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Supplementary material

Szwedo J, Drohojowska J: A swarm of whiteflies – first record of gregarious behavior from

Eocene Baltic amber

Fossil record of Aleyrodidae

If the numbers of described species are an accurate guide, then Aleyrodomorpha Chou,

1963 are by far the least speciose of the sternorrhynchous Hemiptera infraorders, with around

1,550 currently valid recent species (Ouvrard and Martin 2016). This figure may be compared

with about 8,200 Coccidomorpha Heslop-Harrison, 1952 (García et al. 2016), over 5,000

Aphidomorpha Becker-Migdisova et Aizenberg, 1962 (Favret 2016), and over 3,800 Psyl-

lodea Latreille, 1807 (Ouvrard 2016). In respect to fossils the state of knowledge of these

groups is very uneven. For the moment only fossil aphids are covered by recent review (Heie

and Wegierek 2011), fossils are also included in Aphid Species File (Favret 2016). Also fossil

scale insects are covered by ScaleNet (García et al. 2016). For the moment there are no com-

prehensive lists of fossil psyllids and their relatives and whiteflies.

The earliest fossil record of Aleyrodidae Westwood, 1840 (the sole family of the Hemi-

ptera Linné, 1758 suborder Sternorrhyncha Amyot et Serville, 1843 and infraorder Aleyrodo-

morpha Chou, 1963) comes from the Upper Jurassic (Shcherbakov 2000), little older Aleyro-

didae were found in the Middle Jurassic deposits of Daohugou Fossil Lagerstätte in north-

eastern China (unpublished data). A few other fossils are reported from the Lower Cretaceous,

Upper Cretaceous, Palaeogene and Neogene rock deposits and fossil resins (Schlee 1970, Ri-

etschel 1983, Poinar 1992, Shcherbakov 2000, Azar 2007, Schmidt et al. 2010, Drohojowska

and Szwedo 2011a, b, 2013a, b, 2015, Drohojowska et al. 2015), but only a few have been

formally described. The specimen from Miocene amber of Iquitos, Peru (Antoine et al. 2006:

13596, Fig. 3E), presumed male of Aleyrodidae, is in fact winged male of scale insect

(Coccoidea). Several fossils ascribed to Aleyrodidae are of bearing taxonomic problems (see

below).

Annotated list of fossil Aleyrodidae

Family Aleyrodidae Westwood, 1840

Supplementary material p. 1

Subfamily Aleurodicinae Quaintance et Baker, 1913

Aretsaya Drohojowska et Szwedo, 2015: 377

Type species: Aretsaya therina Drohojowska et Szwedo, 2015, by original designation

Aretsaya therina Drohojowska et Szwedo, 2015: 378, Figs. 7A-D, 8A-F, 9A-F

Lebanese amber – Lower Cretaceous, Valanginian-Aptian (135-125 Ma); Ain Dara,

Lebanon; holotype, specimen No. AD25, Azar collection, deposited in the Natural History

Museum of the Lebanese University; amber inclusion, imago, male, right fore and hind

wing destroyed, impressed thorax.

Clodionus Drohojowska et Szwedo, 2013b: 324

Type species: Clodionus fizoli Drohojowska et Szwedo, 2013b: 324, by original

designation and monotypy

Clodionus fizoli Drohojowska et Szwedo, 2013b: 7, Figs. 15-25

Oise amber – Lowermost Eocene, Ypresian (Sparnacian) (55-53 Ma); Department Oise,

France; holotype, specimen MNHN PA 5149 deposited in Museum national d’Histoire

Naturelle, Paris; amber inclusion, imago, male.

Gapenus Drohojowska et Szwedo, 2013a: 101

Type species: Gapenus rhinariatus Drohojowska et Szwedo, 2013: 101, by original desig-

nation and monotypy

Gapenus rhinariatus Drohojowska et Szwedo, 2013a: 105, Figs. 1-14

Lebanese amber – Lower Cretaceous, Valanginian-Aptian (135-125 Ma); Hammana/

Mdeirij, Lebanon; holotype, specimen No. 1568, Azar collection, deposited in the Natural

History Museum of the Lebanese University; amber inclusion, imago, male, no recogniz-

able thoracal structures.

Isaraselis Drohojowska et Szwedo, 2013b: 333

Type species: Isaraselis cladiva Drohojowska et Szwedo, 2013, by original designation

and monotypy

Isaraselis cladiva Drohojowska et Szwedo, 2013b: 336, Figs. 58-72

Szwedo J, Drohojowska J: Supplementary material p. 2


France; holotype, specimen MNHN PA 16445, Museum national d’Histoire naturelle,

Paris; amber inclusion, imago, male, thorax covered with gas bubble.

Lukotekia Drohojowska et Szwedo, 2013b: 328

Type species: Lukotekia menae Drohojowska et Szwedo, 2013, by original designation

Lukotekia menae Drohojowska et Szwedo, 2013b: 328, Figs. 26-46


France; holotype, specimen MNHN PA 2349, Museum national d’Histoire naturelle, Paris;

amber inclusion, imago, female.

Lukotekia sp. Drohojowska et Szwedo, 2013b: 332, Figs. 47-57


France; holotype, specimen MNHN PA 1671 3/4, Museum national d’Histoire naturelle,

Paris; amber inclusion, imago, probably female, poor condition.

Milqartis Drohojowska et Szwedo, 2015: 373

Type species: Milqartis azari Drohojowska et Szwedo, 2015, by original designation and

monotypy

Milqartis azari Drohojowska et Szwedo, 2015: 377, Figs. 4A-D, 5A-F, 6A-D


Mdeirij, Lebanon; holotype, specimen No. 1620 A, Azar collection, deposited in the Natu-

ral History Museum of the Lebanese University; amber inclusion, imago, male, displaced

head, partly damaged, thorax strongly damaged.

Oisedicus Drohojowska et Szwedo, 2013b: 321

Type species: Oisedicus maginus Drohojowska et Szwedo, 2013, by original designation

and monotypy

Oisedicus maginus Drohojowska et Szwedo, 2013b: 322, Figs. 1-14


France; holotype, specimen MNHN PA 2493, Museum national d’Histoire Naturelle, Paris;

amber inclusion, imago, male, mesothorax and metathorax covered with gas bubble, apical

part of abdomen and hind wing covered with gas bubbles.


Paernis Drohojowska et Szwedo, 2011b: 662

Type species: Paernis gregorius Drohojowska et Szwedo, 2011, by original designation

and monotypy

Paernis gregorius Drohojowska et Szwedo, 2011b: 663, Figs. 1-20

Baltic amber – Eocene (Lutetian-Priabonian) (47-37.7 Ma); holotype, specimen MIBUG

5418 [MAI_5418], Museum of Amber Inclusions, University of Gdańsk, Gdańsk; amber

inclusion, imago, male, partly covered with milky veins and gas bubbles.

Shapashe Drohojowska et Szwedo, 2015a: 370

Type species: Shapashe aithiopa Drohojowska et Szwedo, 2015, by original designation

and monotypy

Shapashe aithiopa Drohojowska et Szwedo, 2015: 373, Figs. 2A-D, 3A-I



History Museum of the Lebanese University; amber inclusion, imago, female, partly dam-

aged.

Rovnodicus Drohojowska et Szwedo, 2015 in Drohojowska et al. 2015: 261

Type species: Rovnodicus wojciechowskii Drohojowska et Szwedo, 2015, by original des-

ignation and monotypy

Rovnodicus wojciechowskii Drohojowska et Szwedo, 2015 in Drohojowska et al. 2015: 264

Baltic amber – Eocene (Lutetian-Priabonian) (47-37.7 Ma); holotype, specimen No. SIZK-

K-75070 [K-7050]; Klesovo deposit, Baltic (Rovno) amber, coll. Schmalhausen Institute of

Zoology, Ukrainian Academy of Sciences, Kiev; amber inclusion, imago, female, partly

covered with milky veins and gas bubbles.

Yamis Drohojowska et Szwedo, 2015: 379

Type species: Yamis libanotos Drohojowska et Szwedo, 2015, by original designation and

monotypy

Yamis libanotos Drohojowska et Szwedo, 2015: 382, Figs. 10A-D, 11A-J, 12A-D




History Museum of the Lebanese University; amber inclusion, imago, male, partly dam-

aged, fore and hind wings overlapped, ventral side covered with gas bubble.

Aleurodicinae indet.: Drohojowska et Szwedo, 2013b: 337, Figs. 73-77


France; specimen MNHN PA 17081, Museum national d’Histoire Naturelle, Paris; amber

inclusion, imago, male.

Subfamily Aleyrodinae Quaintance et Baker, 1913

Aleurochiton Tullgren, 1907: 14

Type species: Chermes aceris ovatus Geoffroy, 1762, a rejected trinominal and a synonym

of Coccus aceris Modeer, 1778, by monotypy

Aleurochiton petri Rietschel, 1983: 98, Figs. 1a, b

Kiesgrube Fr. Bauer; Pliocene, Piacenzian (3.6-2.58 Ma); Neu-Isenburg, Hessen, Germa-

ny; holotype, specimen SMF 33 030, Senckenberg Museum, Frankfurt; imprint of pupal

case, 4th instar, on Acer leaf.

Baetylus Drohojowska et Szwedo, 2011a: 181

Type species: Baetylus kahramanus Drohojowska et Szwedo, 2011, by original designation

Baetylus kahramanus Drohojowska et Szwedo, 2011a: 4, Figs. 1-22



History Museum of the Lebanese University; amber inclusion, imago, male, body with

wings.

Snotra Szwedo et Drohojowska, 2016 (this paper)

Type species Snotra christelae Szwedo et Drohojowska, 2016, by original designation and

monotypy

Snotra christelae Szwedo et Drohojowska, 2016 (this paper)

Baltic amber – Eocene (Lutetian-Priabonian) (47-37.7 Ma); holotype, female (Figs. 1, 2, 3

& 7, specimen No. 1), specimen No. CCHH 7149 Christel and Hans-Werner Hoffeins col-

lection, to be deposited in Senckenberg Deutsches Entomologisches Institute (SDEI); am-


ber inclusion, imagines, female holotype - specimen No. 1; paratypes males - specimens

Nos. 3, 7, 9, 12; paratypes females - specimens Nos. 2, 5, 6, 8, 11; paratypes, sex not to be

recognized - specimens Nos. 4, 10.

Subfamily Bernaeinae Shcherbakov, 2000: 31Bernaeidae Zherikhin, 1980: 80, nomen nudum

Bernaea Schlee, 1970: 18

Type species: Bernaea neocomica Schlee, 1970, by monotypy

Bernaea neocomica Schlee, 1970: 18, Figs. 13-26.

Lebanese amber – Lower Cretaceous, Valanginian-Aptian (135-125 Ma); Jouar Ess Sous,

Bkassine, Jezzine, Lebanon; holotype, collection Staatliches Museum für Naturkunde,

Stuttgart; amber inclusion, imago, female.

Note. In original paper Schlee (1970), no data about the locality or closer data about the

specimen acquisition place and registration number were given. The data on the locality

and amber were provided in Schlee and Dietrich (1970) and verified by Professor Dany

Azar (personal communication).

Burmoselis Shcherbakov, 2000: 33

Type species Burmoselis evelynae Shcherbakov, 2000, by original designation and mono-

typy

Burmoselis evelynae Shcherbakov, 2000: 35, Figs. 10, 11.

Burmese amber – earliest Upper Cretaceous, early Cenomanian (99.7-94.3 Ma); holotype,

specimen NHM In. 20193, Natural History Museum, London; amber inclusion, imago,

female.

Heidea Schlee, 1970: 9

Type species: Heidea cretacica Schlee, 1970, by monotypy

Heidea cretacica Schlee, 1970: 9, Figs. 2-12.

Lebanese amber, Lower Cretaceous, Barremian (130.0-125.5 Ma); Jouar Ess Sous, Bkas-

sine, Jezzine, Lebanon; holotype, collection Staatliches Museum für Naturkunde, Stuttgart,

Germany; amber inclusion, imago, male, venation not visible.


Note. In original paper Schlee (1970), no data about the locality or closer data about the

specimen acquisition place and registration number were given. The data on the locality

and amber were provided in Schlee and Dietrich (1970) and verified by Professor Dany

Azar (personal communication).

Juleyrodes Shcherbakov, 2000: 32

Type species: Juleyrodes gilli Shcherbakov, 2000, by original designation

Juleyrodes gilli Shcherbakov, 2000: 32, Figs. 2-5.

Hotont (Khotont) locality, outcrop 354/7 – Lower Cretaceous, early Aptian (125.5-122.5

Ma); Hotont Formation, Ara-Hangayn aymag, 6 km W of Hotont somon, northern part of

Uhaa Mt., Mongolia; holotype, specimen: PIN 4307/231, Paleontological Institute, Russian

Academy of Sciences, Moscow, an imprint of the forewing.

Note. Palaeoentomological papers often place the locality close to Jurassic-Cretaceous

boundary (Tithonian-Berriasian), but if it is correlative with Baissa and Turga it is likely

early Aptian (possibly Barremian).

Hutel-Hara (Khutel-Khara) locality, outcrop 300 – Lower Cretaceous, Berriasian (145-

139.8 Ma); Lower Tsagaan Tsav Formation, East-Gobi aymag, 70 km SW of Saynshand

somon, eastern Hara-Hutul Range, Mongolia; specimen PIN 3965/445, Paleontological

Institute, Russian Academy of Sciences, Moscow, left forewing (part and counterpart; an-

terodistal wing segment folded back).

Note. Hutel-Hara outcrop is dated presumably to earliest Cretaceous and yields more than

3000 fossil insects, that reveal Cretaceous rather than Jurassic affinities (Rasnitsyn and

Zherikhin 2002)

Juleyrodes visnyai Shcherbakov, 2000: 33, Figs. 6, 7.

Mikhailovka – Upper Jurassic, Callovian (164.7-155.7 Ma), Karabastau Formation,

Karatau Range, Kazakhstan; Type specimen: PIN 2997/3837, Paleontological Institute,

Russian Academy of Sciences, Moscow; imprint, imago, body with wings.

Juleyrodes sp.: Shcherbakov 2000: 33, Fig. 8.

Mikhailovka – Upper Jurassic, Callovian (164.7-155.7 Ma), Karabastau Formation,

Karatau Range, Kazakhstan; Specimen: PIN 2997/5071, Paleontological Institute, Russian

Academy of Sciences, Moscow; imprint, imago, male (?), body with wings.


Juleyrodes sp.: Shcherbakov 2000: 34

Purbeck – Lower Cretaceous, Berriasian-Barremian (145-125 Ma); middle Purbeck, Eng-

land; imprint, forewing.

Bernaeinae gen. indet.: Shcherbakov 2000: 34, Fig. 9

Mikhailovka, Upper Jurassic, Callovian (164.7-155.7 Ma); Karabastau Formation, Karatau

Range, Kazakhstan; Specimen: PIN 2239/532, Paleontological Institute, Russian Academy

of Sciences, Moscow; imprint, imago, male (?), body with wings.

?Bernaeid incertae sedis: Whalley and Jarzembowski 1985: 394, Figs. 12A-D, 13

Sierra del Montsec – Lower Cretaceous, early Barremian (130-125.5 Ma); La Pedrera de

Rubies Formation, Lérida, Spain; Specimen In. 60600 collection W. Ball and F.M. Wonna-

cott, Natural History Museum, London; imprint of puparium.

Note. The specimen was only provisionally placed in Bernaeinae, based on opinion, that

the Mesozoic fossil whiteflies known to these Authors represent this subfamily. The speci-

men was subsequently listed by Shcherbakov (2000: 35), but its taxonomic placement was

not commented and remains unclear.

Subfamily Udamoselinae Enderlein, 1909

unidentified Udamoselinae: Shcherbakov 2000: 34

Lebanese amber – Lower Cretaceous, Valanginian-Aptian (135-125 Ma).

Note. The taxonomic status of Udamoselinae as distinct subfamily is debatable (Martin

2007).

Undetermined Aleyrodidae

Aleyrodidae indet.: Drohojowska and Szwedo 2013b: 340, Fig. 78


France; Specimen MNHN PA 15401, Museum national d’Histoire Naturelle, Paris; amber

inclusion, imago, forewing and hind wing preserved.


Aleyrodidae indet.: Drohojowska and Szwedo 2013b: 340, Fig. 79.

Oise amber – Lowermost Eocene (55-53 Ma), Ypresian (Sparnacian); Department Oise,

France; MNHN PA 1297, Museum national d’Histoire Naturelle, Paris; amber inclusion,

imago, body deteriorated, fore and hind wing preserved.



France; MNHN PA 2287, Museum national d’Histoire Naturelle, Paris; “ghost inclusion”,

imago, venation of wings, body barely visible; microscopic slide, preserved in Canada bal-

sam.



France; MNHN PA 2474 2/3, Museum national d’Histoire Naturelle, Paris; amber inclu-

sion, imago, male, body poorly preserved; microscopic slide, preserved in Canada balsam.




imago, female, quite well preserved, presentation excluding closer identification.




imago, imago, weakly visible in amber.



France; MNHN PA 420, Museum national d’Histoire Naturelle, Paris; cast of body, imago,

female, imprint on surface of amber, body features not preserved.





imago, female, details weakly visible, wings, poorly preserved.




imago, male, weakly visible, ventral view.

Aleyrodidae indet.: Drohojowska and Szwedo 2013b: 340.



imago, specimen poorly preserved.

Aleyrodidae indet: Jarzembowski and Coram 1997: Fig. 6

Poxwell – Lower Cretaceous, Berriasian-Barremian (145-125 Ma); lower Purbeck, Eng-

land; specimen MNEMG 1996.299 Jarzembowski collection, Maidstone Museum, Maid-

stone; imprint of puparium.


Durlston Bay – Lower Cretaceous, Berriasian-Barremian (145-125 Ma); middle Purbeck,

England; specimen MNEMG 1996.300 Coram collection, Maidstone Museum, Maidstone;

imprint of puparium.


Auclaye Brickworks – Lower Cretaceous, Berriasian-Barremian (145-125 Ma); Upper

Weald Clay, England; specimen MNEMG 1996.3001 Goodman collection, Maidstone Mu-

seum, Maidstone; imprint of puparium.

Aleyrodidae indet: Jarzembowski and Ross 1994: 218, Fig. 2

Bembridge Marls – latest Eocene, Priabonian (~32.4 Ma); Bouldnor Formation, Insect

Limestone, Isle of Wight, UK; specimen from A. Mitchell collection, Maidstone Museum,

Maidstone; imprint of puparium.


Note. This specimen was also mentioned by Shcherbakov (2000: 35)

Aleyrodidae indet: Rasnitsyn and Ross 2000: 22

Burmese amber – earliest Upper Cretaceous, early Cenomanian (99.7-94.3 Ma); specimen

In. 20703, Natural History Museum, London.

Aleyrodidae indet: Schlee 1970: 34

Denmark, Baltic amber – Eocene (Lutetian-Priabonian) (47-37.7 Ma)

Note. Schlee (1970) listed 13 specimens from the collection of the Zoological Museum of

the Copenhagen University, giving some details about their morphometrics and features,

but no conclusive descriptions. This collection contains 5 males, 4 females and 4 speci-

mens with sex not recognized, collected by various collectors at Danish seashore. The ma-

terial is under survey at the moment.

Taxa incertae sedis and nomina nuda

Aleurochiton Tullgren, 1907: 14

Type species: Chermes aceris ovatus Geoffroy, 1762, a rejected trinominal and a synonym

of Coccus aceris Modeer, 1778, by monotypy

‘Aleurochiton’ eozaenicus Weigelt, 1940: 347 – nomen nudumAleurochiton’ eozaenicus Weigelt, 1940: Drohojowska et al. 2015: – nomen nudum

Geiseltal – Middle Eocene (48-41 Ma), (Lutetian); Geiseltal Fossillagerstätte, Sachsen-

Anhalt, Germany; imprint, puparium.

Note. Weigelt (1940) mentioned a finding of puparium on the leaf surface, from the Middle

Eocene Fossil Lagerstätte Geiseltal in Germany. Unfortunately, he gave no diagnosis or

other features allowing recognition of the identity of this fossil, but used the name for it -

Aleurochiton eozaenicus. This name was recognized as nomen nudum by Drohojowska et

al. (2015).

Aleurodicus Douglas in Morgan, 1892: 32

Type species: Aleurodicus anonae Morgan, 1892: 32 (considered a synonym of Aleyrodes

cocois Curtis, 1846: 284), by subsequent designation by Quaintance 1908: 8.

‘Aleurodicus’ burmiticus Cockerell, 1919: 241, Fig. 1


Burmese amber– earliest Upper Cretaceous, early Cenomanian (99.7-94.3 Ma); holotype,

specimen In. 19134 (In. 19133-4), R.J.C. Swinhoe collection, Natural History Museum,

London; inclusion in amber, imago, male.

Note. Cockerell (1919) did not make any subfamily placement of this fossil. Schlee (1970)

argued placement of this fossil to subfamily Aleurodicinae (treated as separate family

Aleurodicidae in his paper). This opinion was challenged by Shcherbakov (2000), who

stated that hindwing venation and genitalia confirm the subfamily placement in Aleurodici-

nae, but generic assignment of this species remains doubtful. Evans (2008), listing the

Aleurodicus species, noted that Martin (2008) transferred the species native to Oriental or

Australasian Region, previously placed in the Aleurodicus Douglas, 1892, to the genera

Palaealeurodicus (erroneously spelled Paleoaleurodicus [sic!]) and Aleuroctarthrus Mar-

tin, 2008, and that the fossil may belong to one of these two genera.

Aleyrodes Latreille, 1796: 93

Type species: Phalaena (Tinea) proletella Linnaeus, 1758: 537, by subsequent designation

and monotypy

‘Aleyrodes’ aculeatus Menge, 1856: 18

Baltic amber – Eocene (Lutetian-Priabonian) (47-37.7 Ma); imago; specimen (female?)

probably lost.

Note. The original description is extremely poor, without any figure, and diagnosed as:

‘similar to living A. chelidoni Latr., except for the tip of abdomen being acuminate and

with two small pointed processes, which are found in both sexes by Burmeister (Entomol.

II: 82)’. Aleyrodes chelidonii Latreille, 1807 is synonym of Aleyrodes proletella (Linnaeus,

1758), but the characters given by Menge (1856) do not allow to make any generic assign-

ment of the fossil. Evans (2007, 2008) listed this species in the genus Aleurodicus Douglas,

1892, but gave no reasons for such transfer.

‘Minutabythus’: Kaddumi 2005a: 107

Type species: ‘Minutabythus jordanicus’: Kaddumi 2005, by monotypy

‘Minutabythus jordanicus’: Kaddumi 2005a: 107, Figs. 90-92Minutabythus jordanicus: Kaddumi 2005a: 107, 109

Minotabythus jordanicus: Kaddumi 2005a: 107, 109

Minutabythus jordanicus: Kaddumi 2005b: 146, 148, Figs. 146-148

Minotabythus jordanicus: Kaddumi 2005b: 146, 148


Minutabythus jordanicus Kaddumi, 2005: Kaddumi 2007: 192, Figs. 146-149

Minotabythus jordanicus: Kaddumi 2005: Kaddumi 2007: 192, 194

Jordanian amber – Lower Cretaceous (Albian) (112-99.7 Ma), Kurnub sandstone forma-

tion, Zarqa river basin, Jordan, specimens from Hani Faig Kaddumi collection, Eternal

River Museum of Natural History, Jordan.

Note. Kaddumi (2005a: 107) firstly named the species Minutabythus jordanicus (generic

name spelled also ‘Minotabythus’) and placed it in the newly established by him family

Alerollidae (spelled also as Allerollidae). Family name is not based on the generic name,

and must be regarded as invalid according to the rules of the International Code of Zoolog-

ical Nomenclature Article 11.7., Article 29.1 and Article 29.3. (ICZN 1999). The names

“Alerollidae” and “Allerollidae” do not follow these rules and must be regarded as nomina

nuda (Drohojowska and Szwedo 2011a). The invalid names “Alerollidae” and “Alleroli-

dae” appeared also in the second (Kaddumi 2005b) and third (Kaddumi 2007) editions of

the book. This invalid family name in form ‘Alerollidae Kaddumi, 2005’ is also listed in

Palaeobiology Database (PaleoBioDB 2016). The generic name Minutabythus appeared for

the first time in Kaddumi 2005a, p. 107, with a species Minutabythus jordanicus. This

species should be regarded as type species because of monotypy, according to the Article

63.3. of the Code (ICZN 1999). Kaddumi (2005a, b) did not designate any specimen as a

type, however probably he designated the type specimen ERMNH 188-3 (Kaddumi 2007,

fide PaleoBioDB 2016). This action resulted in establishment of the name and date of its

creation as Minutabythus jordanicus Kaddumi, 2007, making hence the earlier references

nomina nuda. However, the specimen is deposited in private collection (it is not following

Recommendation 16C and Recommendation 72F; ICZN 1999), any of the editions of Kad-

dumi’s work (2005a, 2005b, 2007), which were published privately is not available in rec-

ognized and indicated publicly accessible libraries (see Article 8 and Recommendation 8C;

ICZN 1999); the books are not provided with ISBN number, indexed in WorldCat, the

work and taxa indexed in Zoological Record or ZooBank. All these put the value and valid-

ity of the name ‘Minutabythus jordanicus’ in strong doubts and should result in treatment

of name ‘Minutabythus jordanicus’ as nomen nudum rather than as a valid name.

The specimen figured (Kaddumi 2005a, b, 2007) clearly represents Aleyrodidae, however

on the basis of available figures (drawing and photos) as well as description given it is hard

to decide to which subfamily it could belong. The differential diagnosis of this fossil is

missing and description is giving no data on important to taxonomy of Aleyrodidae fea-


tures, which makes the closer identification and comparison of this fossil with contempora-

neous fossils from Lebanese amber impossible. Forewing venation suggests relationships

with Aleurodicinae or Udamoselinae, while very simple venation of hind wing, with a sole

vein relates it rather with Aleyrodinae (Drohojowska and Szwedo 2013a). No word is

given about wax plates, their presence or absence, as well as no data on paronychia or

other tarsal structures. The 9-segmented antennae seems to be plesiomorphic condition as

number of antennal segments is up to 11 in Bernaeinae, while in recent Aleyrodinae the

number of antennomeres could vary from 4 to 7, but among the fossils the taxa with 10

antennomeres are to be found, in recent Aleurodicinae it is variable from 4 to 7, but up to

10 antennomeres among fossil taxa, in Udamoselinae antennae consist on 7 antennomeres.

The male genital structures make this fossil resembling Bernaeinae or Aleurodicinae, but

no conclusive characters are available from the original paper.

Baltic amber, its age, origin, floristic composition of Baltic amber forests and whiteflies

diversification

The longstanding debates on Baltic amber age and origin, origin of its deposits in various

regions, are unfinished for number of reasons. The biggest concentration of amber in the de-

posit, in the Gulf of Gdańsk (Sambian Peninsula to Chłapowo), is definitively secondary one;

the other deposits, i.e. Górka Lubartowska amber (Lublin Region, Poland), Ukrainian amber

(deposits near Klesov, Rovno and Zhitomir; Bogdasarov 2010, Perkovsky et al. 2010), Bitter-

feld amber (Rascher et al. 2008, Wolfe et al. 2016) are variously aged and secondary as well.

The same type of resin as Baltic amber was found also as far North as Spitsbergen and as far

North-West as Axel Heiberg Island in the Canadian Arctic (Azar et al. 2011; Wolfe et al.

2009) and aged as older than Gulf of Gdańsk deposits. The origination of the Baltic amber

forest is often coincided with the Middle Eocene Climatic Optimum, but this opinion is not

universally accepted. Baltic amber was aged within the range of 37–47 Ma (Ritzkowski 1997;

Perkovsky et al. 2007; Bogdasarov 2010). Absolute dating analyses of glauconites from Sam-

bia Peninsula showed that the “blue earth” formation (amber bearing Prussian Formation) is

allocated to the Middle Eocene (Lutetian: 44.1 ± 1.1 Ma) and is thus significantly older than

previously assumed (Wappler 2003, 2005). Limnic sediments of Eckfeld Maar, aged 44.3 ±

0.4 Ma, correlate with K-Ar radiometric data from the Sambia Peninsula and contain insect

genera known only from Baltic amber (Wappler 2005). Weitschat and Wichard (2010) men-


tioned that two additional amber-bearing horizons in the underlying beds of “blue earth” in-

dicate that amber had already been transported to secondary deposits about 50 million years

ago. Assumptions on the Middle Eocene age of Baltic amber was argued by Perkovsky et al.

(2007), and the Upper Eocene (Bartonian/Priabonian: 37.7 ± 3 Ma) age of Prussian Formation

was suggested by these authors. The age, similarities and dissimilarities of inclusions from

various deposits of Baltic amber (Bitterfeld amber, amber form Gulf of Gdańsk area, amber

from Ukraine) were discussed by Perkovsky et al. (2007), Szwedo and Sontag (2013), phys-

ical and chemical properties of amber from various deposits were discussed by Matuszewska

(2010), Sodhi et al. (2012) and Wolfe et al. (2016). It is obvious that after nearly 250 years of

scientific investigations on amber there are still numerous areas clouded and questions un-

answered.

Several types of the forests were present on the Earth during the Eocene epoch (Willis and

McElwain 2002, Kvaček 2010), and these forests were probably denser than those of the

Cretaceous, because of absence of large grazing animals (Gould 1993). Amber producing

forests covered vast areas, and were differentiated due to geographical, topographical, clima-

tological and hydrological factors, changing during the times of their presence (at least 10

million years), variable in plant composition at various scales – this diversity is reflected to

some extent also in amber preserved in the deposits and its inclusions. Reconstruction of “am-

ber forests” environments was presented by Kohlman-Adamska (2001), based on palaeo-

botanical data. Over 750 species of plants have been described from the Baltic amber (Selden

and Nudds 2004), morphological research has allowed to identify 250 species of spore-bear-

ing, herbaceous and arborescent plants from the amber producing forests (Pielińska 2008).

The forests of Scandinavian Peninsula were formed by various trees: firs Abies Miller, spruces

Picea A. Dietrich, larches Larix Miller and numerous representatives of Cupressaceae: Cali-

fornian cedar Calocedrus decurrens Torrey (Florin), Thujopsis Siebold et Zuccarini ex End-

licher, Chamaecyparis Spach and Thuja Linné; umbrella-pines Sciadopitys Siebold et Zuccar-

ini occurred in the mountain forests of the higher positions. Open forest-steppes of the lower

parts of the mountains were created primarily by various species of pines Pinus Linné, but

also many species of oaks Quercus Linné, both shedding leaves and wintergreen ones, as well

as palms Palmaceae. Other trees were: beeches Fagus Linné, chestnuts Castanea Miller,

maples Acer Linné, cycads of the genus Zamia Linné, shrubs such as magnolias Magnoli-

aceae), laurels (Lauraceae); the undergrowth was composed mainly by grasses (Poales). Wa-

terlogged riverside forests were composed inter alia by Glyptostrobus pensilis (Staunton ex D.


Don) K. Koch, shrubs of willow (Salicaceae), Myricaceae, Clethraceae and herbaceous plants

of the spiderwort family Commelinaceae. The forests of the southern banks of the Eocene

epicontinental Paratethys Sea which have been exuding resin transformed into amber depos-

ited in Ukraine and eastern Poland, represents notophyllous evergreen forests (Kvaček 2010).

The amber forest of the Ukraine is considered as representing flora of more xeric environ-

ments compared to that of the Baltic amber forest. Amber producing tree(s) was the same as

for the Baltic amber (Perkovsky et al. 2010), the flora as mixture of tropical and subtropical

plants with some temperate zone floristic elements, including laurels Lauraceae, myrts Myrta-

ceae, proteas Proteaceae and mulberrys Moraceae. The Bitterfeld amber forest seems to

present more questions than answers (Rascher et al. 2008, Wolfe et al. 2016). The amber was

discovered in the deposits assigned a Miocene age (Barthel and Hetzer 1982), more recent

geochronological efforts (Knuth et al. 2002) place these sediments in the late Oligocene

(Chattian; 23.0–28.1 Ma). Some authors interpret it as redeposited Baltic amber (Weitschat

1997, Perkovsky et al. 2007), others treat it as a separate type of amber, of a considerably dif-

ferent, Oligocene age (Knuth et al. 2002, Fuhrmann 2005). The most recent opinion formu-

lated by Wolfe et al. (2016), based on geochemical analyses of composition and amber prop-

erties, consider amber from Gulf of Gdańsk and from Bitterfeld broadly contemporaneous,

but originating from different sectors of the Eocene palaeo-North Sea Basin (north-western

Paratethys) drainage system: Baltic amber from the north and Bitterfeld from the south. Very

little is known about the floristic composition of Bitterfeld amber producing forests, but it

seems to represent notophyllous evergreen type of forest (Kvaček 2010). Very far in the north

in the polar deciduous to mixed mesophytic amber forest were characterized by a dominant

broad-leaved deciduous component, usually with a large size of the leaf lamina, with rare

maidenhair Ginkgo Linné, mesophytic conifers Amentotaxus Pilger and extinct cupressoids

(Utescher and Mosbrugger 2007, Kvaček 2010). The Canadian Arctic amber forests were a

mosaic of taxodiaceous swamp (dawn redwood Metasequoia Hu et Cheng dominant with or

without swamp cypress Glyptostrobus Endlicher), a mixed coniferous community, and alder

Alnus Linné fern bogs (Greenwood and Basinger 1994, Jahren 2007). There are also convin-

cing faunistic similarities between taphocoenoses preserved as inclusions in Baltic amber

from the Gulf of Gdańsk, Ukraine, Bitterfeld as well as this from Belarussiya and Lithuania,

and it seems more likely that the differences observed result from geographic and habitat dif-

ferences (Dlussky and Perkovsky 2002, Dlussky and Rasnitsyn 2009, Sontag and Szadziewski

2011, Szwedo and Sontag 2013).


The mother plant of Baltic amber is still under debate, and various opinions were presented

(Szwedo 2010). It was thought since the paper of Göppert (1836) that the resin that became

amber was produced by the tree with formal, scientific name Pinites succinifer Göppert, 1836,

later (Conwentz 1890) emended to Pinus succinifera (Göppert, 1836) Conwentz, 1890. This

name denotes collectively amber-producing trees, mainly pines. More recently, it has been

proposed that conifers of the family Sciadopityaceae were responsible (Wolfe et al. 2010,

Sadowski et al. 2015), but the Pinaceae are not to be excluded (Dolezych et al. 2011).

The Aleyrodidae occurred in the Eocene Baltic amber producing forests, preserved as inclu-

sions are represented nowadays by four taxa (Menge 1856, Drohojowska and Szwedo 2011a,

Drohojowska et al. 2015, Szwedo and Drohojowska - this paper). The taxonomic status of

‘Aleyrodes’ aculeatus Menge, 1856 remains unclear, as type material seems to be lost. Two

taxa represent subfamily Aleurodicinae – Paernis gregorius Drohojowska et Szwedo, 2011,

from the Gdańsk Bay deposits and Rovnodicus wojciechowskii Drohojowska et Szwedo, 2015

from deposits of amber in Ukraine, Klesovo. The taxon described above, Snotra christelae sp.

n. is the first unambiguous representative of subfamily Aleyrodinae among inclusions of

Baltic amber.

The Aleyrodidae use plant sap as their diet and most of them are believed to be closely re-

lated with particular host-plants (Gerling 1990; Grimaldi and Engel 2005). However, the asso-

ciations of modern whiteflies and their host plants are still not well known and understood

(Dubey and Ko 2006, Manzari and Quicke 2006). Most of Aleyrodidae utilize angiosperms as

the host plants, so it is likely that their relatively rich record from the Cretaceous, reflects the

evolutionary transformation of biota and available host plants of these times (Drohojowska

and Szwedo 2015). The diversification of the whiteflies during the Cretaceous is well evi-

denced. Both recent subfamilies, Aleyrodinae and Aleurodicinae are documented from the

various early and late Cretaceous deposits, and concurrently, the declining Jurassic and Creta-

ceous Bernaeinae were still present. The angiosperm divergences before the Barremian (be-

fore 130 Ma) were sporadic, but they became both numerous and frequent in the late Creta-

ceous times and tapering off by the early Palaeogene (Ruban 2012, Magallón et al. 2015), and

the species richness may have come much later (Sanderson 2015). The fossils of whiteflies

allowed postulate that the periods of Palaeocene-Eocene Thermal Maximum (PETM) and

Eocene Thermal Maximum (ETM2) triggered diversification of these insects (Drohojowska

and Szwedo 2015), and this ascertainment match well to the observed pattern of angiosperms

diversification. Most of the recent Aleyrodidae utilize as host plants two advanced according


to recent system of plant classification groups of angiosperms, i.e. asterids and rosids (Stevens

2008, APG III 2009). The asterids lineage diversified during the Palaeogene, while rosids

during the late Cretaceous (Stevens 2008, Magallón et al. 2015). Representatives of both plant

lineages are reported from taphocoenoses of Baltic amber from various outcrops, thus, these

plants could serve as suitable food source for the Aleyrodidae during the Eocene, and evolu-

tionary traits of these insects are to be correlated with evolutionary changes of their host

plants. We can assume that the early, Cretaceous separation of the whiteflies subfamilies,

fruited in the rapid diversification of both lineages in concordance with evolution and diversi-

fication of host plants available. The subfamily Aleurodicinae is highly diversified with 13

tribes recognized (Manzari and Quicke 2006), alas such analysis for Aleyrodinae is not avail-

able. This diversity seems to result at least partly, from the long evolutionary history of the

group. For the moment, too little is known about the diversity of Baltic amber Aleyrodidae,

and presenting strong conclusions seem to be premature. The known fossils clearly differ

from the recent representatives of the family, still presenting some characters recognized as

plesiomorphic. On the other hand these inclusions witnessed major global and local changes,

climatic conditions and types of habitats no longer existing in the modern world, giving the

unique opportunity to peep ancient biotas existing over 40 million years ago.

An open question is, if the resin mother plant was or could be a host plant for Snotra chris-

telae sp. n. and this could be the reason for aggregate preservation in amber. The great major-

ity of whiteflies is not highly host-specific, but colonize only dicotyledonous angiosperms,

mainly woody shrubs and trees, and a smaller, but significant part, feed on monocots, particu-

larly grasses and palms (Mound and Halsey 1978; Carver and Reid 1996). The oldest known

whiteflies are reported from the Late Jurassic (Shcherbakov 2000), they were differentiated

into subfamilies during the Lower Cretaceous, before drastic vegetational changes in mid-

Cretaceous times, during which the gymnosperm- and fern-dominated flora were replaced by

angiosperm-dominated communities (Drohojowska and Szwedo 2011a, 2015). Then, their

early evolution must to be related with gymnosperm forests, which diversified during the Late

Jurassic and Early Cretaceous (Rees et al. 2000; Taylor et al. 2009), or with pro-angiosperm

plants of these times (Liu and Wang 2015). Whiteflies diversification after the early Creta-

ceous times was probably strongly influenced by the events of the period of intense faunistic

turnover of insect assemblages, the “mid-Cretaceous biocoenotic crisis” (Rasnitsyn 1988;

Zherikhin 1978, 1993, 2002; Krassilov 2003). It was endogenous community crisis evoked by

competitive replacements in the early successional vegetation (Zherikhin 1993, 2002). The


periods of Palaeocene-Eocene Thermal Maximum and Eocene Thermal Maximum were prob-

able triggers for diversification and origination of modern Aleyrodidae fauna (Drohojowska

and Szwedo 2015), which is also suggested by molecular data (Campbell et al. 1994, Boykin

et al. 2013). The periods of Palaeocene-Eocene Thermal Maximum and Eocene Thermal

Maximum were probable triggers for diversification and origination of modern Aleyrodidae

fauna (Drohojowska and Szwedo 2015), which is also suggested by molecular data (Campbell

et al. 1994, Boykin et al. 2013). It cannot be excluded that Snotra christelae sp. n. is the rem-

nant of ancient gymnosperm-related Aleyrodinae, thus it was caught into amber during the

mating flight near its host-plant (amber producing tree). Numerous gymnosperms and an-

giosperms co-occurred in the Eocene Baltic amber forest (Czeczott 1961), and it cannot be

excluded, that Snotra christelae sp. n. as well as other Aleyrodidae reported from amber were

trophically related to angiosperm host plants, richly present in the differentiated Eocene Baltic

amber forests. In such case the swarm entombing of newly described above taxon in amber

should be regarded as accidental.

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