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2023Lagriodidae

January 2023
Annales Zoologici 73(2):261-292

Authors:

Richard A. B. Leschen

New Zealand Arthropods Collection

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THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH,

STAT. NOV. (COLEOPTERA: TENEBRIONOIDEA), WITH

DESCRIPTIONS OF TWO NEW AUSTRALIAN SPECIES

JOHN F. LAWRENCE1,2, RICHARD A. B. LESCHEN3,

MARIO ELGUETA4, NICK PORCH5

and ADAM ŚLIPIŃSKI1*

1Australian National Insect Collection, CSIRO, GPO Box 1700, Canberra, Australia

261 Glenbar Rd., The Palms, Queensland 4570, Australia

3Manaaki Whenua Landcare Research, New Zealand Arthropod Collection,

Auckland, New Zealand

4Museo Nacional de Historia Natural (MNHN), Área de Entomología. Santiago, Chile

5School of Life and Environmental Sciences, Deakin University, Australia

*Corresponding author: e-mail: Adam.slipinski@csiro.au

Citation: Lawrence, J.F., Leschen, R.A.B., Elgueta, M., Porch, N., Ślipiński, A. 2023. The

family Lagrioididae Abdullah & Abdullah, stat. nov. (Coleoptera: Tenebrionoidea), with

descriptions of two new Australian species. Annales Zoologici, 73: 261–292.

doi: 10.3161/00034541ANZ2023.73.2.009

Received: 27 March 2023 Accepted: 30 May 2023 Printed: 30 June 2023

Abstract.— Relationships of the anthicid subfamily Lagrioidinae are discussed in detail

and evidence presented supporting treatment of this group as a distinct family,

Lagrioididae, stat. nov., of Tenebrionoidea, unrelated to the family Anthicidae. Lagrioida

rufula Fairmaire & Germain, 1860 is designated as the type species of Lagrioida

Fairmaire & Germain, 1860. Two new Australian species, L. norfolkensis Lawrence

& Ślipiński sp. nov. (Norfolk Island) and L. tasmaniae Lawrence & Ślipiński sp. nov.

(Tasmania and Victoria) are described and illustrated. All remaining species are

redescribed and key to the species is provided.

Key words.— Anthicidae, Australia, Chile, New Zealand, Brazil, new species, phylogeny,

ocean dunes

INTRODUCTION

The genus Lagrioida Fairmaire & Germain (1860)

was placed by its describers in the family Lagriidae

(Tenebrionidae: Lagriinae), but Champion (1890)

excluded it from that group, redescribed the genus

based on specimens collected by J. J. Walker on sand

dunes at Coquimbo and Valparaiso, Chile and noted the

resemblance of Lagrioida to several ‘melandryid’

genera, such as Eurypus Pascoe and Physcius Cham-

pion (now included in Mycteridae: Eurypinae). This

association with Mycteridae was followed in catalogues

by Blair (1928) and Blackwelder (1945). Crowson

(1955) placed the Lagrioida in his family Cononoti-

dae, along with Cononotus LeConte and Agnathus

Germar. Abdullah & Abdullah (1968) considered this

ANNALES ZOOLOGICI (Warszawa), 2023, 73(2): 261-292

PL ISSN 0003-4541 © Museum and Institute of Zoology PAS

doi: 10.3161/00034541ANZ2023.73.2.009

group of genera to form a new tribe, Lagrioidini, with-

in the anthicid subfamily Eurygeniinae, and later

(Abdullah 1974) elevated that tribe to subfamily level

within Anthicidae. As discussed in detail below, the

phylogenetic relationships of the genus Lagrioida

continue to be uncertain. The present study re-exam-

ines the phylogenetic position of Lagrioida within

Tenebrionoidea, elevates the group to family rank,

describes two new Australian species, and provides

a key to world species.

MATERIALS AND METHODS

With the following exceptions or modifications,

morphological terms used here are discussed and/or

illustrated in Lawrence et al. (2010b, 2011) and

Lawrence & Ślipiński (2013). As discussed in a previ-

ous work (Lawrence, 2019), punctation and vestiture,

particularly of the dorsal surfaces, may have a differ-

ent appearance when observed in dried specimens

under incident light (most dissecting microscopes)

than when observed in cleared specimens in fluid

under transmitted light (dissecting microscopes with

transmitted light stage or compound microscopes). The

term megasetae is used for the dominant form of dor-

sal vestiture in this group. The term microsetae is used

for the very fine, erect setae, which occur in some but

not all taxa; these were referred to as ‘tactile setae’ by

Werner & Chandler (1995) and appear to be some kind

of mechanoreceptor. Head length is measured from the

anterior edge of the clypeus to the centre of the poste-

rior edge above the occipital foramen and width is

always that just behind the eyes but before the neck.

Intercoxal processes are not included in the length

measurements of prosternum, mesoventrite, metaven-

trite and first abdominal ventrite. Metaventrite length

in this sense is equivalent to the shortest distance

between the mesocoxal and metacoxal cavities. Hind

wing terminology follows that in Lawrence et al. (2021,

2022). The wing length is measured from the base of

the subcostal vein to apical edge, the width is the great-

est width and the length of the apical field is measured

from cross-vein r4 to the apical edge.

New Zealand regions are those of Crosby et al.

(1998). The following acronyms are used to denote

sources of specimens and type repositories (Australian

unless otherwise noted):

AMS – Australian Museum, Sydney, NSW, Australia;

ANIC – Australian National Insect Collection, CSIRO,

Canberra, ACT, Australia;

CAS – California Academy of Sciences, San Francis-

co, CA, USA;

MGC – Marcelo Guerrero Collection, Santiago, Chile;

MNHN – Museum National d’Histoire Naturelle, Paris,

France;

MHNS – Museo Nacional de Historia Natural, Santia-

go, Chile;

MVM – Museum of Victoria, Melbourne, VIC, Aus-

tralia;

NHML – Natural History Museum, London, UK;

NZAC – New Zealand Arthropod Collection, Landcare

Research, Auckland, New Zealand;

PVC – Pedro Vidal Collection, Santiago, Chile;

QDAF – Queensland Department of Agriculture and

Forestry, Brisbane, QLD, Australia;

QMB – Queensland Museum, Brisbane, QLD, Aus-

tralia;

SAM – South Australian Museum, Adelaide, SA, Aus-

tralia;

TMAG – Tasmanian Museum and Art Gallery, Hobart,

TAS, Australia;

VMD – Victor M. Diéguez Collection, Santiago, Chile.

Family Lagrioididae, stat. nov.

Lagrioidini Abdullah & Abdullah, 1968: 73.

Diagnosis

Adults differ from those all other Tene-

brionoidea with respect to the following combination

of characters: 1) head constricted to form broad neck,

2) antennal insertions concealed, 3) antennae relative-

ly long with weak club; 4) eyes relatively large, slightly

transverse, entire and coarsely facetted; 5) mandibular

mola well-developed but without fine transverse ridges;

6) apical maxillary palpomere securiform, 7) base of

pronotum distinctly narrower than combined elytral

bases, 8) lateral pronotal carinae absent, 9) procoxae

strongly projecting with concealed trochantins, 10) me -

sanepisterna meeting or almost meeting at midline,

11) mesocoxal cavities partly closed laterally by

mesepimera, 12) metacoxa with narrow-based internal

apophysis, 13) metendosternite with well-developed

laminae, 14) hind wing with medial fleck, 15) penulti-

mate tarsomere strongly bilobed, 16) first two abdomi-

nal ventrites connate, and 17) aedeagus of inverted

tenebrionoid type with ventral tegmen. Larvae differ

from those of other tenebrionoids with respect to the

following character combination: A) five pairs of stem-

mata, B) frontal arms U-shaped with short endocarina

at base of each, C) antennal sensorium dome-like and

about half as long as antennomere 3, D) frontoclypeal

suture present but very weakly impressed, E) mandi -

bular mola with transverse ridges, F) mala cleft with

three small apicomesal teeth, G) hypostomal rods very

short, H) urogomphi narrowly separated with small,

mesobasal teeth almost meeting, I) interurogomphal

pit or pits absent, J) sternite IX with a pair of basolat-

eral teeth and K) thoracic spiracles annular-biforous

with short accessory tubes, basal abdominal spiracles

annular-biforous with long accessory tubes and most

abdominal spiracles annular.

262 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 263

Figure 1. Lagrioida species in natural habitats. (A) L. australis from Tasmania, © Simon Grove; (B–F) L. brounii from New Zealand.

(B) © Ormond Torr; (C) © James Bailey; (D, E) © Arnim Littek, (F) © Lisa Bennett. All from iNaturalist.

Family relationships

As mentioned above, Lagrioida was tentatively

placed by Crowson (1955) in the family Cononotidae,

also containing the genera Agnathus and Cononotus

and apparently related to Salpingidae and Mycteridae

on the one hand and the ‘Anthicid group’ on the other.

Abdullah & Abdullah (1968) placed Lagrioida,

Cononotus and Agnathus in a new tribe, Lagrioidini,

within the anthicid subfamily Eurygeniinae, and Abdul-

lah (1974) elevated the group to subfamily level within

Anthicidae (including Pedilinae, Steropinae and

Macratriinae), differing from other subfamilies in the

connation of the first two abdominal ventrites, the

broad neck (absent in Agnathus and Cononotus),

‘nearly open’ mesocoxal cavities and widely separated

metacoxae (not true in Lagrioida). The association

with Anthicidae was questioned based on descriptions

of the larvae of Agnathus and Cononotus by Mamaev

(1976) and Doyen (1979), respectively; and those two

genera were placed in the pyrochroid subfamily

Agnathinae Lacordaire, 1859 (a senior synonym of

Cononotinae LeConte, 1862). Although Lagrioida

remained in Anthicidae in some works (Costa et al.

1995; Lawrence & Newton 1995; Werner & Chandler

1995; Klimaszewski & Watt 1997), there has always

been some doubt about its phylogenetic relationships,

and Lagrioidinae was considered to be Tenebrionoidea

incertae sedis by Lawrence et al. (2010a).

Costa et al. (1995) described the first known Lagri-

oida larva (that of L. nortoni Costa et al.), noting that

it resembled typical anthicid larvae in having a penicil-

lus at the base of the mandible and the absence of

interurogomphal pits. The larva differed from those of

typical Anthicidae in having 5 pairs of stemmata and

a frontoclypeal suture. According to the same authors

Lagrioida adults are similar to those of Anthicidae in

1) the possession of a weak antennal club, 2) lack of

pronotal carinae, 3) securiform apical maxillary

palpomere, 4) mesanepisterna meeting in front of

mesoventrite, 5) metendosternite with well-developed

laminae, 6) metacoxa with narrow-based apophysis

and 7) penultimate tarsomere lobed beneath, but differ

from anthicids in having A) the first two ventrites con-

nate, B) a weakly constricted head with a wide neck,

C) concealed antennal insertions, D) inverted aedea-

gus, E) penis with two basal struts, and F) phallobase

without struts. The last feature refers to the sclerotised

lateral edges of the basale in some Anthicinae.

Costa et al. (1995) also noted that adult Lagrioida

shared the following features with Pyrochroidae (then

referred to as Pedilidae, and excluding Agnathus and

Cononotus): 1) weak antennal club, 2) lack of pronotal

carinae, 3) securiform apical maxillary palpomere,

4) meeting of mesanepisterna in front of mesoventrite,

5) well-developed metanepisternal laminae, 6) narrow-

based metaocoxal apophysis, and 7) lobed penultimate

tarsomere, while differing in A) connation of first two

abdominal ventrites, B) head with wide neck, C) con-

cealed antennal insertions, D) inverted aedeagus, and

E) paired basal penile struts. Larvae were said to share

the following features with those of Pedilidae (sensu

stricto): mandible with 1) bidentate mandibles and

2) tridentate scissorial area, 3) three small teeth at

malar apex, while differing in A) the absence of uro-

gomphal pits and B) absence of anterior asperities on

sternum IX. This last difference may apply to L. nor-

toni, but larvae of L. australis have a single pair of

asperities at the anterolateral angles of sternum IX.

Lagrioida was also compared with the genera

Agnathus and Cononotus; Lagrioida adults resem-

ble those of both genera in having 1) the first two

abdominal ventrites connate and 2) the aedeagus of the

inverted type, while they differ in having A) weakly

clubbed antennae (not filiform), B) deeply lobed penul-

timate tarsomeres (not lobed), C) mesocoxal cavities

partly closed laterally by mesepimera (not by meeting

of ventrites). Lagrioida larvae resemble those of

Agnathus in having 1) 5 stemmata on each side,

2) bidentate mandiles with 3) tridentate cutting edge,

and 4) small teeth at the malar apex, while differing in

A) presence of frontoclypeal suture and B) absence of

paired interurogomphal pits. They resemble the

Cononotus larva in having a cleft mala but differ in

having 5 stemmata and lacking paired interurogom-

phal pits.

The first molecular analysis to include Lagrioida,

that of McKenna et al. (2015), based on DNA sequence

data from eight nuclear genes, found Lagrioida to

form a clade with Prostomis Latreille (Prostomidae)

and two genera of Oedemeridae (Calopus Fabricius

and Ditylus Fischer von Waldheim) under Bayesian

inference but to form a clade with three pyrochroid

genera (Morpholycus Lea, Pedilus Fischer von

Waldheim and Pyrochroa Geoffroy) under maximum

likelihood.

Morphological analysis

Forty-nine taxa of Tenebrionoidea representing 28

families were scored for 63 adult and 40 larval charac-

ters (Appendices 1 and 2). The morphological matrix

was constructed in Mesquite v3.61 (Maddison and Mad-

dison 2019). Parsimony searches were conducted in

TNT v1.5 (Goloboff & Catalano 2016) using 1000 repli-

cates with default settings in a Traditional Search, with

all characters with equal weights and unordered. Node

support was evaluated using 1000 replicates of the

symmetric resampling option in TNT with default set-

tings. Maximum parsimony analysis of the morpholog-

ical dataset yielded three equally parsimonious trees

with 1071 steps showing two radically different posi-

tions for Lagrioida (Fig. 2) and para- or polyphyletic

Anthicidae. The strict consensus of these trees (L 1245;

Ci=13; Ri=31) and the tree obtained from the 1000 re -

plicates of symmetric resampling are almost identical

and show negligent support for some of the branches

with the backbone, including the position of Lagrioida

not resolved.

Based on the profound morphological differences

between Lagrioida and those families in which it was

previously placed, and the results of the above phyloge-

netic analyses, we believe that Lagrioida cannot be

placed in any of the existing tenebrionoid families and

are here elevating the subfamily Lagrioidinae to the

status of independent family Lagrioididae stat. nov.

264 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 265

Figure 2. Results from parsimony analyses of morphological data in TNT. (A, B) two shortest trees (L = 1071) from Traditional search in TNT;

TAXONOMY

Lagrioida Fairmaire & Germain, 1860

Lagrioida Fairmaire & Germain, 1860: 3–4. Type species, here desig-

nated: Lagrioida rufula Fairmaire & Germain, 1860.

Lagrioida: Fairmaire & Germain,1863: 234–235; Werner & Chandler

1995: 25.

Lagrioda: Pascoe 1876: 58, Hudson 1923: 384, misspelling.

Diagnosis

As for the family.

Adult description

Length 2.2–6.75 mm; body

2.3–2.75 × as long as wide, yellow or yellowish-brown

to black in colour. Dorsal punctation consisting of

large, shallow megapunctures with gradual edges,

appearing to be very coarse and dense under incident

light, enclosing smaller, more clearly defined, oval

micropits, each containing a longitudinal groove and

associated with an inclined or decumbent megaseta;

dorsal vestiture usually consisting of white or yellow,

decumbent megasetae, often somewhat thickened, and

sometimes a second set of less abundant, very fine,

erect microsetae not associated with micropits; inter-

spaces smooth and shiny; punctation of ventral sur-

faces somewhat finer and more uniform than that of

dorsal surfaces, with circular megapunctures uniform-

ly dense, with similar micropits.

Head not or slightly declined, gradually constricted

posteriorly to form short, gradual temples and broad

neck. Frontal region not or only slightly, gradually

declined. Eyes protuberant, entire, coarsely facetted,

with short interfacetal setae. Antennal insertions

slightly concealed from above; subantennal groove

absent. Frontoclypeal suture absent. Anterior edge of

clypeus straight to convex. Labrum slightly transverse,

its anterior edge broadly rounded. Antennae 11-seg-

mented, with weak, 3-segmented club; antennomeres

1–8 narrow and elongate, antennomere 10 usually

transverse. Mandible about 1.5 × as long as wide with

outer edge almost straight for basal two-thirds, then

somewhat abruptly curved mesally to bidentate apex,

basal two-thirds of outer edge forming slightly concave

subtriangular area; incisor edge with two or three

teeth; mola subtriangular, with a somewhat irregular

surface of very fine tubercles and microtrichia; pros-

theca large, membranous, lined with long hairs apical-

ly; base of mesal surface with short membranous lobe.

Maxilla with distinct, setose galea and lacinia, the

latter without uncus; maxillary palpomere 2 elongate,

3 short, triangular, 4 large, asymmetrically expanded

and securiform. Ligula laterally expanded and apically

truncate; labial palps widely separated, with apical

palpomeres fusiform. Mentum transverse, widest at

middle, sides angulate. Submentum fused to gula to

form gulamentum. Gular sutures widely separated.

Corpotentorium narrow; cervical sclerites slender and

sinuate.

Pronotum 1.03–1.25 × as long as wide, widest ante-

riorly; sides slightly rounded or sinuate, not explanate,

base distinctly narrower than combined elytral bases.

Lateral pronotal carinae absent; anterior angles

absent; posterior angles broadly rounded; posterior

edge more or less straight; disc without paired basal

impressions, sometimes with one or two pairs of short,

oblique curved sulci extending anteroventrally from

the sides of base. Prosternum in front of coxae longer

than shortest diameter of procoxal cavity, flat to mod-

erately convex. Prosternal process very narrow at mid-

dle, not extending to edges of coxae, concave at middle

and subacute to narrowly rounded or truncate at apex.

Notosternal sutures absent. Procoxae projecting well

below prosternum, with moderately long concealed

lateral extensions and concealed trochantins. Procoxal

cavities circular, narrowly separated, without antero-

lateral incisions, externally broadly open, internally

closed. Pterothorax. Scutellar shield not abruptly ele-

vated, anteriorly simple, posteriorly broadly rounded

to angulate. Elytra 1.7–2.1 × as long as combined width

and 2.6–4.0 × as long as pronotum, disc with humeri

weakly developed and located at lateral edge, conceal-

ing anterolateral angles from above; punctation irregu-

lar, without scutellary striole; apices meeting at suture

or independently rounded and slightly separated, often

exposing at least part of pygidium; epipleuron narrow

and incomplete. Mesoventrite triangular separated by

straight complete sutures from mesanepisterna, which

are narrowly separated to contiguous at midline, each

with a transverse procoxal rest; anterior edge of disc

often modified; mesoventrital process complete, nar-

row, slightly convex, its apex broadly rounded with

subapical cavity. Mesocoxae not or only slightly pro-

jecting, with exposed trochantins. Mesocoxal cavities

subcircular, narrowly separated, partly closed lateral-

ly by mesepimeron. Junction of meso- and metaventrite

forming a monocondylic (tongue-and-groove) joint.

Metaventrite with relatively short discrimen; postcoxal

lines absent; metanepisternum long and narrow, with

outer edge straight; exposed portion of metepimeron

extending well beyond metanepisternal apex. Metacox

ae narrowly separated, not extending laterally to

meet elytra; plates absent, with narrow-based internal

apophysis. Metendosternite with lateral arms, moder-

ately to very long laminae, no anterior process and

anterior tendons on lateral arms or not apparent. Ful-

ly developed hind wing (Fig. 8A–F) with apical field

long and devoid of veins or sclerotisations, radial cell

reduced with pigmented lumen and truncate apical

edge, basal portion of RP very short, radio-medial loop

acute, MP1+2 well-developed with short medial spur;

medial field with four free veins and medial fleck,

wedge cell well developed with truncate apex, and anal

embayment absent; wings sometimes reduced with

reduction or loss of apical field. Legs moderately well

266 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

developed, slender; prothoracic legs somewhat shorter

than meso- or metathoracic legs; trochanterofemoral

joint strongly oblique with base of femur separate from

coxa; tibial spurs well developed and paired on all legs;

tarsi 5-5-4 in both sexes; penultimate tarsomere with

ventral lobe deeply divided and extending beyond mid-

dle of ultimate tarsomere; pretarsal claws simple.

Abdomen with five ventrites, the first two connate;

ventrite 1 usually subequal in length to 2; postcoxal

lines absent; intercoxal process acute; ventrites 3 to 5

each shorter than one preceding it and ventrite 5 (ster-

nite VII) short and broad, distinctly wider than tergite

VII (pygidium). Functional spiracles on segments I–VII,

located in pleural membrane, those on segments VI and

VII enlarged and the latter often visible from above on

either side of pygidial base. Anterior edge of segment

IX in male with pregenital ring subacute anteriorly; ter-

gites IX and X often fused and membranous. Aedeagus

of inverted tenebrionoid type (with ventral tegmen),

symmetrical; tegmen divided, with basale and apicale;

apicale acute at apex, without accessory lobes or

tegminal struts; anterior edge of penis with paired

struts. Sternite VIII in female with long, curved, basal-

ly articulated spiculum ventrale. Ovipositor moderate-

ly elongate and lightly sclerotized, except for bacula;

paraprocts at least twice as long as coxites, which are

subdivided into short basal and longer apical lobes,

densely setose, with styli well developed. Internal

female tract with spermatheca trilobed and bursa with-

out armature. Foregut with well-developed crop and

simple proventriculus.

Larva description.Based primarily on intermedi-

ate instars of Lagrioida australis Champion, 1895

(see data below) supplemented by published larval illus-

trations of Lagrioida nortoni in Costa et al. (1995).

Body elongate, almost parallel-sided, but slightly

wider at middle, slightly flattened; lightly sclerotized

except for clypealia and tips of pretarsal claws,

mandibles and urogomphi. Dorsal surfaces smooth;

ventral surfaces of laterotergites with patches of

minute tubercles; vestiture consisting of short to very

long, erect setae, the longest almost as long as average

abdominal segment.

Head capsule 0.85 × as long as wide with posterior

edge slightly emarginate and sides subparallel. Epicra-

nial stem absent; frontal arms more or less U-shaped,

their bases contiguous with underlying basal endocari-

nae. Stemmata five on each side, close together in

obliquely vertical rows of 3 and 2. Frontoclypeal suture

present but vaguely indicated; clypeus 0.46 × as long

as wide, with sides straight and anteriorly converging

and division between postclypeus and anteclypeus not

clearly delimited. Labrum 0.68 × as long as wide, with

sides curved and apex broadly rounded. Antennae

about 0.4 × as long as head width, 3-segmented with

well-developed antennifer, antennomere ratio 1.67:

2.33: 1.00, sensorium dome-like and half as long as

antennomere 3. Mandibles asymmetrical, 1.5 × as long

as wide, with broad base and narrowly bidentate apex;

outer edge with basal, triangular area flattened, lightly

sclerotised and setose; incisor edge with three teeth;

mola well developed, with fine, transverse ridges, with

small tuft of hairs at base; prostheca absent. Ventral

mouthparts retracted; maxillary articulating area lon-

gitudinally oval. Cardo divided by internal ridge, trans-

verse to slightly oblique; stipes longer than wide; mala

obliquely truncate, cleft, with three teeth at inner api-

cal angle and a number of apical and mesal spines;

maxillary palp 3-segmented, with distinct palpifer,

ratio of palpomere lengths 1.00: 1.13: 1.63. Labium con-

sisting of prementum, mentum and gulamentum; ligula

shorter than labial palp, with broadly rounded apex;

palps 2-segmented, separated by more than width of

first palpomere. Hypopharyngeal sclerome molar-like,

transverse. Hypostomal rods very short, parallel or

barely diverging posteriorly; ventral epicranial ridges

absent. Gular sutures separate; gula wider than long.

Thorax slightly more than twice as long as wide;

abdomen about three × as long as thorax; terga ex -

tending laterally slightly beyond edges of sterna.

Proventer with relatively large, subpentagonal prester-

num, but without other distinct armature. Thoracic

spiracle annular-biforous (bicameral) with accessory

chambers shorter than peritreme diameter, not located

on spiracular tube. Legs moderately well developed,

5-segmented, more or less equal in length, almost as

long as thoracic width; coxae separated by more than

one but less than two diameters; each coxa somewhat

projecting, about 0.7 × as long as wide ; trochanter

0.67 × a long as coxa and 1.33 × as long as wide; femur

1.25 × as long as trochanter and1.67 × as long as wide;

tibiotarsus 1.20 × as long as femur and 2.4 × as long

as wide; pretarsus 3.0 × as long as wide and 0.75 × as

long as tibiotarsus, slightly broader and lightly sclero-

tised basally and abruptly narrowed, narrowly acute

and well sclerotised apically. All legs relatively sparse-

ly clothed with fine hairs; pretarsus with two setae,

lying more or less side by side. Meso- and metaterga

and abdominal terga I–IX each with weak, slightly sin-

uate, transverse carina, without patches or rows of

asperities. Protergum 0.8 × as long as wide, with sides

almost parallel; mesotergum 0.73 × as long as proter-

gum and 0.57 × as long as wide; metategum 0.82 × as

long as protergum and 0.61 × as long as wide; first

abdominal tergum 0.95 × as long as metatergum and

0.53 × as long as wide; abdominal terga II–VII each

about 0.6 × as long as wide; tergum VIII slightly longer

than VII and 0.67 × as long as wide.

Abdominal terga I–VIII extending laterally onto

ventral surface; abdominal sterna transversely

quadrate with rounded angles, with few setae; sternum

I 0.46 × as long as wide, II–VIII about 0.8 × as long as

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 267

wide and VIII 0.65 × as long as wide. Tergum IX 0.89 ×

as long and 0.8 × as wide as tergum VIII, 0.73 × as long

as wide at base, with sides more or less parallel to

about middle. Urogomphi 0.44 × as long as basal tergal

width, separated by 0.11 × the basal width of one,

basal half of each lightly pigmented, posteriorly direct-

ed with sides slightly converging and apical half heavi-

ly pigmented, anterodorsally directed, with sides more

strongly converging to narrowly acute apex; basal sec-

tion of each with several setose tubercles, the largest

and most obvious being one lateral near base, one

mesal near apex and one mesal at base, the last almost

meeting that on the opposing urogomphus; interuro-

gomphal pit or pits absent; apex of tergum IX almost

vertical but slightly extending onto ventral surface; not

forming articulated plate. Sternum IX 0.63 × as long as

wide, widest at base, which is strongly curved, apex

angulately emarginate, surface apically declined, with

small tubercle at each anterolateral angle; segment X,

transversely oval with weak indications of short, broad

pygopods. First pair of abdominal spiracles annular-

biforous, with accessory chambers longer than per-

itreme diameter; spiracles on following abdominal seg-

ments annular and those on segment VIII about the

same size as others on abdomen, facing laterally, with-

out spiracular tubes

Biology.Almost all known Lagrioida have been

collected on, in, or behind sand dunes on ocean beach-

es (Fig. 1E) above the high tide line and in association

with a variety of beach grasses (Fig. 1F) and other

dune plants at night. A few of the Australian records

for L. australis Champion, however, are not from

coastal localities, but from inland areas (e.g., Kelso on

the Richmond River) suggesting that psammophilous

environments bordering fresh water may, additionally,

be suitable habitat. Adults of L. australis may be col-

lected on stems, leaves or spikelets of various grasses

(Poaceae), but they have also been found under logs, in

wet seaweed on the beach, or in the sand among the

roots of herbs or grasses in sand dunes. Adults of

L. nortoni Costa et al. were observed feeding on grass

spikelets and larvae of the same species were collected

by sifting sand at the bases of grasses or herbs and

apparently feed on dead leaves and root debris. Adults

of L. rufula Fairmaire & Germain were collected in

dunes under associated Mesembryanthemum L.

(Aizo aceae) and were also reported from Senecio

bahioides Hook. & Am., Astragalus L. and Cristaria

Cav. Champion (1895) reported L. australis adults

from the roots of ‘grass and herbage’. Lagrioida

brounii (Fig. 1B–D) is a nocturnal species found scur-

rying on sand (pers. obs.), beneath and under bark of

driftwood (Pascoe 1876, Emberson 1998), amongst

dry leaf litter in foredunes (pers. obs.), and collected

by pulling grasses and shaking their roots over

a sheet (Hudson 1923). It may also occur on flowers of

introduced plants (Hodge 2017) and is one of the most

common species found year-round in marine strand-

lines (Hodge et al. 2019) and in decaying kelp (Macro-

cystis pyrifera (L.) C. Ag.) (Inglis 1989). Lagrioida

nortoni adults were found on leaves, spikelets and

stems of Paniceum racemosum Spr. and Spartina

ciliata Kunth (Poaceae) and larvae in the laboratory

fed on dead leaves and root debris. Relatively large

numbers of adult Lagrioida tasmaniae sp. nov. and

a lesser number of L. australis were collected, pri-

marily at night, from marram grass (Ammophila

arenaria (L.) Link) and Spinifex sericeus R. Br. on

several Tasmanian beaches.

Key to the species of Lagrioida

1. Megasetae of upper surfaces inclined to decumbent,

slender, cylindrical and relatively widely spaced,

each one slightly overlapping the one behind it and

well separated from that on either side; fine,

erect microsetae often present but varying in abun-

dance; hind wing with anterior edge declined just

beyond radial bar, so that apical field is absent;

colour highly variable, ranging from yellow to black

and sometimes with yellow spots on darker re-

gions of elytra (Fig. 3); widely distributed in New

Zealand . . . . . . . . . . . . . . . . . . . . . L. brounii Pascoe

–. Megasetae of upper surfaces decumbent to

appressed, often somewhat thickened, flattened and

narrowly spaced, each one more strongly overlap-

ping the one behind it and closer to that on either

side; erect microsetae variable, but often rare or

absent; hind wing usually with well-developed apical

field, but sometimes very short; colour yellow to

black, but never with elytral spots . . . . . . . . . . . . . . 2

2. Elytra with numerous erect microsetae (Fig. 4D);

length usually less than 4.00 mm; antennomere 9 short-

er than 10; elytra less than 2.0 × as long as wide; ante-

rior edge of mesoventrite without elevated area or

carina; upper surfaces brown to black and legs some-

times yellowish-brown (Fig. 4E); Australia: TAS and

VIC . . . . L. tasmaniae Lawrence & Ślipiński sp. nov.

–. Elytra without or with very few erect microsetae;

antennomere 9 always longer than 10; IF length less

than 4.00 mm THEN elytra more than 2.0 × as long

as wide and anterior edge of mesoventrite with

curved carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Anterior end of mesoventrite with triangular or

V-shaped, heavily sclerotised elevation (Fig. 6D);

total length usually between 3.5 and 5.5 mm with a

mean length between 4.38 and 4.48 mm; elytral

megasetae more or less thickened and flattened;

antennae more than 2.4 × as long as head width;

apical field of hind wing well-developed in both sex-

es; tegmen less than 4 × as long as wide; penis less

than 5 × as long as wide with converging basal

struts; Australia and Norfolk Island . . . . . . . . . . . . 4

268 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

–. Anterior end of triangular mesoventrite without

sclerotised elevation; IF total length greater than

4.0 mm THEN antenna less than 2.3 × as long as

head width, apical field of hind wing reduced or

absent in males of some species; tegmen more than

4 × as long as wide and penis more than 5 × as long

as wide with parallel basal struts; southern South

America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4. Raised area on mesoventrite in the shape of an

inverted V (Fig. 6D); antennomere 3 less than 1.5 ×

as long as 2; apical antennomere more than 1.4 × as

long as wide; apical labial palpomere narrowly

rounded at apex; pronotal disc with one pair of

curved, basolateral sulci; hind wing with wedge cell

more than 3 × as long as wide and apically trun-

cate; tegmen with basale asymmetrically constricted

near base; gonocoxites less than 3 × as long as

combined width; colour variable but often reddish-

brown to dark brown; elytral megasetae tending

to form clusters, creating a mottled appearance

(Fig. 4A,B); southeastern coastal Australia and Tas-

mania . . . . . . . . . . . . . . . . . . . L. australis Champion

–. Raised area on mesoventrite in the shape of a trian-

gle; antennomere 3 more than 1.6 × as long as 2; api-

cal antennomere less than 1.4 × as long as wide; api-

cal labial palpomere truncate at apex; pronotal disc

with two pairs of curved, basolateral sulci; hind wing

with wedge cell less than 3 × as long as wide and

apically acute; basale not constricted at base; gono-

coxites more than 3.5 × as long as combined width;

colour yellowish-orange to reddish-orange (Fig.

4C,F); elytral megasetae not forming clusters;

Norfolk Island . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . L. norfolkensis Lawrence & Ślipiński sp. nov.

5. Total length greater than 4.5 mm; antennae less

than 2.3 × as long as head width; pronotal disc with

two pairs of short, curved basolateral sulci; elytra

more than 3.15 × as long a pronotum; apical field of

hind wing reduced in males so that radial cell and

medial fleck both very close to wing apex; tegmen

more than 4.5 × as long as wide; penis more than

0.85 × as long as tegmen and more than 8 × as

long as wide; gonocoxae more than 2.0 × as long as

their combined width; colour yellow to grey or black;

Chile . . . . . . . . . . . . L. rufula Fairmaire & Germain

–. Total length less than 4.0 mm; antennae more than

2.3 × as long as head width; pronotal disc without

short, curved, basolateral sulci; elytra less than 3.15

× as long as pronotum; apical field of hind wing well

developed in both sexes; tegmen less than 4.0 × as

long as wide; penis less than 0.75 × as long

as tegmen and less than 5 × as long as wide; gono-

coxae less than 2.0 × as long as their combined

width; colour .yellow to brownish-yellow; southern

Brazil . . . . . . . . . . . . . . . . . . . L. nortoni Costa et al.

Lagrioida australis Champion

(Figs 1A; 4A, B; 5A; 6A, 7A, B; 8F)

Lagrioida australis Champion, 1895: 238. Type locality: Sandy Bay,

near Hobart, Tasmania, Australia.

Diagnosis. This species resembles L. norfolkensis

in having the anterior end of the mesoventrite with

a sclerotised elevation and the hind wing fully devel-

oped with a long apical field in both sexes. It differs

from L. norfolkensis in that the elytral megasetae

form clusters creating a mottled effect, the hind wing

has an apically truncate wedge cell and the basale is

asymmetrically constricted near base, It differs from

L. tasmaniae is its larger size, very few erect elytral

microsetae and antennomere 9 longer than 10. The two

South American species are relatively small, lack the

sclerotised elevation of the mesoventrite and often

have the apical field of the hind wing reduced or absent

in male.

Redescription. Length 3.30–5.60 (4.48 ± 0.51) mm

(n = 12). Body 2.30–2.71 (2.52) × as long as wide;

colour yellowish-brown to grey or black; dorsal sur-

faces densely clothed with slightly thickened, whitish,

decumbent megasetae with irregular patches on elytra

closer together producing a mottled effect (Fig. 4A,B);

microsetae absent or rare.

Head 1.33 × as long as wide; neck 0.85 × as wide

as head. Eye 0.44 × as long as head width. Labrum 0.6

× as long as wide, with sides weakly curved and slight-

ly converging to subtruncate apex. Antenna 2.6 × as

long as head width; third antennomere 1.38 × as long

as second; apical antennomere 1.33 × as long as and

0.64 × as wide as 10 and 1.78 × as long as wide, widest

at middle and narrowly rounded apically. Apical maxil-

lary palpomere with outer edge 1.67 × as long as inner

edge. Apical labial palpomere 2.67 × as long as wide,

widest at middle and narrowly rounded at apex. Men-

tum 0.54 × as long as wide, with angulate sides and

subtruncate apex. Submental peduncle 0.4 × as long as

wide. Gula 0.8 × as long as wide, sutures slightly

curved outwardly.

Pronotum 1.09–1.25 (1.14) × as long as wide,

widest at anterior third; sides weakly constricted at

posterior fourth; anterior edge subtruncate; posterior

edge broadly rounded with narrow marginal bead; disc

with single pair of curved basolateral sulci; megapunc-

tures coarse and dense, circular to slightly longitudi-

nally oval, with interspaces usually less than a third as

great as a puncture diameter. Prosternum in front of

coxae 1.43 × as long as mid length of procoxal cavity,

moderately convex. Prosternal process with narrowest

point 0.14 × as wide as mid length of coxal cavity,

slightly expanded and truncate apically.

Pterothorax. Elytra 1.73–2.08 (1.91) × as long as

combined width and 2.82–3.29 (3.13) × as long as

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 269

pronotum; almost conjointly rounded at apex; humeri

weakly developed and concealing broadly rounded

anterolateral angles; disc with megapunctures (Figs

5A, 6A) not or slightly larger than those on pronotum

and equally dense. Mesoventrite 0.61 × as long as

wide, widest posteriorly and acute anteriorly, with

mesanepisterna contiguous at midline; anterior region

of disc with small. subapical, V-shaped raised area;

mesoventrital process 0.68 × as long as body of

ventrite and 0.18 × as wide as shortest diameter of

a mesocoxal cavity. Metaventrite 0.57 × as long as

wide; discrimen 0.50 × as long as metaventrite; poste-

rior edge obliquely sinuate on either side of angular

median emargination. Metanepisternum 4.1 × as long

as wide, widest at apical end and narrowing to truncate

apex. Metacoxae separated by 0.75 × shortest (longitu-

dinal) diameter of one, each coxa 0.25 × as long as

wide. Hind wing about 2.5 × as long as wide; apical

field about 0.5 × total wing length; radial cell 0.1 ×

wing length, 2.4 × as long as wide, with strongly

oblique inner edge, broadly obtuse inner basal angle;

cross-vein r3 short, oblique and fused for most of its

length with r4, which is strongly curved towards wing

base; radio-medial loop acute; base of RP very short

and oblique; medial spur strongly curved posteriorly

and ending well before wing margin; medial field with

four free veins and divided medial fleck; wedge cell 1.83

× as long as medial spur, 3.67 × as long as wide, api-

cally truncate; anal lobe well-developed with one vein.

Abdomen 1.46 × as long as wide; ventrite 1 slight-

ly shorter than 2 and 0.62 × as long as metaventrite

length; ventrite 5 (sternite VII) 0.33 × as long as wide,

broadly rounded at apex. Tergite VII (pygidium) 0.85 ×

as long as wide, strongly rounded apically, apical half

of disc moderately sclerotised. Sternite VIII in male

0.40 × as long as wide, widest at base with sides dis-

tinctly converging to slightly emarginate apex; disc

with transverse, sub-basal carina. Tergite VIII in male

0.67 × as long as wide, strongly concave at base, sub-

angulate apically, lightly sclerotised at apex only. Seg-

ment IX with pregenital ring wide at base and subacute

apically. Tegmen (Fig. 7A) 3.2 × as long as wide, with

basale 1.35 × as long as wide, widest at apex, asym-

metrically constricted near subtruncate base, sides

subparallel from basal third to apex, which is weakly

biconcave; apicale 1.26 × as long as basale and 2.0 ×

as long as wide, widest at base with sides gradually

converging to apical fourth, then more strongly so to

acute apex. Penis (Fig. 7B) 0.53 × as long as tegmen,

2.92 × as long as wide, widest at base, sides slightly

converging to apical fourth, then abruptly narrowed to

form short, apical lobe, sides of which are subparallel

for basal half then strongly converging to acute apex;

base with anteriorly converging struts, each 0.3 × as

long as body of penis. Sternite VIII in female 1.14 × as

long as wide, strongly biemarginate at base with

sclerotised rim; sides slightly converging to about mid-

dle, then straight and converging to narrow, emar-

ginate apex. Tergite VIII in female 0.8 × as long as

wide, strongly concave basally, with sides straight and

subparallel to truncate apex. Ovipositor 7.0 × as long

as wide, paraprocts 1.96 × as long as gonocoxites,

which are 2.25 × as long as combined width; gonocox-

ite divided into small proximal lobe and distal lobe four

as long with narrowly rounded apex; gonostyli terminal

but slightly subapical, 0.2 × as long as gonocoxite, par-

allel-sided and 5 × as long as wide.

Types

Syntype. Hobart Tasmania J.J. Walker

Lagrioida australis Cham./ G.C. Champion Coll. B.M.

1927-409/ NHMUK15011453 plus QR Code” (NHML, lec-

totype here designated, left specimen of two mounted

on the same card). Paralectotype: same data as the lec-

totype (NHML, right specimen on the mounting card

with NHMUK15011453 and QR Code).

Specimens examined

NSW: Bateman’s Bay,

12.i.1966, sand plants, E. Britton (4, ANIC); Botany

21.i.1901 (3, MVM); Brindle Creek, Wiangaree St. For.,

740m, 29.ii–3.iii.1980, A. Newton, M. Thayer (1, ANIC);

Broulee, 24.vi.1973, Z. Liepa (1, ANIC); Broulee, beach

at Candagan Ck., 27.x.1982, sand dunes, J. T. Doyen, J.

F. Lawrence (10, ANIC, QMB); Bundjalung N. P., Black

Rock rest area, 21.xi.1982, J. T. Doyen (1, ANIC); Coff’s

Harbour, 4.iii.1980, in and under seaweed on sandy

beach, A. Newton, M. Thayer (4, ANIC, FMNH);

Coledale (34.17’S, 153.09’E), 17.i.1987, under wet sea-

weed on beach, A. Newton, M. Thayer (6, ANIC, FMNH);

Congo, 8 km SbyE Moruya, xii.1981–ii.1982, window-

pane gutter trap, M. S. Upton (24, ANIC); Durras

Beach, N of Batemans Bay, 14.x.1973, Z. Liepa

(2, ANIC); Forster, 27.xii.1968, beach dunes, I. C. Yeo

(1, QDAF); Lord Howe Island, 12.i.1985, G. Borne-

missza (1, ANIC); Lord Howe Island, 21.i.1985, J. Bal -

derson (1, MVM); Lord Howe Island, Blinky Beach,

2.xii.1966, beach vegetation, E. Britton (1, ANIC);

Malua Bay, McKenzies Beach, 12.x.1983, seaweed on

beach, R. B. Halliday (2, ANIC); Murramarang N. P., nr.

Batemans Bay, 22.xii.1986, FMHD #86-657, in and

under rotten seaweed on beach, A. Newton, M. Thayer

(5, ANIC, FMNH); Newport, 31.vi.1924, H. J. Carter (3,

ANIC); No specific locality, French Collection (4, MVM);

No specific locality, Blackburn Collection (1, MVM);

Pambula River estuary, 3.viii.1973, Z. Liepa (3, ANIC);

Redcliffe?, ii.1931, Wassell (1, ANIC); Sapphire Beach,

9km N Coff’s Harbour (30.14’S, 153.09’E), 1m, 13.i.1987,

in and under moist fresh seaweed on beach, A. Newton,

M. Thayer (1, ANIC); Smith’s Lake, 20 km S Forster,

1m, 27.viii.1982, SBP116, beach drift, S. & J. Peck (7,

ANIC, CMN); Stanwell Park, iii.19094, HJC (3, MVM);

Station Creek, Yurargir N.P., 20.xi.1982, at light, J. & E.

Doyen (1, ANIC); Sydney, J. Armstrong (2, ANIC);

Sydney, K.K. Spence (3, ANIC); Sydney, sea beach

(2, QDAF); Sydney, Lea (2, MVM); Terrigal, 9.xii.1976,

270 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 271

Z. Liepa (2, ANIC); Tomakin Beach, 26.x.1982, sand

dunes at night, J. Doyen, J. Lawrence (2, ANIC); Waian-

garee St. For., Brindle Creek, 740m, 29.ii–3.iii.1980,

subtrop. rainforest, A. Newton, M. Thayer (1, ANIC);

Yarra Bay, x.1931, K. K. Spence (2, ANIC); Yarra

Beach, K. Spence (2, ANIC); Yuragir N. P., Station

Creek, 20ix, 1982, at light, J. T. Doyen (1, ANIC). Rock-

dale nr Sydney, 6-9.viii.1905, G.E. Bryant (13, NHML).

QLD: Bundaberg, Koebele Colln. (1, CAS); Fraser Is -

land, 21.iv.1973, A. Postle (1, QDAF). VIC: Carrum,

19.xii.1920, C. Oke (2, MVM); Chelsea, 29.ix.1919, F. E.

Wilson (4, MVM); Frankston, 21.iv.1918, H. Pottinger

(5, QDAF); Mordialloc, JCG (1, MVM); St. Kilda, 26.v.

1923, C. Oke (1, MVM) TAS: Bellerive Beach, Bellerive

Dunes (42.878857S, 147.377399E), 23.iv.2022, marram

grass, day, L/ Forster (3, TMAG); Blythe River, i.1937,

C. Davis (1, ANIC); Hobart, J. J. Walker, Fenyes Colln.

(CAS); Hobart, Lea (3, MVM); Snug Beach, 22.iv.2022,

marram grass, night, L. Forster (1, TMAG); Stony

Head, Seaview Rd. sentry hut (41.0153S, 146.9617E),

21.iii.2021, beach sand among edge dune plants, L. For -

ster (1, TMAG); Kelso, beach (2, ANIC); No specific

locality (2, MVM); Hobart, J.T. Walker (NHML).

Larval material examined

New South Wales:

Tomakin Beach, 3.x.1982, beach sand, reared in lab,

J. T. Doyen (7 larvae, 2.8–6.0 mm; ANIC); same data,

taken with adults on sand at night, J. F. Lawrence

(1 larva, 5 mm; ANIC).

Notes.In the original description the length was

given as 3.25–5.25 mm and the author noted that the

numerous examples varied greatly in size and colour.

Very small specimens were thought to be males, but

they actually belong to L. tasmaniae described below.

Lagrioida brounii Pascoe

(Figs 1B–F; 3A–E; 5D, 6B, 7C, 8E)

Lagrioida brounii Pascoe, 1876: 58. Type locality: Tairua, Coroman-

del Peninsula, North Island, New Zealand.

Lagrioida brouni: Werner & Chandler 1995: 25.

Diagnosis

In addition to their restriction to New

Zealand, specimens of L. brounii differ from those of

all other Lagrioida in having the megasetae of the

upper surfaces slender, cylindrical and relatively wide-

ly spaced, each one well separated from that on each

side and only slightly overlapping those on the same

row; the colour is also highly varied and the hind wing

may be reduced, lacking an apical field.

Redescription

Length 4.20–6.10 (4.85 ± 0.66) mm

(n = 14). Body 2.31–2.61 (2.49) × as long as wide; col-

o ur highly varied (Fig. 3), yellow to dark brown or black,

elytra sometimes yellowish-brown with yellow spots;

dorsal surfaces moderately densely clothed with de -

clined or decumbent, slender, cylindrical, yellow or pale

megasetae and scattered very fine, erect microsetae.

Head 1.14 × as long as wide; neck 0.83 × as wide

as head. Eye 0.48 × as long as head width. Labrum 0.5

× as long as wide, with sides weakly curved and slight-

ly converging to subtruncate apex. Antenna 2.55 × as

long as head width; third antennomere 1.33 × as long

as second; apical antennomere 1.33 × as long as and

0.86 × as wide as 10 and 1.6 × as long as wide, widest

at middle and narrowly rounded apically. Apical maxil-

lary palpomere with outer edge 1.5 × as long as inner

edge. Apical labial palpomere 2.25 × as long as wide,

widest at middle and narrowly rounded at apex. Men-

tum 0.53 × as long as wide, angulate laterally, with

apex subtruncate. Submental peduncle 0.33 × as long

as wide. Gula 0.70 × as long as wide, sutures slightly

curved and anteriorly converging.

Pronotum 1.07–1.19 (1.13) × as long as wide;

widest at anterior fourth; sides weakly constricted at

posterior fourth; anterior edge broadly rounded; poste-

rior edge weakly curved with narrow marginal bead;

disc with two pairs of short, slightly curved, basolater-

al sulci; megapunctures moderately coarse and dense,

circular to slightly longitudinally ovate and usually

separated half a puncture diameter. Prosternum in

front of coxae 1.32 × as long as mid length of procoxal

cavity, moderately convex. Prosternal process at nar-

rowest point 0.13 × as wide as mid length of coxal cav-

ity, barely expanded and subtruncate apically.

Pterothorax. Elytra 1.73–1.95 (1.85) × as long as

combined width and 2.50–3.20 (2.87) × as long as

pronotum; widest at middle with sides moderately

curved and apices conjointly rounded; humeri weak,

flattened and concealing broadly rounded anterolater-

al angles; disc with megapunctures (Figs 5D, 6B) slight-

ly larger, more circular than those on pronotum and

separated by less than a third of a puncture diameter.

Mesoventrite 0.6 × as long as wide, widest posteriorly

and narrowly rounded anteriorly, with mesanepisterna

contiguous at midline; anteror region of disc with nei-

ther curved carina nor elevated area; mesoventrital

process 0.75 × as long as body of ventrite and 0.25 ×

as wide as shortest diameter of mesocoxal cavity.

Metaventrite 0.35 × as long as wide; discrimen 0.49 ×

as long as metaventrite, posterior edge obliquely sinu-

ate on either side of angular median emargination.

Metacoxae separated by 0.7 × shortest (longitudinal)

diameter of one, each coxa 0.34 × as long as wide. Me -

tanepisternum 4.7 × as long as wide, widest anterior-

ly, sides straight and apex subtruncate. Hind wing

about 3.2 × as long as wide; anterior edge abruptly

narrowed at apical fourth; apical field absent; wing

apex broadly rounded, consisting entirely of medial

field; radial cell absent; cross-veins r3 and r4 absent;

radio-medial loop more or less acute; RP base absent;

medial spur short and straight; medial field with three

free veins and large, medial fleck occupying entire apex

of field; wedge cell usually present; anal lobe absent.

272 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

Figure 3. Lagrioida brounii, habitus, representing various colour variants.

Abdomen 1.35 × as long as wide; ventrite 1 slight-

ly shorter than ventrite 2 and 1.22 × as long as

metaventrite length; ventrite 5 (sternite VII) 0.36 × as

long as wide, broadly rounded apically. Tergite VII

(pygidium) relatively lightly sclerotised, 0.81 × as long

as wide, widest at base with sides straight, slightly con-

verging to apical fifth, then more strongly convergin to

broadly rounded, setose apex. Sternite VIII in male 0.4

× as long as wide, widest at base, which is deeply con-

cave, with paired transverse, slightly curved struts not

meeting at midline; sides strongly converging to apex,

which is weakly emarginate forming two rounded

lobes; disc with median membranous strip. Tergite VIII

in male 0.91 × as long as wides, subtruncate at base

with sides straight and slightly converging and apex

produced and subangulate. Segment IX with pregenital

ring moderately broad at base but narrowly acute at

anterior apex. Tegmen (Fig. 7C) 3.16 × as long as wide;

basale 1.23 × as long as wide, widest at apex, subtrun-

cate at base with sides slightly diverging to basal

fourth and then subparallel to apex; apicale 2.07 × as

long as basale and 1.58 × as long as wide, widest at

base with sides very weakly sinuate and slightly con-

verging to apical fifth, then more abruptly converging

to narrowly acute apex lined laterally with numerous

spicules. Penis 0.6 × as long as tegmen, 3.16 × as long

as wide, widest near base with sides subparallel to api-

cal fifth, then more abruptly narrowed to form sub-

acute apical lobe; base with slender, anteriorly con-

verging struts 0.28 × penile length. Sternite VIII in

female 0.75 × as long as wide with biemarginate base

lined with sclerotised rim and broadly rounded apex.

Tergite VIII in female 0.50 × as long as wide; base

emarginate with two oblique basal struts not meeting

at midline. Ovipositor 5.0 × as long as wide; paraprocts

2.2 × as long as gonocoxites, which are 1.35 × as

long as combined width; gonocoxites divided into

small proximal lobe, as long as wide with baculum dis-

tinctly cvurved, and distal lobe four × as long, gradual-

ly narrowing to subacute apex; gonostyli terminal,

0.14 × as long as gonocoxites, parallel-sided and

4.6 × as long as wide; ventral sclerite 0.64 × as long as

gonocoxite.

Types

“Lectotype/ Type/ N.Z., Tairua/ Lagrioida

Brounii type Pasc/ Pascoe Coll. 93-60/ Lagroida

Brounii, Pasc/ NHMUK14088849 + QRCode”(NHML,

lectotype and paralectotype; selected by F. G. Werner

in Werner & Chandler (1995)).

Specimens examined. New Zealand: North

Island: Northland: Cooper’s Beach, near Manganui,

Doubtless Bay, 16.xi.1931, grasses (4, ANIC); Motuopao

Island, 3.xii.1983, Phormium tenax, J. C. Watt

(3, NZAC); Omamari Beach, N of Dargaville, 5.iii.1969,

J. E. Tobler (1, CAS); Ruakaka Beach, 13.xii.1951, A. E.

Brookes (2, NZAC); Spirits Bay, xi.1967, sand dunes,

J. McBurney (2, NZAC); Sweetwater, 90-Mile Beach,

2.ii.1962, B. M. May (2, NZAC); Taipa, 5.vii.1983,

J. Doyen (2, CAS); Whangarei Heads, Ocean Beach

(35.50’S, 174.34’E), 13.ii.2000, at night on sand, G. Hall,

R. Leschen (1, NZAC); Auckland: Helensville, 26.x.

1992, sandy sea beach, under logs, A. Larochelle, M.-C.

Lariviere (1, NZAC); Karekare Beach, 28.iii.1975, in

sand under kelp, B. A. Holloway (1, NZAC); Muriwai

Beach, iii.1968, under log, J. C. Watt (1, NZAC); Muri-

wai Beach, NW of Auckland, 12.ix.1932 (1, ANIC); Tola-

ga Bay, 2.iii.1972, G. Kuschel (2, NZAC); Waipoua

Beach, 13.x.1967, under seaweed, storm drift line, J. C.

Watt (1, NZAC); Waipou Cove, 6.xii.1961, G. Kuschel

(1, NZAC); Whatipu, xi.1963, sand dunes, lupin litter,

B. M. May (1, NZAC); Bay of Plenty: Oroiti Beach,

10.iii.1993, J. S. Dugdale (1, NZAC); Whangaparoa,

23.xi.1992, sandy silty seaside under wood debris,

A. Larochelle (1, NZAC); Whangaparoa Beach, 25–26.

xi. 1992, B. I. P. Barratt (4, NZAC); Waikato: Waikato

Heads, 1.i.1959, B. M. May (1, NZAC); Taranaki: Waiiti

Beach, 28.x.1968, beating flax, K. S. Fox (2, NZAC);

Hawkes Bay: Ocean Beach, 24.i.1.1991, pit traps, C.

Duffy (1, NZAC); Pourerere Beach, 20.iv.1984, ex dry

sand, J. C. Watt (5, NZAC); Whakaki Beach, 9.i.1962,

G. Kuschel (2, ZNAC); Wanganui: Himatangi Beach,

2.i.1975, under logs, J. C. Watt (1, NZAC); Wellington:

Tirahi Bay, 7.xii.1941, D. Spiller (5, NZAC); South

Island: Nelson: Farewell Spit, 2 km beyond light

house, 28.i.1967, A. K. Walker (2, NZAC); Heaphy

Track, Heaphy Hill, 6.xi.1999, on tree trunks at night,

R. Leschen & G. Hall (1, NZAC); Patarau, Sandhills Ck.,

26.x.1968, sand dunes, J. I. Townsend (1, NZAC); Pat-

terus Rock, 15.i.1966, A. K. Walker (1, NZAC); Pohara

Beach, Takaka, 18.i.1973, G. Kushel (3, NZAC); Rabbit

Island, 1.ii.1961, W. P. Thomas (1,, NZAC); Tahunanui

Beach, 16.iii.1960, E. S. Gourlay (1♂, ANIC); Tahu-

nanui, 28.x.1971, J. C. Watt (5, NZAC); Westport Beach,

20.i.1935, E. S. Gourley (1, NZAC); Buller: Mouth of

Buller River (41.43’S, 172.33’E), 11.ii.1999, on beach, R.

Hoare (4, NZAC); Mid Canterbury: Banks Peninsula,

Balys Road, sand dune spit, 14.xii.1977, E. Schlinger

(1, NZAC); Spencer’s Beach, 10.x.1973, sand beach,

under logs, D. S. Horning (1, NZAC); Westland:

Jackson’s Bay, Neils Beach, 28.ii.1966, G. Kuschel

(1, NZAC); Dunedin: Aramoana, Otago Harbour, 15.xii.

1993, Stop 93-11B, sandspit, D. H. Kavanaugh (2, CAS);

Taieri Mouth, 24.xii.1995, underside of beach drift-

wood, J. Nunn (1, NZAC); Southland: Kaka Point, near

Owara, 3.xii.1996, J. Nunn (1, NZAC); Fiordland: Blue-

cliffs Beach, Waitutu, 20.ii.2003, J. Nunn (1, NZAC);

Chatham Islands: Awatotara, 600’, 19.ii.1967, J. S.

Dugdale (1, NZAC); Hapupu, 27–28.ii.1967, collected at

night, G. W. Ramsay (2, NZAC); Maunganui Beach,

3.iii.1967, G. W. Ramsay (1, NZAC); Northern Beach,

28.ii.1967, E. W. Valentine (1, NZAC); Owenga,

25,28.ii.1967, sand dunes, kelp, A. K. Walker (6, ANIC);

Waitangi, 9,ii, 13.ii, 24.ii.1967, sand dunes at night,

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 273

J. S. Dugdale, G. Kuschel, A. K. Walker (25, NZAC,

ANIC); Waitangi, dunes, 8.iii.1991, R. Craw (4, NZAC).

Note. This species is found along the entire coast-

line of New Zealand and is considered here as a single

species, though it is possible that it represents a clus-

ter of closely related species. Based on external exam-

ination of the specimens noted above, it is not possible

to separate discrete elements, and specimens of differ-

ent sized and colour patterns often geographically

overlap. From the small number of complete dissec-

tions made, it was noted that the hind wings in both

sexes are reduced (Fig. 8E). Although it is possible that

some individuals are fully winged, this does suggest

reduced gene flow between populations.

Lagrioida norfolkensis Lawrence & Ślipiński sp. nov.

(Figs 4D, E; 5F, 6C, D, G, 8A)

Diagnosis

This species is most similar to L. aus-

tralis, both of which have a triangular or V-shaped ele-

vation at the anterior end of the mesoventrite, elytral

setae somewhat thickened and flattened and the hind

wing with a well-developed apical field. It differs from

L. australis in the shape of the raised area at the ante-

rior end of the mesoventrite, longer third antennomere,

shorter apical antennomere and elytral megasetae not

forming clusters. L. tasmaniae is a much smaller

species with numerous erect microsetae, while the two

South American species have no sclerotised mesoven-

trital elevation.

Description

Length 3.60–5.60 (4.38 – 0.59) mm

(n = 10). Body 2.34–2.63 (2.50) × as long as wide;

colour (Fig. 4C,F) uniformly yellow to brownish-yellow

or with head and pronotum slightly darker than elytra

or undersides; dorsal surfaces densely setose, with

slightly thickened, whitish, decumbent megasetae of

more or less uniform thickness and not forming a pat-

tern on elytron; microsetae very rare or absent Head

1.23 × as long as wide; neck 0.92 × as wide as head.

Eye 0.48 × as long as head width. Labrum 0.50 × as

long as wide, sides very slightly converging to subtrun-

cate apex. Antennae 2.60 × as long as head width;

third antennomere 1.73 × as long as second; apical

antennomere 1.25 × as long and 0.8 × as wide as

preapical one, 1.25 × as long as wide, widest at middle

and broadly rounded at apex. Apical maxillary

palpomere with outer edge 2.5 × as long as inner edge,

outer apical angle acute and inner apical angle broad-

ly obtuse. Apical labial palpomere 2.25 × as long as

wide, widest near base, with subtruncate apex. Men-

tum 0.52 × as long as wide, widest and angulate at mid-

dle, with apex truncate. Submental peduncle 0.35 × as

long as wide. Gula 0.90 × as long as wide, sutures

slightly curved and converging anteriorly. Pronotum

1.05–1.19 (1.12) × as long as wide; widest at anterior

third; sides weakly constricted at posterior fourth;

anterior edge weakly rounded or subtruncate; posteri-

or edge very slightly curved with fine margin; disc with

two pairs of short, curved, basolateral sulci; megapunc-

tures circular to slightly oval and separated by less

than a third of a puncture diameter. Prosternum in

front of coxae 1.43 × as long as mid length of procoxal

cavity. Prosternal process at narrowest point 0.11 × as

wide as mid length of procoxal cavity, slightly concave

at middle with subtruncate apex.

Pterothorax. Elytra 1.76–2.01 (1.90.) × as long as

combined width and 3.00–3.26 (3.12) × as long as

pronotum; disc with megapunctures (Fig. 5F) slightly

larger than those on pronotum and usually circular but

equally dense. Mesoventrite 0.66 × as long as wide,

widest posteriorly and subacute anteriorly, with

mesanepisterna very narrowly separated at midline;

anterior region of disc with small, raised triangular

area; mesoventrital process 0.68 × as long as main

body of ventrite and 0.25 × as wide as shortest diame-

ter of mesocoxal cavity. Metaventrite 0.69 × as long as

wide, widest posteriorly; posterior edge sinuate on

either side of angulate emargination; discrimen 0.37 ×

as long as metaventrite. Metanepisternum 4.67 × as

long as wide, widest apically and obliquely truncate at

apex. Metacoxae separated by 0.67 × shortest (longitu-

dinal) diameter of one, each coxa 0.28 × as long as

wide. Hind wing about 2.7 × as long as wide; apical

field about 0.5 × total wing length; radial cell 0.06 ×

wing length, cross-vein r3 short, oblique and fused for

most of its length with r4, which is strongly curved and

complete; base of RP very short and oblique; medial

spur strongly curved and ending well before wing mar-

gin; medial field with four free veins and elongate-oval,

divided medial fleck; wedge cell 1.4 × as long as medi-

al spur, 2.63 × as long as wide, apically truncate; anal

lobe reduced.

Abdomen 1.3 × as long as wide; ventrite 1 slightly

shorter than 2 and 0.65 × as long as metaventrite

length; ventrite 5 (sternite VII) 0.36 × as long as wide,

broadly rounded at apex. Tergite VII (pygidium) 0.9 ×

as long as wide, with lateral edges curved with sclero-

tised rim and apex broadly rounded and setose. Stern-

ite VIII in male 0. 5 × as long as wide, widest at base,

sides converging to slightly emarginate apex. Tergite

VIII 0.67 × as long as wide, concave at base, broadly

rounded and densely setose apically. Segment IX with

pregenital ring broad at base, abruptly narrowed ante-

riorly and subacute at apex. Tegmen (Fig. 7D) 3.0 × as

long as wide; basale 1.26 × as long as wide, subtrun-

cate at base, with sides slightly curved and diverging to

apex; apicale 1.5 × as long as basale and 2.10 × as

long as wide, widest near base with sides slightly con-

verging to apical fourth, then more strongly so to acute

apex, lined laterally with numerous spicules. Penis 0.55

× as long as tegmen, 2.8 × as long as wide, widest at

274 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 275

Figure 4. Lagrioida species, habitus. (A, B) L. australis; (C, F) L. norfolkensis; (D, E) L. tasmaniae.

middle with sides slightly converging from middle to

apical fourth, then abruptly converging to form narrow

lobe, which is parallel-sided to about middle, then

abruptly narrowing to acute apex; base with slender,

converging struts 0.27 × as long as body of penis. Ster-

nite VIII in female 0.79 × as long as wide, with base

weakly biemarginate with sclerotised rim; sides sub-

parallel to about middle, then converging to narrowly

rounded apex; spiculum ventrale long, curved and

basally articulated. Tergite VIII in female 0.66 × as

long as wide, with base deeply emarginate with sclero-

tised rim; sides slightly converging to middle, then

abruptly converging to narrowly rounded apex. Ovipos-

itor 7.7 × as long as wide, paraprocts 1.94 × as long as

gonocoxites, which are 4.1 × as long as combined

width; coxites not clearly divided into lobes; coxital

bacula curved and basal; gonostyli terminal, 0.2 × as

long as gonocoxites, parallel-sided and 6 × as long as

wide.

Types

Holotype, ♂, “29.045S 167.58E, NORFOLK

ISLAND, Point Hunter Res., 29 Nov. 1984, T. A. Weir”

(ANIC).

Paratypes. Norfolk Island: Kingston, 7.xi.1967,

above high tide level, on grass at night, G. Kuschel

(47, NZAC, ANIC). Point Hunter Reserve, 29.xi.1984,

T. A. Weir (10, ANIC).

Lagrioida nortoni Costa, Vanin & Ide

(Figs 5B, 8C)

Lagrioida nortoni Costa, Vanin & Ide, 1995: 114. Type locality: Rio

Grande do Sul: Rio Grande, Praia do Cassino, Brazil.

Diagnosis

This species is about the same size as

L. tasmaniae but differs from that species in having

few, if any, erect elytral microsetae, antennomere 9

longer than 10, elytra more than twice as long as wide

and with a curved carina at the anterior edge of the

mesoventrite. It differs from both L. australis and

L. norfolkensis, not only ion size but in the lack of

a sclerotised elevation at the anterior edge of the meso -

ventrite. Unlike the Chilean L. rufula, the apical field

of the hind wing is well-developed in both sexes and

both aedeagus and ovipositor are shorter and broader.

Redescription

Length 3.10–3.60 (3.28 – 0.17) mm

(n = 4). Body 2.67–2.73 (2.97) × as long as wide; colour

uniformly yellow to brownish-yellow or with head and

pronotum slightly darker than elytra or undersides;

dorsal surfaces densely setose, with slightly thickened,

whitish, decumbent megasetae of more or less uniform

thicknes. not forming a pattern on elytron, and

microsetae moderately abundant but very short.

Head 1.20 × as long as wide. Neck 0.84 × as wide

as head. Eye 0.45 × as long as head width. Antenna

2.56 × as long as head width; third antennomere 0.91

× as long as second; apical antennomere 1.65 × as

long and 0.9 × as wide as preapical one, 1.38 × as long

as wide and widest at middle. Labrum 0.54 × as long as

wide, sides slightly curved, apex broadly rounded. Api-

cal maxillary palpomere with outer edge 2.28 × as long

as inner edge. Apical labial palpomere 2.67 × as long

as wide, widest at middle with narrowly rounded apex.

Mentum 0.44 × as long as wide, widest at middle, sides

subangulate with angle rounded and preceded by

notch; apex truncate. Submental peduncle 0.48 × as

long as wide. Gula 0.92 × as long as wide, sutures

slightly curved and converging anteriorly. Pronotum

1.07–1.16 (1.10) × as long as wide, widest at anterior

edge; sides gradually converging posteriorly; not con-

stricted at posterior fourth; anterior edge subtruncate;

posterior edge very slightly curved with fine margin;

disc without basolateral sulci; megapunctures usually

slightly longitudinally oval and separated by less than

a third of a puncture diameter; Prosternum in front of

coxae 1.11 × as long as mid length of procoxal cavity;

anterior edge weakly biconcave. Prosternal process at

narrowest point 0.08 × as wide as mid length of pro-

coxal cavity, slightly concave at middle with subtrun-

cate apex.

Pterothorax. Elytra 2.00–2.02 (2.02.) × as long as

combined width and 2.76–3.15 (2.97) × as long as

pronotum; disc with megapunctures (Fig. 5B) slightly

smaller than those on pronotum, but equally dense,

more or less circular and somewhat more variable in

size. Mesoventrite 0.67 × as long as wide, widest pos-

teriorly and subacute anteriorly, with mesanepisterna

narrowly separated at midline; anterior region of disc

with curved, transverse carina. Mesoventrital process

0.65 × as long as main body of ventrite and 0.23 × as

wide as shortest diameter of a cavity. Metaventrite 0.53

× as long as wide, widest posteriorly; discrimen 0.49 ×

as long as metaventrite; posterior edge very slightly

concave on either side of angulate emargination. Meta-

coxae separated by 0.67 × shortest (longitudinal)

diameter of one, each coxa 0.33 × as long as wide.

Metanepisternum 4.3 × as long as wide, with truncate

apex. Hind wing about 2.8 × as long as wide; apical

field about 0.5 × total wing length; radial cell 0.08 ×

wing length; cross-vein r4 strongly curved, fused at

base with r3; base of RP absent; medial spur slightly

curved and ending well before wing margin; medial

field with four free veins and elongate-oval, divided

medial fleck; wedge cell narrowly truncate or subacute;

anal lobe reduced.

Abdomen 1.2 × as long as wide; ventrite 1 slightly

shorter than 2 and 0.81 × as long as metaventrite

length; ventrite 5 (sternite VII) 0.3 × as long as wide,

broadly rounded at apex. Tergite VII (pygidium) 0.7 ×

as long as wide and 0.88 × as wide as ventrite 5 (ster-

nite VII), widest at truncate base, sides gradually con-

verging to broadly rounded apex; sides with sclerotised

rim on basal half, disc moderately sclerotised and

276 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 277

Figure 5. Lagrioida species, elytral punctures and setae as visible on slide mounted specimens. (A) L. australiae; (B) L. nortoni;

setose. Sternite VIII in male 0.68 × as long as wide,

widest at deeply emarginate base with sides converg-

ing to slightly emarginate apex. Tergite VIII in male

0.86 × as long as wide, widest at concave base, sides

very slightly converging to apical third, then more

strongly so to weakly angulate apex. Segment IX with

pregenital ring broad at base, gradually narrowed

anteriorly and subacute at apex. Tegmen 3.75 × as

long as wide; basale 1.4 × as long as wide, broiadly

rounded at base with sides subparallel from basal third

to subtruncate apex; apicale 1.67 × as long as basale

and 2.33 × as long as wide, widest at base with sides

gradually converging to subacute apex. Penis 0.7 × as

long as tegmen, 3.33 × as long as wide, widest at base

with sides gradually converging to narrowly acute

apex; basal struts parallel and 0.2 × as long as body of

sternite. Sternite VIII in female 0.73 × as long as wide

at biemarginate base, with sides gradually converging

to subacute apex. Tergite VIII in female 0.6 × as long

as wide, base deeply emarginate with sclerotised rim

obsolete at middle, sides parallel and apex truncate.

Ovipositor 4.2 × as long as wide, paraprocts 1.82 × as

long as gonocoxites, which are 1.22 × as long as com-

bined width; coxites each divided into small proximal

lobe, with slightly curved baculum, and distal lobe

four × as long; gonostyli 0.13 × as long as gonocoxites,

parallel-sided and 5 × as long as wide.

Types

“Brasil, Rio Grande do Sul: Rio Grande,

Praia do Cassino, dunas sobre Panicum racemosum,

16.IV.1993, Exp. MZSP col.” (MZSP). Holotype male,

not examined.

Specimens examined. Brazil: Male and female

paratypes from type locality (ANIC).

Lagrioida rufula Fairmaire & Germain

(Figs 6E–H)

Lagrioida rufula Fairmaire & Germain, 1860: 4. Type locality: San

Antonio, Chile. Fairmaire & Germain 1863: 235; Telnov & Degio-

vanni 2021: 146.

Lagrioida obscurella Fairmaire & Germain, 1860: 4. Type locality:

San Antonio, Chile. Champion 1895: 238 (syn.); Telnov & Degio-

vanni 2021: 146.

Diagnosis

Differing from L. tasmaniae in the

absence or near absence of erect microsetae on the ely-

tra and antennomere 9 always longer than 10 and from

L. australis and L. norfolkensis in the lack of a scle-

rotised elevation at the anterior end of the mesoven-

trite. If differs from L. nortoni in its larger size (more

than 4.5 mm), shorter antennae, and reduced apical

field of the hind wing in males.

Redescription

Length 4.10–8.00 (5.83 ± 1.14) mm

(n = 24). Body 2.27–2.76 (2.49) × as long as wide.

Colour yellowish-brown or reddish-brown to dark

brown or black; dorsal surfaces densely clothed with

slightly thickened, whitish, decumbent setae; microse-

tae short and relatively sparse.

Head 1.25 × as long as wide; neck 0.9 × as wide as

head. Eye 0.5 × as long as head width. Labrum 0.6 ×

as long as wide, with sides weakly curved and slightly

converging to subtruncate apex. Antenna 2 × as long

as head width; third antennomere 1.5 × as long as sec-

ond; apical antennomere 1.5 × as long as and equal in

width to preapical one, 1.9 × as long as wide, widest at

middle and narrowly rounded at apex. Apical maxillary

palpomere with outer edge 1.5 × as long as inner one.

Apical labial palpomere 2 × as long as wide, widest at

middle and narrowly rounded at apex. Mentum 0.45 ×

as long as wide, with sides angulate at basal third and

narrowing beyond this to truncate apex. Submental

peduncle 0.4 × as long as wide. Gula 0.8 × as long as

wide, sutures and slightly curved outwardly and con-

verging anteriorly.

Pronotum 1.06–1.21 (1.15) × as long as wide;

widest near anterior end; sides weakly constricted at

posterior fourth; anterior edge subtruncate; posterior

edge broadly rounded with narrow marginal bead; disc

with two pairs of short, curved, basolateral sulci; mega-

punctures usually separated by less than a third of

a puncture diameter. Prosternum in front of coxae

1.4 × as long as mid length of procoxal cavity, moder-

ately convex. Prosternal process at narrowest point

0.08 × as wide as mid length of coxal cavity, parallel-

sided and narrowly rounded apically, but concave at

middle and often concealed by procoxae.

Pterothorax. Elytra 1.74–2.17 (1.93) × as long as

combined width and 3.00–3.96 (3.49) × as long as

pronotum; disc with megapunctures (Fig. 5E) slightly

larger and more circular than those on pronotum, but

equally dense. Mesoventrite 0.6 × as long as wide,

widest posteriorly and subacute anteriorly, with

mesanepisterna almost contiguous at midline; anterior

region of disc with curved, transverse carina; mesoven-

tral process 0.7 × as long as body of ventrite, 0.25 × as

wide as diameter of a mesocoxal cavity. Metaventrite

0.5 × as long as wide; discrimen 0.5 × as long as

metaventrite, posterior edge obliquely sinuate on

either side of angular median emargination. Metacox-

ae separated by 0.7 × shortest (longitudinal) diameter

of one, each coxa 0.33 × as long as wide. Metanepister-

num 4.3 × as long as wide with truncate apex. Hind

wing in male about 2 × as long as wide, in female about

2.6 × as long as wide; apical field 0.25 (male) or 0.6

(female) × total wing length; radial cell 0.05 × wing

length; cross-vein r3 fused with r4 except for very short

stub; base of RP very short; medial spur straight and

ending well before wing margin; medial field with four

free veins and elongate-oval, divided medial fleck;

wedge cell 2.3 × as long as medial spur and 2.8 × as

long as wide, with truncate apex; anal lobe highly

reduced.

278 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 279

Figure 6. Lagrioida species. A–C: elytral punctures and setae as visible on slide mounted specimens. (A) L. australis; (B) L. brounii;

(C), L. norfolkensis; D–F anterior part of mesoventrite: (D) L. norfolkensis; (E) L. australis; (F) L. rufula. (G) L. L. norfolkensis, molar

surface.

Abdomen 1. 4 × as long as wide; ventrite 1 as long

as 2 and 0.65 × as long as shortest distance between

meso- and metacoxal cavities; ventrite 5 (sternite VII)

0.4 × as long as wide, broadly rounded at apex. Tergite

VII (pygidium) 0.9 × as long as wide and 0.7 × as wide

as pygidium, strongly rounded apically, apical half of

disc moderately sclerotised and densely setose. Ster-

nite VIII in male 0.55 × as long as wide, widest at base

with sides distinctly converging to slightly emarginate

apex; base distinctly emarginate with sclerotised rim

incomplete at middle. Tergite VIII in male 1.05 × as

long as wide, widest at deeply, angulately emarginate

base, with sclerotised rim not meeting at midline; sides

curved and converging to narrowly rounded apex. Seg-

ment IX with pregenital ring moderately wide at base,

abruptly narrowed at middle and almost parallel-sided

to acute apex. Tegmen (Fig. 7G,H) 5.0 × as long as

wide; basale 2.0 × as long as wide, broadly rounded at

base with sides almost parallel-sided to truncate apex;

apicale 1.2 × as long as basale and 2.15 × as long as

wide, widest at base with sides gradually converging to

subacute apex. Penis 0.9 × as long as tegmen, 6 × as

long as wide, widest near base, sides slightly converg-

ing to apical ninth, then more abruptly converging to

very narrowly rounded apex; base with subparallel

struts 0.9 × as long as body of penis. Sternite VIII in

female 1.1 × as long as wide; widest at base, which is

base deeply biemarginate with sclerotised rim; sides

slightly conve4rging to middle, then more strongly con-

verging to narrowly rounded, slightly emarginate apex.

Tergite VIII in female 0.7 × as long as wide at base,

which is deeply emarginate with sclerotised rim; sides

gradually converging to subtruncate apex. Ovipositor

7.0 × as long as wide, paraprocts 2.0 × as long as

gonocoxites, which are 2.6 × as long as combined

width; coxites each divided into smaller proximal lobe

and distal lobe 2.5 × as long; gonostyli terminal, 0.07 ×

as long as gonocoxites, parallel-sided and 3 × as long

as wide.

Types

Series of specimens from San Antonio, Chile

(MNHN). Not examined.

Specimens examined

Chile (region: province,

from north to south): Atacama: Copiapó: Bahia Blan-

ca, Caldera, 16–24.ix.2000, V. M. Diéguez (2, VMD); Puer-

to Viejo, S. Caldera (27°48’32”S, 70°19’31”W), 12.iii.

1997 (2, PVC). Atacama: Huasco: Huasco (dunas), 28-

29.ix.1981, M. Elgueta (1, MHNS). Coquimbo: Elqui: La

Herradura, 18.ix.1964, L. E. Peña (2, MHNS); La Sere-

na, 15.iv.1995, V. M. Diéguez (2, VMD); La Serena,

7.xii.1950, on beach, Ross & Michelbacher (5, CAS); La

Serena, 10km N, 21.ix.1999, M. Guerrero (MGC); Los

Choros, 5.i.2007, Fermin Kong (2, VMD); Morrillos,

(30°08’57.8”S, 71°22’15.5”W), 45 msnm, 12.xi.2010,

dunas, Astragalus, Cristaria, M. Elgueta (1, MHNS);

Punta Lengua de Vaca (30°17’42”S, 71°36’17”W),

18.iv.1997 (2, PVC); Punta de Choros, 10.ix.2004, M.

Guerrero (2, MHNS); Tongoy, 2.v.2002, M. Guerrero (3,

MGC); Tongoy, iii.1972, En plantas dunas (1, MHNS).

Coquimbo: Choapa: Chigualoco, 11.xi.1967, Senecio

bahioides Hook. & Arn., C. W. O’Brien (1, ANIC); Los

Vilos, 17.ix.1984, G. Carrasco (2, MHNS); Los Vilos, 18-

26.vii.1981, G. Arriagada (5, MHNS); Los Vilos,

25.viii.1966, E. Schlinger, M. Irwin (1, CAS). Los Vilos,

3km N, 28.x–1.xi.2012, pitfall c/cerveza, Diéguez & Ar -

riagada (1, VMD); Pichidangui, 1.iii.1970, G. Arriagada

(1, MHNS); Valparaíso: Petorca: Caleta la Ligua

(32°24’44.2”S, 71°.24’45.0”W), E. Pullalli Dunas,

4msnm, 30.xii.2012, Ambrosia, doca, M. Elgueta

(2, MHNS). Valparaíso: Valparaíso: Concon, 13.xii.

1970, J. Solervicens (2, MHNS); ConCon, X.1884 (6,

MHNS); Viña del Mar, ii.1897 (12, MHNS); Viña del Mar,

ix.1895, Edwyn P. Reed (4, CAS). Valparaíso: San

Antonio: Algarrobo, 2.vii.1951, G. Kuschel, L. Peña (4,

MHNS); Bucalemu, Yali, 18.x1998, P. Vidal (4, PVC);

Las Cruces, 10 mnsm Ambrosia (33°29’34.3”S,

73°38’11.8”W), 27.xii.2012, dunas, doca ciperáceas, M.

Elgueta (2, MHNS); Las Cruces, 19.ix.1974, G. Arriagada

(2, MHNS); Las Cruces, 20.vii & 24.viii.1959, Noodt (2,

MHNS); Matanzas, 16.ii.1998, P. Vidal G.H. (2, PVC); Pla-

ya Santo Domingo (33°36’26”S, 71°.37’56”W), 10.viii.

1997, P. Vidal (1, PVC); Rocas de Santo Domingo, 23.xi.

1968, sifting sand dunes, C. W. & L. O’Brien (1, ANIC);

Santo Domingo, 26.iii.2005, P. Vidal G.H. (1, PVC); Santo

Domingo, ii.1982, G. Kuschel (3, NZAC). Maule: Curicó:

Iloca, 4.xi.1983, M. Elgueta (2, MHNS). Los Rios: Valdi-

via: Chaihuin (40.03’40.5”S, 77°39’33.3”W), 26m,

15.i.2006, J. Mondaca E. (2, MHNS). Los Lagos: Chiloé:

Chepu, 5.x.1958, G. Kuschel (1, MHNS); Punta Tique,

29.i.1987, J. Solervicens (2, MHNS).

Note. Champion (1895) noted that the smaller,

darker specimens and larger, lighter ones were usual-

ly taken together, and considered the smaller species

(L. obscurella) to represent males of L. rufula.

Lagrioida tasmaniae Lawrence & Ślipiński sp. nov.

(Figs 4D, E; 5C; 7E, F; 8D)

Diagnosis. This species differs from all others in

the presence of numerous erect elytral microsetae, and

in the combination of small size, shorter ninth anten-

nomere and anterior edge of mesoventrite without an

elevated area or curved carina.

Description. Length 2.90–4.60 mm: males 2.90–

3.80 (3.25 ± 0.027) mm (n =10); females 3.50–4.60 (3.97

± 0.83) mm (n = 10). Body length/elytral width: males

2.39–2.63 (2.48); females 2.40–2.62 (2.49). Greatest

depth/elytral width: males 0.65–0.73 (0.69); females

0.65–0.74 (0.70). Colour (Fig. 4D,E) reddish-brown to

dark brown or black above, with legs, antennae and

mouthparts somewhat lighter, males darker than

females; dorsal surfaces densely clothed with slightly

thickened, whitish, decumbent megasetae interspersed

with scattered microsetae (Fig. 4D).

Head 1.11 × as long as wide. Neck 0.78 × as wide

as head. Eye 0.44 × as long as head width. Labrum 0.6

× as long as wide, with sides almost parallel-sided and

apex subtruncate. Antennae 2.48 × as long as head

width, third antennomere 1.39 × as long as second;

apical antennomere 1.6 × as long as and as wide as

preapical one, 2.0 × as long as wide, widest at middle

and narrowly rounded apically. Apical maxillary

280 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

palpomere with outer edge 1.13 × as long as inner

edge. Labial palpomere 2.25 × as long as wide, widest

at middle and narrowly rounded at apex. Mentum 0.43

× as long as wide, widest and angulate at middle with

truncate apex. Submental peduncle 0.36 × as long as

wide. Gula 0.61 × as long as wide, sutures curved and

converging anteriorly.

Pronotum: males 1.00–1.07 (1.05), females 1.03–

1.14 (1.07) × as long as wide, widest at anterior end,

not constricted at posterior fourth; anterior edge sub-

truncate; posterior edge broadly rounded with narrow

marginal bead; disc without basolateral sulci; mega-

punctures more or less ovate and usually separated by

less than a third of a puncture diameter. Prosternum in

front of coxae 1.32 × as long as mid length of procoxal

cavity, moderately convex. Prosternal process at nar-

rowest point 0.05 × as wide as mid length of coxal cav-

ity, narrowly rounded at apex.

Pterothorax. Elytra: males 1.76–1.96 (1.84), fe -

males 1.76–1.94 (1.86) × as long as combined width

and males: 2.77–3.18 (2.91), females 2.64 3.20 (2.97) ×

as long as pronotum; disc with megapunctures (Fig.

5C) slightly larger than those on pronotum but equally

dense. Mesoventrite 0.63 × as long as wide, widest pos-

teriorly and narrowly truncate anteriorly, with

mesanepisterna narrowly separated at midline; anteri-

or region of disc with neither curved carina nor elevat-

ed area; mesoventrital process 0.68 × as long as body

of ventrite and 0.2 × as wide as diameter of a mesocox-

al cavity. Metaventrite 0.53 × as long as wide; dis-

crimen 0.5 × as long as metaventrite, posterior edge

obliquely sinuate on either side of angular median

emargination. Metacoxae separated by 0.67 × shortest

(longitudinal) diameter of one, each coxa 0.36 × as

long as wide. Metanepisternum 4.0 × as long as wide,

truncate at apex. Hind wing about 2.25 × as long as

wide; apical field 0.35 × total wing length; radial cell

not clearly indicated; cross-vein r3 completely fused

with r4; base of RP absent; medial spur moderately

long and straight; medial field with four free veins and

elongate-oval, divided medial fleck; wedge cell twice as

long as medial spur, 3.75 × as long as wide and trun-

cate at apex; anal lobe highly reduced.

Abdomen 1.36 × as long as wide; ventrite 1 slight-

ly shorter than 2 and 0.82 × as long as metaventrite

length; ventrite 5 (sternite VII) 0.33 × as long as wide

and broadly rounded at apex. Tergite VII (pygidium)

0.83 × as long as wide and 0.85 × as wide as ventrite

5, strongly rounded apically, apical half of disc moder-

ately sclerotised and setose. Sternite VIII in male 0.76

× as long as wide, widest at base, which is deeply

emarginate. Tergite VIII in male 0.78 × as long as wide,

widest at base, which is deeply biemarginate with

sclerotised rim, incomplete at middle; sides gradually

converging to broadly rounded apex. Segment IX with

pregenital ring moderately wide at base but narrowing

to acute apex. Tegmen (Fig. 7E,F) 3.78 × as long as

wide with basale 1.56 × as long as wide, widest at

apex, broadly rounded at base with sides curved basal-

ly and subparallel apically; apicale 1.43 × as long as

basale and 2.22 × as long as wide, widest at base with

sides gradually converging to subacute apex. Penis

0.69 × as long as tegmen, 4.7 × as long as wide, widest

at middle, sides slightly converging to apical fifth, then

more abruptly narrowed to subacute apex; base with

anteriorly converging struts 0.2 × as long as body of

penis. Sternite VIII in female 0.92 × as long as wide,

widest at base which is emarginate but without sclero-

tised rim; sides subparallel to apical third, then con-

verging to slightly emarginate apex. Tergite VIII in

female 0.69 × as long as wide, widest at base which is

deeply emarginate with complete rim; sides converging

to subtruncate apex. Ovipositor 6.0 × as long as wide,

paraprocts 1.83 × as long as gonocoxites, which are

1.71 × as long as combined width; coxites each divided

into small proximal lobe and distal lobe 3.3 × as long;

gonostyli terminal, 0.15 × as long as gonocoxites, par-

allel-sided and 6 × as long as wide.

Types

Holotype, male. “42.908668 147.352220 /

TAS: Nutgrove Beach / Sandy Bay 222.iv.2022 / dunes,

marram grass / night L. Forster” (TMAG).

Paratypes. Australia: Tasmania: Bellerive Beach,

Bellerive Dunes (42.878857S, 147.337399E), 23.iv.2022,

marram grass, day, L. Forster (2, TMAG); Bruny

Island: Adventure Bay beach, east end (43.3686S,

147.3475E), 20.i.2022, spotlighting at night, S. J. Grove

(2, TMAG); Calvert’s Beach, South Arm Dunes

(43.023771S, 147.492336E), 23.iv.2022, marram grass,

night, L. Forster (8 TMAG; 8 ANIC); Musselroe, Stone-

house (40.7537’S, 148.0192’E), 9.ix.2019, beach & low

dunes, S. J. Grove (1, TMAG); Nubeena, White Beach

(43.1167’S, 147.735’E), 30.vii.2019, L. G. Forster

(1, TMAG); Nutgrove Beach, Sandy Bay Dunes

(42.908668S, 147.352220E), 17.i.2022, marram grass at

night, L. Forster (5, TMAG; 5, ANIC); Nutgrove Beach,

Sandy Bay Dunes (42.908668S, 147.35222 E), 22.iv. 2022,

marram grass, L. Forster (2, TMAG); Pieman Heads

(41.6695’S, 144.6695S, 144.9158’E), 24.i. 2021, S. J. Grove

(1, TMAG); Strahan: Macquarie Heads (42.215’S,

245.225 E), 28.xii.2019, beach & dunes, S. J. Grove (12,

TMAG); St. Helens, 1–4.i.1916, F. M. Littler (6, NZAC,

ANIC); St. Helens: Peron Dunes (41.2902S, 148.3416E),

31.i.2021, S. J. Grove (9, TMAG). Victoria: Barwon

Heads, 4.xi.1961, E. Smith (5, ANIC); Venus Bay No. 5

beach (-38.6783°S 145.7791°E), 23.ix. 2018, M. Weston,

pitfall trap in sand dunes (1, ANIC, DNA voucher).

Note. As noted above part of the original series of

L. australis described from Tasmania by Champion

(1890) consisted of smaller and darker specimens he

considered to be males. However we have found the

these actually belong to a new species, L. tasmaniae

described above.

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 281

282 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

Figure 7. Lagrioida species, male genitalia. (A,B) L. australis, tegmen; (B) tegmen and penis; (C) L. brounii, tegmen and penis;

(D) L. norfolkensis, tegmen and penis; (E) L. tasmaniae, tegmen and penis; (F) L. tasmaniae, tegminal apicale and penis apex; (G) L. rufula,

tegminal apicale and penis apex; (H) tegminal basale and penis base.

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 283

Figure 8. Lagrioida species, wings. (A) L. norfolkensis; (B) L. rufula; (C) L. nortoni; (D) L. tasmaniae; (E) L. brounii; (F) L. australis.

Wing venation according to Lawrence et al. 2022.

ACKNOWLEDGMENTS

We are grateful to Dmitry Telnov, Max Barclay and

Keita Matsumoto for help during AS visit to the NHML

and for providing pictures of the type specimens.

Bronte Sinclair and Lingzi Zhou (ANIC) helped with

photography and logistics. The following curators and

their institutions are acknowledged for providing mate-

rial for this study: Derek Smith (AMS), Chris Grinter

(CAS), Marcelo Guerrero (MGC), Azadeh Taghavian-

Azari (MNHN), Simon Hinkley (MVM), Karin Koch

(QM), Pedro Vidal (PVC), Justin Bartlett (QDAF), Ben

Parslow (SAM), Simon Grove and Lynne Forster

(TMAG), and Victor M. Diéguez (VMD). James Bailey,

Lisa Bennett, Ormond Torr, Arnim Littek and Simon

Grove are sincerely acknowledged for providing pic-

tures of Lagrioida in natural habitats deposited at

iNaturalist. RABL was funded by Strategic Science

Investment Funding for Crown Research Institutes

from the Ministry of Business, Innovation and Employ-

ment’s Science and Innovation Group.

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THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 285

Appendix 1. Taxa used for morphological analysis. Alphabetical by family and subfamily.

LYMEXYLIDAE: Melittommatinae (OUTGROUP)

Melittomma sericeum (Harris)

ADERIDAE

Syzeton sp.

ANTHICIDAE: Anthicinae

Anthicus spp.

ANTHICIDAE: Eurygeniinae

Pergetus campanulatus (LeConte)

ANTHICIDAE: Lemodinae

Lemodes sp.

ARCHEOCRYPTICIDAE

Enneboeus barrocolorado Merkl & Enneboeus sp.

BORIDAE: Borinae

Boros unicolor Say & B. schneideri (Panzer)

Lecontia discicollis (LeConte)

CHALCODRYIDAE

Chalcodrya variegata Redtenbacher

CIIDAE: Sphindociinae

Sphindocis denticollis Fall

ISCHALIIDAE

Ischalia spp.

LAGRIOIDIDAE

Lagrioida australis Champion

MELANDRYIDAE: Melandryinae

Melandrya striata Say & Melandrya caraboides (Linnaeus)

MELOIDAE: Eleticinae)

Iselma flavipennis Haag-Rutenberg & Iselma sp.

MYCETOPHAGIDAE: Mycetophaginae

Nototriphyllus araucania Lawrence et al.

MYCTERIDAE: Eurypinae

Lacconotus pinicola Horn

OEDEMERIDAE: Calopodinae

Calopus angustus LeConte & Calopus serraticornis (Linnaeus)

PROMECHEILIDAE

Hydromedion sp.

Sirrhas limbatus (Champion) & Sirrhas variegatus Lawrence

286 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

Appendix 1. Taxa used for morphological analysis. Alphabetical by Family and subfamily (continued).

PROSTOMIDAE

Prostomis intermedius Blackburn & Prostomis sp.

PYROCHROIDAE: Agnathinae

Agnathus decoratus Germar

Cononotus bryanti VanDyke & Cononotus sericans LeConte

PYROCHROIDAE: Pedilinae

Pedilus spp.

PYROCHROIDAE: Pilipalpinae

Morpholycus apicalis (Macleay) & Morpholycus costipennis Lea

PYROCHROIDAE: Pyrochroinae

Pyrochroa coccinea (Linnaeus)

PYTHIDAE

Anaplopus tuberculatus Blackburn

Ischyomius spp.

Priognathus monilicornis (Randall)

Pytho niger Kirby

Sphalma quadricollis Horn

PYTHIDAE ?

Istrisia rufobrunnea Lewis (Pythidae?: Istrisiinae)

RIPIPHORIDAE: Pelecotominae

Pelecotoma fennica (Paykull)

SALPINGIDAE: Agleninae

Aglenus brunneus (Gyllenhal)

SALPINGIDAE: Othniinae

Elacatis umbrosus (LeConte) & Elacatis delusa Pascoe

SALPINGIDAE: Inopeplinae

Euryplatus dimidiatus

SALPINGIDAE: Prostominiinae

Ocholissa sp.

SCRAPTIIDAE: Anaspidinae

Anaspis spp.

STENOTRACHELIDAE

Stenotrachelus aeneus (Fabricius)

SYNCHROIDAE

Synchroa punctata Newman

TENEBRIONIDAE: Lagriinae

Isopteron sp.

TENEBRIONIDAE: Tenebrioninae

Tenebrio molitor Linnaeus

TENEBRIONIDAE ?

Homocyrtus dromedarius (Guérin-Méneville) & H. bonni (Germain)

TETRATOMIDAE: Penthinae

Penthe obliquata (Fabricius)

TETRATOMIDAE: Piseninae

Pisenus humeralis (Kirby)

TRICTENOTOMIDAE

Trictenotoma sp.

ULOLDIDAE: Merycinae

Meryx aequalis Blackburn & Meryx rugosa Latreille

ZOPHERIDAE: Colydiiunae

Pristoderus saccharatus (Pascoe) & Pristoderus antarcticus White

ZOPHERIDAE: Monommatinae

Hyporhagus gilensis Horn

ZOPHERIDAE: Zopherinae

Phellopsis obcordata (Kirby)

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 287

Appendix 2. Lagrioida morphological analysis: list of characters. 09/01/20.

# 1. posterior (occipital) edge of head capsule

1. without or with pair of small, well separated impressions

2. with pair of large, narrowly separated impressions

# 2. Head

1. slightly, gradually narrowed posteriorly, without distinct neck

2. strongly and more abruptly narrowed posteriorly to form neck

# 3. eyes1. entire or barely emarginate

2. distinctly emarginate

# 4. eyes1. strrongly protuberant

2. not or only weakly protuberant

3. absent

# 5. antennal insertions

1. concealed from above by frontal ridges

2. exposed from above

# 6. antennal insertions

1. not located within emarginations of eyes

2. located within emarginations of eyes

# 7. antennal club

1. moderately to strongly developed

2. weakly developed

3. absent

# 8. frontoclypeal suture

1. absent

2. distinct at least laterally

# 9. apex of labrum

1. rounded

2. truncate

3. emarginate

# 10. mandible

1. less than 1.5 times as long as wide

2. between 1.5 and 2 times as long as wide

3. more than 2 times as long as wide

# 11. mandible

1. unidentate

2. bidentate

# 12. incisor edge of mandible

1. simple

2. with one tooth

3. with two or more teeth

# 13. mandibular mola

1. well developed with fine, transverse ridges

2. well developed but without fine transverse ridges

3. reduced or absent

# 14. apical maxillary palpomere

1. more or less symmetrical, widesrt at middle or near base

2. symmetrical or slightly asymmetrical but widest apically (triangular, securiform)

3. strongly asymmetrical and widest at or near apex (securiform or cultriform)

# 15. labial gland opening in male

1. absent

2. present on mentum

3. present on submentum

# 16. apex of ligula

1. convex, truncate or weakly emarginate

2. deeply emarginate or bilobed

# 17. cervical sclerites

1. present

2. absent

# 18. prothorax widest

1. anteriorly

2. at about middle

3. posteriorly

288 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

# 19. lateral edge of pronotum

1. with complete carina

2. with incomplete carina

3. without carina

# 20. base of pronotum

1. without raised margin

2. with raised margin

# 21. pronotal disc

1. without paired bsasl impressions

2. with paired basal impressions

# 22. anterior edge of pronotumm

1. without collar

2. with collar

# 23. base of pronotum

1. without transverse groove

2. with transverse groove

# 24. lateral portion of pronotum

1. without deep pit

2. with deep pit

# 25. prosternum in front of coxal cavities

1. less than 0.5 times as long as mid length of coxal cavity

2. between 0.5 and 1.0 times as long as mid length of coxal cavity

3. longer than mid length of coxal cavity

# 26. procoxal cavity

1. without or with minute notch at base of notosternal suture

2. with distinct notch at base of notosternal suture

# 27. procoxal cavities externally

1. broadly open

2. narrowly open

3. closed

# 28. procoxal cavities internally

1. open

2. closed

# 29. apical edge of prosternal process

1. acute or narrowly rounded

2. broadly rounded or truncate

3. emarginate

# 30. shortest distance between procoxal cavities

1. less than 0.4 times shortest diameter of cavity

2. greater than 0.4 times but less than shortest diameter of cavity

3. greater than shortest diameter of cavity

# 31. prosternal process

1. complete, extending behind coxae

2. incomplete, not extending behind coxae

# 32. apex of prosternal process

1. not expanded laterally

2. slightly or gradually expanded laterally

3. distinctly and abruptly expanded laterally

# 33. procoxae

1. not or only slightly projecting

2. strongly projecting

# 34. prothoracic trochantin

1. exposed

2. concealed

# 35. ratio of internal extension to externally visible portion of procoxa

1. less than 0.25

2. 0.25 to 0.50

3. greater than 0.50

# 36. apex of scutellar shield

1. acute

2. broadly angulate

3. rounded

4. truncate or emarginate

Appendix 2. Lagrioida morphological analysis: list of characters. 09/01/20 (continued).

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 289

# 37. elytral punctation

1. confused, unordered

2. seriate

# 38. anterior edge of mesoventrite

1. truncate

2. broadly rounded

3. more or less acute

# 39. mesanepisterna

1. well separated from one another

2. very narrowly separated or meeting at a point

3. broadly joined at midline in front of mesoventrite

# 40. meso- and metaventrital processes

1. distinctly separated from one another or absent

2. overlapping 3. joined by ball and socket joint

# 41. ratio of distance between mesocoxal cavities to diameter of one of them

1. less than 0.1

2. 0.1 to 0.3

3. greater than 0.3

# 42. mesocoxal cavity laterally

1. open, partly closed by mesanepisternum and mesepimeron

2. open, partly closed by mesepimeron but not mesanepisternum

3. closed by meeting of mesoventrite and metaventrite

# 43. metathoracic discrimen

1. more than half as long as shortest distance between meso- and metacoxal cavities

2. less than half as long as shortest distance between meso- and metacoxal cavities

3. absent

# 44. metacoxae

1. separated by less than shorter (longitudnal) coxal diameter

2. separated by more than shorter (longitudinal) coxal diameter

# 45. metendosternal laminae

1. well developed and broad

2. reduced and narrowed

3. absent

# 46. apical field of hind wing

1. less than 0.25 times total wing length

2. more than 0.25 times total wing length

# 47. radial cell of hind wing

1. well developed and elongate

2. short and broad

3. incomplete or absent

# 48. wedge cell of hind wing

1. apically truncate

2. apically acute

3. absent

# 49. medial fleck of hind wing

1. absent

2. present

# 50. number of veins in medial field of hind wing

1. five

2. four

3. three

4. two or fewer

# 51. ventral lobes on tarsomeres

1. absent

2. present on penultimate tarsomere only

3. present on penultimate and antepenultimate tarsomeres

4. present on antepenultimate tarsomere only

# 52. pretarsal claws

1. without distinct basal tooth

2. with distinct basal tooth

# 53. abdominal ventrites

1. all free and equally articulated

2. 1 and 2 connate

3. 1 to 3 connate

4. 1 to 4 connate

Appendix 2. Lagrioida morphological analysis: list of characters. 09/01/20 (continued).

290 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

# 54. abdominal intercoxal process

1. acute

2. narrowly rounded

3. broadly rounded

4. absent or very weakly developed

# 55. ventral portion of abdominal segment IX in male

1. forming pregenital ring with sides always separated to anterior edge

2. with sides fused to form spiculum gastrale

# 56. aedeagus

1. normally oriented, with tegmen dorsal and penis ventral

2. inverted, with tegmen ventral and penis dorsal

# 57. tergite X (proctiger) in male

1. absent or entirely membranous

2. lightly sclerotised at middle with paired lateral slcerites or bacula

3. more or less evently sclerotised

# 58. tegmen

1. not transversely divided, basale and apicale fused together

2. transversely divided into basale and apicale

# 59. ratio of length of apicale to that of basale

1. greater than 0.6

2. 0.6 or less

# 60. accessory lobes of apicale

1. absent

2. present

# 61. shape of tegminal apex

1. acute or angulate

2. rounded or truncate

3. emarginate

# 62. tegminal struts

1. absent

2. present

# 63. penis

1. without basal struts

2. with basal struts

# 64. larva: posterior edge of head capsule (dorsal)

1. not or only slightly emarginate

2. distinctly emarginate

# 65. larva: epicranial stem

1. very short or apparently absent

2. moderately long

# 66. larva: frontal arms

1. lyriform

2. V-shaped or U-shaped

# 67. larva: ratio of antennal length to head width

1. less than 0.15

2. 0.15 to 0.50

3. greater than 0.50

# 68. larva: sensorium on preapical antennomere

1. less than 0.33 times as long as apical antennomere

2. more than 0.33 times as long as apical antennomere

# 69. larva: sensorium on preapical antennomere

1. conical or palpiform

2. dome-like

3. complex (C-shaped, undulate)

# 70. larva: frontoclypeal suture

1. absent or vaguely indicated

2. distinct

# 71. larva: apex of mandible

1. with single lobe or tooth

2. bilobed or bidentate

3. trilobed or tridentate

# 72. larva: incisor edge of mandible

1. simple

2. with one subapical tooth

3. with two or more teeth or serrate

Appendix 2. Lagrioida morphological analysis: list of characters. 09/01/20 (continued).

THE FAMILY LAGRIOIDIDAE ABDULLAH & ABDULLAH, STAT. NOV. 291

# 73. larva: mandibular mola

1. well developed

2. reduced or absent

# 74. larva: molar surface

1. simple

2. irregularly tuberculate or asperate

3. with numerous fine, transverse ridges

4. with few coarse ridges

# 75. larva: apex of mala

1. simple

2. cleft or distinctly emarginate

# 76. larva: apex of mala

1. rounded

2. truncate

# 77. larva: innter apical angle of mala

1. with one to several small teeth

2. without teeth or uncus

3. with distinct lobe or uncus

# 78 larva: ligula

1. shorter than labial palp

2. as long as or longer than labial palp

# 79. larva: labial palps

1. contiguous or separated by less than width of first palpomere

2. separated by more than width of first palpomere

# 80. larva: hypopharyngeal sclerome

1. consisting of single sclerite shaped like a molar tooth

2. consisting of transverse bar only

3. absent

# 81. larva: hypostomal rods

1. very short or absent

2. moderately to very long and subparallel

3. moderately to very long and diverging

4. moderately to very long and converging

# 82. larva: ventral epicranial ridges

1. absent

2. present

# 83. larva: gula

1. longer than wide

2. not longer than wide

# 84. larva: patches of dorsal asperities

1. absent

2. on several body segments

# 85. larva: thoracic terga

1. without transverse rows of asperities or carinae

2. with transverse row of asperities or carina on mesothorax and metathorax

# 86. larva: mesocoxae

1. separated by less than one basal coxal diameter

2. separated by one to two basal coxal diameters

3. separated by more than two basal coxal diameters

# 87. larva: larva: pretarsal setae

1. double, lying side by side or obliquely situated

2. double, lying in row, one distal to the other

3. single

# 88. larva: thoracic spiracles

1. annular

2. annular-uniforous

3. annular-biforous

4. annular-cribriform

5. annular-multiforous

# 89. larva: abdominal terga

1. without rows of asperities

2. with transverse rows of asperities on several segments

3. with open or closed circles of asperities on several segments

Appendix 2. Lagrioida morphological analysis: list of characters. 09/01/20 (continued).

292 J. F. LAWRENCE, R. A. B. LESCHEN, M. ELGUETA, N. PORCH and A. ŚLIPIŃSKI

# 90. larva: abdominal segment IX, excluding appendages

1. shorter than segment VIII

2. as long as or longer than segment VIII

# 91. larva: abdominal tergum IX

1. not forming articulated plate

2. forming articulated plate

# 92. larva: abdominal tergum IX

1. without paired processes or urogomphi

2. with paired processes or urogomphi

# 93. larva: abdominal tergum IX

1. without interurogomphal pits

2. with single, undivided interurogomphal pit

3. with single, longitudinally divided interurogomphal pit

4. with two separated interurogomphal pits

# 94. larva: ratio of length of urogomphus to basal width of tergum IX

1. less than 0.2

2. between 0.2 and 1.0

3. more than s1.0

# 95. urogomphi

1. neither bifurcate nor with accessory processes

2. bifurcate, with or without accessory processes

3. with one or more small accessory processes

# 96. larva: urogomphi separated by

1. less than one basal width

2. between one and two basal widths

3. more than two basal widths

# 97. larva: abdominal sternum IX

1. without armature

2. with one basal asperity on each side

3. with two to four basal asperities on each side

4. with 5 or more basal asperities on each side, usually forming transverse row

5. with row of apical asperities

6. with two apicomesal teeth

7. U-shaped with apicolateral teeth

# 98. larva: abdominal sternum IX

1. not enclosed by sternum VIII

2. partly to almost completely enclosed by sternum VIII

# 99. larva: row of basal asperities on sternum IX

1. straight or very slightly curved posteriorly

2. strongly curved posteriorly

3. strongly, doubly curved forming a double „U”

# 100. larva: row of basal asperities on sternum IX

1. more or less continuous or narrowly interrupted at middle

2. broadly interrupted at middle

# 101. larva: paired pygopods on segment X

1. absent

2. present

# 102. larva: anterior abdominal spiracles

1. annular

2. annular-uniforous

3. annular-biforous

4. annular-cribriform

5. annular multiforous

# 103. larva: spiracles on abdominal segment VIII

1. about the same size as others on abdomen

2. distinctly larger than others on abdomen

Appendix 2. Lagrioida morphological analysis: list of characters. 09/01/20 (continued).

ResearchGate has not been able to resolve any citations for this publication.

Jan 1969

Omamari Beach
Of Dargaville

Omamari Beach, N of Dargaville, 5.iii.1969, J. E. Tobler (1, CAS);

Auckland: Helensville, 26.x. 1992, sandy sea beach, under logs

Jan 2000

Whangarei Heads, Ocean Beach (35.50'S, 174.34'E), 13.ii.2000, at night on sand, G. Hall, R. Leschen (1, NZAC); Auckland: Helensville, 26.x. 1992, sandy sea beach, under logs, A. Larochelle, M.-C.

Jan 1968

Muriwai Beach

Muriwai Beach, iii.1968, under log, J. C. Watt (1, NZAC);

sand dunes, lupin litter, B. M. May (1, NZAC); Bay of Plenty: Oroiti Beach, 10.iii

Jan 1963

Xi Whatipu

Whatipu, xi.1963, sand dunes, lupin litter, B. M. May (1, NZAC); Bay of Plenty: Oroiti Beach, 10.iii.1993, J. S. Dugdale (1, NZAC);

Ocean Beach, 24.i.1.1991, pit traps

Hawkes Bay

Hawkes Bay: Ocean Beach, 24.i.1.1991, pit traps, C. Duffy (1, NZAC);

Heaphy Hill, 6.xi.1999, on tree trunks at night

Heaphy Track

Heaphy Track, Heaphy Hill, 6.xi.1999, on tree trunks at night, R. Leschen & G. Hall (1, NZAC);

26.x.1968, sand dunes

Sandhills Patarau
Ck

Patarau, Sandhills Ck., 26.x.1968, sand dunes, J. I. Townsend (1, NZAC); Patterus Rock, 15.i.1966, A. K. Walker (1, NZAC);

Jan 1971

Tahunanui

Tahunanui, 28.x.1971, J. C. Watt (5, NZAC);

Horning (1, NZAC); Westland: Jackson's Bay, Neils Beach, 28.ii.1966, G. Kuschel (1, NZAC); Dunedin: Aramoana, Otago Harbour, 15.xii

Jan 1993

Spencer's Beach, 10.x.1973, sand beach, under logs, D. S. Horning (1, NZAC); Westland: Jackson's Bay, Neils Beach, 28.ii.1966, G. Kuschel (1, NZAC); Dunedin: Aramoana, Otago Harbour, 15.xii. 1993, Stop 93-11B, sandspit, D. H. Kavanaugh (2, CAS);

Southland: Kaka Point, near Owara, 3.xii.1996

Jan 1995

Taieri Mouth, 24.xii.1995, underside of beach driftwood, J. Nunn (1, NZAC); Southland: Kaka Point, near Owara, 3.xii.1996, J. Nunn (1, NZAC); Fiordland: Bluecliffs Beach, Waitutu, 20.ii.2003, J. Nunn (1, NZAC);

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