(PDF) The North American Winnertziini (Diptera: Cecidomyiidae: Winnertziinae). A photographic and illustrated guide to genera and species.

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THE NORTH AMERICAN WINNERTZIINI

(DIPTERA: CECIDOMYIIDAE: WINNERTZIINAE)

A PHOTOGRAPHIC AND ILLUSTRATED GUIDE TO GENERA AND SPECIES.

COPYRIGHT @ 2022.

OF THIS BOOK MAY BE USED OR REPRODUCED IN ANY MANNER WHATSOEVER

WITHOUT WRITTEN PERMISSION OF THE AUTHOR EXCEPT IN CASE OF

QUOTATIONS IN SCIENTIFIC ARTICLES OR REVIEWS.

FIRST EDITION

September 2022

Plakidas, John D.

The North American Winnertziini (Diptera: Cecidomyiidae: Winnertziinae).

A photographic and illustrated guide to genera and species.

Correct Citation: Plakidas, J. D. 2022. The North American Winnertziini (Diptera:

Cecidomyiidae: Winnertziinae). A photographic and illustrated guide to

genera and species. Loyalfield Publishing. Pittsburgh, Pennsylvania. 115 pp.

Author’s e-mail: johnplakidas@verizon.net

Printed in the United States of America.

Cover design: Rhipidoxylomyia volitans n. sp., female paratype.

ISBN-13: 978-1-929094-06-6

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Table of contents.

Abstract and Introduction …………………………………………………………… 3

Collection site data ………………………………………………………………….. 4

Larval diagnosis …………………………………………………………………….. 6

Pupal diagnosis ……………………………………………………………………… 10

Adult diagnosis ……………………………………………………………………… 14

Key to genera ……………………………………………………………………….. 18

Introduction to genera and species …………………………………………………. 18

Kronomyia Felt 1911 ……………………………………………………………….. 20

Rhipidoxylomyia Mamaev 1964 ……………………………………………………. 29

Winnertzia Rondani 1860 …………………………………………………………… 46

Concluding remarks …………………………………………………………………. 111

Acknowledgements …………………………………………………………………. 111

Literature cited ………………………………………………………………………. 112

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Abstract. The North American Winnertziini (Diptera: Winnertziinae: Cecidomyiidae) are

represented with three extant genera: Kronomyia Felt, 1911; Rhipidoxylomyia Mamaev, 1964;

and Winnertzia Rondani, 1860 and one extinct genus Cretowinnertzia Gagné, 1977 which is not

treated here. Four new species are described: Rhipidoxylomyia volitans n. sp., Winnertzia

fenestra n. sp., Winnertzia hortensis n. sp., and Winnertzia vernalis n. sp. With the exception of

Winnertzia arizoniensis Felt, 1908 all of the North American Winnertziini have been collected

from the northeastern United States. Type material and voucher specimens are deposited in The

National Museum of Natural History, Washington, D. C. (USNM).

Introduction.

During the early pioneer phase of systematic entomology Johannes Winnertz described

Cecidomyia lugubris 1853 which would ultimately become the type species for the genus

Winnertzia Rondani, 1860. According to several sources (Osten Sacken, 1904; Panelius, 1965;

Spungis, 1992) the type series for Winnertzia lugubris (Winnertz, 1853) is lost and presently

unidentifiable. However Winnertz’s original description and illustrations still exist which

provide the necessary verification for the reconciliation of this species (see my discussion on

page 46). I have no doubt that the wing illustration (Winnertz, 1853, here as Fig. 72) which

Winnertz sketched is that of a Winnertzia. In the years that followed it was not until 1911 that

Ephraim Felt described Kronomyia Felt, 1911 and more than 50 years elapsed before the

description of Rhipidoxylomyia Mamaev, 1964. Both of these genera are closely related to

Winnertzia. As an assemblage of three extant genera in North America the Winnertziini

forms a natural group based on the following morphological traits. Wing veination with M4

and CuA separate; scales occur on legs and halters; male genitalia displaying an aedeagal

prominence or aedeagal bulge (Jaschhof & Jaschhof 2020); female abdomen is telescopic from

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segments 6 – cercus and the disticerci have clavate setae; larval spatula almost always unidentate

and a pair of terminal spines on the 9th segment; the pupa typically has 4-papillae on the clypeal

apotome, two of which are setiform.

All known larvae (Spungis 1992; Plakidas 1999) live in decaying habitats which include soil,

damp leaf litter, rotting vascular wood, beneath loose rotting bark and under moss. The larval

diet consists exclusively of fungal mycelia. Winnertziini larvae do not ingest detritus as a result

the inner gut lining is always free of debris. The larvae are typically 2-3 mm long, white, pinkish

or yellow in color living in sibling groups making them relatively easy to collect if one wishes to

rear adults.

As for the species composition of Winnertziini living in North America there are 25 available

names including the 4 new species described here. In Parnell’s revision of the North American

Porricondylinae (1971) he proposes Winnertzia aceris Felt 1913, Winnertzia calciequina Felt

1907, and Winnertzia pectinata Felt 1911 as synonyms of Winnertzia solidaginis Felt 1907

without providing evidence for his systematic actions. In light of my findings I am proposing that

all of Felt’s names remain valid until more adult, larva and pupal material becomes available for

future study.

Collection site data.

The North American Winnertziini have been collected (Maps 1 & 2) from the state of New York

at Caroga, Keene Valley, Karner, Albany and Nassau. In Pennsylvania from the Pittsburgh area

found in Allegheny County, in Maryland from the Baltimore area, in New Jersey from Plainfield

and Strom Virginia which is located south of Washington D.C. (not shown on the map). The

only species collected outside of the northeastern United States is Winnertzia arizoniensis Felt

1908, which came from Williams, Arizona. The remainder of the North American continent

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including Alaska and Canada remain unknown and uncollected for the Winnertziini. That’s a

fact.

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Systematic Entomology.

Diptera: Linnaeus, 1758 (on page 584).

Family: Cecidomyiidae, Newman, 1834.

Subfamily: Winnertziinae, Panelius, 1965.

Tribe: Winnertziini: Panelius, 1965.

Here I present a holistic systematic model based on larval, pupal and adult traits. For additional

reading on this tribe see Panelius (1965), Jaschhof & Jaschhof (2013, 2020) and Spungis (1992).

For a newly described extinct genus of this tribe see Fedotova et al. (2022). For a complete list of

available names and proposed synonyms in Winnertziini see Gagné & Jaschhof (2021).

Larval Diagnosis.

LARVA.- Habitus: head, neck, three thoracic and nine abdominal segments (Fig. 1). The ninth

segment possesses a prominent anal pore ventrally, and is equipped with a broad sclerotized

plate with 2-spines on the posterior margin (Figs. 53, 66) or two well defined separate spines

(Fig. 104). Larvae are typically 4 to 5 times longer than wide and cylindrical in appearance.

Head capsule hemicephalic (Figs. 2, 3, 4), meaning it is reduced in size and sclerotization,

except in Winnertzia fungicola Felt which possesses heavy armature anteriorly (Fig. 3). The head

is narrower than the neck segment and capable of being fully retracted into the body cavity. The

mandibles are lance-shaped, have a smooth leading edge and are capable of piercing fungal

mycelia. (By way of comparison the eucephalic head capsule of Bibio sp. is heavily sclerotized

with large serrated mandibles and incapable of retracting into the body cavity). Internally the

sclerotized pharynx is clearly visible and aids in supporting the esophagus. The salivary duct

enters the oral opening just below the esophagus and functions by secreting enzymes useful in

extralimital digestion of mycelia (Solinas, 2011). Cephalic apodemes are lacking from the

posterior rim of the head capsule.

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Respiratory system peripneustic, with eighteen functional spiracles. Two spiracles are found

on the prothorax and first eight abdominal segments, on the dorso-lateral margins.

Color.- Pigmented cuticle: examples are pink, yellow, white and in Kronomyia white with a

brown mid-gut. The cuticle is longitudinally furrowed in Winnertzia and Rhipidoxylomyia,

in Kronomyia the cuticle is smooth and lacks furrowing. The smooth cuticle in Kronomyia is one

of the most reliable traits separating this genus from Rhipidoxylomyia and Winnertzia.

There is a spatula (Fig. 5) on the ventral prothorax which has an undivided anterior margin in all

genera except in Rhipidoxylomyia cinerea (Plakidas, 2019) where the spatula appears tridentate

(Fig. 65). The posterior margin of the ninth segment always has either a pair of well-defined

spines as in Winnertzia and Rhipidoxylomyia, platelets as in Kronomyia, or a broadly sclerotized

area posteriorly terminating in 2-spines (Fig. 66), as in Rhipidoxylomyia cinerea (Plakidas,

2019).

Papillae.- There is one basic type of papilla, the circular papilla which is flush with the body

wall or slightly elevated. Circular papillae appear to be membranous and are either nonsetose

(fig. 6), or possess a micro seta (figs. 5, 34) which occur on the ventral thoracic segments.

Elongate macro setae are absent in Winnertziini larvae.

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Head capsule (Figs. 2, 3, 4). Reduced in size and narrower than the neck and other body

segments. The genae comprise the largest surface area of the head and harbor the mouth parts

(feeding apparatus) located anteroventrally. The cardo is fused with the subgenal carina. The

maxillae are fleshy, lack serrated teeth and are fused with the maxillary palps; the mandibles are

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tightly aligned between the maxillae and the labrum. The mandibles always have a smooth

leading edge (Figs. 2, 4). The labrum is a membranous, dorsoventrally compressed lobe.

The dorsal surface of the head capsule possesses the 2-segmented antenna (Fig. 3), which is

globular in Winnertzia and Rhipidoxylomyia, and cylindrical in Kronomyia. Both the dorsal and

ventral surfaces of the head possess circular non-setose papillae as shown in (Figs. 2, 3, 4), these

papillae may be nonfunctional in that they appear to be vestigial.

Neck.- Wider than the head capsule, retractable into the body cavity and with 2-non-setose

papillae ventrally.

Prothorax.- Dorsal: a pair of spiracles occupy the medial-lateral margins. Ventral prothorax:

possess a sclerotized spatula at mid-line (Fig. 5). It usually extends almost the entire length of

the segment. The spatula is a sclerotized structure which is comprised of an anterior tooth

followed by an elongate shaft and anchored to the membrane by a posterior apophysis. The

anterior margin of the denticle is undivided (unidentate), rarely with three lobes (tridentate). On

either side of the tooth are the sternal nonsetose papillae (Fig. 5). The ventral thoracic lateral

papillae occur in four triplets, each triplet has 2-microsetiform and 1-nonsetose papilla.

Mesothorax and metathorax.- Both of these segments have the same papillae arrangement as

the prothorax, but neither segment has spiracles or a spatula. Each thoracic segment may possess

as many as 22 papillae.

Abdominal segments 1-7 dorsal.- A spinule field is found on the anterior margin, followed by a

pair of spiracles on the dorso-lateral margins with six nonsetose papillae between the spiracles.

Laterally there are 2- papillae per side.

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Abdominal segments 1-7 ventral. - There are 2- micro sternal papillae along the anterior margin

of the spinule fields (Fig. 6), 4-anterior ventral nonsetose papillae and 4- posterior nonsetose

ventral papillae as shown occur outside of the spinule field, a Winnertziinae larval trait.

Eighth abdominal segment dorsal. - A pair of spiracles occur dorso-laterally, always with four

nonsetose papillae between the inner margins of the spiracles (Fig. 53).

Eighth abdominal segment ventral. - With 4-nonsetose papillae located near the posterior margin.

Ninth abdominal segment.- Shorter and narrower than the eighth segment. The ventral surface

has 6 nonsetose papillae. The anal pore occurs on the ventral side of the ninth segment in all

larvae which I have examined. There is a variety of sclerotization which occurs on the posterior

margin of the 9th segment which is best addressed in the species descriptions. However the

sclerotization is in the form of spines or platelets.

Spinule fields. - These are minute spicules which are found on both the dorsal and the ventral

anterior margin of the meso and metathorax and first seven abdominal segments.

Pupal diagnosis.

Pupa. – Exerate (Fig. 7); the leg sheaths free from the body and the abdomen is capable of

movement. Fourteen functional spiracles are present. Two each on the dorsal lateral margins of

the mesothorax and abdominal segments 2-7. None of the pupae I have examined possess

spiracles on the first abdominal segment.

Habitus: head, three thoracic and nine abdominal segments. The pupa when fully developed is

dark brown or a yellow-brown in color.

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Head. - Ventral (Figs. 8, 9, 10): at the apex of the head are the antennal sheaths. These run

laterally along the outer rim of the eye sheaths and contour the outer margin of the palpal sheaths

then terminate at the juncture with the wing sheath and body wall. Small sclerotized horns occur

at or near the apex of the antennal sheaths in Winnertzia and Rhipidoxylomyia (Figs. 8, 9).

However these horns are absent in Kronomyia (Fig. 10). In some species the nonsetose antennal

papillae (Elsayed et al 2020) are visible (Fig. 8) however these papillae are not always present

(Fig. 10). At the lateral anterior end of the eye sheaths are the hemispherical lobes (Figs. 8, 9).

These lobes are common in most Mycetophiliformia pupae and are an identifying feature for this

group. At the posterior end of the face is the clypeal sheath which is bordered laterally by the

palpal sheaths. Anterior to the clypeal sheath is the clypeal apotome. This sclerite contains from

2-4 papillae. These papillae are either setiform or nonsetose. The clypeal apotome lies between

the eye sheaths. At the inner lateral margins of the palpal sheaths are the posterior orbital

papillae (Fig. 10).

Dorsal thorax.- Slightly arched in lateral view (Fig. 7). The dorsal thorax is comprised of the

pronotum, mesonotum and the metanotum. The pronotum harbors the pronotal setae (Figs. 9,

10). These setae are located on elevated tubercles. The mesonotum comprises the greatest

surface area of the dorsal thorax and has a pair of spiracles located on the medial-lateral margins

just above the wing sheaths. The thoracic spiracles are either long or tapering at the apex (Figs.

11, 12), as in Winnertzia and Rhipidoxylomyia, or cup-shaped (Fig. 13) in Kronomyia. Running

along the entire dorsomedial ridge of the mesonotum is the ecdysial suture. This is a line of

weakness which opens during ecdysis and allows the adult to emerge from the exuviae.

The metanotum (Fig. 77) is a narrow segment which abuts to the first abdominal segment.

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Abdomen.- The abdomen is comprised of nine segments. Typically all segments are covered

with spicules except segments 1 and 9 which have a sparse covering of spicules. Functional

spiracles occupy the medial lateral margins of segments 2-7.

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Winnertziini adult diagnosis.

Figure 14. Winnertzia sp. male, lateral view. Light yellow-brown with 11-flagellomeres.

Male and female diagnosis. Body coloration for adults is typically yellow-brown for

Winnertzia (Fig. 14) and Rhipidoxylomyia while Kronomyia adults are darker brown, with

darker scales on the legs. Body size ranges from approximately 1.5 – 5.5 mm. Eyes holoptic, eye

bridge typically with facets. Antenna: 10-12 flagellomeres. Flagellomeres consist of a basal node

and a stem (Figs. 15, 16, 17). Sensoria which are found on the nodes have multiple pore

attachments in Winnertzia (Fig. 15), hyaline sensoria in Kronomyia (Fig. 17) with

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single pore attachment and digitate sensoria in Rhipidoxylomia (Fig. 16) each sensoria with one

large pore attachment to the flagellar node. Palpus 2-4 segmented, segments covered with

microtrichia and sparsely covered with setae and shorter blade-like setae. Thorax (Fig. 18):

anterior pronotum with setae, proepisternum nonsetose, except in W. rubricola (Fig. 166);

katepisternum and anepisternum bare; anepimeron bare in Winnertzia and Rhipidoxylomyia; with

6 – 8 long setae in one Kronomyia sp. female. Dorsocentral and dorsolateral setae present; setae

also occur on the scutellum. The halters are shorter than the height of the thorax. The stem with

setae, the knob with scales and setae. Legs covered with short setae, scales and setiform scales;

first tarsomere with or without a spiniform projection; tarsal claws simple or with a basal tooth,

empodium shorter or longer than the claw. Wing (Fig. 19): R5 and R1 veins with setae; Sc close

to the R1; R5 joins the C at or beyond wing apex, with a break at its juncture. The M4 and CuA

do not join. In some specimens the M4 is only slightly visible.

Abdomen: tergites and sternites covered with microtrichia; in Winnertzia, Rhipidoxylomyia and

Kronomyia tergites 1 – 6 have small translucent windows. Tergites with a complete or nearly

complete posterior row of setae on segments 1 – 8; in the male, tergite 9 in some species nearly

as long as the genitalia.

The male reproductive apparatus in the Winnertziini is a very complex arrangement of

morphological features and internal sclerotization which undoubtedly contribute to each species

reproductive success. For example, in the genus Winnertzia, when the male genitalia is viewed

laterally (Figs. 180, 181) we find the following morphological features in this order. The

aedeagal prominence is situated ventral to the ejaculatory apodeme and is attached to the

gonocoxites anteriorly, the aedeagus is compressed between the ejaculatory apodeme and the

tegmen and narrows distally to its apex, the tegmen is a lightly sclerotized structure which is the

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fusion of the parameres and provides physical support and protection for the aedeagus. Dorsally

the cercus and the 9th tergite provide additional support to the reproductive apparatus. The

gonocoxites act as trusses which provide lateral stability for the reproductive apparatus and

finally the gonostyli are movable appendages capable of grasping the female during copulation.

Internally there is a rigid sclerotized framework which stabilizes all of the membranous features.

The female abdomen in the Winnertziini has segments 1 – 5 unmodified with the tergites and

sternites immovable i.e. not telescopic. Beginning with segment 6 through segment 8, these

segments are almost entirely membranous, narrow and are elongated allowing for a greater

degree of telescoping. Segment 9 consists of a sclerotized, immovable sternite and a

membranous tergite. The 10th segment is the shortest of all abdominal segments and is entirely

membranous, covered with setulae and sparsely covered with setae. The 2-segmented paired

cerci originate from the 10th segment. This arrangement results in two basicerci and two

disticeri. Both segments of the cercus are covered with setulae, setae and each segment has 3-5

clavate setae. The clavate setae are wider and more blunt-ended than the thin-walled sharply

pointed setae and are easily recognizable. Because of their large size, clavate setae have wider

and more prominent bases. Two obvious internal features which are noted, are the sclerotized

spermathecae and the genital rod.

Additional notes on male genitalia. Gonocoxites are always densely covered with microtrichia,

sparsely covered with setae. The inner gonocoxal emargination which is an area of union

between the two gonocoxites, is typically U – shaped or V – shaped. The gonocoxal

emargination is sometimes called the ventral plate. Distally each gonocoxite has a gonostylus

which is capable of independent movement and acts as a grasping appendage. Each gonostylus

has two attachments points with its gonocoxite, an anterior apophysis with a muscle attachment

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at the outer edge of the gonocoxites, and a second attachment point which is closer to the inner

margin. The apex of the gonostylus has a pectinate comb-like claw in Winnertzia (Fig. 182 ), and

lacks a pectinate claw in Rhipidoxylomyia (Fig. 45), and Kronomyia (Fig. 28).

Winnertziini adult key to genera.

1 Flagellar sensoria with multipore attachments in ♂ & ♀ (Figs. 15, 79, 80);

female post abdomen with 1 or 2 sclerotized spermathecae (Figs. 160, 106);

male gonostylus with a pectinate comb-like distal or sub distal

claw (Figs. 182, 110); ……………………………………………….. Winnertzia, p. 46

- Flagellar sensoria with single pore attachment in ♂ & ♀;

female post abdomen without sclerotized spermathecae; male

gonostylus without a pectinate claw ………………………………………..…………. 2

2 Flagellar sensoria arranged in pairs, always 4 sensoria in females,

2-4 in males (Figs. 16, 41)…………………..………………….. Rhipidoxylomyia, p. 29

- Flagellar sensoria hair-like, randomly arranged on nodes (Fig. 20) ….. Kronomyia, p. 20

Introduction to species.

Kronomyia populi Felt, 1911 ………..………………..…….. 20

Rhipidoxylomyia minor Mamaev, 1964 …………………….. 30

Rhipidoxylomyia cinerea (Plakidas, 2019) ……….…………. 36

Rhipidoxylomyia volitans n. sp. ………….…………………. 41

Winnertzia aceris Felt, 1913 ……..………………………….. 49

Winnertzia ampelophila (Felt, 1907a) …….………………… 49

Winnertzia arizoniensis Felt, 1908 ………………………….. 52

Winnertzia calciequina Felt, 1907c ………………………….. 53

Winnertzia carpini Felt, 1907c ……………………………….. 59

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Winnertzia fenestra n. sp ……………………………………… 60

Winnertzia fungicola Felt, 1921 …………………………..….. 63

Winnertzia globifera Mamaev, 1963 ………..………………… 67

Winnertzia graduata Spungis, 1992 …………………………… 71

Winnertzia hortensis n. sp. …………………………………….. 75

Winnertzia hudsonici Felt, 1908 …….…………………………. 79

Winnertzia karnerensis Felt, 1908 ……………………………… 80

Winnertzia nigripennis Kieffer, 1896 ………………………….. 81

Winnertzia notmani (Felt, 1919) …………….…………………. 83

Winnertzia padicola Spungis …………………………………… 84

Winnertzia palustris Felt, 1915 …………………….………….. 87

Winnertzia pectinata Felt, 1911 ……..………………………… 90

Winnertzia pinicorticis Felt, 1907b ……………………………. 91

Winnertzia rubida Felt, 1908 ……..…………………………… 93

Winnertzia rubricola Mamaev, 1963 .…………………………. 97

Winnertzia solidaginis Felt, 1907a …….……………………… 102

Winnertzia vernalis n. sp. ……………..………………………. 107

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Genus: Kronomyia Felt 1911: page 476.

Type species Kronomyia populi Felt, by original designation. Holotype female NYSM.

=Trichoxylomyia Mamaev 1964: page 897. Type species, ovalis, original designation.

Felt described his new genus and species, Kronomyia populi Felt, from a single female reared

from “punky poplar wood collected at Nassau, N. Y”, with no rearing date provided. In the brief

description he does include two illustrations (Figs. A, B) which I include here to compare with

more recent material from Maryland and Pennsylvania. It appears safe to assume the holotype

female for K. populi Felt to be conspecific with the newly collected material (compare figure A

with figure 21 and figure B with figure 23). The tenth and eleventh flagellomeres are

conspecific, while the second palpal segments differ only in their distal portions.

Diagnosis. Based on Kronomyia populi Felt, from New York, Maryland and Pennsylvania. Male

and female with 11 flagellomeres (Figs. 20, 21, 22). Nodes have a basal whorl of setae, followed

by numerous translucent sensory hairs in the female and 1-2 sensory hairs in the male (Fig. 22).

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The stems of female flagellomeres are about ⅓ the length of the node (Fig. 20), the 10+11

flagellomeres are fused (Fig. 21). Stems of male flagellomeres are about ½ the length of the

nodes (Fig. 22) and flagellomeres 10+11 fused. The female palpus is 2-segmented (Fig. 23) and

the 2nd segment is elongated and enlarged distally. (In Felt’s description of K. populi he shows a

much larger club shape on the 2nd palpal segment). Male palpus 3-segmented. Lateral thorax as

shown (Fig. 25).

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Anepimeron with long setae, anterior pronotum with setae, proepisternum nonsetose.

Legs: 1st tarsomere with a spiniform projection (Fig. 27), tarsal claws with a basal tooth,

empodium shorter than the claw (Fig. 26). Wing (Fig. 24), veination as shown, typical for the

tribe. Halters shorter than the height of the thorax, covered with microtrichia, stems with dark

brown setae and scales, knobs with a dense covering of broad scales and setiform scales which

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are dark brown (Fig. 39), the cuticle beneath is translucent yellow.

Male abdomen (Fig. 31): tergites 1 – 6 with a posterior row of setae and setae mainly at the

lateral margins, no scales; tergite 7 with only a few posterior setae; tergite 8 appears to be

entirely membranous without setae; tergite 9 deeply bilobed and with setae situated distally on

each lobe (Fig. 28).

Female abdomen (Fig. 32): tergites 1 – 5 with a posterior row of setae and scattered setae

elsewhere; sternites 2 – 5 with posterior row to an intermixed double row of setae and setae

occur along the perimeters. The protrusible portion of the ovipositor is membranous to lightly

sclerotized with scattered setae, the cercus with setulae, setae and clavate setae (Fig. 29).

Male genitalia (Figs. 28 & 30). The overall appearance is compact, gonocoxites about twice as

long as wide covered with microtrichia and sparsely covered with setae, the ventral inner

gonocoxal emargination (Fig. 30) displays an open V-shape; the ejaculatory apodeme is

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sclerotized, sinuous and shorter than the gonocoxites. The tegmen is membranous (Fig. 28) and

extends above the ejaculatory apodeme, the parameral apodemes are well sclerotized. The

aedeagal prominence is triangular and has a dense covering of microtrichia (Fig. 30). The

gonostyli (Fig. 30) are ovoid and have a dense covering of setae on the inner margin.

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Female cercus (Fig. 29) is 2 – segmented both segments of about equal length, both segments

densely covered with setulae and sparsely covered with setae. There are 4 – 5 clavate setae

distally on both segments. Clavate setae are easily recognized by their wide profile which

contrasts them from the more common thin walled setae.

Larva (Figs. 33 – 35) . Description as for the tribe. Several notable characteristics which

standout as identifying features for Kronomyia are the head capsule which has elongate-

cylindrical antennal flagellum (Fig. 33). The flagellum is 3X longer than wide. The spatula (Fig.

34) has an oblong distal tooth which is not much wider than the shaft and the anterior and

posterior apophyses are clearly absent. The 9th abdominal segment has two small spines on the

posterior margin (Fig. 35). In one specimen only one spine is fully developed so there appears to

be variability in this trait. Larvae are found in heavily decayed vascular wood of fallen Tulip

trees and directly beneath rotting bark of fallen Ash trees. Decay type is brown-rot.

Pupa (Figs. 36 – 38). Description as for the tribe. Notable characteristics which stand out as

identifying features for Kronomyia are the four papillae on the clypeal apotome (Fig. 36), two of

which are setose. The posterior orbitals have 1- setiform and 2-3 nonsetose per side. The thoracic

spiracle is shortened forming a cup like aperture with an adjacent seta (Fig. 37). The abdominal

segments are covered with fine spicules and stronger spines are found on dorsum (Fig. 38). The

9th segment clearly is shorter and narrower than the 8th and the genital sheath appears to be

bilobed.

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Taxonomic notes. At least one other Kronomyia sp. occurs in eastern United States. I have

reared two females from decaying wood, neither with larval data. These two flies represent a

much larger species, 3.5 – 5.5 mm body length, with darker brown coloration. Both females have

flagellomeres with long stems and the 10+11 flagellomeres are not fused. Each of these females

were slide mounted with their exuviae and the pupal traits all conform to those of K. populi. Both

of these flies are deposited in the USNM for future study.

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Material studied. One larva, Kronomyia populi Felt, slide mounted in canada balsam, collected

from under Ash bark, Penna. Allegheny Co., Aspinwall, October 15, 2015. One female slide

mounted in canada balsam, Kronomyia populi Felt, 1911, Penna. Allegheny Co., Aspinwall,

reared July 6, 2016. One male, Kronomyia populi Felt, slide mounted in canada balsam, Penna.

Allegheny Co., Aspinwall, reared July 6, 2016.

Two females, Kronomyia sp., on separate slides, Penna. Allegheny Co., Aspinwall, reared

March 6, 2017 and one reared March 10, 2017, from decaying wood.

All of the material studied is deposited in the USNM.

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Genus: Rhipidoxylomyia Mamaev 1964: page 898.

Type species: Rhipidoxylomyia rubella Mamaev, by original designation.

=Cryptoxylomyia Mamaev 1996: page 65. Type species Rhipidoxylomyia excavata Mamaev

1964: page 900.

Three species occur in eastern North America, Rh. cinerea (Plakidas 2019) and Rh. minor

Mamaev 1964, and Rh. volitans n. sp. I have previously identified specimens illustrated on page

68 (Plakidas 2019) as Rh. minor Mamaev 1964. This species appears to be similar to Rh. rubella

Mamaev 1964 with only slight differences between them. They may in fact be synonymous.

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Another taxonomic consideration is that Rhipidoxylomyia material I have studied which includes

larvae, pupae and adults, conform to Winnertzia Rondani 1860 with only two main points of

variability. These being the sensoria found on flagellar nodes and the male gonostylus. In

Winnertzia flagellar sensoria have multiple pore attachments and the male gonostylus has a

chitinous comb-like spine distally (Fig. 182) or sub distally (Fig. 110). In Rhipidoxylomyia

flagellar sensoria have a single pore attachment and the male gonostylus lacks a chitinous spine,

instead there is a dense sole of closely joined setae on the inner margin (Fig. 45), or whip-lash

setae (Fig. C, p. 43). As one can see these two genera are very closely related. However the fact

that Rhipidoxylomyia males lack a gonostylus spine probably accounts for a morphological

reproductive barrier, eliminating the possibility for a Rhipidoxylomyia sp. males to

successfully copulate with a Winnertzia sp. female. It is reasonable to assume that these genera

should remain distinct.

Rhipidoxylomyia minor Mamaev 1964: page 900.

Diagnosis. Adults are light yellow to yellow-brown (Fig. 40); male body length 1.5 – 1.75 mm;

females 1.5 – 3.4 mm. Males with 11 – flagellomeres, the first 9 flagellomeres with 2 – forked

sensoria (Fig. 41) at the distal margin of the nodes; flagellomeres 10+11 lack sensoria (Fig. 42).

Female with 11 – flagellomeres, the last 2 are fused, each node has 4 – sensoria, each with 5-

branches (Fig. 43). The last flagellomere in one female lacks sensoria. Palpus 4 – segmented,

segments progressively longer; eye bridge complete. Wing (Fig. 44) as for the tribe. Thorax:

dorsocentral setae and dorsolateral setae present; lateral sclerites bare. Male abdomen: tergites 1

– 6 with a sparse posterior row of setae and scattered setae elsewhere, no scales; sternites 2 – 8

with a complete posterior row of setae and scattered setae elsewhere, no scales. Male genitalia

(Fig. 45, 46): 9th tergite nearly as long as the gonocoxites, with setae laterally and on the distal

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margins of each lobe; tegmen, ejaculatory apodeme and aedeagal prominence covered with

microtrichia as shown; gonostyli (Fig. 45) ovoid with a dense covering of closely joined setae on

the inner margin. Female abdomen telescopic, cercus (Fig. 47) basicercus and disticercus of

about equal length, both covered with setulae, sparsely covered with setae and clavate setae (Fig.

48) mainly found distally on each segment.

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Larva (Figs. 49 – 53). Larvae were first discovered in decaying vascular wood several

millimeters below the surface in a fallen Tulip tree (Liriodendron tulipifera), April 15, 1995.

Collection site 7Km S Columbia MD. More recently I collected larvae from a rotting branch

fragment lying in the leaf litter on February 1, 2016 in Allegheny Co., PA. Three females and

two males were reared soon after in March 2016. Larvae are typical Winnertziini in general

appearance and have two traits which they share with other Winnertzia larvae. These are the

ovoid flagellum of the antenna (Fig. 51) and the longitudinal furrowing on the cuticle.

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Other noteworthy larval traits are the spatula which has a well-developed posterior apophysis

found in specimens from both Maryland and Pennsylvania; and the large spines on the 9th

segment (Fig. 53) which arise from a common base.

Pupa (Figs. 54 – 55). Face (Fig. 54), clypeal apotome has 4 – papillae, 2 are nonsetose and 2 are

setiform. The posterior orbitals occur in 2 groups of three papillae, with 1 – setose and 2 –

nonsetose papillae in each group. The antennal horns occur at the apex of the antennal sheaths.

They are small and very low profile indicating they provide little in the way of functional value

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in rasping through decayed wood during adult emergence from a decaying substrate. The

thoracic spiracles are long and narrow tapering to a rounded apex (Fig. 55). There is no seta

associated with the spiracle.

The abdominal segments are covered with small spicules except for the dorsum of segments 2 –

6 where we find stronger and longer spines medially.

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Taxonomic notes. Comparison of the European specimens of Rh. Minor Mamaev (Spungis

1992, p. 9) with specimens from Maryland and Pennsylvania reveals strong morphological

similarities in male genitalia’s and male and female flagellomeres. The male genitalia

specifically are so similar that there appears to be no reproductive barriers between European and

North American specimens. In addition, the larvae of European and North American specimens

were both found in decayed wood (Mamaev 1964) (Plakidas 2019). I do note however, that these

conclusions are based on a very small sampling of specimens so future researchers should be on

the lookout for additional larval material to rear adults for study.

Material studied. One larva, Rh. Minor Mamaev, slide mounted in Euparal, Penna. Allegheny

Co., collected February 1, 2018 from rotting branchlet in leaf litter. One male and 1 – female

same data as larva, both reared March 22, 2016. Additional material previously deposited in the

USNM labeled as Rhipidoxylomyia sp. lot # 973. This series includes 1-larva, 1- female and 1-

male all from Columbia MD., April 15, 1995.

All material deposited in The National Museum of Natural History, Washington D. C.

(USNM).

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Rhipidoxylomyia cinerea (Plakidas 2019) page 42.

Originally described as Cryptoxylomyia cinerea based on the excavated gonostylus which is a

shared character with Rhipidoxylomyia excavata Mamaev 1964. However after reviewing the

shared variation amongst species in the genus Winnertzia it is reasonable to assume that one

would expect to find such variation in Rhipidoxylomyia as well. I concur with the findings of

Jaschhof & Jaschhof (2013) that Cryptoxylomyia Mamaev 1992 is a junior synonym of

Rhipidoxylomyia Mamaev 1964.

Excerpts from the original description of Rh. cinerea Plakidas 2019, pages 42 – 47. Male and

female pale gray, eyes black. Body length male, 2.2 mm; female, 3.2 mm. Both sexes with 11

flagellomeres, male flagellomeres have branched sensoria on first 10 flagellomeres (Fig. 56),

females have branching sensoria on all 11 flagellomeres (Fig. 57). The wing (Fig. 58) has a

slightly arched R5 and the CuA is close to the lower margin and not traceable to the C, the M4 is

absent. Tarsal claws are simple in the male and with a basal tooth in females with a reduced

empodium (Fig.63). The abdominal tergites 1 – 6 have a posterior row of setae in the male (Fig.

59) and female and a group of setae laterally, no scales. Male genitalia (Figs. 60, 61): the main

identifying features of the genitalia include the unmodified 9th tergite, the gonostyli lack an

apical spine and the gonocoxal apodemes are shorter than the distance between them (transverse

bridge). The female cercus is 2-segmented (Fig. 62), the disticercus has a rounded apex and is

longer than the basicercus, both with setulae, setae and clavate setae.

The larva (Figs. 64 – 66) is 4.3 – 4.5 mm, white, cuticle with longitudinal furrowing and the 9th

abdominal segment has a large sclerotized area on the posterior margin which terminates in 2

spines (Fig. 66). The head (Fig. 64) has elongated antennae with an ovoid flagellum. The spatula

(Fig. 65) has a well-developed anterior apophysis, remaining papillae as for the tribe.

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The pupa (Figs. 67 – 69): 3.5 mm, dark brown when fully mature. The clypeal apotome has 2

setiform papillae, and there 2 groups of three posterior orbitals, each group with 2 nonsetose and

1 setose papilla (Fig. 67). Thoracic spiracles (Fig. 68) are elongate with a rounded apex.

Abdominal spiracles are on the lateral margins of segments 2 – 7, all segments finely spiculate

lacking strong spines (Fig. 69). The 9th segment has two membranous elongated lobes on its

dorsum (Fig. 69).

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Taxonomic notes. Rhipidoxylomyia cinerea (Plakidas 2019) when compared to other

Rhipidoxylomyia spp. is somewhat of an anomaly in that the R5 vein is longer and curved beyond

midlength joining the C well beyond the wing apex (Fig. 58) and the female disticercus is much

longer than the basicercus (Fig. 62). These two traits are unique among extant

Rhipidoxylomyias. However in one newly described extinct species from Fushan amber (China,

50-53 MA), Fushuniola mai Fedotova & Perkovsky 2022, we find these two traits present. One

may pose the hypothesis that Fushuniola mai is in fact a basal clade to the modern day species of

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Rhipidoxylomyia where Rh. cinerea represents a possible direct lineage to Fushuniola mai

Fedotova & Perkovsky.

Material studied. Holotype male on slide labeled: Penna. Allegheny Co., Fairhill Road,

Sewickley, May 17, 1997, bred from larvae in decaying Black Birch log. Paratypes: male and

female on separate slides same data as holotype; 2 larvae on separate slides, collected November

1996 from decaying Black Birch log. All type material mounted in Euparal and deposited in the

USNM.

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Plate 1. Rhipidoxylomyia volitans n. sp. female paratype.

Rhipidoxylomyia volitans n. sp.

Diagnosis. Plate 1 and figures A - G.

Male and female (Plate 1) pale yellow, thorax darker, legs translucent with a covering of light

brown scales. Male ≈ 1.75 mm, wing 1.4 mm; female 2.6 – 2.75 mm, wing 1.75mm (n=2). Male

11-flagellomeres, the first nine flagellomeres with 2-forked sensoria (fig. A) distally,

flagellomeres 10+11 are fused and lack sensoria. Female with 11-flagellomeres, the first ten with

4- branching sensoria (Fig. B), flagellomeres 10+11 fused. Palpus 3-segmented, segments

progressively longer. Wing (Plate 1) typical for the genus except the R5 is only slightly curved,

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not as strongly arched as in Rhipidoxylomyia cinerea (Fig. 58). Tarsal claws are sickle form with

a strong basal tooth, empodium rudimentary.

Male genitalia (Figs. C, D, E); aedeagal prominence clearly visible, covered with scales and

longer than the tegmen; tegmen longer than the ejaculatory apodeme and each lateral margin

with a row of long spines (Fig. D). The 9th tergite deeply emarginated, the lateral lobes with long

setae. The gonostylus is very distinctive being nearly 3X longer than wide at its widest point and

with long whip-lash setae as shown (Fig. C). The gonocoxites are splayed and the inner

gonocoxal emargination (Fig. E) is membranous and setose.

Female abdomen (Plate 1) widest at segments 2 and 3 then segments 4 – 6 abruptly narrow and

is telescopic from segments 7 – cercus (Fig. F). The disticercus is shorter than the basicercus

both segments are densely covered with setulae, sparsely covered with setae and have narrow

clavate setae.

Pupa as for the genus and the thoracic spiracle (Fig. G) elongated, slightly arched and narrowing

to a pointed apex.

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Larva unknown. When I discovered the 3 – larvae in decaying wood I decided not to slide mount

one of them since there were so few.

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Etymology. The specific name, volitans, Latin feminine singular for hovering.

Type material. Holotype male labeled Penna. Allegheny Co., Aspinwall, reared from decaying

wood April 8, 2022; 2 – paratype females same collection data as the holotype, one reared April

3, 2022 and the other reared April 7, 2022. The female reared April 3 also has its exuviae

mounted with it under the same cover glass. Deposited in the National Museum of Natural

History, Washington, D. C.

Taxonomic notes. The new species, Rhipidoxylomyia volitans n. sp. is very unique from other

Rhipidoxylomyia species in that the male gonostylus is distinctive with long whip-lash setae (Fig

C), compared to the gonostyli of Rhipidoxylomyia cinerea (Plakidas) (Fig. 60) which are

excavated medially. The gonostyli of Rhipidoxylomyia minor Mamaev (Fig. 45 ) are more ovoid

and have a dense tuft of setae on the inner margin.

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Genus: Winnertzia Rondani 1860: page 290.

Type species Cecidomyia lugubris Winnertz, 1853: page 305.

Rondani described Winnertzia 1860 as a new genus based on wing veination compared with

other known genera for that time. He designated Cecidomyia lugubris Winnertz, 1853 as the type

species for the new genus. At this point I would like to revisit Winnertz 1853 page 305 and

review his description. Here Winnertz tells us that he reared the flies from rotting wood and the

mushroom Polyporus versicolor. I’m assuming that the larvae were in the decaying wood and

somehow associated with the mushroom, possibly in the mushroom membrane itself. After he

reared the flies, both male and female, he tells us that “after the flies died the body became

brownish or black-brown, the neck more or less white and the legs pitch colored. The abdomen

remained dusty-yellow.” His complete description is accompanied with four illustrations: the 4-

segmented palpus (not shown), male antenna (Fig. 70) and female antenna (Fig. 71), each with

12-flagellomeres, and an exceptional wing illustration (Fig. 72), which is undoubtedly that of a

Winnertzia. Osten Sacken (1904) who visited Winnertz in 1854 at his home in Crefeld Germany

confirmed Winnertz’s illustration technique to be very accurate (read Osten Sacken’s account on

page 54, 1904). From this account we are reassured that the illustrations for W. lugubris can be

useful in rediscovering this species in the future.

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The type material for W. lugubris is lost (Panelius, 1965) and aside from these illustrations, its

exact morphological identity is presently uncertain. However, Winnertz did leave us with

biological data which will be useful in the future in rediscovering this species. Larvae were

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associated with rotting wood and the mushroom, Polyporus versicolor, so I envision a future

researcher in Europe collecting that mushroom in hopes of securing Winnertzia larvae to rear

adults for study. Here in Allegheny County Pennsylvania I have collected Winnertzia fungicola

Felt larvae in the mushroom membrane of Trichaptum biforme, a polypore mushroom which

grows in clusters on fallen trees. These findings support the claim that the larvae of some species

of Winnertzia live and feed in the living membrane of mushrooms.

Relevant publications for available names of North American Winnertzia species are: Felt

(1915) which includes a key to 12 species on pages 130 – 142. Panelius (1965) which includes

his diagnosis for the new tribe Winnertziini, important data on lost types from Europe which are

unidentifiable, and a key to males of 14 species on pages 113 – 114. Parnell (1971) did a

revision of the Felt types of Winnertzia and included a key to species for males and females on

page 287. At this point I find it appropriate to address Parnell (1971) because of his use of

synonyms without justification. His synonyms were carried over in a subsequent catalog (Gagné

2004) which gives one the impression that these synonyms are valid and I disagree. The

reasoning behind avoiding synonyms at this time is based on two main criteria. First Parnell’s

use of the synonym clause is based on his opinion and he provides no precise systematic rational

to support his findings. Second and most importantly we have only a few specimens with which

to study, therefor, based on a very small sample size of Winnertzia species we cannot make

inferences concerning synonymy with any degree of confidence. At this early stage in

systematics and with very limited material to study, all of Felt’s types must be preserved for

future generations as nomen conservandum, i.e. “a name to be conserved”. As more material

becomes available for study, future researchers will then have greater confidence in naming new

species, or assigning synonyms.

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Winnertzia aceris Felt, 1913: page 213. Holotype male, NYSM.

Parnell (1971, page 290), as a synonym of Winnertzia solidaginis Felt, 1907a.

White larvae were collected from beneath thin decaying bark of a Sugar Maple, Acer saccharum,

at Nassau, N.Y., March 12, 1913. Adults were reared in April 1913. Felt tells us that W. aceris is

closely allied to both W. calciequina Felt 1907 and W. pectinata Felt 1911. However I

demonstrate that W. calciequina (Fig. 85) and W. pectinata may not be synonymous with W.

solidaginis (Fig. 178). Parnell (1971, p. 290) first assigned W. aceris as a junior synonym of W.

solidaginis. This action was clearly invalid while Parnell gave no reasoning for placing W. aceris

with W. solidaginis. From the original description of W. aceris one can easily discern differences

between the two species which are potential reproductive barriers in the male genitalias. Felt

describes the male 9th tergite (dorsal plate) in W. aceris “…long, triangularly emarginated, the

lobes broad, obliquely truncate and sparsely setose.” (Felt 1913, p. 214). This description is

clearly different from the 9th tergite of W. solidaginis (Fig. 179).

Winnertzia ampelophila (Felt, 1907a): page 144. Holotype male, NYSM.

Parnell (1971, page 287).

As Porricondyla ampelophila Felt, 1907a, page 144.

As Porricondyla ampelophila Felt, 1907b, page 48.

Felt described W. ampelophila (Felt 1907) twice in the same year (1907) in two separate

journals. In his 30th report of the State Entomologist (1915), Felt designates the primary

reference for W. ampelophila as 1907a.

Diagnosis. Male: 1.5 mm; antenna with 12 flagellomeres, 11+12 fused, sensoria thin, U-shaped

and free ending (Fig. 73). Palpus 4-segmented, 4th segment twice the length of the 3rd. Tarsal

claws simple, empodium rudimentary. Wing as for the genus. Genitalia (Fig. 74), 9th tergite

horizontal along the distal margin, inner gonocoxal emargination deeply U-shaped, gonocoxal

apodemes short. Gonostylus with an apical claw of closely appressed spines.

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The larva (Fig. 75) is typical for the genus with strong longitudinal furrowing and the spatula

(Fig. 76) has a triangular tooth and well developed shaft. The lateral papillae occur in 2 groups of

triplets per side. The 9th segment has two spines originating from a common base (Fig. 75).

The pupa (Fig. 77) has an elongate thoracic spiracle with a rounded apex, abdominal spiracles

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are found on the lateral margins of segments 2 – 7 and the segments are covered with fine

spicules. Strong spines are absent on the abdominal segments.

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Type material: holotype male collected in Albany, N. Y., on Virginia creeper, Parthenocissus

quinquefolia, July 3, 1906. NYSM. Type Cecid. 450.

Material studied: 1 larva, 1 male, 1 female on slides. Six larvae collected on October 1, 1994,

from decaying vascular wood several millimeters below the surface of a fallen tulip,

Liriodendron tulipifera. Decay color white. Collected 7Km S Columbia, MD. Deposited in the

USNM. The three slide mounted specimens which I deposited in the USNM, believed to be

conspecific with W. ampelophila.

Winnertzia arizoniensis Felt, 1908: page 421. Holotype female NYSM.

The following diagnosis and color description after Felt (1915, p. 134). “Female: 2 mm; antenna

probably extending to the base of the abdomen, thickly haired, fuscous yellow, probably 14

segments, the 5th subcylindric, with a length about one-half greater than its diameter. Palpi; the

first segment short, stout, irregular, the 2nd rather stout, with a length about 3 times its diameter,

the 3rd one fourth longer and more slender and the 4th one half longer and more slender than the

3rd. Mesonotum, scutellum and abdomen a dull reddish brown. Wings narrow, hyaline, Costa

pale straw. Legs yellowish brown; claws long, stout, strongly curved, unidentate, the pulvilli

shorter than the claws. Ovipositor longer than the body, the terminal lobes biarticulate, the 2nd

segment more slender, with a length about twice its diameter, broadly rounded apically. Type

Cecid. 1022.”

Taxonomic notes. The single female holotype was collected by H. S. Barber on June 12 from

Williams, Arizona. The year of collection was not given. The original description for W.

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arizoniensis Felt 1908, consisted of a brief description in a key to the species (p. 421). Felt did a

follow-up key to Winnertzia in 1915 where he provides a more detailed description for W.

arizoniensis without illustrations. This species is represented by a single slide mounted female in

the NYSM. This specimen also represents the only Winnertzia from North America which Felt

identified that was not collected in the North Eastern United States.

Winnertzia calciequina Felt, 1907a: page 161. Holotype male NYSM.

Figure 78. Winnertzia calciequina Felt. Male (distal flagellomeres missing).

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The following diagnosis partially after Felt (1915, pages 138 – 140) and newly reared specimens

from Allegheny County Pennsylvania. Male and female 2.0 – 2.5 mm; antenna 2+12. Male color:

face greenish yellow. Mesonotum dark brown, sparsely clothed with fine silvery hairs; scutellum

and post scutellum dark brown. Abdomen yellowish green basally, the apical segments light

brown. Wings hyaline, Costa dark brown; halters whitish transparent. Coxae, femora and tibiae

mostly pale yellowish, tarsi nearly uniform fuscous; claws stout and strongly curved (Fig. 81)

with a basal tooth, empodium shorter than the claw. Genitalia: gonocoxites very stout, broad,

obliquely truncate; gonostyli stout, broadly rounded and with a conspicuous distal claw (Fig. 85).

Dorsal plate (9th tergite) long, broad, emarginated. Inner gonocoxal emargination broadly

rounded. Male flagellomeres 1 – 11 have stems nearly equal to the height of the nodes (Figs. 79,

83), the sensoria extend from near the base of the node and distally forming a Y-shape (Fig. 79)

or occur more distal on the nodes (Fig. 83). The female flagellomeres 1 – 11 have noticeably

short stems (Fig. 80), with well-developed U-shaped sensoria. Tarsal claws in both sexes have a

basal tooth and the empodium is shorter than the claw (Fig. 81). The wing (Fig. 84), 2.5 mm,

typical for the genus and has noticeably dark brown C and R5 veins. The male genitalia (Fig.

85): the gonocoxites are longer than wide, the 9th tergite is deeply emarginated and has 4-5 setae

along the posterior margin of each rounded lobe; the tegmen is unsclerotized and triangular with

a rounded apex; the aedeagal prominence is very distinctive with a fringe of short hairs along the

outer margin and the inner wall is covered with knob-like scales; the ejaculatory apodeme is

tubular and strongly sclerotized; the gonostylus is more than twice as long as wide with a distal

tooth comprised of many sclerotized comb-like spines. The female cercus (Fig. 82) has an oval

disticercus with at least 3 – clavate setae distally and one long sharply pointed seta near the apex.

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The white larva, 5.5 mm, is very distinctive with a well-developed spatula which has a strong

posterior apophysis and the tooth with a broadly rounded anterior margin (Fig. 86). The 9th

abdominal segment has 2 – strong spines on the posterior margin which arise from separate

bases. The pupa is typical for the genus with one distinct difference. The thoracic spiracle is

circular and nearly flush with the body wall and has a strong spine-like appendage (Fig. 87).

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The pupal thoracic spiracle of W. calciequina Felt appears to be a species defining trait.

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Taxonomic notes. Felt describes this species (1907a, page 161) with only a brief description. In

1915 Felt provides us with a more detailed description including collection data on pages 138 –

140. Here he states that the flies were taken on Pine, probably Pinus strobus, at Albany, N. Y.,

July 24, 1906. At least one male and one female of W. calciequina were collected at that time.

So it is clear that this species was not reared from larvae which would be helpful in attempting to

place this species as a synonym of W. solidaginis. Additionally, I collected larvae for W.

calciequina Felt under moss on a rotting Pinus sp. log which is a piece of biological evidence

supporting my material as being conspecific with W. calciequina Felt.

Panelius (1965) addresses this species and illustrates the male genitalia (Fig. 33, F) on page 117.

Although he places his synonymy in question he did study the female for W. calciequina Felt and

2-♂ and 2-♀ for W. pectinata Felt.

A final piece of evidence which I believe separates W. calciequina from W. solidaginis are the

pupal thoracic spiracles. Winnertzia calciequina has a circular thoracic spiracle with a spine-like

appendage (Fig. 87), whereas W. solidaginis has a cup-shaped thoracic spiracle that lacks a

spine-like appendage (182A).

Material examined. Two larvae, both slide mounted, collected March 2, 2020, from under moss

on a decaying Pine log, Penna. Allegheny, Co., Fairhill Rd., Sewickley; 1 male reared April 14,

2020, whose larva was under moss on a decaying Pine log, Penna. Allegheny Co., Fairhill Rd.,

Sewickley, slide mounted in Canada balsam and deposited in USNM.

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Winnertzia carpini Felt 1907a: page 421. Holotype male NYSM.

The following diagnosis is taken from Felt (1915) and Parnell (1971). This is a light yellow-

brown species. Male: 1.0 mm; antenna 2+12, 11+12 fused; flagellar stems shorter than the nodes,

sensoria U-shaped on the first 10 flagellomeres. Palpus 4-segmented, the 4th segment 1½X

longer than the 3rd. Wing as for the genus; tarsal claws with a basal tooth, empodium shorter

than the claw. Genitalia (Fig. 88): 9th tergite emarginated with rounded lateral lobes; inner

gonocoxal emargination U-shaped; tegmen membranous and slightly longer than the ejaculatory

apodeme. The gonostyli are twice as long as wide with a distal claw of comb-like teeth.

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Winnertzia fenestra n. sp. Holotype male USNM.

Diagnosis. Male: 1.75 mm; antenna 2+10; sensoria thread-like nearly straight with the distal ends

free from the node and occurring on the first 8 – flagellomeres (Fig. 89); flagellomeres 9+10

without sensoria (Fig. 90). Palpus 4-segmented (Fig. 93), first segment globular, 2nd and 3rd of

about equal length, the 4th segment narrower and longer than the 3rd segment. Thorax with a

few dorsolateral setae and two complete rows of dorsocentral setae; the lateral thorax bare. Legs

pale translucent yellow covered with long and shorter setae no scales on leg segments; tarsal

claws (Fig. 91) with a basal tooth, empodium shorter than claw. Halters: knob is white-

translucent with only setae and setiform scales there are no broad plate-like scales on the halters.

Wing (Fig. 92) 1.25 mm, R5 is nearly straight to wing margin. Genitalia: (figs. 94 – 97); as

shown, gonocoxal apodemes about as long as the transverse bridge; tegmen unsclerotized;

parameral apodemes strongly sclerotized; aedeagal prominence not visible; 9th tergite

emarginated (Fig. 97); inner gonocoxal emargination (Fig. 96) U-shaped and without setae;

gonostyli with a distal tooth comprised of comb-like teeth, and about 2X longer than wide.

Etymology. The specific name fenestra, from the Latin meaning window, and referring to the

collection site from a window pane.

Type material. Holotype male, labeled: Penna. Allegheny Co., Aspinwall, collected April 15,

2017 on window pane. Mounted in Canada balsam and deposited in the USNM.

Taxonomic notes. Winnertzia fenestra n. sp. is an anomaly among other North American male

Winnertzia’s with only 10 well defined flagellomeres (Fig. 90).

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Winnertzia fungicola Felt 1921, page 118. Holotype male NYSM.

Diagnosis. Male: 2.0 mm; female 2.5 mm; this is a light yellow-brown species whose larvae are

documented from at least three mushroom species, which include Lenzites saepiaria, Polyporus

betulinus and Trichaptum biforme. The larvae feed in the living mushroom membrane of

Trichaptum biforme (personal observation) where they also pupate. Both males and females have

12 – flagellomeres; 4 – segmented palpus; the apex of the eye bridge without facets; tarsal

claws are thick and with a basal tooth, the empodium is longer than the claw (Fig. 100). The

male genitalia (Figs. 98-99) is typical for the genus except the ejaculatory apodeme is tubular in

appearance and about as long as the gonocoxites. The gonostylus (Fig. 99) is 3X longer than

wide. The female has 2 large circular – ovoid sclerotized spermathecae (Fig. 106).

The larva has an unusual sclerotized appendage on the anterior lateral margins of the head (Figs.

101, 102) which undoubtedly aids in cutting through mushroom tissue. The anterior margin of

the spatula is also unique where we find spine-like appendages extending anteriorly (Fig. 103).

The 9th abdominal segment has 2 – well defined spines which originate separately (Fig. 104).

The pupal face is typical for the genus (Fig. 105) and the non-setose antennal papillae are

visibly prominent.

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Material examined. One male, 2-females + 1-exuviae and 2-larvae. The larvae were discovered

January 17, 2014 in the living tissue of the mushroom cap of Trichaptum biforme. Adults were

reared February 23, 2014. Deposited in the National Museum of Natural History, Washington, D.

C. (USNM).

Taxonomic notes. Winnertzia fungicola Felt is the only species of Winnertzia which I have found

whose larvae live and feed in living mushroom tissue.

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Winnertzia globifera Mamaev 1963. Holotype male HYNM, Japan.

106A

Figure 106A. Winnertzia globifera Mamaev, male.

Diagnosis. Figures 106A – 115. Male 2.0 mm, wing 1.8 mm; female 2.7 mm, wing 1.7 mm.

Male with 11-12 flagellomeres (Fig. 107); females 10-11 flagellomeres (Fig. 108); palpus 3-4

segmented. Thorax and wing (Fig. 106A) as for the genus. Tarsal claws without a basal tooth,

empodium shorter than the claw (Fig. 109). The female foreleg 5th tarsomere has a double row

of blunt spines and is noticeably inflated and larger than the mid and hind leg acropods (Fig.

109). The male genitalia is very distinctive with a subapical claw and a medially inflated

ejaculatory apodeme (Fig. 110). The female cercus has both basicercus and disticercus of about

equal length and each with clavate setae distally (Fig. 111).

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Larva. 3.5 mm, yellow-white; the head capsule (Fig. 112) typical for the genus. The spatula has a

broadly rounded anterior tooth (Fig. 113), a lightly sclerotized shaft and a well-developed

posterior apophysis. In two of the slide mounted larvae the shaft is less visible and the posterior

apophysis is not clearly defined. This indicates that sclerotization of the spatula is variable.

The posterior margin of the 9th segment has 2 well defined spines originating from separate

bases.

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Pupa (Fig. 114), 3.0 mm and dark brown when fully matured. There are two main distinguishing

features with the pupa. First is the abdomen has fine spicules on both the dorsum and ventrum

and the thoracic spiracle (Fig. 115) is elongated.

Material examined. Three larvae, 2-males and 1-female on slides. Larvae were collected

February 27, 2017 under loose decaying bark of a fallen Ash tree, Fraxinus sp., Allegheny CO.,

Aspinwall, Penna. All on slides and mounted in Canada balsam. Deposited in the USNM.

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Winnertzia graduata Spungis 1992. Holotype male LASS, Latvia.

Diagnosis. Figures 116 – 125. Male 2.1 mm, wing 1.9 mm; female ≈ 2.5 mm, wing 2.1 mm.

Male with 12-flagellomeres (Fig. 116), the 12th flagellomere the shortest; female with 12-

flagellomeres (Fig. 117), the 12th flagellomere as long as 11. Sensoria on all 12-flagellomeres in

both male and female. Palpus 4-segmented, segments progressively longer. Thorax and wing

(Fig. 119) as for the genus. The first tarsomere with a spiniform projection; tarsal claws with

a basal tooth, empodium shorter than the claw (Fig. 118). Genitalia (Fig. 120): 9th tergite

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with a deep V-shaped emargination; cercus longer than the gonocoxites; gonostylus with a distal

comb-like tooth. The ejaculatory apodeme is tubular (Fig. 121), aedeagus clearly visible,

however the aedeagal prominence is not discernable possibly due to cover glass compression.

Female cercus with both segments of nearly equal length and clavate setae distally on basi- and

disticercus.

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Larva. 2.7 mm, white. The spatula has a triangular anterior margin (Fig. 122); and the 9th

abdominal segment with distinctive coniform spines on the posterior margin (Fig. 123).

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Pupa. 2.7 mm, dark brown when fully developed. The apex of each antennal sheath has a

nonsetose papilla (Fig. 124), and the clypeal apotome has 4-papillae, 2-nonsetose and 2-

microsetiform. The thoracic spiracle (Fig. 125) is very prominent and elongated.

Biology. The larvae develop in decaying wood beneath rotting bark and are found in Prunus

serotina, Black Cherry. Decay color brown.

Material examined. Two males; 2-females and 1-larva on slides in Euparal. Larvae collected

December 20, 2007 from decaying Black Cherry log, Aspinwall, Allegheny Co., Penna. Adults

reared March 15, 2008. Deposited USNM.

Taxonomic notes. The adult males which I reared are similar with the description and

illustrations provided by Spungis (1992, p. 28).

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Winnertzia hortensis n. sp. Holotype male USNM.

Diagnosis. Figures 126 – 131. Male ≈ 2.0 mm, wing 1.7 mm; female damaged.

Male with 12-flagellomeres, stems slightly shorter than nodes, sensoria on all flagellomeres as a

single thread (Fig. 126). Female, although damaged does have 12-flagllomeres with large U-

shaped sensoria on all 12 flagellomeres (Fig. 127). Palpus 4-segmented. Thorax and wing as for

the genus. First tarsomere without a spiniform projection; tarsal claws with a basal tooth,

empodium shorter than claw. Legs have a variety of scale shapes including broad scales. Male

genitalia (Figs. 128-130): is very distinctive having a rectangular shaped tegmen with a slightly

pointed apex (Figs. 128, 130); the aedeagal prominence is longer than the ejaculatory apodeme

and is translucent; the gonostylus claw is comprised of one strong tooth and several shorter

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spines (Fig. 128); the cercus lobes are triangular and longer than the gonocoxites. The 9th tergite

has two broadly rounded lobes with a distinctive shallow U emargination (Fig. 129).

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The female cercus (Fig. 131) is typical for the genus with clavate setae positioned distally on

both cercus segments and the disticercus with several longer setae at or near the apex. There is

one sclerotized ovoid spermatheca.

Larva, 3.4 mm, white and typical for the genus.

Pupa (Figs. 132, 133), 2.5 – 2.7 mm. There are two main distinguishing features I find with the

pupa which help separate this species from other Winnertzia species for which I have pupae. The

short tubular thoracic spiracles (Fig. 132). The second defining trait is the presence of strong

spines on the anterior-medial margins of abdominal segments 2 – 6 (Fig. 133). These traits found

in this combination are unique among Winnertzia’s.

Biology. Larvae are found feeding beneath decaying Oak bark.

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Etymology. The specific name, hortensis, from the Latin hortus, which means garden, and refers

to being collected in my backyard.

Type material. Holotype male Pennsylvania Allegheny Co., Aspinwall, reared May 25, 2020;

paratype female (damaged) same data as holotype; 3 – paratype larvae all collected March 2020,

living under decaying bark of an Oak branchlet, in leaf litter. Deposited USNM.

Taxonomic notes. Winnertzia hortensis is placed with the W. solidaginis group by virtue of the

tegmen flaps, 12-flagellomeres in both sexes, and dorsolateral thorax setae in a single group.

Winnertzia hortensis differs from W. solidaginis in the shape of the tegmen, and the 9th tergite.

The female has only one sclerotized spermatheca, whereas W. solidaginis females have two

sclerotized spermathecae.

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Winnertzia hudsonici Felt 1908, page 422. Holotype female NYSM.

Diagnosis. Female: 1.5 mm; antenna with 12-flagellomeres, sensoria U-shaped on all

flagellomeres. Palpus 4-segmented, segments successively longer, the 4th segment about twice

as long as the 3rd. The foreleg first tarsomere longer than the fore tibia; tarsal claws with a basal

tooth, empodium shorter than the claw. Wing hyaline, Costa light brown, veination as for the

genus. Halters pale yellow, fuscous subapically. Coxae, femorae and tibiae mostly light fuscous

yellow, tarsi mostly dark brown. Mesonotum dark brown, sub median lines pale yellowish.

Abdomen fuscous yellow, with 2-sclerotized spermathecae.

Taxonomic notes. Felt (1908) describes Winnertzia hudsonici Felt on the basis of a single

female reared “from a jar containing leaves of Crataegus bearing greenish, stout, cup-shaped,

fimbriate galls.” The larva of Winnertzia hudsonici presumably inhabited decaying vegetable

matter in the rearing jar. The fly was reared in Albany, N. Y. July 25, 1907.

Additionally, if one is promoting synonyms (Gagné and Jaschhof 2021), W. hudsonici Felt is a

junior synonym of W. karnerensis Felt. As is evident in the 1908 key to species, W. karnerensis

appears before W. hudsonici Felt. Parnell (1971) misrepresented his use of synonyms by placing

W. hudsonici Felt as the senior name to W. karnerensis Felt. This erroneous error was then

carried over into the present world catalog (Gagné and Jaschhof 2021).

However I do not consider W. hudsonici Felt and W. karnerensis Felt to be synonymous. One

will never know with certainty since both species are represented by a single female. In the

absence of larva, pupa and males these two names must remain nomen conservandum.

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Winnertzia karnerensis Felt 1908, page 422. Holotype female NYSM.

Diagnosis. Female 2.0 mm; antenna with 12-flagellomeres, U-shaped sensoria on all

flagellomeres. Palpus 4-segmented, segments successively longer, the 4th segment nearly twice

as long as the 3rd segment. The foreleg first tarsomere longer than the fore tibia; tarsal claws

with a basal tooth, empodium shorter than the claw. Scutum dark brown; scutellum fuscous;

abdomen pale yellow-brown. Wings hyaline, Costa dark brown, veination as for the genus.

Halters slightly fuscous; coxae and femorae brown, tibiae darker, tarsi gray. Two sclerotized

spermathecae.

Taxonomic notes. Felt (1908) described W. karnerensis Felt on the basis of a single female

collected “on hard pine, Pinus rigidae, at Karner, N. Y. June 26, 1906. Winnertzia karnerensis

Felt is described on page 422 (Felt 1908) and appears in the key to species before W. hudsonici

Felt. Therefore, if one is using synonyms, W. karnerensis Felt is the senior name to W. hudsonici

Felt.

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Winnertzia nigripennis Kieffer 1896, page 36. All type material lost.

Presently unidentifiable.

I provided a brief description (page 71) and illustration (page 74) of Winnertzia nigripennis

Kieffer in Plakidas (2019) from Allegheny county Pennsylvania. I based my findings on Panelius

(1965, p. 116) and a brief description and an illustration in Spungis (1992, p. 26). However after

revisiting the original description and illustrations in Kieffer (1896), his illustrations here in Plate

II, I believe my identification in Plakidas (2019) of W. nigripennis to be inaccurate. Two main

differences occur. First, the 9 tergite in Kieffer (Plate II, Fig. F) is shorter and narrower that

gonocoxites and in the wing the basal portion of the Costa is inflated and the R5 is strongly

curved beyond midlength reaching the C beyond the wing apex (Plate II, Fig. D). In Plakidas

(2019, Fig. 156, shown below) my illustration clearly shows the 9th tergite longer and as wide as

the gonocoxites. The specimen which I presented is clearly not that of W. nigripennis and is

noted as a misidentification. In fact, on the basis of

the short and narrow 9th tergite and the gonocoxal lobe (Plate II, Fig. F), W. nigripennis Kieffer

is more closely allied to W. rubricola Mamaev (Fig. 167) in that regard. However, since the type

material for W. nigripennis Kieffer is lost we will never know its exact identity. Especially the

internal sclerotized anatomy of the male genitalia.

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Winnertzia notmani (Felt 1920), page 281. Holotype female NYSM.

=Parwinnertzia Felt 1920, page 281. Type species notmani Felt by original designation.

Diagnosis. Female 1.5 mm; antenna broken with 10-flagellomeres remaining, U-shaped sensoria

on all 10-flagellomeres. Palpus 2-segmented. Wing longer and narrower than other Winnertzia’s,

about 2.5 times longer than wide and the M4 vein absent. Scutum and scutellum reddish-brown;

abdomen, halters and legs fuscous yellow. Tarsal claws simple, empodium rudimentary. First

tarsomere with a spiniform projection. Spermathecae membranous.

Taxonomic notes. Mr. Howard Notman donated this point mounted fly to the New York State

Museum. He collected the specimen on July 16, 1917 at Keene Valley, N. Y. It can be assumed

that Felt derived his color description from a dried specimen, which was later cleared and slide

mounted in Canada balsam.

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Winnertzia padicola Spungis 1992, page 32. Holotype male LASS, Latvia.

Diagnosis. Figures 134 – 143. Male ≈ 2.0 mm; with 12-flagellomeres (Figs. 134, 135, 136);

female ≈ 2.4 mm; with 12-flagellomeres (Figs. 137); palpus 4-segmented. Thorax and wing as

for the genus. First tarsomere with a blunt projection; tarsal claws with a basal tooth, empodium

shorter than the claw. Male abdomen with brown pigmented transverse bands; male genitalia

(Figs. 138, 139), 9th tergite with an emargination; the tegmen is nearly transparent and slightly

longer than the ejaculatory apodeme; the aedeagus and aedeagal prominence are not visible in

my specimens since they were cleared in KOH as this process dissolves away delicate

membranous tissues. The female has one sclerotized spermatheca.

Larva 3.3 mm; white. The spatula (Fig. 140) has a broadly rounded anterior margin and the

posterior apophysis is not developed. The 9th abdominal segment has two sclerotized spines on

the posterior margin with their bases in contact (Fig. 141).

Pupa: with long pronotal setae (Fig. 142) and a very distinctive thoracic spiracle (Fig. 143)

which is cup-shaped and only slightly elevated above the body wall. There is one nonsetose

papilla associated with the spiracle. This appears to be a species defining trait.

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Biology. Larvae have been collected from decaying vascular wood of a fallen Oak tree, several

millimeters below the surface. The decay color was white. The larvae were collected March 10,

1995, 10 Km SE of Columbia Maryland.

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Material examined. One larva, 1-male and 1-female on slides, deposited in the USNM.

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Winnertzia palustris Felt 1915, page 133. Holotype female NYSM.

Diagnosis. Figures 144 – 149. Female: ≈ 2.1 mm; wing 1.5 mm. Eleven flagellomeres with thin

U-shaped sensoria (Fig 145) with the distal ends extending beyond the nodes and stems .2 to .28

the length of the nodes. Sensoria occur on all 11-flagellomeres. Palpus 4-segmented. Thorax and

wing (Fig. 144) as for the genus. First tarsomere with a short blunt projection (Fig. 146); the

acropod on the foreleg is larger than the mid and hind leg acropods and has intermixed rows of

sharply pointed spines or possibly setae (Fig. 147), and the tarsal claws are falcate, lack a basal

tooth, the empodium is rudimentary. Abdomen yellow-green (Plakidas 2019, figure 157) or

yellow-brown (Fig. 144), tergites and sternites darker. Tergites 1-6 with a posterior row of long

brown setae, tergites 2-6 with lateral setae. Two large ovoid spermathecae are present internally,

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one is located in segment 6, the other spermatheca in 7. The cercus has sharply pointed clavate

setae mainly near the distal margins of both segments.

Larva (Figs. 148, 149). Length 4.1 – 5.5 mm; white; spatula when fully developed (Fig. 148)

dark brown, with a broadly rounded anterior margin and posterior apophysis undeveloped. The

9th segment (Fig. 149) displays two prominent brown spines, their bases not in contact.

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Biology. The larvae of Winnertzia palustris Felt were collected on January 15, 2014 from rotting

Birch bark. Two females were reared soon after, one on January 30 and the other on February 15,

2014.

Material examined. Three larvae and 2-females; 1-larva and 1- female deposited in the USNM.

Taxonomic notes. Felt described Winnertzia palustris from a single female collected on July 10,

1909 on low vegetation at Canada Lake, Town of Caroga, New York. The fly was collected by

C. P. Alexander. Although this species is represented by a single specimen there are three main

identifying traits which were useful in aligning my specimens with W. palustris. First the inflated

foreleg acropod with a tarsal claw without a basal tooth, the 11-flagellomeres with U-shaped

sensoria and the color description which included the greenish-yellow abdomen.

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Winnertzia pectinata Felt 1911, page 478. Lectotype male (Parnell 1971) NYSM.

Diagnosis. Male and female with 12-flagellomeres; palpus 4-segmented. Wings as for the genus,

Costa dark brown. Halters translucent yellow; legs translucent yellow tarsi darker. Tarsal claws

stout, with a basal tooth, empodium short. Male genitalia: gonostyli with a strong pectinate distal

claw; 9th tergite long, broadly and slightly emarginated, the lobes broadly rounded. The inner

gonocoxal emargination V-shaped, the distal gonocoxal lobes broad, truncate and sparsely

covered with setae.

Biology. Winnertzia pectinata Felt was reared from partially decayed Chestnut bark (Castanea

dentata) collected at Nassau, New York, April 26, 1911.

Taxonomic notes. Winnertzia pectinata Felt is presently considered to be a junior synonym of

Winnertzia solidaginis Felt 1907 by Gagné and Jaschhof (2021). Ephraim Felt allied W.

pectinata Felt 1911 to W. calciequina Felt 1907 and Panelius (1965, p. 118) placed the two

species in synonymy. Parnell (1971, p. 290) carried over the synonyms and placed both W.

calciequina and W. pectinata as junior synonyms of W. solidaginis Felt 1907.

However, it is my firm belief that with only a few specimens with which to study the use of

synonyms here is premature.

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Winnertzia pinicorticis Felt 1907. Holotype male USNM.

Diagnosis. Male: 1 mm; 12-flagellomeres, sensoria U-shaped on first 11 flagellomeres, the 12th

flagellomere ovoid, without sensoria. Palpus 4-segmented. Legs missing from holotype; wing

typical for the genus, Costa light brown. Halters pale yellow. Genitalia (Fig. 150): gonocoxites

stout, inner gonocoxal emargination narrow and deeply U-shaped; gonocoxal apodemes long and

thin, transverse bridge thin and lightly sclerotized; parameral apodemes thin and diverging

laterally; ejaculatory apodeme tubular; 9th tergite as long as the gonocoxites and slightly

emarginated. Gonostyli inflated almost 2X as long as wide; distal claw comprised of at least one

strong spine and numerous comb-like teeth.

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Taxonomic notes. Winnertzia pinicorticis Felt was reared from larvae found under the bark of

Pinus virginiana from Strom Virginia, February 16, 1892. Parnell (1971, p. 288) remarks that W.

pinicorticis “is close to W. carpini but differs from it in the number of antennal flagellomeres

and the slight differences in the post abdominal structures.” The 9th tergite in W. carpini is more

deeply emarginated and the gonocoxal apodemes are shorter in length (Fig. 88) compared to W.

pinicorticis (Fig. 150).

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Winnertzia rubida Felt 1908, page 422. Holotype male NYSM.

Diagnosis. Male: 1.25 – 2.5 mm; female with fully extended ovipositor: 2.4 – 3.7 mm. Antenna:

male with 12-flagellomeres (Fig. 152), sensoria only on the first 11 flagellomeres; female with

11-flagellomeres (Fig. 153), sensoria on all flagellomeres. Palpus 4-segmented, segments

progressively longer; tarsal claws with a basal tooth, empodium rudimentary (Fig. 154); wing

(Fig. 151) as for the genus, wing base (Fig. 155) clearly shows the Sc vein close to the R vein.

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Halters fuscous-yellow. Genitalia: 9th tergite (Fig. 159) deeply emarginated with rounded lobes;

gonocoxal apodemes long, slender and longer than the distance between them (Fig. 156).

Inner gonocoxal emargination broadly U-shaped (Fig. 158); ejaculatory apodeme as long as the

gonocoxites and the gonostylus (Fig. 156) with a distal claw of comb-like teeth. Tegmen flaps

(Fig. 156) very well developed in this specimen, and strongly indicates its affinity with

Winnertzia solidaginis Felt. Female telescopic portion of abdomen Figs. 160, 161) with one

sclerotized circular spermatheca at the anterior portion of abdominal segment 6. Both the

basicercus and the disticercus have clavate setae near their distal margins (Fig. 161).

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Material examined. One male, 1-female reared from a rotting Birch log, March 15, 2013. The

male specimen clearly within interspecific variation with the holotype for W. rubida Felt.

One male, one female deposited in the USNM.

Taxonomic notes. Winnertzia rubida Felt has three distinct morphological traits separating it

from W. solidaginis Felt. In the male genitalia W. rubida has long hook-shaped parameral

apodemes and a deeply emarginated 9th tergite. The female has 1-sclerotized spermatheca.

Winnertzia solidaginis males have less prominent parameral apodemes and a slightly

emarginated 9th tergite, while the female has 2-sclerotized spermathecae.

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Winnertzia rubricola Mamaev 1963, page 564. Holotype male HYNM, Japan.

Diagnosis. Male: ≈2.5 mm; female: ≈2.5 – 3.0 mm. Male (Figs. 162, 163) and female (Fig. 164)

with 11-flagellomeres, sensoria on all eleven flagellomere for both sexes. Wing (Fig. 165);

lateral thorax bare (Fig. 166); tarsal claws with a basal tooth, empodium rudimentary.

Proepisternum has 2-setae in both male and female specimens. The presence of these

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setae on the proepisternum may be a species defining trait.

Male genitalia (Fig. 167): the 9th tergite is deeply emarginated with broadly rounded lateral

lobes. Gonostylus (Fig. 168) strongly arched beyond midlength and with a distal claw of closely

joined teeth. Female ovipositor portion of abdomen (Fig. 169) has 2-sclerotized spermathecae,

one large circular spermatheca in segment 6 and one smaller spermatheca in segment 7.

Larva (Figs. 170 – 173): head capsule (Fig. 170) lightly sclerotized and typical for the genus.

The spatula (Fig. 171) has a single tooth and only a partially developed shaft anteriorly. The

anterior portion of the shaft is membranous in appearance, as one follows the outline of the shaft

posteriorly its outline deteriorates and is nearly invisible at the posterior end.

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The two terminal spines (Fig. 172) originate from a common basal area of sclerotization. The

ventral 9th segment has an unusual feature where there are 2- membranous spines (Fig. 173)

situated posterior to the anal pore.

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Pupa (Fig. 174): is typical for the genus, the thoracic spiracle (Fig. 174) is elongated and has

three setiform papillae near its base.

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Biology. Larvae were discovered under rotting Ash bark, Fraxinus sp., on January 19, 2015,

Aspinwall Penna. Allegheny Co. Two females and one male were reared on February 22, 1015.

Spungis (1992) reports larvae found in darkish decaying wood of Lime and Oak in bracket fungi.

Material examined. One larva, 1-male and 2-females, Penna. Allegheny Co., Aspinwall, slide

mounted in Euparal, deposited USNM.

Taxonomic notes. The male specimen which I reared from Aspinwall closely resembles the male

genitalia illustrated by Spungis (1992: Fig. 44, p. 20).

Winnertzia solidaginis Felt 1907a, page 149. Holotype NYSM.

Winnertzia solidaginis Felt 1907b, page 53.

Felt described this species twice in two separate publications in 1907 and designated 1907a as

the primary reference (1915, page 136).

Diagnosis. Based on recently reared specimens and field collected specimens from Allegheny

County, Pennsylvania. Male: 2.2 mm (n=2); female: 3.0-3.5mm (n=2); larva: 3.5 mm (n=2);

pupa: 3.5-4.0 mm (n=2). Light brown species, tarsi darker as a result of a dense covering of dark

brown scales. Antenna with 12-flagellomeres in male (Fig. 175) and female (Fig. 176). Male

sensoria irregularly U-shaped. Female sensoria originating near midlength on the nodes and

deeply U-shaped. Sensoria occur on all flagellomeres in male and female. Palpus 4-segmented,

4th segment the longest (Fig. 177); thorax and wing as for the genus. First tarsomere with a blunt

projection; tarsal claws with a basal tooth, empodium rudimentary. Legs: scales are mainly

setiform while others are wider in appearance. Scales are light brown on femorae and tibiae, and

darker brown on all tarsal segments. Halters translucent with setae and narrow setiform scales.

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Male genitalia (Figs. 178 – 182): ejaculatory apodeme longer than the height of the

gonocoxites; tegmen lightly sclerotized and nearly equal to the height of the ejaculatory

apodeme, with well-defined flaps (Fig. 178); aedeagal prominence triangular, covered with what

appear to be scales and in lateral view appears to originate from, or is attached to the gonocoxites

(Figs. 178, 180, 181). The 9th tergite is as long as the height of the gonocoxites and slightly

emarginated (Fig. 179), the posterior margin darkly pigmented; cerci longer than 9th tergite and

the inner gonocoxal emargination U-shaped (Fig. 179). The gonocoxites have a distal setose lobe

and the gonostyli are twice as long as wide with a distinct pectinate claw (Fig. 182) comprised of

comb-like teeth. The reproductive apparatus clearly shows the aedeagal prominence ventral to

the ejaculatory apodeme, distally the aedeagus is compressed between the ejaculatory apodeme

and the tegmen. Dorsally the 9th tergite appears to provide support for the reproductive

apparatus.

Female disticercus has 4-5 clavate setae and one long sharply pointed setae distally. There are 2-

sclerotized round spermathecae in my specimens.

Larva: typical for the genus, spatula with a triangular to rounded distal tooth, the posterior

apophysis is not developed in my specimens. The 9th segment has 2-terminal spines originating

from a common base.

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Pupa: typical for the genus with four papillae on the clypeal apotome, 2 of which are setiform

and 2 non-setose; the non-setose antennal papillae are easily recognized; the thoracic spiracle is

cup-shaped (Fig. 182A) with a flattened apex and there are 3-setose

papillae adjacent to the spiracle. The abdominal segments are partially covered with fine

spicules, strong spines are absent.

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Material examined. Four microscope slide mounts from Penna. Allegheny Co.: 1-larva from

decaying Birch log collected April 1, 2013 and 2-males + 1-exuviae on one slide, reared April

15, 2013. From 2018: 1-male + 1-female + 1- exuviae (siblings) on slide, reared April 25, 2018;

1-male reared April 25, 2018 on slide. Deposited in USNM.

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Winnertzia vernalis n. sp. Holotype male USNM.

Figures 183 – 191.

Diagnosis. Male ≈ 2.0 mm, wing 1.75 mm; female ≈ 2.5 mm, wing 1.6 mm; larva 2.8 mm. Both

male and female light brown, legs with darker brown setiform scales. Male with 12-

flagellomeres, stems shorter than nodes (Fig. 183), 11+12 closely joined. Female with 12-

flagellomeres, stems short, sensoria deeply U-shaped (Fig. 184) on all 12 flagellomeres. Wing as

for the genus; tarsal claws with a basal tooth, empodium as long as the bend in claw (Fig. 185).

Male genitalia (Figs. 188, 189), the notable feature is the aedeagal prominence which is

completely translucent and covered with light brown hair-like scales. The tegmen is shorter than

the ejaculatory apodeme and has a horizontal distal margin. The 9th tergite is emarginated and

the inner gonocoxal emargination is difficult to discern as a result of slide mounting distortion.

The female abdomen from segments 6-cercus is telescopic (Fig. 186), 2-ovoid spermathecae are

present, the disticercus has three prominent clavate setae laterally and one long seta distally (Fig.

187). Larva typical for the genus and very similar to the larva of W. solidaginis, especially with

regard to the shape of the spatula (Fig. 190), the head capsule and the terminal spines originating

from a common base (Fig. 191). Pupa unknown.

Etymology. The specific name vernalis, Latin for springtime, and refers to the early spring adult

emergence.

Type material. Holotype male, on slide, Penna. Allegheny Co., Aspinwall, reared March 30,

2022, larva from decaying Oak branchlet; paratype female, reared March 30, 2022, same data as

holotype; paratype larva, collected March 2022 from Oak Branchlet in leaf litter.

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Taxonomic notes. Winnertzia vernalis n. sp. is unique when compared to other Winnertzia’s

with the horizontal distal margin of the tegmen and the conical aedeagal prominence covered

with light brown scales (Figs. 188, 189).

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Concluding remarks.

The present study of the North American genera and species of Kronomyia Felt,

Rhipidoxylomyia Mamaev and Winnertzia Rondani was conducted by reviewing all of Felt’s

original descriptions and his 30th report of the State Entomologist (1915). I also closely

reviewed Parnell’s (1971) work on the Felt types. In addition I had in my possession numerous

Winnertziini which I collected and reared adults for over thirty years.

Three conclusions result from this study.

One: in some cases the use of synonyms are premature due to the lack of material with which to

study. I propose avoiding synonyms at this time.

Two: twenty five valid species are known to exist in North America, mainly in the northeast.

Three: with such a small area of North America explored, we can only image the great number

of species yet to be discovered.

Acknowledgements. I extend my appreciation to Dr. Z. A. Fedotova, All-Russia Scientific-

Research Institute of Plant Protection, Pushkin, St. Petersburg, Russia for her review of the

manuscript.

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Literature cited.

Elsayed, A. K., S. Fujita & M. Tokuda 2020. Description of the larva and pupa of Diadiplosis

hirticornis (Diptera: Cecidomyiidae), a predator of mealybugs. Japanese Journal of Systematic

Entomology, 26(1): 43-47.

Fedotova, Z. A., E. E. Perkovsky, A. J. Ross & Q. Zhang 2022. A new genus and species of

gall midges the tribe Winnertziini (Diptera, Cecidomyiidae, Porricondylinae) from Lower

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Felt, E. P. 1907a. 22nd report of the State Entomologist on injurious and other insects of the

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of New York 1907. New York State museum Bulletin 124: 5 – 541, 44 plates.

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Felt, E. P. 1913. Descriptions of gall midges. Journal of the New York Entomological Society

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Felt, E. P. 1915. 30th report of the State Entomologist on injurious and other insects of the State

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Gagné, R. J. 1977. Cecidomyiidae (Diptera) from Canadian amber. Proceedings of the

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Jaschhof, M. & C. Jaschhof 2020. Reevaluation of species richness in Winnertzia (Diptera,

Cecidomyiidae, Winnertziinae), with description of 37 new species from Sweden, Peru and

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largely on the examination of the Felt types. Miscellaneous Publications of the Entomological

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Plakidas, J. D. 2018. Identification of the Porricondylinae and Winnertziinae larvae (Diptera:

Cecidomyiidae sensu stricto). Loyalfield Publishing, Pittsburgh, Pennsylvania. 88 pages.

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from Allegheny County Pennsylvania. Loyalfield Publishing, Pittsburgh, Pennsylvania. 89

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The North American Winnertziini (Diptera: Cecidomyiidae: Winnertziinae). A photographic and illustrated guide to genera and species.

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