Tydides, Kuzne so ia o Venilia) bu hey a e lis ed he ea e . In con as ,
synonymic gene a, junio and senio synonyms, a e d awn sepa a ely (e.g. Quad o ydeus and
B achy ydeus). Mos synonymies a e no based on ypes (homo ypic synonymy) bu esul
om a compa ison o da a eco ded in he li e a u e (Fig. 1). Gene a a e lis ed he ea e in
alphabe ical o de , excep Tydeus he ype genus. The numbe o componen species (possibly
be o e synonymiza ion) po en ially ollowed by he numbe o species inqui endae is indica ed
be ween squa e b acke s.
(1) Tydeus [58+46]. The his o y o he genus is complex and con olu ed and summa ized by
And é (2005). The ype species designa ed by he In e na ional Commission on Zoologi-
cal Nomencla u e (ICZN. 2008) is Tydeus spa hula us Oudemans, 1929 (consequen ly, i
is also he ype species o he amily and supe amily).
(2) Acan hodides [1]. Monospeci ic genus e ec ed by Kaźmie ski (1996b). The ype species
is A. ja ema by o iginal designa ion.
(3) A idiolo yia [3]. Genus e ec ed by Kaźmie ski (1996b). The ype species, Lo yia
a icana Bake , 1965, was designa ed in he o iginal publica ion.
(4) A o ydeus [5]. I was e ec ed by Bake (1970) as subgenus o Tydeus o accommoda e
h ee species: muns e i,kenyensis, and meye ae. These h ee species exhibi h ee di e -
en leg chae o axies. The ype species designa ed by Bake (1970: 165) is Tydeus mun-
s e i and no A. kenyensis as epo ed by And é (1980: 107), Kaźmie ski (1998a: 354)
and Sil a e al. (2016: 8). The species desc ibed by Meye and Ryke (1959) and s udied
by And é (1980: 126) and by Uecke mann and G ou (2007) has 6 geni als [and no 4 as
no ed by Bake (1970: 164)] and exhibi s he same phane o axy as he nominal subgenus
Tydeus.
(5) And elo yia [1]. This genus was e ec ed by Khaus o (2022). Type species: A. hajiqan-
ba i by o iginal designa ion.
(6) Apolo yia [1]. Genus desc ibed by And é (1980) wi h Lo yia congoensis Bake , 1970
as ype species by o iginal designa ion.
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 152
Figu e 1 . Diach onic iew o old gene a and subgene a desc ibed in Tydeinae. Only o mal synonymies a e indica ed, no he new combina ions
o ype species.
(7) B achy ydeus [220+9]. The genus was e ec ed by Tho (1931) wi h Tydeus c ucia us
as he ype species. The ype species was edesc ibed by And é (2005) in Oudemans’
collec ion. Synonymized wi h Tydeus by Bake and Wha on (1952: 191) and by Bake
(1965: 99). The genus was no lis ed by And é (1980) and by Kaźmie ski (1998a). I
was es o ed by And é (2005: 995) and ca alogued by Sil a e al. (2016: 9). Roughly,
Lo yia sensu Kaźmie ski co esponds o B achy ydeus (And é, 2005: 995; Temp li e
al., 2015: 944; Akba i e al., 2015a: 424).
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 153

(8) Calo ydeus [15+4]. The genus was e ec ed by Oudemans and he ype species is Calo y-
deus c oceus by o iginal designa ion (Oudemans, 1937: 922). Synonymized wi h Tydeus
by Bake and Wha on (1952: 191), synonymiza ion epea ed by Bake (1965: 100). The
genus was no lis ed by And é (1980), Kaźmie ski (1998a) and Sil a e al. (2016) bu was
es o ed by And é (2005: 995, 998). Among o he species, he genus comp ises he ype
species s udied by Oudemans and h ee species desc ibed by Bake (namely a hu bake i,
cumb ensis,shawi). Pseudolo yia was synonymized wi h Calo ydeus by And é (2005:
995). A neo ype om Oudemans’ collec ion was selec ed by And é (2005: 996).
(9) Edlo yia [1]. The ype species by o iginal designa ion o he genus e ec ed by Kaźmie -
ski (1996b: 201) is based on a unique specimen o Bake (1968), Lo yia undado ensis,
which is a deu onymph he chae o axy o which was pa ly published bu comple ed by
Kaźmie ski (1996b: 201).
(10) Eo ydeus [1]. Subgenus e ec ed by Kuzne so (1973) a e s udy o C imean specimens
and aised o ull genus ank by And é (1980: 113) a e examina ion o a Canadian spec-
imen. No lis ed by Kaźmie ski (1998a) and Sil a e al. (2016). The ype species, Tydeus
(Eo ydeus)mi abilis, was assigned o Lo yia by Kaźmie ski (1998a: 336) and o B achy-
ydeus by Sil a e al. (2016: 19).
(11) Homeo ydeus [2]. Genus e ec ed by And é (1980: 116–117) bu wi hou ixing a ype
species, which did no acco d wi h A . 13.3 o ICZN (”To be a ailable, e e y new genus-
g oup name published a e 1930 … mus … be accompanied by he ixa ion o a ype
species in he o iginal publica ion″). Ano he disc epancy wi h he code ( he inclusion o
he ype species o ano he genus) was ou lined p e iously. Genus ea ed by Kaźmie ski
(1989). The 3 species desc ibed by Bake (namely a hu bake i,cumb ensis,shawi) we e
ans e ed o Calo ydeus by And é (2005: 995, ambiguous ema ks), he wo Belgian
species ha e a se a on coxa I and emained in he so-called genus. Synonymized wi h
Calo ydeus by And é (2021: 1026).
(12) Homoio ho ydeus [1]. Genus p oposed by Schiess (1981: 84) and synonymized wi h
Tydeus by Kaźmie ski (1998a: 342).
(13) Idiolo yia [2]. The ype species o he genus e ec ed by And é (1980: 117) is Lo yia
macquillani Bake , 1968, by o iginal designa ion.
(14) Kenlo yia [1]. Monospeci ic genus desc ibed by Kaźmie ski (1996b) wi h Kenlo yia
masaii as ype species by o iginal designa ion.
(15) K an zlo yia [1]. Monospeci ic genus e ec ed by And é (1980: 117) wi h Lo yia g ewia
Bake , 1968 as ype species by o iginal designa ion.
(16) Kuzne so ia [7]. Replacemen name p oposed by Kamme e (2006: 269) o Venilia,
name al eady used o molluscs, insec s, bi ds… Type species: Pa alo yia libe a Li shi z,
1973. Venilia is a junio synonym o Tydeus acco ding o And é (1980: 154) o junio syn-
onym o B achy ydeus acco ding o And é (2005: 995). Senio homonym o Kuzne so ia,
eplacemen name ad anced by Doweld (2016) o a C us acea.
(17) Kuzne so ydides [1]. Replacemen name p oposed by And é (2021: 1030) o Tydides,
name al eady used o insec s. The genus was e ec ed by Kuzne so (1975) wi h Tydides
ul e Kuzne so , 1975 as ype species by o iginal designa ion.
(18) Lasio ydaeus [?]. Fi s , monospeci ic genus e ec ed by Be lese (1908: 15) wi h Lasio y-
daeus glycyphaginus as ype species. Desc ip ions a e sho : 6 lines o he genus, 3 lines
o he species collec ed in Flo ence (I aly). The p odo sum illus a ed by Be lese (1910:
pla e 18, ig. 6) has only 3 pai s o se ae and i is di icul o conclude i i is p o- o e-
cu ed; howe e , he h ee se ae a e in line as in mammila is ( ig. 8, p odo sum ecu ed)
and hei alignmen con as s wi h he adjacen igu e ( ig. 7) in which he p odo sum
is p ocu ed wi h 4 se ae a anged in a zigzag. Las ly, he do sal opi hosomal pa e n
d awn on ha igu e is simple, i consis s o se e al ans e se ows o 4 se ae each and
shows no ace o he do sal mig a ion o he second la e al se ae (his L2) e oked by Bake
(1965). I was di ided in 2 subgene a by Be lese (1910: 211), he nominal subgenus wi h
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 154
2 species and Melano ydaeus wi h 5 species. The genus was synonymized wi h Tydeus by
Bake and Wha on (1952: 191; wi h a ques ion ma k). The genus—monosubgene ic—
was es o ed by Bake (1965: 101) bu was based on a Cali o nian species (L. k an zi).
The p odo sum o he Cali o nian species desc ibed by Bake (1965: 101-102) and And é
(1980: 118) is p ocu ed, he opi hosomal pa e n is di e en om ha obse ed in ydeid
mi es, he a sus I has 12 se ae ( s 8 in Tydeinae), he Cali o nian mi e is no congene ic
wi h he I alian ype species (which seems o be a eal Tydeidae e en i i is impossible
o know wha he genus ac ually is) and appea s o be an Iolinidae Tydaeolinae genus
inqui endus.
(19) Lo yia [1]. Monospeci ic genus e ec ed by Oudemans (1925) wi h L. supe ba as ype
species. The e y sho desc ip ion o he ini ial pape was comple ed in a subsequen
a icle (Oudemans, 1928a). The whole s o y is epo ed by And é (2023). The genus
was no lis ed by And é (1980). The holo ype and unique specimen o he ype species
is los . Monospeci ic and, e en ecen ly, con used o synonymized wi h B achy ydeus
(e.g. Mondin e al., 2016; Khaus o e al., 2020; Nu olini e al., 2020)
(20) Melano ydaeus [?]. Melano ydaeus was i s desc ibed as a subgenus o Lasio ydaeus
(Be lese, 1910: 211) and was aised o ull gene ic le el by Tho (1933: 48). I was di-
ided in o wo subgene a, Melano ydaeus and S ylo ydeus and synonymized wi h Tydeus
by Bake and Wha on (1952: 191). I was es o ed by Bake (1965: 102) bu was no
keyed ou . “The placing o Melano ydaeus as a synonym o Tydeus by Bake and Wha -
on mus a p esen be conside ed o be in e o ” commen ed Bake (1965: 102) who
concluded “The ype, in he Be lese collec ion, is in ex emely poo condi ion, and i is
impossible o know wha he genus ac ually is”. Genus inqui endus.
(21) Melisso ydeus [3]. Genus e ec ed by And é (1985a: 244) wi h M. mac osolenus as ype
species by o iginal designa ion.
(22) Me alo yia [7]. Genus ad anced by And é (1980: 118) wi h Lo yia a maghensis Bake ,
1968 as ype species by o iginal designa ion.
(23) Momenia [1]. Genus p oposed by Kaźmie ski (1996b) wi h Tydeus longichelus Momen
& El-Bagou y, 1989 as ype species by o iginal designa ion.
(24) Neoapolo yia [5]. Genus e ec ed by El-Bagou y and Momen (1990) wi h N. aegyp ica
as ype species by o iginal designa ion.
(25) Neolo yia [3]. Genus ad anced by And é (1980: 127) wi h Re e ydeus boycei Bake ,
1944 as ype species by o iginal designa ion.
(26) Nudilo yia [6]. Genus e ec ed by Kaźmie ski (1996b) wi h N. pa a e ula as ype species
by o iginal designa ion.
(27) O a ep ydeus [1]. Monospe ic genus e ec ed by Uecke mann and G ou (2007) o O.
s epheni, ype species by o iginal designa ion.
(28) O ho ydeus [?]. Genus e ec ed by And é (1980: 127–128) wi h Tydeus goe zi as ype
species by o iginal designa ion. Howe e , And é (1980: 128) mis akenly placed Tydeus
muns e i, he ype species o A o ydeus, in O ho ydeus. Kaźmie ski (1989) p oposed
a new combina ion, Lo yia. The genus was synonymized wi h Tydeus by And é (2005:
996).
(29) Pa alo yia [5]. Genus e ec ed by Bake (1965) o ecei e Tydeinae wi h longi udinal
s iae be ween se ae D2. Type species by o iginal designa ion: Lo yia cumb ensis Bake ,
1944. Kaźmie ski (1989) p oposed a new combina ion. The genus was lis ed by Kaźmie -
ski (1998a: 340) and by Sil a e al. (2016: 32) (Fig. 1).
(30) Pe a o ydeus [1]. Monospeci ic genus p oposed by And é (1980: 142) wi h Tydeus
(A o ydeus)meye ae Bake , 1970 as ype species by o iginal designa ion.
(31) Pe ydeus [4]. This subgenus o Tydeus e ec ed by And é and Naudo (1965) and ecog-
nized by Bake (1970: 164). I was no lis ed by And é (1980), Kaźmie ski (1998a) and
Sil a e al. (2016) (Figu e 1). The ype species by o iginal designa ion, Tydeus schus e i,
was assigned o Lo yia by Kaźmie ski (1998a: 336) and o B achy ydeus by Sil a e al.
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 155
(2016: 24). No o mulae we e gi en in he desc ip ion bu he o iginal igu e 3 allows o
iden i y he mi e as belonging o he genus Tydeus (Fe II wi h 2 se ae, Fe III wi h a single
se a).
(32) Pseudolo yia [11]. The genus was e ec ed by Kaźmie ski (1989) and synonymized wi h
Calo ydeus by And é (2005: 995). Type species: P. edwa dbake i by o iginal designa ion.
(33) Quad o ydeus [1]. Monospeci ic genus ad anced by Momen and Lundq is (1996), syn-
onymized wi h Lo yia by Kaźmie ski (1998a: 293). The genus is lis ed by Sil a e al.
(2016: 34) bu he ype species, Q. sleipne i, was assigned o Quad o ydeus on page 34
and o B achy ydeus on page 25. The ype species by o iginal designa ion, Q. sleipne i,
was assigned o B achy ydeus by Khaus o and Khaus o (2023: 268).
(34) Quasi ydeus [2]. Genus e ec ed by Kaźmie ski (1996b) wi h Tydeus (T.) icensis Bake ,
1970 as ype species by o iginal designa ion.
(35) Raphi ydeus [1]. Monospeci ic subgenus o Lo yia e ec ed by Tho (1933: 54) o Lo -
yia (Melano ydaeus) aphigna hoides Be lese 1910, ype species by mono ypy. Di i-
sion no accep ed by Bake (1944: 215) and ecalled in Bake (1968: 987). Again, syn-
onymized wi h Lo yia by Bake (1965: 105) and wi h B achy ydeus by And é (2005:
995).
(36) Re e ydeus [5]. Genus e ec ed by Tho (1931) wi h R. ca enula us as he ype species.
Las used in mi e desc ip ions by Bake (1944) o boycei and doddsi. The genus was
lis ed— o he las ime!—by Bake and Wha on (1952: 192), wi h ca enula us as ype
species. A synonymiza ion wi h Lo yia was ad anced by Bake (1965: 105). Al hough
he ype species, ca enula us, was ne e edesc ibed om No way, i was included in he
genus B achy ydeus by Sil a e al. (2016: 13). The specimens o R. i ipa us s udied by
G andjean (1938) belong o B achy ydeus.
(37) S ylo ydeus [2]. Subgenus e ec ed by Tho (1933: 50) and synonymized wi h Tydeus by
Bake and Wha on (1952: 191). No ype species designa ed. The wo componen species
(b e is ylus and s ylige ) a e species inqui endae.
(38) Tydides [1]. Monospeci ic genus e ec ed by Kuzne so (1975) o T. ul e . Junio homonym
o he genus Tydides S ål, 1865 (Insec a, Redu iidae). Replacemen name: Kuzne so y-
dides p o ided by And é (2021: 1030).
(39) Tydulosus [4]. The genus was e ec ed by Bake (1965: 103) o species wi h a baske -
wea e pa e n. The ype species designa ed by Bake , Tydeus g anulosus Canes ini,
1886a, is a species inqui enda and has no ype. Synonymized wi h Tydeus by And é
(1980: 154). The species ini ially assigned o Tydulosus a e included in B achy ydeus
and Tydeus by Sil a e al. (2016).
(40) Venilia [7]. Genus e ec ed by Kuzne so (1979) comp ising 7 species e iewed by Kaźmie -
ski (1981). Junio synonym o Tydeus acco ding o And é (1980: 154), junio synonym
o Lo yia acco ding o Kaźmie ski (1989) and junio synonym o B achy ydeus acco d-
ing o And é (2005: 995). Junio homonym o a mollusc, successi e eplacemen names:
Kuzne so ia and Kuzne so enilia. Type species: Pa alo yia libe a Li shi z, 1973.
Gene a in Aus alo ydaeinae and P e ydeinae
(41) Aus alo ydaeus [1]. Monospeci ic genus p oposed by Spain (1969) o A. ki s enae, ype
species by o iginal designa ion.
(42) No zelo yia [1]. Monospeci ic genus e ec ed by Kaźmie ski (1996b) o N. dese a, ype
species by o iginal designa ion.
(43) P elo yia [2]. Genus e ec ed by And é (1980: 142-143) wi h Lo yia indonesiensis
Bake , 1970 as ype species by o iginal designa ion.
(44) P e ydeus [12]. Genus p oposed by And é (1980: 143) wi h Lo yia ke ani Ma shall,
1970 as ype species by o iginal designa ion.
(45) Uecke mannia [1]. Monospeci ic genus ad anced by Kaźmie ski (1996b) o Pa alo yia
g ewiae Uecke mann, 1988, ype species by o iginal designa ion. The ype species is
desc ibed in Uecke mann and Smi h Meye , 1988: 46-47.
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 156
Figu e 2 . Cumula i e genus desc ip ion cu es in Tydeidae, wi h and wi hou co ec ions ollowing subsequen synonymies and es o a ions.
Desc ip ion yea s a e g ouped by decade. Whi e numbe s in his og am indica e he numbe o gene a lis ed in he publica ions indica ed below.
As e isk e e s o he es ima ion by Zhang e al. (2011). The las poin ep esen s an incomple e decade (2021-2023). Subgene a a e no
included.
Numbe o gene a in Tydeidae
The p ac ical ag eemen which exis s be ween specialis s as o he delimi a ion o gene a a ies
om one zoological g oup o ano he . In many g oups, his ag eemen be ween specialis s is
poo , and his esul s in a g ea ins abili y o he gene ic classi ica ion (Dubois, 1988: 16).
Figu e 2 gi es a his og am based on he monog aphs, books and ca alog lis ed p e iously
and wo cumula i e genus desc ip ion cu es since 1831. The i s cu e p o ides he numbe
o all new gene a and subgene a e ec ed du ing each decade while he second akes in o
accoun synonymies and es o a ion. The maximum numbe o gene a and subgene a desc ibed
in Tydeidae is 43 (a o al o 45 names – 2 eplacemen names). A e synonymiza ion and
es o a ion, he cu en numbe o gene a and subgene a in Tydeidae is 30 (Fig. 2). The
ins abili y e oked by Dubois (1988) is disce nable h ough he gap be ween he wo cu es.
Depending on he cu e selec ed, he numbe o gene a a he beginning o he 2011-2021
decade amoun s o 29-43 and is close o he es ima ion by Zhang e al. (2011: 130), 30 gene a.
Bo h cu es appa en ly begin o la en du ing he las 40 yea s.
Numbe o species in Tydeidae
A common app oach o es ima ing he o al numbe o ex an species in a axonomic g oup is o
ex apola e om he empo al pa e n o known species desc ip ions (Bebbe e al. 2007). This
pe spec i e is no new in mi es and was used by Wha on (1964) o mi es and T ombiculidae
and by And é and N’D i (2013) o mi es and Tydeoidea. Ye , such an app oach is no
ecommended due o he da a weakness: only 433 ydeid species ha e been desc ibed, igu e 3
shows he beginning o a logis ic cu e and he e is no long- e m end sensu Edie e al. (2017).
Ne e heless, a u ning poin is appa en om igu e 3 and is indica ed by a double a ow.
Bo h cumula i e cu es sugges a change in he desc ip ion a e mani es ough he slopes
es ima ed be o e and a e he 1951-1960 decade. This is con i med by he decade end in
species desc ip ion wi h a maximum numbe o 83 species desc ibed in a single decade. The
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 157
Figu e 7 . The dis ibu ion o s udy si es o ydeid ypes be ween A – 1901 o 1920, B – 1951- o 1970, and C – om 2001 o 2020. The in luence
o Be lese (I aly) and Tho (Sweden) is appa en in A.
As hei colleagues, aca ologis s who deal wi h Tydeoidea ha e a limi ed knowledge o
global aca o auna. E en i he Eu ocen ism appa en in Figu e 7A ends o disappea la e ,
some majo a eas ha e been li le explo ed: A ica (no ydeid mi e in he ca alog by Mwase
and Bake , 2006), Aus alia (Halliday (1998) does no ci e a single iden i ied species), China
(only 27 de e mined species in Lin and Zhang, 2010: 54), Middle Eas (17 de e mined species
in Kamali e al., 2001: 150-152; only 1 in Halliday e al., 2018: 133), No h Ame ica (only
30 species in Beaulieu e al. (2019: 97)… This limi ed knowledge o Tydeidae may be
pe cei ed as a cogni ion s ill a a nascen s age (S epanyan and Za ikian, 2024: 217). Ano he
indica o o limi ed knowledge is he absence o ecap u e o a ypical species desc ibed long
ago: Aus alo ydaeus ki s eneae desc ibed by Spain (1969) and Lo yia supe ba desc ibed by
Oudemans (1925) bu collec ed in 1923.
Las ly, molecula da a a e indica i e o c yp ic o undesc ibed species. Beaulieu e al.
(2019: 97) ci ed only 30 ydeid mo phospecies known om Canada agains 217 species based
on molecula da a; i.e. mo phospecies would ep esen a mos 14% o all species li ing in
Canada.
Objec i i y and objec i e synonymies
No only he “ancien ” species a e neglec ed, bu also he ypes. The p inciple o ypi ica ion
s a es ha any named axon, in he amily g oup, genus g oup o species g oup, has a name-
bea ing ype which allows he applica ion o he name o he axon o be objec i ely applied. Ye ,
name-bea ing ypes a e a ely eques ed o be he e e ence objec o a necessa y edesc ip ion
o an accu a e iden i ica ion. Mos synonymies be ween gene a a e no based on he ype
ma e ial (Fig. 1). Synonymies may e en esul om synonymiza ion wi hou commen s as
done by Bake and Wha on (1952: 191) because he amily had “been di ided in he pas in o
oo many gene a based on such cha ac e s as size, wid h, leng h o se ae, ype o se ae, and
p esence o absence o eye spo s.” The absence o any explana ions is also no iceable when
Bake (1970: 163) p oposed new combina ions o species o iginally placed in Tydeus.
Wha e e he sense a ibu ed o objec i i y h ough his o y (Das on and Galison, 2007;
La ou , 2012), ypes in ICZN p o ide an objec i e e e ence o he alloca ion o a nomen o
any axon and designa e objec s (Dubois, 2011: 21).
Inadequa ely desc ibed axa
Inadequa ely desc ibed axa is ano he p oblem emphasized by Lindquis (2001) and ende ed
mo e complica ed by he absence o edesc ip ions. Ye , Bake (1965: 95) al eady w o e ha
“i will be necessa y o e-collec a ype locali ies o de e mine he s a us o mos Eu opean
species”. Mos Tydeus desc ibed by Koch om Regensbu g (Ge many) we e inadequa ely
desc ibed, ha e ne e been edesc ibed and he e is no neo ype. Tydeus b e iculus edesc ibed
by Kaźmie ski (1998a: 305) is based on Polish specimens, no on opo ypes. I s synonymy
wi h Tydeus cu us Be lese, 1910 is he e o e suspec .
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 164

The i s comple e o gano axy was p o ided by G andjean (1938), which means ha ea lie
desc ip ions a e se iously inadequa e and conce n 72 species (i.e. 17% o cu en species).
Leg chae o axies published be o e Bake ’s (1965) seminal pape we e qui e unusual and such
an absence conce ns ca 95 species (22%). Redesc ip ions a e excep ional (less han 1% o
desc ibed species) and only include ha o Tydeus i alicus by Kaźmie ski (1998a: 310–311)
and hose o T. c oceus (L.), T. c ucia us Koch and T. spa hula us Oudemans by And é (2005),
all om Oudemans’ collec ion.
Inadequa e desc ip ions and he absence o subsequen edesc ip ions also conce n impo an
species such as “Lo yia o mosa” i s collec ed on ci us in No he n A ica. The o iginal
desc ip ion did no men ion he o gano axy o legs and did no illus a e se e al se ae (P2, H1
o H2), he e icula ion pa e n is jus ske ched in ig. 15. The chae o ac ic o mulae deduced
om o iginal igu es ( igs 7-10), I(6-3-2-2-0) II(6-2-2-2-0) III(5-2-1-1-0) IV(5-2-0-1-?), do
no co espond o o mulae o Table 1. Wha does Lo yia o mosa au ho em designa e? Is i
he same species as ha desc ibed by Coo eman (1958)? Me alo yia magdalenae desc ibed
by Ge son (1968), also wi h a e icula ed pa e n, was also collec ed om ci us in he same
clima ic zone; wha a e he di e ences wi h he species desc ibed by Coo eman (1958)?
A simila p oblem a ises wi h Tydeus kochi Oudemans (1928b: 377) men ioned a species
wi h bulbous sho se ae (“kolbige ku ze Bo s e” in Ge man) while Bake (1970: 170–171, his
ig. 26) d ew a species wi h nude se ae said o be “o medium leng h, lanceola e, se a e”. Wha
does Tydeus kochi au ho em designa e?
Adequa e ( e)desc ip ions
“Ancien ” species a e o en neglec ed. I he o namen a ion (s ia ion pa e n, e icula ion,
baske wea e…) is dis ega ded a he gene ic le el, i is o pa amoun impo ance ( oge he o
he s a e, shape, leng h o se i o m o gans) o delinea e species. A help ul sys em o such was
p oposed by Kaźmie ski (1998a: 288-289). Howe e , he o namen a ion is di icul o ep esen
wi h d awings and illus a es he p oblems encoun e ed when we look back in o he pas wi h
mode n echniques and equipmen (Ra cli , 2009). In Akba i e al. (2015b), he habi us o
Tydeus shabes a iensis is illus a ed bo h wi h a d awing ( hei igu e 1) and wi h a pho og aph
( hei igu e 3). The compa ison o pho os and line d awings e ealed ha do sal body se ae
seem hicke and mo e se a ed in d awings han in pho os. Ano he example is p o ided by
Ripka e al. (2013) who desc ibed se e al new species o a bo eal Tydeidae om Hunga y.
They p o ided phase con as pho os o some species. The compa ison o pho os and line
d awings again e eals some inadequacy o line d awings. The pho o o Lo yia sanc ikingai
clea ly illus a es a he hick and blun - ipped do sal idiosomal se ae ( hei ig. 18) while line
d awings ( hei ig. 15) sugges hin and poin ed se ae. This ecalls he old deba e be ween
d awings and pho og aphs in science. “Bu he ‘pho og aphic eye’ does no app oach he
objec s only in a mo e hones and unbiased way, i is o en able o g asp hem mo e di ec ly and
mo e sha ply.” (“Abe das ‘pho og aphische Auge’ i an die Gegens ände nich nu eh liche
und o u heilslose he an, es e mag dieselben auch iel ach unmi elba genaue und schä e
zu e assen.”) ad anced F aenkel and P ei e (1892: 1). S a e al. (1896: i) added: “Bu all
such d awings a e necessa ily impe ec and in ol e a pe sonal elemen o in e p e a ion.”
Pho og aphs o he s ia ion pa e n in Tydeoidea was in oduced by And é and Duca me
(2003). In he u u e, adequa e mic og aphs mus be pa o any ( e)desc ip ions, be hey
phase con as mic og aphs (e.g. Escoba -Ga cia e al. 2023: Figs 1, 5 & 6); Khaus o and
Khaus o , 2023: Fig. 1) o scanning elec on mic oscope (SEM) mic og aphs (e.g. Khaus o
and Khaus o , 2023: Figs 4 & 5; We gin e al., 2000).
The edesc ip ion o any species, howe e p ecise i is, mus be based on mi es o he
same p o enance as he ype se ies. This migh bea se ious p oblems as emphasized abo e
wi h Lasio ydaeus. The species “Lo yia o mosa” desc ibed inadequa ely as demons a ed
abo e, o e s ano he example. The species was ne e edesc ibed om he ype se ies, no
om specimens o he same p o enance (ci us in no he n A ica). The specimens o Lo yia
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 165
o mosa desc ibed by Jeppson e al. (1975, ig. 85) a e p obably Ame ican and bea he same
binomen as he A ican species desc ibed by Coo eman (1958). Ye , hey di e on se e al
poin s: he e icula ion pa e n as well as he leng h, shape ( ounded s sha p end) and inse ion
o some se ae (P3, L1).
On ogeny and sexual polymo phism
In he pas , gene a we e e ec ed and based on misiden i ied imma u e specimens: he so-called
genus Hypopus by Dugès (1834: 20) is a amous example. In Tydeidae, a nymph in pupa ion
was illus a ed, wi hou unde s anding wha i was, by Be lese (1910: igs 5, 5a): Tydaeus
(Tydaeolus)a omus had ou ichobo h ia.
Tydeus muns e i, he species desc ibed by Meye and Ryke (1959: 413), s udied by And é
(1980: 128) and by Uecke mann and G ou (2007: 2062-2064) and designa ed as he ype
species o A o ydeus, has 6 geni als and no 4 as no ed by Bake (1970: 164) and exhibi s he
same phane o axy as he nominal subgenus Tydeus. The con usion be ween a i onymph wi h
4 geni als and he imago wi h 6 geni als migh explain he misunde s anding.
The hypo hesis sugges ed by Uecke mann and G ou — he specimen s udied by Bake was
a i onymph—is all he mo e a ac i e as he con usion imago- i onymph was also made
by him when he desc ibed he holo ype o meye ae Bake (1970: 165), he ype species o
Pe a o ydeus. The holo ype was examined by And é (1980: 142) who s a ed ha he holo ype
was no a emale bu a i onymph in pupa ion.
A p oblem also conce ns he genus Edlo yia based on a unique specimen o Bake (1968),
a deu onymph he chae o axy o which was incomple ely published.
Addi ionally, labo a o y colonies a es ed hely oky in ydeid mi es (He nandes e al. 2006;
Sil a e al. 2014) and con i med he sex a io obse ed in he ield; i is an open ques ion
whe he his end is compa able o hose obse ed in O iba ida (Hee ho e al. 2009). A las
p oblem conce ns he s ia ion pa e n which could a y be ween males and emales as shown
by Khaus o (2023), such a seconda y sexual dimo phism complica es he species iden i ica ion
and ecalls he he e omo phic males obse ed in Iolinidae by Kaźmie ski (1998b: 43).
G oup o species s complex o species
As al eady p oposed by A hias-Hen io (1975), he de elopmen o iden i ica ion ools has
no hing in common wi h sys ema ic esea ch, clea ly isible mo phological ai s a e su icien
o his app oach. Cu en ly, he genus B achy ydeus con ains 220 species and i is di icul o
disc imina e hem. G ouping ydeid species wi h simila mo phology is an old adi ion al eady
used by Koch (1850: 71), Oudemans (1929: 480), Bake (1968: 987) and Kaźmie ski (1980:
89) and migh help o make accu a e iden i ica ions.
As no iced by Zachos (2016), species names a e disc e e uni s in a con inuous wo ld wi h
uzzy bounda ies and species close o one ano he migh be g ouped in species complexes as
al eady done in E iophyidae species associa ed wi h pa icula c op sys ems (de Lillo e al.,
2018). Con a ily o species g oup whe e simila i y is a key poin , phylogene ic c i e ia a e
impo an and migh help o de ec close o c yp ic species, o ins ance hose li ing in he same
ag oecosys em such B achy ydeus o mosus collec ed on ci us. Such complexes in Tydeidae
a e e oked by Pa ke (1982: 123). As demons a ed by Halliday (2010) and Saccagi and
Uecke mann (2024), close species may be a sou ce o axonomic and nomencla u al con usion
ha po en ially complica es pes managemen and biosecu i y.
Un esol ed hypo heses
Only un esol ed hypo heses ela ed o species o ag onomic impo ance a e lis ed he ea e .
Tydeus cali o nicus (Banks) is suspec ed o be a junio synonym o T. spa hula us by
Uecke mann and G ou (2007: 2364). Simila ly, Uecke mann e al. (2019: 505) a e ques ioning
he iden i y o he Tu kish specimens o Tydeus cali o nicus (Banks) as he expanded caudal
And é H. M. (2025), Aca ologia 65(1): 149-172. h ps://doi.o g/10.24349/l5 b-767l 166
se ae (spa ula e) di e s om specimens om he USA (lanceola e). The ue iden i y o
Tu kish specimens is again ques ioned by Akyazı e al. (2024: 1039). Tydeus goe zi Sch u
was i s iden i ied as T. cauda us by Akyazı e al. (2017: 12). Tydeus cauda us auc o um
migh designa e T. goe zi, a leas in some cases (And é, 2021: 1031).
Las ly, Tydeus ibu ni Koch has ne e been eco ded since he o iginal desc ip ion in 1838.
Howe e , ibu ni was lis ed wi h Te anychus u icae by P i cha d and Bake (1955: 437).
Tydeus ibu ni migh be he old name o T. goe zi since bo h mi es a e equen on Vibu num
(And é, 2011: 77–80). The hypo hesis was ad anced by And é (2021: 1031).
Conclusions
All ydeid gene a a e now de ined by me is ic cha ac e s (chae o axy, solenidio axy…) and no
based on he o namen a ion (s ia ion pa e n, e icula ion, baske wea e…). Some ai s a e
neglec ed such as he numbe o opis hosomal se ae (9 s 10) o he epime al se ae. The new
nomencla u e is simple han in he pas ( he e we e 3 synonymies and 1 homonymy o Venilia
and 3 chae o axies o he same genus, A o ydeus) and acco ds wi h ICZN. A ype species is
ixed o all gene a. The numbe o gene a (31) and non-nominal subgene a (3) emains almos
he same as in he pas , e en a e he nomencla u al changes abo e-men ioned.
Any classi ica ion—wha e e i is—is a axonomy based on ou cu en knowledge o
mi es and no a classi ica ion o mi es hemsel es. Ou knowledge is so meage (3.72% o
mo phospecies, s ill less i c yp ic species a e conside ed) ha i allows o explo e e olu iona y
ends o some ai s bu no o de ine a eal phylogeny o species. Las ly, he classi ica ion
p oposed does no e lec he in a-popula ion a ia ions, no he a ia ions bo h wi hin and
be ween species. Species names a e disc e e uni s in a con inuous wo ld wi h uzzy bounda ies,
a o io i ha applies o genus names. Aca ology is s ill in i s in ancy…
Acknowledgemen s
I wish o hank D. Nicolson and G.D. Owe o p o iding he basic da abase as well as K.-H.
Schmid o sugges ing he amendmen . Special PDFs we e sen by A.A. Khaus o , I. Łaniecka,
R. Łaniecki, R. Ochoa. I am g a e ul o hose who ead and analyzed d a s o he ms: G.W.
K an z who supe ised in 1968 my mas e ’s hesis on Tydeidae, F.A. He nandes, A.A. Khaus o ,
R. Ochoa, A.W. Ulsame and he wo e iewe s selec ed by he jou nal.
ORCID
Hen i M. And é h ps://o cid.o g/0000-0003-4889-8983
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