New non-biting midges (Diptera: Chironomidae) from Lower Cretaceous Wealden amber of the Isle of Wight (UK)

Accepted Manuscript New NON-BITING midges (Diptera: CHIRONOMIDAE)FROM lower Cretaceous Wealden amberof the isle of Wight (UK) Viktor Baranov, Wojciech Giłka, Marta Zakrzewska, Edmund Jarzembowski PII:

S0195-6671(18)30368-9

DOI:

https://doi.org/10.1016/j.cretres.2018.11.012

Reference:

YCRES 4015

To appear in:

Cretaceous Research

Received Date: 10 September 2018 Revised Date:

2 November 2018

Accepted Date: 18 November 2018

Please cite this article as: Baranov, V., Giłka, W., Zakrzewska, M., Jarzembowski, E., New NON-BITING midges (Diptera: CHIRONOMIDAE)FROM lower Cretaceous Wealden amberof the isle of Wight (UK), Cretaceous Research, https://doi.org/10.1016/j.cretres.2018.11.012. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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NEW NON-BITING MIDGES (DIPTERA: CHIRONOMIDAE) FROM LOWER CRETACEOUS WEALDEN AMBER

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OF THE ISLE OF WIGHT (UK)

Viktor Baranov1*, Wojciech Giłka2, Marta Zakrzewska2*, Edmund Jarzembowski3,4 1

LMU Munich Biocenter, Department of Biology II Großhaderner Str. 2, 82152 Planegg-Martinsried,

Germany

University of Gdańsk, Faculty of Biology, Department of Invertebrate Zoology and Parasitology,

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2

Laboratory of Systematic Zoology; Wita Stwosza 59, 80-308 Gdańsk, Poland 3

State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and

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Palaeontology and Center for Excellence in Life and Paleoenvironment, Chinese Academy of Sciences, Nanjing 210008, China 4

Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK

*corresponding authors: [email protected], [email protected]

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ABSTRACT. Non-biting midges (Chironomidae) from Lower Cretaceous Wealden amber of the Isle of Wight (lower Barremian, ca. 128 Ma) are reviewed. As a result, Dungeyella gavini Jarzembowski, Azar et Nel, 2008, the only chironomid species

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known from this amber deposit, is for the first time recognised from the adult male, and the systematic position of Dungeyella within the subfamily Buchonomyiinae is

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established. Libanodiamesa simpsoni sp. nov. (Prodiamesinae), now found in Wealden amber, is the second species of the genus previously only recorded from Lower Cretaceous Lebanese amber. A detailed morphological analysis revealed characters (wing venation patterns, genital apparatus structure) defined as unique for the two genera, the diagnoses of which are amended. Biogeographical features of the Wealden amber Chironomidae are also discussed against the background of their fossil records from the Cretaceous.

ACCEPTED MANUSCRIPT KEY WORDS: Chironomidae, Mesozoic, amber, systematics, biogeography

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1. INTRODUCTION

Non-biting midges (Chironomidae) are among the most abundant and diverse aquatic insects, with over 7000 described extant species worldwide (Ferrington, 2007, Pape et

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al., 2011). The geological history of non-biting midges stretches from the Late Triassic onwards, with the oldest known chironomid being Aenne triassica Krzemiński et

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Jarzembowski, 1999. Since chironomid larvae in different genera are closely associated with specific types of habitat and environmental conditions, the family is often used in environmental monitoring and paleoenvironmental reconstruction. Seredszus and Wichard (2007) and Grund (2006) have pioneered the approach of paleohabitat

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reconstruction based on the generic composition of chironomid species complexes in amber. This approach works best when extant genera are present in the fauna, so it is practically limited to the Cenozoic; however, some paleoenvironmental information can

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be inferred from extinct taxa as well (Azar et al., 2008, Jarzembowski et al., 2008, Baranov et al., 2017). Thus studies of fossil chironomid faunas are not only providing

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us with new taxonomic and phylogenetic information, but also allow an insight into the palaeoenvironment in which the insects lived. Adult chironomids were amongst the first inclusions found in Cretaceous amber

in the UK (Jarzembowski, 1995). These were from Wealden amber ('chiltonchineite') in the Wessex Formation exposed by marine erosion on the southwest coast of the Isle of Wight (IoW) in southern England and are some 128 million years old (Jarzembowski, 1999). Unlike in the Wealden rock fauna, chironomids dominated the small amber

ACCEPTED MANUSCRIPT assemblage considered to have been preserved in a climatically controlled debris deposit (Jarzembowski et al., 2008). The first chironomid inclusion to be described from this bed, Dungeyella gavini Jarzembowski, Azar et Nel, 2008, was a female clearly belonging to an extinct fauna unlike Cenozoic chironomids.

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In this paper, using subsequently found inclusions in the Wealden amber of the IoW, we describe a new species of the subfamily Prodiamesinae as well as elucidate the taxonomic position of the genus Dungeyella Jarzembowski, Azar et Nel, 2008, the only

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chironomid so far described from 'chiltonchineite'.

2. MATERIALS AND METHODS

Geological context

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2.1.

The amber bed is actually a plant-debris horizon (Lignite 6, Fig. 1A) exposed by the sea on the Channel coast of the IoW near a small valley called Chilton Chine (Sweetman,

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2011: text-Fig. 4.17, Fig.1B, C). The amber occurs occasionally in the mass of plant debris considered to have been deposited from sheet flooding on a river floodplain

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following seasonal wildfire and rainstorms on wooded higher ground (loc. cit.: text-Fig. 4.9). Arthropod inclusions are uncommon, but at least four orders of insects and spiders have been found including other Diptera, although chironomids comprise nearly half of the total specimens (Jarzembowski et al., 2008). The amber is of early Barremian age and generally considered to be of extinct coniferous origin (Bray and Anderson, 2008).

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Material collection and preparation

The material was collected in June 2001 and is deposited in the Natural History Museum (NHM), London, UK. The amber specimens were prepared as described in

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Jarzembowski et al. (2008). All photographs were taken using an Euromex Oxion light microscope and attached digital camera. Drawings were produced by line-tracing photos in the Inkscape software (Burington, 2017). New nomenclatural acts and the paper were in

Zoobank,

in

accordance

with

ICZN

requirements

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registered

(LSID:urn:lsid:zoobank.org:pub:1A707206-B6DF-439F-820E-10CFC2567D61).

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Measurements of specimens are in micrometres, except for the total body length and wing length (in millimetres, rounded off to the first decimal place). The body length was measured from the antennal pedicel to the end of the gonostylus/cercus, and the wing from the arculus to the tip. Measurements were rounded off to the nearest 10 µm (legs

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segments) and 1 mm (flagellomeres and palpomeres). The antennal and leg ratio (AR, LR) were calculated to the second digit after the decimal point. According to Sæther (1980), morphological terminology and abbreviations are as follows: R1, R2+3, R4+5 -

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radial veins, MCu - medial-cubital crossvein; RM - radial-medial crossvein; Sc subcostal vein; Pm - maxillary palp, pm1-5 - palpomeres 1-5, fl1-14 - antennal

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flagellomeres 1-14; Dc - dorsocentral setae; Pa - prealar setae; Scts - scutellar setae; p1p3 legs of fore- through hind pair; fe - femur, ti - tibia, ta1-5 - tarsomeres 1-5; AR antennal ratio.

3. RESULTS

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Systematic palaeontology

Diptera Linnaeus, 1758 Chironomidae Newman, 1834

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Buchonomyiinae Brundin et Sæther, 1978 Dungeyella Jarzembowski, Azar et Nel, 2008

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Diagnosis (redefined). Adults (both sexes). MCu vein present, R1 and R4+5 very short, terminating at 0.65 of wing length from base, R2+3 absent, Sc well developed, setae on

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brachiolum and axillary sclerite present. Adult male. Hypopygium with bilobed gonostylus: ventral lobe club-shaped, dorsal lobe banana-shaped, not divided.

Dungeyella gavini Jarzembowski, Azar et Nel, 2008

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Figs 2-3

Material examined. Adult male (Fig. 2A) in flat piece of amber, ca. 3 mm thick

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(depository number: NHMUK PI II. 3109 [CC 3/3]. Wealden amber, plant debris bed above Chilton Chine sandstone, Wessex Formation, SW coast of IoW; lower Barremian

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(ca. 128 Ma). Amber collected by Mr. Martin Ian Simpson during the winters of 19932001; the specimen found during sorting the material between 10 and 17th of June 2001. Diagnosis: as for the genus. Description Adult male (n = 1). Total body length 1.3 mm (Fig. 2A). Wing length 0.8 mm, wing width 0.3 mm. Total length/wing length ratio 1.62; wing length/width ratio 2.6.

ACCEPTED MANUSCRIPT Colouration. Thorax, abdomen legs and head brown, abdominal tergite yellow. Head. Eyes bare, without dorsomedial extension. Setae on pedicel not observed. Antenna with 14 flagellomeres, flagellomeres fl1-fl14 length (in µm): 24, 16, 13, 12, 10,

of palpomeres pm1-5 (in µm): 11, 27, 42, 47, 72.

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11, 12, 9, 12, 14, 10, 15, 13, 142, 27; AR 0.96, plume well-developed (Fig 2B). Length

Thorax. Dc 7, strong; Pa at least 2, strong; Scts 6; postnotum bare (Fig. 2B).

Wing. Extension of costa absent. MCu vein present, R1 and R4+5 very short, terminating

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at 0.65 of wing length from base, R2+3 absent, Sc well developed. First axillary sclerite and brachiolum bearing numerous setae of the subequal size. Wing membrane with

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numerous microtrichia. Squama with 7 setae. Anal lobe not pronounced (Fig. 2C, D). Legs. Mid leg tibia with prominent and slim spur, setae-like without denticles, 10 µm long (Fig. 2E). Strong pseudospurs on mid leg tarsomeres 1-3 (Fig. 2E). Hind leg tibia with smooth spur, 10 µm long, tibial comb absent, hind leg tarsomeres 1-3 with

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pseudospurs. Pulvilli absent, claws simple (Fig. 2E). Length of leg segments as in Table 1.

Hypopygium (Fig. 3A-D). Gonocoxite cylindrical, ca. 50µm long. Gonostylus bilobed:

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ventral lobe club-shaped, dorsal lobe banana-shaped (Fig. 3A, B). Volsella elongated, cylindrical, about 2/3rds of length of gonocoxite. Tergite IX bearing long setae on the

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margin (Fig. 3B-D).

Remarks

In the material examined for this paper, we have discovered a chironomid male which appears to be conspecific with Dungeyella gavini. Attribution was done based on the identical shape and arrangement of the tibial spurs, similar length proportions of the tarsomeres (ta4 shorter than ta5), and the essentially identical wing venation pattern and

ACCEPTED MANUSCRIPT shape. The female and the male of D. gavini have similar wing length/width ratios, with usual sexual differences: female 2.5, male 2.6 (Fig. 2C, D) (cf. Jarzembowski et al., 2008). The genus Dungeyella was described based on a single female from the IoW

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Wealden amber. Lack of additional material in the Leng collection, as well as crucial morphological features such as the genitalia being obscured in the amber, led to the genus being unplaced within the family Chironomidae (Jarzembowski et al., 2008).

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These authors hypothesized that Dungeyella gavini belongs to either the subfamily Buchonomyiinae or Podonominae Thienemann, 1937. Based on the following

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combination of characters: brachiolum and axillary sclerite of the wing bearing setae, MCu in basal position, together with the absence of R2+3, led the authors to suggest affinity with the extant subfamily Buchonomyiinae. However, due to the antennae of D. gavini having only 11 flagellomeres, in contrast to 14 in extant Buchonomyiinae, they

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also suggested a possible affiliation of D. gavini with the subfamily Podonominae. In the light of the new material from the Simpson collection, we are set to reevaluate Dungeyella's taxonomic position. Analysis of the characters visible on the

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newly found male allows us to redefine the systematic position and especially the diagnosis of Dungeyella, pointing with certainty towards it being a representative of the

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Buchonomyiinae. These characters are as follows: scale-like setae on the brachiolum and axillary sclerite of the wing, MCu present, R2+3 absent, male antennae with 14 flagellomeres, flagellomere 13 much longer than flagellomere 14, hypopygium with bilobed gonostylus and elongated cylindrical volsella. Dungeyella can be easily distinguished from the two other genera of the Buchonomyiinae - Buchonomyia Fittkau, 1955 and Furcobuchonomyia Baranov, Góral et Ross, 2017 by the RM and FCu veins being closely aligned and the female flagellum

ACCEPTED MANUSCRIPT bearing only 11 flagellomeres. D. gavini therefore represents the third Cretaceous species

of

the

otherwise

mid-Cretaceous

Buchonomyiinae,

together

with

Furcobuchonomyia saetheri Baranov, Góral et Ross, 2017 and F. pankowskii Giłka et

Prodiamesinae Sæther, 1976

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Libanodiamesa Veltz, Azar et Nel, 2007

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Zakrzewska, 2017.

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Diagnosis [adult female]. Females of this genus can be easily distinguished from any extant and fossil Chironomidae based on the following combination of characters: no ovipositor at segment X, gonocoxite IX (GC IX) well developed and separated from tergite IX (T IX), GC IX in lateral view longer than T IX, cerci rhombic with tapering

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finger-like protrusion; wing vein R2+3 present, MCu situated in basal third of wing, FCu separated from RM by at least three times length of RM. The above characters

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supplement the original diagnosis of the genus based on males only.

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Libanodiamesa simpsoni sp. nov. Fig. 4

Type material. Holotype, adult female (Fig. 4A) in flat piece of amber (depository number: NHMUK PI II. 3108 [CC 3/2]. Wealden amber, plant debris bed above Chilton Chine sandstone, Wessex Formation, SW coast of IoW; lower Barremian (ca. 128 Ma).

ACCEPTED MANUSCRIPT Amber collected by Mr. Martin Ian Simpson during the winters of 1993-2001; the specimen found during sorting the material between 10 and 17th of June 2001.

Derivation of name. The species is named in honour of Mr Martin Simpson,

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palaeontologist and fossil collector, who generously donated the amber for our study.

Diagnosis [adult female]. The new species is easily separable from the only known

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species of the genus Libanodiamesa - L. deploegi Veltz, Azar et Nel, 2007 (described based on adult male) by the more basal position of MCu (situated at 0.31 of FCu stalk

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length in L. simpsoni vs. 0.44 in L. deploegi), cubital and anal veins broadly separated in L. simpsoni (cubital and anal veins run in close proximity in L. deploegi), as well as different wing length/width ratio (2.6 in L. simpsoni female vs. 2.3 in L. deploegi male).

Adult female (n = 1).

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Description

Total body length 1.2 mm (Fig. 4A, B). Wing length 1.3 mm, wing width 0.5 mm. Wing

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length/total length ratio 1.1 (Fig. 4B); wing length/width ratio 2.6. Colouration. Whole specimen brown, wings hyaline, without markings.

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Head. Ocelli absent. Eyes bare, with slight dorsomedial extension (Fig. 4C). Pedicel bare. Antenna with 7 flagellomeres, flagellomeres fl1-fl7 length (in µm): 34, 29, 32, 31, 31, 33, 102; AR 0.56. Maxillary palp with 5 palpomeres, only pm4 and pm5 clearly visible (lengths in µm: 113 and 146 respectively). Clypeus square, with 10 clypeals (Fig. 4C). Thorax. Scutellum and postnotum with setae.

ACCEPTED MANUSCRIPT Wing. Bare. R2+3 weak, Sc absent, MCu very faint, situated at wing base at 0.31 of FCu stalk length (Fig. 4D E). Cubital and anal veins broadly separated. Costal extension absent (Fig. 4E). Legs. Tibiae of all legs bearing a straight short spur (length ca. 10 µm), tarsi without

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pseudospurs, all tarsomeres cylindrical. Leg segment lengths as in Table 2.

Abdomen. Tergite VIII simple. Gonocoxite IX well separated from tergite IX, longer than tergite IX in lateral view. No ovipositor at segment X. Cerci rhombic, with

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digitiform projection (Fig. 4F).

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Remarks

Libanodiamesa simpsoni is included in the subfamily Prodiamesinae based on the following combination of characters: ovipositor absent, simple R2+3 present, ta4 longer than ta5 on fore-and hind legs, all tarsi cylindrical, gonocoxite IX large and well

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separated from tergite IX. It is attributed to the genus Libanodiamesa based on the extremely basal position of MCu, FCu distal to RM, and wing membrane and eyes bare (Fig. 4B-E). The extremely basal position of MCu is a unique character state in

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Prodiamesinae, which can be considered as an autapomorphy of the genus Libanodiamesa (Sæther, 1989, Veltz et al., 2007).

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As a rule, in Chironomidae adults the wing length/width ratio in females is lower

than that in males, since the male wing is slender/elongated, whereas the female wing is broader and/or shorter (e.g. see the previous species, Fig. 2C, D). Opposite wing proportions found in the two compared Libanodiamesa species (Fig. 4E, G) supports the concept of the newly described L. simpsoni as being a distinct species. Other discrepancies observed in these species (see the diagnosis) significantly exceed any expected span of sexual dimorphism in Prodiamesinae (Sæther, 1977, 1989).

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4. DISCUSSION

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The Wealden amber of the Isle of Wight is nationally important because it the first in situ amber deposit in the UK to yield insect inclusions. Moreover, it contains true flies of which chironomids constitute a large proportion, interpreted as evidence of seasonal

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resin production in the Early Cretaceous of southern England, as in the younger (Eocene) Baltic amber (Jarzembowski et al., 2008). The Early Cretaceous chironomids

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from the UK are important because they can be compared with the well-studied early Barremian chironomid fauna of the Lebanese amber which is of similar age. It is notable that the Lebanese and Wealden ambers share at least one genus of Chironomidae - Libanodiamesa Veltz, Azar et Nel, 2007. Shared extinct genera of

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Chironomidae between different fossil faunas are extremely rare and suggestive of ancient biogeographical connections between the regions (Lukashevich and Przhiboro, 2011). The occurrence of Libanodiamesa in Wealden amber suggests that other taxa

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described from Lebanese amber should be sought in the British fossil record. Several species of Prodiamesinae are known from the fossil record. Cretodiamesa

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taimyrica Kalugina, 1976 is known from the uppermost Cretaceous amber from the Taimyr peninsula in Russia, Cretodiamesa arieli Veltz, Azar et Nel, 2007, Libanodiamesa deploegi and Paicheleria magnifica Azar et Nel, 2010 are described from Barremian Lebanese amber, as well as the new species of Libanodiamesa from the IoW. In addition to Cretaceous taxa, three records of the Prodiamesinae are known from the Eocene: Prodiamesa indeserta Seredszus et Wichard, 2009 from Baltic amber, Monodiamesa guglielmia Doitteau et Nel, 2007 from Paris basin (Oise) amber, and an

ACCEPTED MANUSCRIPT undescribed Prodiamesinae male cf. Prodiamesa from the Indian Cambay amber (Stebner et al., 2017). Unfortunately, extant members of the subfamily Prodiamesinae are eurybiontic inhabiting a variety of habitats from brackish lagoons to mountainous

Libanodiamesa (Moller Pillot, 2013).

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streams, which makes it difficult to make any inference regarding the palaeoecology of

Another significant biogeographical feature of the Wealden amber Chironomidae is the occurrence of Buchonomyiinae consistent with the importance of this subfamily

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clade in Cretaceous ecosystems. There are three described species in mid-Cretaceous ambers and, moreover, Buchonomyiinae comprise the bulk of Chironomidae in

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Burmese amber (Zakrzewska and Giłka, 2018; Baranov, unpublished data). By contrast, Buchonomyiinae, are extremely rare in the extant fauna, suggesting a much more prominent role in past insect communities. Extant Buchonomyiinae are represented by the single genus Buchonomyia and three species B. thienemanni Fittkau, 1955, B.

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burmanica Brundin et Sæther, 1978 and B. brundini Andersen et Sæther, 1995, distributed in Europe, Myanmar and Costa Rica respectively. It is hypothesized that the extreme rarity of the modern Buchonomyiinae is caused by their unusual lifestyle,

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namely being parasites of psychomyiid caddisflies (Ashe et al., 2015). Fossil Buchonomyiinae were found together with Psychomyiidae (i.e. Burmese amber

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Furcobuchonomyia) co-occurring with the earliest geological records of the Psychomyiidae (Baranov et al., 2017). Since no pre-Barremian Psychomyiidae are known, it is difficult to infer when the association of the Buchonomyiinae and their potential hosts developed (op. cit.). It is hard to hypothesize why Buchonomyiinae were more diverse and abundant in the Cretaceous, but probably as a result of a Jurassic/Cretaceous radiation in caddis behaviour (Jarzembowski et al., 2016) and it might also be related to the large shift in the chironomid faunas which occurred in the

ACCEPTED MANUSCRIPT late Cretaceous (Kalugina, 1974). The shift was manifested by the replacement of the typical Mesozoic chironomid species complexes, including many lacustrine Podonominae and Aenneinae, by the more extant-type species complexes rich in Orthocladiinae and Chironominae. Kalugina (1974) has hypothesized, that this shift was

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prompted by the emergence of the angiosperm-dominated deciduous forests, which led to the enhanced transport of the allochthonous organic matter into the water bodies. It is possible that this shift has led to the extinction of the majority of the Buchonomyiinae

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species; however, until more fossils from sedimentary and amber deposits are found,

5. CONCLUSION

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this will remain a speculation.

This study once again highlights the importance of fossil chironomids for

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reconstructing the history of the recent fauna as was first shown in Brundin's seminal monograph on transantarctic biogeographical connections (Brundin, 1966), in reverse of the famous saying of the father of modern historical geology Charles Lyell (quoting

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James Hutton) „the present is the key to the past".

6. ACKNOWLEDGMENTS

This research was supported by the State Key Laboratory of Palaeontology and Stratigraphy (Nanjing Institute of Geology and Palaeontology, CAS) 183101 and is a Leverhulme Emeritus Fellowship contribution for EAJ. VB is grateful to the organizing committee of the international symposium “Amber. Science and Art” at AMBERIF

ACCEPTED MANUSCRIPT 2018, whose financial assistance allowed him to visit the WG lab in Gdańsk. Sincere thanks are due to Dr. Darren Naish and Dr. Sam W. Heads for their kind permission to reproduce the geological maps and stratigraphical schemes from their respective works. Comments on the manuscript by the Editor-In-Chief Dr. Eduardo Koutsoukos and two

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anonymous reviewers are greatly appreciated.

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ACCEPTED MANUSCRIPT Figure legends Fig. 1. Geological context of the Wealden Group of the Isle of Wight. A) Stratigraphic scheme of the Wealden on the Isle of Wight showing the distribution and lithology of insect-bearing horizons (modified from Heads, 2008, by permission). B) Isle of Wight

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geological map showing the location of the Wealden. C) Chilton Chine’s position on the Isle of Wight coast (marked with a red dot). Modified from Martill and Naish (2001),

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with author’s’ permission.

Fig. 2. Dungeyella gavini Jarzembowski, Azar et Nel, 2008, male (A, B, D, E) and

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female (C). A: habitus, lateral view. B: head and thorax. C, D: wing (C: modified from Jarzembowski et al., 2008). E: apex of midleg tibia and tarsus. Scale bar = 100 µm.

Fig. 3. Dungeyella gavini Jarzembowski, Azar et Nel, 2008, male hypopygium

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photographed (A, C) and drawn (B, D), in ventrolateral (A, B) and dorsolateral view (C, D). Red: gonocoxite; blue: gonostylus, ventral branch; green: gonostylus, dorsal branch;

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purple: volsella. Scale bar = 100 µm.

Fig. 4. Libanodiamesa simpsoni sp. nov., holotype female (A-F) and L. deploegi Veltz,

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Azar et Nel, 2007, holotype male (G). A, B: habitus in dorsolateral view, photographed (A) and drawn (B). C: female genitalia, ventral view. D: head, ventral view. E: wing base, with crossvein MCu. F, G: wing (G: modified from Veltz et al., 2007). Scale bar = 100 µm.

P1

fe

ti

ta1

ta2

ta3

ta4

ta5

LR

340

315

140

80

50

45

55

0.44

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405

190

165

115

65

50

55

0.87

P3

365

345

150

110

75

40

55

0.43

Table 1. Leg segment lengths (µm) and leg ratios of male Dungeyella gavini

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Jarzembowski, Azar et Nel, 2008. p1-3: pairs of legs 1-3, fe: femur, ti: tibia, ta1- 5:

ti

ta1

ta2

ta3

ta4

ta5

LR

P1

355

400

345

175

130

95

80

0.86

P2

525

385

310

150

110

75

75

0.81

P3

425

455

145

110

115

80

85

0.32

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fe

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tarsomeres 1-5, LR: leg ratio.

Table 2. Leg segment lengths (µm) and leg ratios of female Libanodiamesa simpsoni

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sp. nov. p1-3: pairs of legs 1-3, fe: femur, ti: tibia, ta1- 5: tarsomeres 1-5, LR: leg ratio.

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